Blackwell Science, LtdOxford, UKBOJBotanical Journal of the Linnean Society0024-4074The Linnean Society of London, 2002

140

Original Article

LEAF ANATOMY IN ALSTROEMERIACEAEO. B. LYSHEDE

Botanical Journal of the Linnean Society, 2002, 140, 261–272. With 32 figures

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Comparative and functional leaf anatomy of selected
Alstroemeriaceae of mainly Chilean origin
OLE B. LYSHEDE*
Department of Plant Biology, The Royal Veterinary & Agricultural University, Thorvaldsensvej 40,
DK-1871 Frederiksberg C, Denmark

Received May 2002; accepted for publication July 2002

The resupinate leaves of 16 species of Alstroemeriaceae were examined using light- and scanning electron micros-
copy. The leaf anatomy is described for all of the species, that of the petiole and stem for selected species. The mes-
ophyll consists of chlorenchyma and includes idioblasts that contain raphides. Dorsiventral, isolateral or isobilateral
leaf types were observed. Petioles are always isolateral. Two epidermal types are commonly observed: type I contains
jigsaw puzzle-like intercostal cells and longitudinally elongated cells above the veins; type II contains only longitu-
dinally elongated cells, usually longer, above the veins. Some species have an epidermis which differs from the main
types. All species show adaptation to resupination by having an inverted anatomy. Due to the twist of the leaves,
adaxial indicates the lower surface and abaxial the upper. Stomata are found on both surfaces. Palisade cells, when
they occur, are always on the abaxial surface. Most species are mesomorphic in spite of the dry environments in
which they grow; a few show xeromorphic features. The leaves are compared and discussed with relation to ecological
conditions. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society, 2002, 140, 261-272.

ADDITIIONAL KEYWORDS: Alstroemeria – Bomarea – epicuticular wax – Leontochir – mesomorphic –

resupinate leaves – xeromorphic.

INTRODUCTION 1886; Goebel, 1920; Kugler, 1928). This affects the leaf
anatomy of the different species is various ways. Sev-
The family Alstroemeriaceae, chiefly distributed in
eral early workers studied the anatomy of resupinate
South America, contains five genera: Alstroemeria,
leaves. Dufour (1886) briefly discussed a few species
Bomarea, Leontochir, Schickendantzia and Taltalia.
of Alstroemeria, while Colozza (1898, 1901) compre-
However, this number can vary according to one’s
hensively described several species of Bomarea and
taxonomic viewpoint. If accepted as separate genera,
Alstroemeria. Later, Tetley (1936) described leaf devel-
Leontochir, Schickendantzia and Taltalia are all
opment in A. aurantiaca (= A. aurea). More recently,
monotypic. Based on chromosome studies Bayer
Bomarea edulis (Tussac) Herb. from Argentina was
(1988) separated Leontochir as monotypic. Following
briefly described by Sanso & Xifreda (1995).
their morphological and anatomical studies on subter-
The distribution of Alstroemeria is largely confined
ranean organs, pollen and seeds, Sanso & Xifreda
to two areas: (1) Chile/Argentina and (2) Brazil, with
(2001) concurred with Bayer, but included Schicken-
very few species found in other South American coun-
dantzia and Taltalia in Alstroemeria. Except for
tries (Aker & Healy, 1990). With a few exceptions, only
Bomarea which has a wider distribution, the other
Chilean material has been available from the botani-
genera are mainly found in Chile, Argentina and
cal gardens for this study. This is unfortunate, because
Brazil. Most members of the family have resupinate
there are distinct morphological and ecological differ-
leaves, i.e. the petiole is permanently twisted in such a
ences between the two groups (see the key in Baker,
way that the abaxial lamina of the leaf is turned
1888) which may be reflected in their leaf anatomy.
upwards and the adaxial surface downwards (Dufour,
Bayer (1987) described the morphology of the Chilean
species in a comprehensive monograph which included
*E-mail: obl@kvl.dk the species distribution; the information provided on

© 2002 The Linnean Society of London, Botanical Journal of the Linnean Society, 2002, 140, 261–272 261
262 O. B. LYSHEDE

the ecological habitats was, however, patchy. Alstroe- Table 1. Alstroemeriaceae. Abbreviations: BH: Botanical
meria pulchella, originally from Brazil, was intro- Garden of University of Copenhagen. KVL: Botanical
duced to Argentina; Sanso (1996) gives some notes on Section, The Royal Veterinary & Agricultural University,

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its habitats there. Copenhagen
Little recent information on the leaf anatomy of the
Alstroemeriaceae is available; the early descriptions, List of species Source Origin
written in Italian and Spanish, provide few illustra-
tions. The purpose of this paper is therefore to com- Alstroemeria KEW Chile
angustifolia Herb.
pare and illustrate the resupinate leaf anatomy of the
A. aurea Graham KEW Chile/
examined species and correlate it with the ecological
(A. aurantiaca D. Don) Argentina
habitats. In the descriptions which follow, adaxial
A. exserens Meyen BH Chile
indicates the lower surface and abaxial the upper, due
A. hookeri Schult. BH Chile/
to the twist characteristic of resupination. The possi-
ssp. hookeri Argentina
ble cause of the resupination is briefly discussed. A. ligtu L. ssp. ligtu BH Chile
A. ligtu L. ssp. simsii KEW Chile/North
MATERIAL AND METHODS (Spreng.) Bayer Brazil
(A. haemantha Ruiz et Pav.)
Plant material was kindly provided by The Royal A. magnifica Herb. BH Chile
Botanic Gardens, Kew, The Botanical Garden, Univer- A. pallida Graham KEW Chile
sity of Copenhagen (BH), and the Section of Botany, A. pelegrina L. BH Chile
The Royal Veterinary & Agricultural University, A. presliana Herb. BH Chile
Copenhagen (KVL). Fully grown leaves of several A. pulchella L.f. BH North Brazil/
Alstroemeria species, of one species of Bomarea and of (A. psittacina Lehm.) Argentina
Leontochir (Table 1) were examined using light- and A. pulchra Sims BH Chile
scanning electron microscopy. A. versicolor Ruiz et Pav. BH Chile
A. sp. (ref. OL nn1) KEW Chile (?)
Bomarea multiflora KEW Venezuela
LIGHT MICROSCOPY (L.f.) Mirb.
Small sections of leaf laminas taken from midribs, Leontochir ovallei Phil. KVL Chile
leaf edges and petioles were fixed in formalin-acetic
acid-alcohol (FAA) 5 : 5 : 90, dehydrated in alcohol and
embedded in Tecnovit resin. After sectioning on a sputtered with gold/palladium before examination in a
Microm HM 340 E rotary microtome, the sections were JEOL JSM 840 A scanning electron microscope.
variously stained with either PAS-Aniline Blue Black,
Fuchsin-Safranin-Astra Blue (FSA), or Sudan 4 prior
to mounting in DPX. Cross-sections of the leaf laminas RESULTS
of all species, together with longitudinal sections of
GENERAL LEAF AND STEM ANATOMY
the leaf laminas and cross-sections of petioles and
stems of selected species were examined. The lamina mesophyll of the leaf is chlorenchymatous
throughout. In the petioles the number of chloroplasts
per cell is often much diminished. A general feature
SCANNING ELECTRON MICROSCOPY is the presence of idioblasts containing raphides in a
Sections of leaf laminas, petioles and selected shoot polysaccharide matrix (Fig. 1). As a rule, these are
apices were fixed as above and after dehydration found more frequently along the edges than elsewhere
transferred to 100% acetone, critical point dried, and in the leaf. Several mesophyll cells have giant nuclei,


Figures 1–8. Fig. 1. Bomarea multiflora. Two idioblasts with raphides embedded in polysaccharide matrix in leaf meso-
phyll. Scale bar = 1 mm. Fig. 2. Alstroemeria aurea. Binucleate mesophyll cell. Scale bar = 1 mm. Figs 3-8. SEMs. Fig. 3.
A. pulchella. Adaxial epidermis, Type I with jigsaw puzzle intercostal area. To the right, epidermal cells above vein. Scale
bar = 100 mm. Fig. 4. A. exserens. Abaxial epidermis,Type II with stomata. Scale bar = 100 mm. Fig. 5. A. ligtu ssp. simsii.
Abaxial epidermis, modified Type II with coarsely sinuous side walls; without stomata. Scale bar = 100 mm. Fig. 6. Bomarea
multiflora. Adaxial epidermis with stoma and epicuticular wax. Scale bar = 10 mm. Fig. 7. Bomarea multiflora. Cross-
section of stem. Scale bar = 100 mm. Fig. 8. Alstroemeria hookeri. Shoot apex with asymmetrical leaf primordia. Scale
bar = 100 mm.

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264 O. B. LYSHEDE
suggesting polyploidy; they are sometimes binucleate
(Fig. 2). The vascular bundles in most species are sur-
rounded by parenchymatous bundle sheaths which
often contain chloroplasts. The epidermal cells of the
ridges above the vascular bundles are in general
larger than the intercostal epidermal cells; the latter
vary considerably in shape and size between species.
The guard cells are small.
The leaf epidermis as seen in SEM consists (with
some exceptions) of two main types. In Type I (Fig. 3)
the intercostal epidermal cells are short and tesse-
lated, with interlocking sinuous walls. The stomata
appear in rows and are orientated parallel with the
leaf axis, as in most monocots. The cells above the
veins are longitudinally elongated, the walls not sin-
uous. This type is most often found on the adaxial
surface. In Type II (Fig. 4) the epidermal cells are lon-
gitudinally elongated; those above the veins are
usually longer. The cells may have straight or sinuous
anticlinal walls (Fig. 5). This type is most often found
on the abaxial surface, but occurs on both surfaces in
some species and may be with or without stomata. The
epidermis will henceforth be described as Type I or II;
any variation will be pointed out, as in some species
the epidermis deviates completely from the two types.
The epidermal cells of the petioles are all longitudi-
nally elongated and may be with or without stomata.
Epicuticular wax is found on both surfaces and ranges
from prominent vertical platelets (Fig. 6) to minute
globular bodies.
The stem is atactostelic. The single-layered epider-
mis has a thin cuticle and stomata. The cortex consists
of a 3–4-layered chlorenchyma and includes idioblasts
with raphides. In Bomarea the peripheral part of the
stele consists of a 4–5-layered fibre sheath bordering
the cortex. Collateral vascular bundles are dispersed
over the stelar area, as seen in cross-section. In the
centre, however, are five bundles containing larger
xylem vessels arranged in a ring (Fig. 7). The stem
anatomy of B. multiflora is very similar to that of
other Bomarea species described by Colozza (1898)
and Sanso & Xifreda (1995). The stem anatomy of
A. aurea differs by having a fibre sheath with only 1–
2 layers; it also lacks the central group of bundles.
Instead, there is a central water-storing parenchyma
of thin-walled cells. Its stem diameter is about double
that of B. multiflora.
In the shoot apex the youngest primordial leaves
show tendencies towards asymmetry (Fig. 8) which
may indicate incipient twisting of the leaves.
Figures 9–16. Cross-sections of leaves of Alstroemeria. Fig. 9. A. angustifolia. Note idioblast, stoma and large epidermal
cell at the edge. Fig. 10. A. aurea. Fig. 11. A. exserens. Fig. 12. A. hookeri. Fig. 13. A. ligtu ssp. ligtu. Fig. 14. A. ligtu ssp.
simsii. Fig. 15. A. magnifica. Fig. 16. A. pallida. All scale bars = 20 mm, except Fig. 13 = 10 mm.
© 2002 The Linnean Society of London, Botanical Journal of the Linnean Society, 2002, 140, 261–272
LEAF DESCRIPTIONS
Alstroemeria angustifolia Herb
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The leaf is isolateral with a 6–8-layered chlorenchyma;
the layer along the abaxial epidermis appears slightly
denser (Fig. 9). It is ridged on the adaxial surface above
the veins, smooth on the abaxial except above the mid-
dle nerve which protrudes on both sides. Stomata are
numerous on the adaxial surface, much less frequent
on the abaxial. A single row of prominent thick-walled
epidermal cells constitutes the leaf margin.
SEM. The adaxial epidermis is Type I, jigsaw puzzle-
like; the abaxial epidermis is Type II, but the cells are
rectangular with sinuous anticlinal walls intercostally
and with straight lateral walls above the veins. The
adaxial epidermis is densely covered by small vertical
wax platelets, the abaxial with small dotted wax bod-
ies. The petiole has long pointed cells with straight lat-
eral anticlinal walls and has stomata on both surfaces.
Ecology. Mountains, hills, gorges and canyons. The
leaf is mesomorphic.
Alstroemeria aurea Graham ( syn. A. aurantiaca
D. Don)
The leaf is isolateral with a dense uniform meso-
phyll as seen in cross section (Fig. 10), although there
are larger plicate mesophyll cells in the abaxial row. In
longitudinal section several multilobed mesophyll
cells can be seen. Giant nuclei are common in this spe-
cies, together with many binucleate cells (Fig. 2).
SEM. The adaxial epidermis is Type I, but the inter-
costal cells are elongated longitudinally with coarsely
wavy anticlinal walls. The abaxial epidermis is
Type II, with a few stomata. Vertical fine wax platelets
cover the abaxial intercostal cells.
Ecology. Mountains, Nothofagus forests, exposed
gravel and grass slopes. The leaf is mesomorphic.
Alstroemeria exserens Meyen
The leaf is isolateral with the mesophyll consisting of
mainly roundish cells of equal size as seen in cross-
section (Fig. 11). The veins are positioned quite close
to each other with a regular alternation between
smaller and larger. This makes the leaf finely ridged
on both surfaces. The epidermal cells on both surfaces

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12

14

16
13

15
11
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266 O. B. LYSHEDE
are slightly larger above the veins. The stomata are
slightly sunken.
SEM. Both surfaces are Type II with a regular and
rather small distance between the costal and intercos-
tal zones (Fig. 4). The intercostal cells are rectangular
with more or less straight lateral anticlinal walls, the
costal cells similar, but longer. Stomatal density is
high on both surfaces. The wax layer is delicate and
continuous on both surfaces, although rather irregular
on the adaxial surface.
Ecology. Stony slopes and river banks. The leaf is
mesomorphic.
Alstroemeria hookeri Schult. ssp. hookeri
The leaf blade is slightly dorsiventral with a layer of
blunt palisade like cells towards the abaxial surface
(Fig. 12). The petiole is isolateral with markedly fewer
chloroplasts in its mesophyll, some in the middle of
the twist and very few at the base.
SEM. The adaxial epidermis is Type I, but the inter-
costal cells are rectangular; the abaxial epidermis is
Type II, with stomata. Epicuticular wax is present as
delicate thin strands orientated perpendicular to
the longitudinal axis of leaf. In the shoot apex the
youngest leaf primordia show a slight asymmetry
(Fig. 8).
Ecology. Sandy areas, dunes. The leaf is mesomorphic.
Alstroemeria ligtu L. ssp. ligtu
The leaf is isolateral in both the lamina and petiole,
but with fewer chloroplasts in the latter (Fig. 13). The
mesophyll includes several large idioblasts.
SEM. The adaxial epidermis is Type I with short blunt
intercostal cells; the abaxial epidermis is Type II, asto-
matous, with finely sinuous side walls intercostally.
The epicuticular wax is very thin on both surfaces. On
the abaxial, it consists of simple or branched rodlets;
on the adaxial, of minute globules. As in A. hookeri,
the youngest leaf primordia at the apical meristem
show a tendency towards asymmetry.
Ecology. Mountains,among trees and shrubs,roadsides
rocks, canyons and hills. The leaf is mesomorphic.
Alstroemeria ligtu L. ssp. simsii (Spreng.) Bayer
(syn. A. haemantha Ruiz et Pav.)
The dorsiventral leaf has a distinct palisade layer
below the abaxial surface among which several bi- or
trilobed cells appear (Fig. 14). The spongy tissue is
3-layered in the lamina, but multilayered along the
© 2002 The Linnean Society of London, Botanical Journal of the Linnean Society, 2002, 140, 261–272
midrib and in the petiole. The leaf margin is bluntly
shaped by tiers of three large cells.
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SEM. The adaxial epidermis is Type I, with richly
branched intercostal cells; the abaxial epidermis is
Type II, with coarsely sinuous intercostal side walls,
but without ridges and stomata (Fig. 5).
Ecology. Coastal areas, mountains, hills, gorges and
brushy slopes. The leaf is mesomorphic.
Alstroemeria magnifica Herb.
The leaf is dorsiventral with one palisade-like layer
beneath the abaxial surface; there are several bilobed
cells (Fig. 15). The cells towards the adaxial surface
are rounded or longitudinally elongated. The epider-
mal cells on both surfaces are larger above the veins
than above the intercostal areas, making both sur-
faces ridged. Stomata are more numerous on the adax-
ial surface.
SEM. The adaxial epidermis is Type I, with the inter-
costal cells rather short and almost rectangular; the
abaxial epidermis is Type II, but with numerous inter-
costal stomata. The epidermal cells are covered by
minute wax bodies on both surfaces.
Ecology. Coastal areas, hilly areas between ocean
and desert, rocky slopes and thickets. The leaf is
mesomorphic.
Alstroemeria pallida Graham
The leaf is isolateral with a quite dense mesophyll of
6–9 cell layers (Fig. 16). The epidermal cells on both
surfaces are slightly larger above the veins.
SEM. Both epidermises are Type II, with stomata on
both surfaces. The wax on the abaxial surface is rather
thick and consists of confluent vertical platelets;
on the adaxial, it consists of thin vertical platelets,
mainly seen in the furrows over the anticlinal epider-
mal walls.
Ecology. High mountains, canyons and calderas. The
leaf is mesomorphic.
Alstroemeria pelegrina L.
The leaf is dorsiventral with one palisade layer below
the abaxial surface and a rather dense spongy tissue
of 4–5 layers with numerous chloroplasts in the cells
(Figs 17–20). There are distinct idioblasts. The abaxial
epidermis consists of a single layer of giant cells with
large vacuoles. This layer is equal to or exceeds the
mesophyll in thickness. The adaxial epidermis con-
sists of giant cells above the veins and small cells
in the intercostal zones where the stomata occur,

slightly sunken into the epidermal surface. The large
epidermal cells often almost occlude the intercostal
zones (Fig. 18).
SEM. The adaxial epidermis is a strongly modified
Type I - rectangular cells with small lateral out-
growths near to the stomatal complexes in the inter-
costal zones which interchange with the zones above
the veins with the large cells (Fig. 19). The adaxial
petiole epidermis is Type II. The abaxial epidermal
cells of the lamina are fusiform hexagonal in outline
with straight walls (Fig. 20) while those of the petiole
taper. Both epidermises are covered by minute wax
platelets. The leaf primordia at the shoot apex appear
to be symmetrical.
Ecology. Coastal dunes, steep coastal cliffs. The suc-
culent epidermis with the protected stomata and the
dense mesophyll with numerous chloroplasts charac-
terize the leaf as xeromorphic.
Alstroemeria presliana Herb.
The leaf is dorsiventral with a single palisade layer
and 3–4-layered spongy parenchyma with large inter-
cellular spaces (Fig. 21). The epidermal cells are more
or less equal in size as seen in cross section.
SEM. The two epidermises are both Type II and fairly
similar in appearance, with numerous stomata. The
epidermal cells are rectangular with slightly undulat-
ing lateral anticlinal walls. The epicuticular wax lay-
ers are inconspicuous.
Ecology. Mountains, gravelled road-sides, Nothofagus
forest. The leaf is mesomorphic.
Alstroemeria pulchella L.f. (syn. A. psittacina Lehm.)
The leaf is distinctly dorsiventral with a well devel-
oped palisade layer towards the abaxial surface. This
layer constitutes nearly half of the mesophyll thick-
ness and includes many two- to four-lobed palisade
cells as seen in cross section (Fig. 22). In longitudinal
section, most palisade cells appear multilobed
(Fig. 23). The petioles are isolateral.
SEM. The adaxial epidermis is Type I with multilobed
intercostal cells (Fig. 3) and covered with a dense layer
of vertical wax platelets. The abaxial epidermis is
Type II, with coarsely undulating side walls in the
intercostal cells. Stomata are not present on this sur-
face. The petiolar epidermal cells are long and with
lobed cell walls.
Ecology. Railways, sheltered locations. The leaf is
mesomorphic.
© 2002 The Linnean Society of London, Botanical Journal of the Linnean Society, 2002, 140, 261–272
LEAF ANATOMY IN ALSTROEMERIACEAE 267
Alstroemeria pulchra Sims
The leaf is dorsiventral, but with rather blunt pali-
sade-like cells abaxially, several of which are multi-
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lobed (Figs 24, 25). The spongy parenchyma cells
towards the adaxial epidermis are large and surround
large intercellular spaces. The abaxial epidermis con-
sists of cells of uniform size and without stomata -
very similar to that of A. pelegrina. The adaxial epi-
dermis consists of flat cells in the intercostal areas,
but with distinctly larger succulent cells above the
veins.
SEM. The adaxial epidermis is Type I with the inter-
costal cells slightly lobed, elongated perpendicular to
the leaf axis, and with stomata at regular intervals
(Fig. 25). The cells above the veins are larger than
those of the lamina and inflated. The astomatous
abaxial epidermis consists of long, rather broad cells,
rectangular or with bluntly pointed ends. The cell sur-
face is densely ridged by narrow cuticular striations
and covered with minute wax bodies.
Ecology. Mountains, hills, canyons and grassy slopes.
The leaf is mesomorphic, although the succulent epi-
dermis is a xeromorphic feature.
Alstroemeria versicolor Ruiz et Pav.
The leaf is isobilateral with a palisade layer towards
both surfaces (Fig. 26). The mesophyll is 4–5-layered
and rather compact. The epidermal cells above the
veins are only slightly larger than the intercostal epi-
dermal cells, as seen in cross section.
SEM. The adaxial intercostal cells are rectangular
with undulating lateral walls and with a larger
density of stomata than on the abaxial surface. Both
belong to Type II. The abaxial surface wax consists of
delicate strands, the adaxial of minute bodies.
Ecology. Mountains, canyons and slopes. The leaf is
mesomorphic.
Alstroemeria sp. (ref. OL nn1)
This unidentified species is included because of its
interesting and deviating leaf anatomy (Figs 27 and
28). Due to an accident during consignment its label
disappeared and attempts to re-identify the species
have unfortunately proved unsuccessful.
The leaf is dorsiventral with a single palisade layer
of wedge-shaped cells towards the abaxial surface. The
spongy mesophyll is 2-layered with many large inter-
cellular spaces. The vascular bundles are surrounded
by a fibre sheath 2–4 cell layers thick. The epidermal
cells appear uniform in cross section and are covered
by a thin cuticle. The stomata are sunken. The petiole
is isolateral and consists of 8–10 mesophyll layers.
268 O. B. LYSHEDE

SEM. The adaxial epidermis consists of zones of lon- more regular cells and has in general smaller inter-
gitudinally elongated cells with cuticular micropapil- cellular spaces. An interesting feature is that the chlo-
lae above the veins alternating with intercostal zones roplasts in the petiole and leaf occur along the parts of

of isodiametrical papillate cells. It is covered with
dense layers of small wax bodies (Fig. 28). Stomata are
not visible from the surface but are placed at the base
of some of the papillate cells. The abaxial epidermis is
astomatous and consists of longitudinally stretched
cells covered by cuticular knobs.
Ecology. The thin leaf blade, wedge shaped palisade
cells, spongy tissue and large intercellular spaces are
hygromorphic features that point to a plant growing in
shade. The fibre sheaths, sunken stomata and surface
relief are xeromorphic features. The anatomy is very
interesting; it is regrettable that the species so far
remains unknown.
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cell wall opposite to the intercellular spaces and not
along those bordering neighbouring cells (Fig. 31).
SEM. The adaxial epidermis is Type I with blunt, few-
branched cells with cuticular striations and many
stomata (Fig. 32). The astomatous abaxial surface is
smooth and consists of longitudinally elongated cells
with finely striated cuticular sculpturing perpendicu-
lar to the long axis. There is no distinct difference
between the costal and intercostal areas on the abax-
ial surface. Both surfaces are covered by tiny wax
platelets, denser on the adaxial surface and formed by
stellate bodies and small platelets. All epidermal cells
of the petiole are longer.

Bomarea multiflora (L.f.) Mirb. Ecology. Desert. The leaf is mesomorphic.

The leaf is isolateral with a slight tendency towards a
palisade-like cell layer beneath the abaxial surface
(Fig. 29). The mesophyll is generally compact, with
small intercellular spaces; it commonly includes idio-
blasts with raphides (Fig. 1). The larger vascular
bundles are capped by wide-celled fibres while the
smaller ones are surrounded by parenchymatous bun-
dle sheaths. In longitudinal section, many mesophyll
cells are elongated and multilobed. The stomata are
slightly sunken.
SEM. The adaxial epidermis is Type I, but with only
moderately branched cells which are rather blunt,
irregular in shape and frequently alternating with the
stomata in longitudinal rows. The abaxial epidermis is
Type II, with slightly undulating walls in the intercos-
tal areas. Both surfaces are covered by well developed,
mainly vertical wax platelets (Fig. 6). On the abaxial
surface the platelets are positioned perpendicular to
the leaf axis, whilst on the adaxial surface they are
denser and not orientated in any specific direction.
Ecology. High mountains, hedges and edges of woods,
cloud forests.The leaf is mesomorphic.
Leontochir ovallei Phil.
The isolateral leaf has a mesophyll of 4–5 layers of
irregular cells with large intercellular spaces
(Figs 30–32). The epidermal cells are large compared
with those of the mesophyll. The petiole consists of
DISCUSSION
In all the species examined the leaf anatomy is
inverted in accordance with resupination, bringing the
phloem above the xylem in the vascular bundles. Dor-
siventral species have the palisade layer towards the
abaxial surface and the stomata are always found in
the adaxial epidermis (i.e. usually Type I, sometimes
Type II, and sometimes neither). In species where sto-
mata are present on both surfaces there are generally
fewer on the abaxial surface and so belong to Type II.
The general foliar anatomy of the species investigated
shows a mesophyll consisting of chlorenchyma cells
and large idioblasts with raphides. Idioblasts were
observed in all the species studied and have been
described in Bomarea edulis (Sanso & Xifreda, 1995);
however, they are generally not characteristic for
families within the Liliales (Rudall et al., 2000). The
vascular bundles are in general surrounded by paren-
chymatous bundle sheaths, which may be with or
without chloroplasts. Exceptions are found in the uni-
dentified Alstroemeria where they are surrounded by
strong fibre sheaths (Fig. 27); they are also supported
by fibres in Bomarea multiflora, confirming the
early observation of Colozza (1898) in B. oligantha
Baker (= B. m). Vascular fibres were also described in
B. edulis (Sanso & Xifreda, 1995). The cells in the epi-
dermis are often larger above the veins; this reaches


Figures 17–24. Leaves of Alstroemeria. Fig. 17. A. pelegrina. Cross-section. Scale bar = 100 mm. Fig. 18. A. pelegrina.
Cross-section. Scale bar = 20 mm. Fig. 19. A. pelegrina. Adaxial surface with giant epidermal cells above veins. SEM.
Scale bar = 100 mm. Fig. 20. A. pelegrina. Abaxial surface. SEM. Scale bar = 10 mm. Fig. 21. A. presliana. Cross-section.
Scale bar = 20 mm. Fig. 22. A. pulchella. Cross-section. Scale bar = 10 mm. Fig. 23. A. pulchella. Longitudinal section. Scale
bar = 10 mm.Fig. 24. A. pulchra. Cross-section. Scale bar = 100 mm.

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an extreme in A. pelegrina where the cells on the The epidermis in the studied species is described as
adaxial surface partly conceal the intercostal areas being either Type I (Fig. 3) or Type II (Figs 4, 5).
(Figs 18, 19). The abaxial surface of the leaf of this Type I, with its ‘jigsaw-puzzle’ appearance, always

species (Figs 17, 20), of A. pulchra (Fig. 24) and of
Leontochir (Fig. 30) is composed of large uniform epi-
dermal cells with no structural differences above the
vascular bundles. The small stomata found on one or
both surfaces may be sunken in various ways, most
pronounced in A. pelegrina (Fig. 20) and in A. sp.
(Figs 27, 28). Trichomes were not found in any of the
species studied, but have been observed in Bomarea
edulis from Argentina (Sanso & Xifreda, 1995). The
twist of the petiole may occur through an asymmetri-
cal development during leaf growth. In some of the
species the youngest leaf primordia were observed to
be asymmetrical and it may be assumed that as
growth continues the asymmetry becomes enlarged
and thus responsible for the resupination. This is in
accordance with the studies of Seybold (1925) in
A. chilensis Lem.
Most of the species studied have isolateral leaves
with a uniform mesophyll structure. Alstroemeria ver-
sicolor (Fig. 26) is slightly isobilateral and the remain-
ing species are dorsiventral with a palisade layer
towards the abaxial surface The palisade layer may be
slightly developed as in A. hookeri (Fig. 12) or com-
posed of distinctly multilobed groups of cells as seen in
A. pulchella (Figs 22, 23), the only Brazilian species in
this investigation. The mesophyll may be rather com-
pact or have large intercellular spaces. There is a clear
difference between the leaf lamina and the petiole.
The latter is always isolateral, with fewer chloroplasts
per cell and larger intercellular spaces; it is also often
thicker. In the leaf and especially the petiole of Leon-
tochir (Figs 30, 31) it is interesting to observe that
the chloroplasts are placed opposite the intercellular
spaces and not bordering the neighbouring cells, a fea-
ture that possibly suggests an efficient uptake of car-
bon dioxide. A further notable feature is the difference
between the two subspecies of A. ligtu; ssp. ligtu is iso-
lateral (Fig. 13) while ssp. simsii is dorsiventral with a
distinct palisade layer (Fig. 14). The epidermises are
of identical types, but with differences in the intercos-
tal branching on the adaxial surface and the sinu-
ousity of anticlinal walls on the abaxial. Ssp. simsii
has formerly had species status as A. haemantha.
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occurs on the adaxial surface, but varies from species
to species with differences in branching and shape of
the epidermal intercostal cells. Particularly distinct
is A. pulchra (Fig. 25), with the epidermal cells elon-
gated in the direction perpendicular to the leaf
longitudinal axis and not parallel to it as in most
other species. The Type II epidermis of longitudinally
stretched cells is generally found on the abaxial sur-
face, but occurs on both surfaces in, for instance,
A. exserens (Fig. 4) and A. pallida. The unidentified
Alstroemeria sp. differs from the remainder in having
an adaxial epidermis with papillate intercostal cells
and stomata invisible from the surface (Fig. 28). Epi-
cuticular wax is present on all the species, although
the shape and amount vary considerably. The dense
layers of vertical wax platelets seen in Bomarea
(Fig. 6) and A. pulchella are particularly noteworthy.
In contrast to these are the minute wax bodies in
A. magnifica.
ECOLOGY
Information concerning the habitats of the studied
species has tended to be scanty. A few are found in
open Nothofagus forests (A. aurea, A. presliana). Most
of the others grow in open land on rocky or grassy
slopes, on hills or in canyons or crevices, while several
grow near the coast (Bullock, 1952; Navas, 1973;
Bayer, 1987; Christensen et al., 1999). Uphof (1940)
described the habitats of A. versicolor and both
A. ligtu subspecies as xerophytic shrubbery.
A. pelegrina and A. hookeri typically grow in dunes
and on coastal cliffs. Several other coastal species
endure long periods of drought (Ørgaard & Kris-
tiansen, 1998). It may therefore seem surprising that
most of the species are clearly mesomorphic in their
leaf anatomy as they lack a thick cuticle, indumen-
tum, supporting tissues, and a well developed photo-
synthetic tissue. Presence of these structural elements
would indicate xerophytism (Fahn & Cutler, 1992). An
explanation may be that most species are geophytes
that either survive the dry period as underground
tubers or rhizomes or obtain a sufficient water supply


Figures 25–32. Fig. 25. Alstroemeria pulchra. Adaxial side of leaf. The intercostal epidermal cells are elongated perpen-
dicular to the leaf axis. SEM. Scale bar = 100 mm. Fig. 26. A. versicolor. Cross-section of leaf. Scale bar = 20 mm. Fig. 27.
A. sp. Cross-section of leaf. Scale bar = 20 mm. Fig. 28 A. sp. Adaxial surface of leaf. Intercostal areas with papillate cells
hiding the stomata at their base. SEM. Scale bar = 100 mm. Fig. 29. Bomarea multiflora. Cross-section of leaf. Scale
bar = 20 mm. Fig. 30. Leontochir ovallei. Cross-section of leaf. Scale bar = 20 mm. Fig. 31. Leontochir ovallei. Cross-section
of petiole. Note the position of chloroplasts surrounding the intercellular spaces (i). Scale bar = 20 mm. Fig. 32. Leontochir
ovallei. Adaxial surface of leaf with modified Type I epidermis; bluntly shaped intercostal cells. SEM. Scale bar = 100 mm.

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272 O. B. LYSHEDE
from these. Bomarea, by contrast, is a climber, but still
has underground storage organs.
The presence of thick layers of epicuticular wax is
likewise often included among the xeromorphic fea-
tures. It is interesting to note that some of the species
which have thick layers are mesomorphic, whereas
some of the xeromorphic species have only thin
epicuticular wax particles. Alstroemeria pelegrina is
the most xeromorphic of the investigated species. The
giant epidermal cells covering the abaxial surface
have a water storage function and may also serve as a
filter, protecting the plant from the intense direct and
indirect light found in coastal dunes and cliffs. The
large cells of the adaxial epidermis in this species may
in addition protect the plant against too much sto-
matal transpiration by almost occluding the inter-
costal parts and their stomata. Further, the dense
chloroplast-rich mesophyll provides efficient photo-
synthesis when stomata are open. However, it may be
noted that A. hookeri growing in similar habitats has
a distinct leaf anatomy without xeromorphic charac-
ters. The abaxial epidermis of A. pulchra and Leon-
tochir is similar to that of A. pelegrina. The stomata in
the unidentified Alstroemeria sp. are sunken, level
with the base of the papillate intercostal epidermal
cells on the adaxial leaf surface; the latter are also
supplied with cuticular micropapillae. The purpose of
this may be to protect the plant by reflecting light from
the surface. There is also a strong fibre sheath sur-
rounding each vascular bundle. These xeromorphic
features contrast with the hygromorphic few-layered
mesophyll with wedge-shaped palisade cells and many
intercellular spaces.
Almost all species in the Alstroemeriaceae have
resupinate leaves. The cause, it may be speculated,
might be some environmental pressure occurring
early in the family’s history. The mesophyll is rela-
tively similar in most of these species, although, as
described above, the epidermis can vary considerably.
This may be due to selection through the course of
their evolution. As already mentioned, two regional
groups of Alstroemeria exist: one in Chile/Argentina
which is recognized as the distribution centre of the
genus (Aker & Healy, 1990), and one in Brazil. The dif-
ferences in morphology may reflect adaptation to hab-
itat. Alstroemeria pulchella, the only Brazilian species
in this study, is distinguished by its well developed
palisade layer with many multilobed cell groups below
the abaxial epidermis, which makes it an atypical
Type II. It may be representative of the different foliar
anatomy of the Brazilian Alstroemerias. Future inves-
tigations may show if the differences in morphology
between the two regional groups are paralleled in leaf
anatomy.
© 2002 The Linnean Society of London, Botanical Journal of the Linnean Society, 2002, 140, 261–272
ACKNOWLEDGEMENTS
I am deeply indebted to Lise Girsel, who carefully
Downloaded from https://academic.oup.com/botlinnean/article-abstract/140/3/261/2433636 by Siksha O Anusandhan University (Bhubaneswar) user on 30 January 2019
prepared the many specimens for microscopy, and to
Alexander Schulz, who read the manuscript.
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