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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.

© 2002 The Linnean Society of London, Botanical Journal of the Linnean Society, 2002, 140, 261–272
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LEAF ANATOMY IN ALSTROEMERIACEAE

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suggesting polyploidy; they are sometimes binucleate LEAF DESCRIPTIONS


(Fig. 2). The vascular bundles in most species are sur-
rounded by parenchymatous bundle sheaths which Alstroemeria angustifolia Herb

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


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.

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266 O. B. LYSHEDE

are slightly larger above the veins. The stomata are midrib and in the petiole. The leaf margin is bluntly
slightly sunken. shaped by tiers of three large cells.

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SEM. Both surfaces are Type II with a regular and SEM. The adaxial epidermis is Type I, with richly
rather small distance between the costal and intercos- branched intercostal cells; the abaxial epidermis is
tal zones (Fig. 4). The intercostal cells are rectangular Type II, with coarsely sinuous intercostal side walls,
with more or less straight lateral anticlinal walls, the but without ridges and stomata (Fig. 5).
costal cells similar, but longer. Stomatal density is
high on both surfaces. The wax layer is delicate and Ecology. Coastal areas, mountains, hills, gorges and
continuous on both surfaces, although rather irregular brushy slopes. The leaf is mesomorphic.
on the adaxial surface.
Alstroemeria magnifica Herb.
Ecology. Stony slopes and river banks. The leaf is The leaf is dorsiventral with one palisade-like layer
mesomorphic. beneath the abaxial surface; there are several bilobed
cells (Fig. 15). The cells towards the adaxial surface
Alstroemeria hookeri Schult. ssp. hookeri are rounded or longitudinally elongated. The epider-
The leaf blade is slightly dorsiventral with a layer of mal cells on both surfaces are larger above the veins
blunt palisade like cells towards the abaxial surface than above the intercostal areas, making both sur-
(Fig. 12). The petiole is isolateral with markedly fewer faces ridged. Stomata are more numerous on the adax-
chloroplasts in its mesophyll, some in the middle of ial surface.
the twist and very few at the base.
SEM. The adaxial epidermis is Type I, with the inter-
costal cells rather short and almost rectangular; the
SEM. The adaxial epidermis is Type I, but the inter-
abaxial epidermis is Type II, but with numerous inter-
costal cells are rectangular; the abaxial epidermis is
costal stomata. The epidermal cells are covered by
Type II, with stomata. Epicuticular wax is present as
minute wax bodies on both surfaces.
delicate thin strands orientated perpendicular to
the longitudinal axis of leaf. In the shoot apex the
Ecology. Coastal areas, hilly areas between ocean
youngest leaf primordia show a slight asymmetry
and desert, rocky slopes and thickets. The leaf is
(Fig. 8).
mesomorphic.
Ecology. Sandy areas, dunes. The leaf is mesomorphic.
Alstroemeria pallida Graham
The leaf is isolateral with a quite dense mesophyll of
Alstroemeria ligtu L. ssp. ligtu 6–9 cell layers (Fig. 16). The epidermal cells on both
The leaf is isolateral in both the lamina and petiole, surfaces are slightly larger above the veins.
but with fewer chloroplasts in the latter (Fig. 13). The
mesophyll includes several large idioblasts. SEM. Both epidermises are Type II, with stomata on
both surfaces. The wax on the abaxial surface is rather
SEM. The adaxial epidermis is Type I with short blunt thick and consists of confluent vertical platelets;
intercostal cells; the abaxial epidermis is Type II, asto- on the adaxial, it consists of thin vertical platelets,
matous, with finely sinuous side walls intercostally. mainly seen in the furrows over the anticlinal epider-
The epicuticular wax is very thin on both surfaces. On mal walls.
the abaxial, it consists of simple or branched rodlets;
on the adaxial, of minute globules. As in A. hookeri, Ecology. High mountains, canyons and calderas. The
the youngest leaf primordia at the apical meristem leaf is mesomorphic.
show a tendency towards asymmetry.
Alstroemeria pelegrina L.
Ecology. Mountains,among trees and shrubs,roadsides The leaf is dorsiventral with one palisade layer below
rocks, canyons and hills. The leaf is mesomorphic. the abaxial surface and a rather dense spongy tissue
of 4–5 layers with numerous chloroplasts in the cells
Alstroemeria ligtu L. ssp. simsii (Spreng.) Bayer (Figs 17–20). There are distinct idioblasts. The abaxial
(syn. A. haemantha Ruiz et Pav.) epidermis consists of a single layer of giant cells with
The dorsiventral leaf has a distinct palisade layer large vacuoles. This layer is equal to or exceeds the
below the abaxial surface among which several bi- or mesophyll in thickness. The adaxial epidermis con-
trilobed cells appear (Fig. 14). The spongy tissue is sists of giant cells above the veins and small cells
3-layered in the lamina, but multilayered along the in the intercostal zones where the stomata occur,

© 2002 The Linnean Society of London, Botanical Journal of the Linnean Society, 2002, 140, 261–272
LEAF ANATOMY IN ALSTROEMERIACEAE 267

slightly sunken into the epidermal surface. The large Alstroemeria pulchra Sims
epidermal cells often almost occlude the intercostal The leaf is dorsiventral, but with rather blunt pali-
zones (Fig. 18). sade-like cells abaxially, several of which are multi-

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lobed (Figs 24, 25). The spongy parenchyma cells
SEM. The adaxial epidermis is a strongly modified towards the adaxial epidermis are large and surround
Type I - rectangular cells with small lateral out- large intercellular spaces. The abaxial epidermis con-
growths near to the stomatal complexes in the inter- sists of cells of uniform size and without stomata -
costal zones which interchange with the zones above very similar to that of A. pelegrina. The adaxial epi-
the veins with the large cells (Fig. 19). The adaxial dermis consists of flat cells in the intercostal areas,
petiole epidermis is Type II. The abaxial epidermal but with distinctly larger succulent cells above the
cells of the lamina are fusiform hexagonal in outline veins.
with straight walls (Fig. 20) while those of the petiole
taper. Both epidermises are covered by minute wax SEM. The adaxial epidermis is Type I with the inter-
platelets. The leaf primordia at the shoot apex appear costal cells slightly lobed, elongated perpendicular to
to be symmetrical. the leaf axis, and with stomata at regular intervals
(Fig. 25). The cells above the veins are larger than
Ecology. Coastal dunes, steep coastal cliffs. The suc- those of the lamina and inflated. The astomatous
culent epidermis with the protected stomata and the abaxial epidermis consists of long, rather broad cells,
dense mesophyll with numerous chloroplasts charac- rectangular or with bluntly pointed ends. The cell sur-
terize the leaf as xeromorphic. face is densely ridged by narrow cuticular striations
and covered with minute wax bodies.
Alstroemeria presliana Herb.
The leaf is dorsiventral with a single palisade layer Ecology. Mountains, hills, canyons and grassy slopes.
and 3–4-layered spongy parenchyma with large inter- The leaf is mesomorphic, although the succulent epi-
cellular spaces (Fig. 21). The epidermal cells are more dermis is a xeromorphic feature.
or less equal in size as seen in cross section.
Alstroemeria versicolor Ruiz et Pav.
The leaf is isobilateral with a palisade layer towards
SEM. The two epidermises are both Type II and fairly
both surfaces (Fig. 26). The mesophyll is 4–5-layered
similar in appearance, with numerous stomata. The
and rather compact. The epidermal cells above the
epidermal cells are rectangular with slightly undulat-
veins are only slightly larger than the intercostal epi-
ing lateral anticlinal walls. The epicuticular wax lay-
dermal cells, as seen in cross section.
ers are inconspicuous.
SEM. The adaxial intercostal cells are rectangular
Ecology. Mountains, gravelled road-sides, Nothofagus with undulating lateral walls and with a larger
forest. The leaf is mesomorphic. density of stomata than on the abaxial surface. Both
belong to Type II. The abaxial surface wax consists of
Alstroemeria pulchella L.f. (syn. A. psittacina Lehm.) delicate strands, the adaxial of minute bodies.
The leaf is distinctly dorsiventral with a well devel-
oped palisade layer towards the abaxial surface. This Ecology. Mountains, canyons and slopes. The leaf is
layer constitutes nearly half of the mesophyll thick- mesomorphic.
ness and includes many two- to four-lobed palisade
cells as seen in cross section (Fig. 22). In longitudinal Alstroemeria sp. (ref. OL nn1)
section, most palisade cells appear multilobed This unidentified species is included because of its
(Fig. 23). The petioles are isolateral. interesting and deviating leaf anatomy (Figs 27 and
28). Due to an accident during consignment its label
SEM. The adaxial epidermis is Type I with multilobed disappeared and attempts to re-identify the species
intercostal cells (Fig. 3) and covered with a dense layer have unfortunately proved unsuccessful.
of vertical wax platelets. The abaxial epidermis is The leaf is dorsiventral with a single palisade layer
Type II, with coarsely undulating side walls in the of wedge-shaped cells towards the abaxial surface. The
intercostal cells. Stomata are not present on this sur- spongy mesophyll is 2-layered with many large inter-
face. The petiolar epidermal cells are long and with cellular spaces. The vascular bundles are surrounded
lobed cell walls. by a fibre sheath 2–4 cell layers thick. The epidermal
cells appear uniform in cross section and are covered
Ecology. Railways, sheltered locations. The leaf is by a thin cuticle. The stomata are sunken. The petiole
mesomorphic. is isolateral and consists of 8–10 mesophyll layers.

© 2002 The Linnean Society of London, Botanical Journal of the Linnean Society, 2002, 140, 261–272
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

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of isodiametrical papillate cells. It is covered with cell wall opposite to the intercellular spaces and not
dense layers of small wax bodies (Fig. 28). Stomata are along those bordering neighbouring cells (Fig. 31).
not visible from the surface but are placed at the base
of some of the papillate cells. The abaxial epidermis is SEM. The adaxial epidermis is Type I with blunt, few-
astomatous and consists of longitudinally stretched branched cells with cuticular striations and many
cells covered by cuticular knobs. stomata (Fig. 32). The astomatous abaxial surface is
smooth and consists of longitudinally elongated cells
Ecology. The thin leaf blade, wedge shaped palisade with finely striated cuticular sculpturing perpendicu-
cells, spongy tissue and large intercellular spaces are lar to the long axis. There is no distinct difference
hygromorphic features that point to a plant growing in between the costal and intercostal areas on the abax-
shade. The fibre sheaths, sunken stomata and surface ial surface. Both surfaces are covered by tiny wax
relief are xeromorphic features. The anatomy is very platelets, denser on the adaxial surface and formed by
interesting; it is regrettable that the species so far stellate bodies and small platelets. All epidermal cells
remains unknown. 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
DISCUSSION
small intercellular spaces; it commonly includes idio-
blasts with raphides (Fig. 1). The larger vascular In all the species examined the leaf anatomy is
bundles are capped by wide-celled fibres while the inverted in accordance with resupination, bringing the
smaller ones are surrounded by parenchymatous bun- phloem above the xylem in the vascular bundles. Dor-
dle sheaths. In longitudinal section, many mesophyll siventral species have the palisade layer towards the
cells are elongated and multilobed. The stomata are abaxial surface and the stomata are always found in
slightly sunken. the adaxial epidermis (i.e. usually Type I, sometimes
Type II, and sometimes neither). In species where sto-
SEM. The adaxial epidermis is Type I, but with only mata are present on both surfaces there are generally
moderately branched cells which are rather blunt, fewer on the abaxial surface and so belong to Type II.
irregular in shape and frequently alternating with the The general foliar anatomy of the species investigated
stomata in longitudinal rows. The abaxial epidermis is shows a mesophyll consisting of chlorenchyma cells
Type II, with slightly undulating walls in the intercos- and large idioblasts with raphides. Idioblasts were
tal areas. Both surfaces are covered by well developed, observed in all the species studied and have been
mainly vertical wax platelets (Fig. 6). On the abaxial described in Bomarea edulis (Sanso & Xifreda, 1995);
surface the platelets are positioned perpendicular to however, they are generally not characteristic for
the leaf axis, whilst on the adaxial surface they are families within the Liliales (Rudall et al., 2000). The
denser and not orientated in any specific direction. vascular bundles are in general surrounded by paren-
chymatous bundle sheaths, which may be with or
Ecology. High mountains, hedges and edges of woods, without chloroplasts. Exceptions are found in the uni-
cloud forests.The leaf is mesomorphic. dentified Alstroemeria where they are surrounded by
strong fibre sheaths (Fig. 27); they are also supported
Leontochir ovallei Phil. by fibres in Bomarea multiflora, confirming the
The isolateral leaf has a mesophyll of 4–5 layers of early observation of Colozza (1898) in B. oligantha
irregular cells with large intercellular spaces Baker (= B. m). Vascular fibres were also described in
(Figs 30–32). The epidermal cells are large compared B. edulis (Sanso & Xifreda, 1995). The cells in the epi-
with those of the mesophyll. The petiole consists of 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

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

© 2002 The Linnean Society of London, Botanical Journal of the Linnean Society, 2002, 140, 261–272
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from these. Bomarea, by contrast, is a climber, but still ACKNOWLEDGEMENTS


has underground storage organs.
I am deeply indebted to Lise Girsel, who carefully
The presence of thick layers of epicuticular wax is

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prepared the many specimens for microscopy, and to
likewise often included among the xeromorphic fea-
Alexander Schulz, who read the manuscript.
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
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giant epidermal cells covering the abaxial surface Baker JG. 1888. Handbook of the Amaryllideae. London:
have a water storage function and may also serve as a George Bell & Sons.
filter, protecting the plant from the intense direct and Bayer E. 1987. Die Gattung Alstroemeria in Chile. Mitteilung
indirect light found in coastal dunes and cliffs. The der Botanischen Staatssammlung München 24: 1–362.
large cells of the adaxial epidermis in this species may Bayer E. 1988. Beitrag zur Cytologie der Alstroemeriaceae.
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© 2002 The Linnean Society of London, Botanical Journal of the Linnean Society, 2002, 140, 261–272