Seasonal Dimorphism in the Mediterranean Cistus incanus L. subsp. incanus
GIOVANNA ARONNE* and VERONICA DE MICCO
Laboratorio di Ecologia Riproduttiva, Dipartimento di Arboricoltura, Botanica e Patologia Vegetale (Sezione
Botanica), UniversitaÁ degli Studi di Napoli `Federico II', via UniversitaÁ 100, 80055 Portici (Napoli), Italy
Received: 20 November 2000 Returned for revision: 5 January 2001 Accepted: 23 February 2001
Mediterranean perennial species are described as being sclerophyllous, or summer deciduous, or seasonally
dimorphic. Field observation in the coastal maquis of Castelvolturno Nature Reserve, southern Italy, showed that
Cistus incanus L. subsp. incanus is a seasonally dimorphic species as it develops brachyblasts with small leaves in
summer, and dolichoblasts with large leaves in winter. Field biometric data con®rmed that winter shoots were 14-
times longer than those developed in summer and had many more leaves. The area of single winter leaves was ®ve-
times that of summer leaves. Anatomical leaf structure also changed with the season: winter leaves were ¯at while
summer leaves had a crimped lamina which was partially rolled to form crypts in the lower surface. Leaves were
covered by considerably more trichomes in summer than in winter. Stomata were uniformly distributed along the
lower epidermis of winter leaves but were only present in the crypts of summer leaves. In summer leaves, a palisade
layer was often found on both sides of the lamina, the mesophyll cells were generally smaller and the intercellular
spaces were reduced. Winter leaves had a dorsiventral structure and larger intercellular spaces. Seasonal dimorphism
is generally reported to be an adaptation to summer drought. However, the morphology and anatomy of C. incanus
L. subsp. incanus showed that the subspecies has not only developed a strategy to survive summer drought, but has
evolved two dierent habits, one more xerophytic than the other, to optimize adaptation to the seasonal climatic
changes occurring in Mediterranean environments.
# 2001 Annals of Botany Company
Key words: Cistus, Cistus incanus L. subsp. incanus, climatic changes, leaf anatomy, leaf dimorphism, Mediterranean
shrubs, phenology, seasonal dimorphism.
INTRODUCTION
The Mediterranean-type climate is characterized by hot, dry
summers alternating with cool, wet winters (
;
). The seasonal ¯uctuations in soil moisture are
considered a limiting factor for growth and productivity of
Mediterranean perennial species (
;
). According to the severity of the summer drought,
Mediterranean ecosystems can be distributed along a
gradient which has maquis with evergreen sclerophylls at
the wet end, and garigue with seasonally dimorphic species
at the dry end (
). Evergreen sclerophylls are
characterized by small, thick, leathery, long-lived leaves
(
). Seasonally dimorphic species are charac-
terized by a seasonal reduction in their transpiring surface
which is achieved by shedding the larger winter and spring
leaves growing on dolichoblasts and developing smaller
summer leaves on new brachyblasts (
Species which exhibit seasonal dimorphism are reported
in dierent Mediterranean-type ecosystems (
,
). As regards the Mediterranean
region, this habit was described for species from the
Greek phrygana (
,
,
The morphology, physiology and leaf anatomy of Cistus
species are reported in several studies (e.g.
). However, to date, no investigation has
de®ned the biometrical and leaf anatomical dierences
between summer and winter habits of Cistus plants. In the
present work, phenology and seasonal changes in shoot
biometry, leaf morphology and anatomy were studied in
Cistus incanus L. subsp. incanus. The nomenclature for
C. incanus L. subsp. incanus follows
.
MATERIALS AND METHODS
Plant material was collected at Castelvolturno Nature
Reserve on the Tyrrhenian coast, north of the Bay of
Naples (southern Italy). The site is situated on stabilized
sand dunes and has a typical Mediterranean climate
(
) with an annual rainfall of
about 1000 mm, but with precipitation concentrated in
autumn and winter, followed by a dry summer. The
vegetation, which is subject to ®re, varies from 0.5 to 3 m
in height and is characterized by a patchwork mosaic of
shrub species and restricted gap areas colonized by
therophytes. The co-dominant species are Phillyrea latifolia
L., Pistacia lentiscus L., Rhamnus alaternus L., Myrtus
communis L., Rosmarinus ocinalis L., Cistus salvifolius L.
and Cistus incanus L. subsp. incanus.
In spring 1996, ®ve plants of Cistus incanus L. subsp.
incanus were randomly selected in the ®eld and branches
were tagged to follow shoot development. On each plant,
ten shoots were sampled for biometric measurements both
Annals of Botany 87: 789±794, 2001
doi:10.1006/anbo.2001.1407, available online at http://www.idealibrary.com on
0305-7364/01/060789+06 $35.00/00
# 2001 Annals of Botany Company
* For correspondence. Fax 39 081 7755114, e-mail aronne@unina.
it
at the end of April, before ¯owering and the dry period, and
in September, before the autumn rains. Winter and summer
shoot elongation and the area of each leaf on the shoot were
measured, and the number of leaves that formed during the
two seasons was also counted. Leaf area was measured by
digitizing leaf images and analysing them with `Plant
Meter', a software program specially devised for measure-
ment of lines and areas.
In both seasons, three leaves per plant were also sampled
for subsequent anatomical observation. The leaves were
®xed in a mixture of 40 % formaldehyde:glacial acetic
acid:50 % ethanol (5:5:90 by volume) for several days, cut
into pieces of approx. 5 5 mm, dehydrated in an ethanol
series and embedded in JB4
1
wax (Polysciences, Warring-
ton, PA, USA). Leaf sections (5±8 mm) were stained with
0.5 % toluidine blue in water (
) and observed
under a transmitted light microscope (BX60, Olympus,
Hamburg, Germany). A digital micrograph of one section
per leaf from the area of the maximum leaf width was
obtained with a digital camera (Olympus, CAMEDIA
C2000). The 30 images (one section three leaves ®ve
plants two seasons) were analysed using the image
analysis system, `Plant Meter'. Hair density (n mm
ÿ1
) and
stomatal density (n mm
ÿ1
) were calculated by counting,
respectively, the number of stalks and stomata present
along the section on both abaxial and adaxial sides, and
measuring the length of the section analysed. Similarly, for
each image, the thickness of the palisade layer as well as the
minimum and maximum distance between the upper and
lower epidermises (thickness) of the lamina were measured.
RESULTS
In C. incanus L. subsp. incanus, each axillary bud grows out
in summer, producing a shoot with short internodes
(brachyblast), small leaves and a leaf terminal bud
(
). At the end of the summer, after the ®rst autumn
rains, summer leaves are shed. The terminal bud begins to
re-grow and develops a stem with long internodes and large
leaves (dolichoblast). This growth ceases in late spring when
the terminal bud becomes an in¯orescence (
Subsequently, the large leaves are shed and new brachyblasts
develop from the axillary buds (
). This cyclic process
determines the round shape of such bushes (
Another dierence between the two habits regards lamina
inclination which is horizontal in winter leaves and almost
vertical in summer leaves.
Field observations were corroborated by biometric
measurements (
). Winter shoots were on average
14-times longer than those developed in summer and had
about four-times the number of leaves. However, leaf
density, i.e. the ratio between leaf number and shoot length,
was almost four-fold lower in winter than in summer
shoots. As regards leaf area, winter leaves were ®ve-times
larger than those developed in summer. Therefore, summer
shoots are shorter with fewer, more densely packed, small
leaves while winter shoots have long stems with many larger
leaves.
Interestingly, no interplant variability was found in any
parameter measured on summer samples, while very signi-
F
IG
. 1. Schematic view of C. incanus growth: summer brachyblasts (A); brachyblasts and dolichoblasts (B); development of new brachyblasts after
shedding of winter leaves (C); development of new dolichoblasts (D).
T
A B L E
1. Biometric data from ®ve plants of Cistus incanus
randomly selected in the ®eld and measured at the end of the
summer and winter growth periods respectively
Summer
growth
Winter
growth
t
Shoot length (cm)
0.8
11.2
P 5 0.001
Number of leaves per shoot
3
11
P 5 0.001
Number of leaves per shoot/shoot
length (n cm
ÿ1
)
3.8
1.0
P 5 0.001
Leaf area (mm
2
)
67
339
P 5 0.001
Mean values and signi®cance of Student's t-test.
790
Aronne and De MiccoÐSeasonal Dimorphism in Cistus incanus L.
®cant dierences were reported when the same parameters
were measured in winter (
Transverse sections of winter and summer leaves also
showed dierences in their anatomy. Winter leaves were
¯at, while summer leaves had a crimped lamina which was
partially rolled to form several crypts in the lower surface
(
Lamina thickness was not uniform either in summer or
winter leaves (
and
): signi®cant dierences were
found between the minimum and maximum distance
between the upper and lower epidermis of each section in
both kinds of leaves. Within each leaf, the variation in
thickness (ratio between maximum and minimum widths)
was signi®cantly greater in summer than in winter leaves.
Therefore, the variation in the lamina thickness of a single
leaf was greater in summer leaves.
Upper epidermal cells were much larger in winter than in
summer leaves (
). The palisade parenchyma was
signi®cantly thicker in winter (84 mm) than in summer
leaves (53 mm). However, in summer leaves the palisade
tissue was often present on both sides of the lamina
(
) and the mesophyll cells were generally smaller
with reduced intercellular spaces. As a result, the whole
structure was more compact. Moreover, in summer leaves,
single palisade cells often had undulating walls forming
inward and outward folds alternately (
Both leaf surfaces were covered by a thick layer of white
trichomesÐstellate hairs consisting of a stalk and eight-18
long branches. In both kinds of leaves there were
signi®cantly more trichomes on the lower than on the
upper surface (
). However, trichome density was
much higher in summer than in winter leaves.
In both kinds of leaves, stomata were present only on the
lower surface (
), being uniformly distributed along the
lower epidermis of winter leaves but found almost
exclusively in the crypts of summer leaves (
and
DISCUSSION
Xerophytes are dry habitat plants with transpiration
decreasing to a minimum under conditions of water de®cit
(
). Certain tissues of xerophytes, particu-
larly leaf tissues, become altered structurally in relation to
the environment, and plant survival depends upon the
ability to withstand desiccation without permanent injury
(
).
Seasonal dimorphism has been reported to be an
adaptive strategy to the seasonal climatic changes occurring
in Mediterranean habitats (
,
;
Christodoulakis et al., 1990; Kyparissis and
;
). We have described
this habit for C. incanus at Castelvolturno, suggesting that
the species is well adapted to the rhythmic ¯uctuation of the
Mediterranean climate. Development of brachyblasts in
summer and dolichoblasts in winter is reported for other
Cistus species (
). In C. incanus, most of the
growth in terms of shoot length, number of leaves and leaf
area occurs in winter. During summer, growth is similar in
the whole population because it is limited by drought to the
minimal values for survival, while in winter no limiting
factors aect plant growth and biometric parameters dier
signi®cantly among individuals.
Steep leaf inclination has been described for many species
in dierent environments and interpreted as a strategy to
reduce the amount of direct solar radiation, resulting in
lower leaf temperature and transpiration rates, and avoid-
ance of damage to the photosynthetic apparatus (
;
;
). Erect summer leaves of C. incanus maximize light
interception in the early morning and late afternoon,
keeping noon interception to a minimum. This allows the
species to tolerate very hot environments by physically
evading the midday sun. By contrast, during autumn, winter
and spring, when drought is not a limiting environmental
factor, horizontal leaves optimize direct solar radiation.
Dierent leaf inclination in summer and winter was also
reported for C. incanus by
. We
suggest that the occurrence of vertical leaves in summer and
horizontal leaves in winter is a strategy that evolved in
C. incanus to obtain the best advantage from solar radiation
in both seasons.
This dual adaptation is also corroborated by leaf
anatomy: summer leaves frequently have a palisade layer
under both epidermises while winter leaves have a typical
dorsiventral structure. The presence of a palisade layer on
both leaf surfaces, together with a mesophyll composed of
smaller cells and reduced intercellular spaces, is reported to
be characteristic of xerophytic species (
). These
traits are found in summer leaves of C. incanus but not in
winter ones which show longer palisade cells only under the
upper epidermis, together with wider intercellular spaces.
The occurrence of anatomical dierences between summer
and winter leaves is in agreement with reports for other
seasonal dimorphic species (
).
Moreover, a lower mesophyll cell density and larger inter-
cellular spaces compared to the sclerophyllous Phillyrea
latifolia and Quercus ilex were reported for leaves of
C. incanus sampled in October (
In summer leaves of C. incanus, we observed palisade
cells with involuted walls. This anatomical feature is well
known in some evergreen conifers and is reported in
Caesalpinioid legumes (
). Although their
real function has not yet been ascertained, it has been
T
A B L E
2. Interplant variability of biometric measurement in
summer and winter among the ®ve marked plants of Cistus
incanus at Castelvolturno
Summer growth Winter growth
Shoot length (cm)
0.376
0.002
Number of leaves per shoot
0.922
0.011
Number of leaves per shoot/shoot
length (n cm
ÿ1
)
0.561
0.019
Leaf area (mm
2
)
0.059
0.000
Signi®cance (P-values) of ANOVA.
Aronne and De MiccoÐSeasonal Dimorphism in Cistus incanus L.
791
F
IG
. 2. Light microscope view of cross-sections of C. incanus winter leaves (A, B), and summer leaves (C±F). E, Large epidermal cells; t,
fragments of the numerous trichomes; c, crypt; s, stomata (s). Bars 100 mm.
792
Aronne and De MiccoÐSeasonal Dimorphism in Cistus incanus L.
speculated that they are an important anatomical adap-
tation to periodic drought. Under water stress these cells
should lose water and shrink, reducing the thickness of the
entire leaf. When water becomes available again, they might
quickly enlarge, causing the leaf to expand in thickness
(
Light intensity is also decreased by the hairy covering of
leaves, which is thicker during the season of greatest solar
radiation. Trichomes are reported to be inferior to the
cutinous coat in reducing transpiration (
),
except in strong sunlight where the cuticle has less
protective value (
). Leaf pubescence is
reported to be an adaptation to the Mediterranean
environment by reducing transpiration, increasing the
probability of water uptake by leaves, maintaining favour-
able leaf temperature, and protecting against UV-B
radiation responsible for photosynthetic inhibition (
). Where a single species exists in a mesophytic
and xerophytic form,
found the latter to be
more hairy. Therefore, in the more hairy summer leaves of
C. incanus, light intensity and transpiration should be lower
than in winter leaves, suggesting the occurrence of two
levels of leaf xeromorphism.
Plants with small leaves are more common in dry habitats
(
). A very common characteristic of xeromorphic
leaves is a reduced external area and a lower surface area to
volume ratio (
). A signi®-
cant reduction in the leaf area occurs in C. incanus during
the drier season, and this, together with the modi®cations in
internal leaf structure, allows the species to optimize
environmental seasonal conditions.
The ¯at structure of the lamina is characteristic of meso-
morphic leaves, but a crimped lamina, folded to form crypts
in which stomata are concentrated, is reported to be an
adaptive strategy to drought (
). Leaf
rolling is described for summer leaves of seasonal dimorphic
Mediterranean species to reduce light interception (
). We have shown that C. incanus
develops ¯at leaves in winter and folded leaves, with wider
variation in lamina thickness, in summer. Therefore, leaf
structure is optimized according to seasonal environmental
changes occurring in the Mediterranean.
Stomatal distribution is dierent between the two leaf
types. Stomata are uniformly distributed on the lower
surface of the ¯at winter leaves, whereas they are concen-
trated in the crypts of summer leaves to reduce evapo-
transpiration, as reported for other Mediterranean species
such as Nerium oleander L. (
).
Large epidermal cells, as well as a strati®ed epidermis, are
described as water storage structures characteristic of
xerophytes and are also present in the evergreen Mediterra-
nean species Nerium oleander L. and Rosmarinus ocinalis
L. (
). In C. incanus, large epidermal
cells of winter leaves would support the populations during
occasional periods of winter drought.
In Mediterranean environments, perennial species are
either sclerophyllous, summer deciduous, or seasonally di-
morphic (
). According to
), the latter is an adaptation to summer drought.
described seasonal dimorphism
for Sarcopoterium spinosum as a major strategy which pro-
duces `seasonally dierent plants' from the same individual,
that can successfully stand the variety of unfavourable
Mediterranean conditions. The overall consideration of
phenology, morphology and leaf anatomy of C. incanus is in
agreement with the conclusion by
. We suggest that C. incanus has evolved two dierent
0
50
100
150
200
250
300
SL
Lamina
thickness
(
µ
m)
WL
F
IG
. 3. Mean values and s.d. of minimum (F) and maximum (h)
thickness of the lamina in summer (SL) and winter (WL) leaves of
C. incanus.
0
2
4
6
8
10
12
SL
Trichomes
(n
mm
1
)
WL
F
IG
. 4. Number of trichomes found along leaf sections of both lower
(F) and upper (h) surfaces. Mean values and s.d. are reported for
summer (SL) and winter (WL) leaves.
0
2
4
6
8
10
WL
SL-out
Stomata
(n
mm
Ð1
)
SL-in
F
IG
. 5. Mean number and s.d. of stomata found along leaf sections of
winter leaves (WL) and summer leaves outside the crypt (SL-out) and
inside the crypt (SL-in).
Aronne and De MiccoÐSeasonal Dimorphism in Cistus incanus L.
793
forms, one more mesophytic, the other more xerophytic, to
optimize adaptation to the seasonal ¯uctuation of environ-
mental conditions throughout the year.
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794
Aronne and De MiccoÐSeasonal Dimorphism in Cistus incanus L.
Document Outline
- Seasonal Dimorphism in the Mediterranean Cistus incanus L. subsp. incanus
- INTRODUCTION
- MATERIALS AND METHODS
- RESULTS
- DISCUSSION
- LITERATURE CITED
- Figures
- Tables
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