1981 by sun leaf thickness − shade leaf thick- nesssun leaf thickness.
2
.
7
. Statistics All statistical tests were performed using a
statistical software package Statistica, Statsoft USA. The distribution of h
1
and a were com- pared to a uniform distribution by x
2
test. The differences of h
l
and a during the day and be- tween winter and summer were tested by one-way
analysis of variance ANOVA and Tukey test for multiple comparison. Diurnal trend of cosi was
compared to the cosi of a horizontal plane depending only on solar movement by x
2
test. Differences in mean morphological and anatomi-
cal leaf traits were determined by t-test, ANOVA and Tukey test for multiple comparison. Linear,
quadratic and logarithmic regression relationships for all analysed leaf traits were tested.
3. Results
3
.
1
. Leaf age The typical evergreen sclerophyllous species, Q.
ilex and P. latifolia, had a leaf life-span LLS of 2 – 4 years: 84 and 16 leaves of Q. ilex are shed
respectively after 2 and 3 years; 41, 54 and 5 leaves of P. latifolia are shed respectively after 2,
3 and 4 years. The seasonal leaf dimorphic shrub C. incanus produced smaller leaves summer
leaves at the beginning of May, persisting through summer and falling mostly in autumn
leaf life-span of 4 months; full leaf expansion was reached at 4 months. Larger leaves winter
leaves were produced at the middle of September, persisting through winter and falling in spring
leaf life-span of 8 months. P. latifolia and Q. ilex full leaf expansion was reached at 1 year old
leaves.
3
.
2
. Leaf orientation, leaf inclination and incident PAR
h
l
depended strictly on phyllotaxy: C. incanus and P. latifolia had opposite pattern and Q. ilex a
whorled pattern of leaf attachment. Opposite leaves of C. incanus and P. latifolia were at 90°,
from one to the other on the azimuthal projec- tion, while Q. ilex leaves were about 72°. To
compare these patterns, leaves were monitored in all four cardinal directions. Azimuth was random
with respect to sun and did not change daily, seasonally Fig. 1 or per different leaf age cohort
the deviation from randomness and the differ- ences of means were not significant.
C. incanus, P. latifolia and Q. ilex leaves showed a constant a throughout the day Fig. 2
the differences of means were not significant. a
changed per different leaf age cohort P B 0.01 Table 2. One year old leaves of P. latifolia
showed the steepest leaf inclination 59 9 7° in summer, decreasing with increasing leaf age P B
0.01 and Q. ilex showed the same trend. C. incanus showed a different a trend P B 0.01 of
winter and summer leaves: a was negative − 37 9 12° in winter leaves while it was positive
44 9 13° in summer leaves. a was significantly correlated with leaf age P B 0.01 Table 3: a
increased until full leaf expansion, then it de- creased. RI showed the same trend, in particular
P. latifolia had the highest RI in summer 47. a was a linear function of SLW P B 0.05. C. in-
canus showed, moreover, a particular trend of leaf margin: winter leaves had a marginal leaf folding
along the midrib axis 29 9 9° higher in summer leaves 76 9 11° P B 0.01 Fig. 3.
Diurnal trends of cosi was similar for the three species and per different leaf age cohort,
depending solely on solar movement P B 0.01, therefore no form of heliotropism was present.
3
.
3
. Leaf morphology The average leaf traits of the analysed species
per different leaf age cohort are shown in Tables 2 and 4.
SLW varied significantly P B 0.01 among the species and it increased until full leaf expansion
was reached maximum SA, decreasing with the increase of leaf age. Fully expanded summer
leaves of C. incanus were 20 smaller than winter leaves P B 0.05. SLW at full leaf expansion was
significantly higher P B 0.01 in Q. ilex and P.
latifolia compared to that of C. incanus. Leaves of P. latifolia were characterised by the smallest SA,
almost three times lower than Q. ilex. SLW sig- nificantly correlated with leaf age P B 0.01, L
P B 0.01, LDI P B 0.01 and a P B 0.05, confi- rming the dependence among the leaf traits Table
Fig. 1. Distribution of leaf orientation h
l
frequency of Cistus incanus, Phillyrea latifolia and Quercus ilex sun leaves during the day in winter 020298 and 060398 and in summer 060798 and 030898. In winter: morning = 09:30 h h
s
= 135°, midday = 12:00
h h
s
= 174°, afternoon = 14:30 h h
s
= 212°. In summer: morning = 07:30 h h
s
= 85°, midday = 12:00 h h
s
= 177°, afternoon =
16:30 h h
s
= 269°. h
s
= sun azimuth. The distribution of leaf orientation was random. Standard error is shown.
Fig. 2. Distribution of leaf inclination a frequency of Cistus incanus, Phillyrea latifolia and Quercus ilex sun leaves during the day in winter 020298 and 060398 and in summer 060798 and 030898. In winter: morning = 09:30 h a
s
= 26°, midday = 12:00
h a
s
= 37°, afternoon = 14:30 h a
s
= 28°. In summer: morning = 07:30 h a
s
= 30°, midday = 12:00 h a
s
= 72°, afternoon =
16:30 h a
s
= 35°. a
s
= sun zenith. P. latifolia a was steeper in summer than in winter P B 0.01. Standard error is shown.
3. LDI showed the same SLW trend increasing until full leaf expansion. In essence, longer leaf
life-span combined with higher SLW, L, LDI and steeper a.
3
.
4
. Leaf anatomy The microscopical analysis of fully expanded
leaves revealed significant variations among the
Table 2 a
°, RI , SA cm
2
, DW mg, SLW mg cm
− 2
, LDI g cm
− 3
and L mm per different leaf age months of Cistus incanus
a
C. incanus RI
SA DW
LA SLW
a L
LDI 13
2.9 9 0.3 25 9 5
8.5 9 0.9 2
b
235 9 16 29 9 8
0.36 9 0.01 32
4.0 9 0.7 59 9 11
44 9 13 14.7 9 1.5
4
b
244 9 15 0.60 9 0.02
17 3.5 9 1.0
41 9 12 3
c
11.9 9 1.7 −
34 9 10 201 9 14
0.58 9 0.03 18
4.7 9 0.8 63 9 13
− 35 9 10
13.3 9 1.6 5
c
216 9 13 0.62 9 0.02
20 5.0 9 0.5
8
c
70 9 16 −
37 9 12 14.0 9 1.3
220 9 14 0.64 9 0.03
a
LA, leaf age; a, leaf inclination; RI, reduction of PAR mmol m
2
s
− 1
incident on the sloping leaf surface; SA, surface area; DW, dry weight; SLW, specific leaf weight; LDI, leaf density index; L, total leaf thickness. Standard error is shown.
b
Summer leaves.
c
Winter leaves.
species Table 5. P. latifolia showed a particularly thick adaxial cuticle, more than two time higer
than C. incanus. Q. ilex leaves showed hairs on the abaxial surface while C. incanus on both the
adaxial and the abaxial surfaces data not shown. On the average Q. ilex and P. latifolia showed a
densely packed mesophyll with few air spaces; on the contrary C. incanus showed well developed
intercellular spaces data not shown. P. latifolia had the highest total leaf thickness Table 4. Xh
of the three species was on the average 0.55 Table 5. Total leaf thickness increased until full
leaf
expansion was
reached, decreasing
progressively with leaf age increase P B 0.01. L significantly
correlated with
SLW P B 0.01
Table 3. The plasticity index Pi, calculated on sun and
shade fully expanded leaves was the highest in Q. ilex 0.65 and the lowest in C. incanus 0.29.
4. Discussion
