Phenomorphology and Eco-morphological Characters of

 Springer 2006
Plant Ecology (2006) 187:227–247
DOI 10.1007/s11258-005-6574-0
-1
Phenomorphology and eco-morphological characters of Rhododendron lauroid
forests in the Western Mediterranean (Iberian Peninsula, Spain)
A.V. Pérez Latorre* and B. Cabezudo
Departamento de Biologı´a Vegetal. Facultad de Ciencias, Universidad de Málaga, Apartado 59, E-29080
Málaga, Spain; *Author for correspondence (e-mail: [email protected]; fax: +34 952131944)
Received 2 November 2004; accepted in revised form 25 April 2005
Key words: Functional types, Growth-forms, Phenophasic patterns, Relict, Rhododendron
Abstract
The evergreen broad-leaved forest of Rhododendron ponticum represents a special type of Mediterranean
vegetation because of their relict nature (allegedly pre-glacial, Southern-Iberian and Pontic) and connection
with Macaronesian-Atlantic flora. The findings of ecomorphological (growth forms) and phenological
(phenology) studies point to characteristics typical of its relict character and its relationship with subtropical lauroid vegetation (greater forest stratification, larger leaves, high percentage of photosynthetic
stems, scarce tomentosity, pre-flowering in a season different to Mediterranean species and closeness of
autumn–winter flowering species). There are, however, links with typical Mediterranean vegetation
(Quercus L. forests) that surrounds the Rhododendron stands, due to its adaptation to Mediterranean
climate (sclerophyll leaves, plant and leaf duration, post-fire regeneration, fleshy fruit and fruit setting-seed
dispersal seasonality). Within the community, different groups of plants show different adaptations to the
same biotope, suggesting their distinct paleo-phytogeographical origins. The results confirm the singularity
of this vegetation within the typically Mediterranean environment where it grows and its connections with
other extra-Mediterranean types.
Introduction
The use of ecomorphology (growth forms) and
phenomorphology to study Mediterranean vegetation and flora was first proposed by Orshan
(1982, 1983, 1986 and 1989). Growth forms provide information on adaptive traits (Mooney 1974;
Le Roux et al. 1984; Pierce 1984), while a phenomorphological study provides data on the complete annual cycle concerning changes in the
organs in relation to seasonal climatic changes and
plant architecture. These methods, which may be
included in the broad definition of ‘functional
types’ (Box 1987), have confirmed excellent for
cataloguing vegetation (Nemani and Running
1996), relating vegetation with climatic parameters
(Box 1996), predicting its dynamism (Noble and
Gitay 1996) even detecting species outside their
ecological context (Herrera 1984, 1987). In the
case of Mediterranean vegetation, the standardisation of methodology not only allows between
ecosystems comparison but also means it can be
used for describing as a function of ecomorphological (growth forms) and phenomorphological
characteristics (Floret et al. 1987, 1990). The
method has been used in Mediterranean regions of
the world, including Australia (Pate et al. 1984),
Chile (Orshan et al. 1984; Montenegro et al.
1989), France (Floret et al. 1987, 1990; Romane
1987), Israel (Danin and Orshan 1990; Keshet
et al. 1990), South Africa (Le Roux et al. 1989)
and Spain (Cabezudo et al. 1992, 1993; Pérez
228
Latorre et al. 1995, 2001; Caritat et al. 1997;
Navarro and Cabezudo 1998; Castro Dı́ez and
Montserrat Martı́ 1998; Pérez Latorre and Cabezudo 2002). Pérez Latorre and Cabezudo (2002)
proposed a synthesis of the Orshan’s method to be
applicable to Mediterranean climate regions in the
world. The synthesis was applied to woodlands
(Quercus suber L., Fagaceae) and shrublands (Cistus L. spp., Cistaceae) in Spain, observing clear
distinctions between the two formations as regards
both ecomorphological (growth forms) and phenological parameters (phenophases and related
indexes).
Here we continue this line of work by applying
the method proposed by Pérez Latorre and Cabezudo (2002) to a type of vegetation of great paleophytogeographical and conservation interest: the
paleomediterranean relict lauroid forests of Rhododendron ponticum L. (Ericaceae). These date
from the end of the Tertiary (Quézel 1985; Mai
1989) and are now relegated to the extreme enclaves of western (Strait of Gibraltar) to eastern
(Bosphorus) Mediterranean, showing climatic
peculiarities in both sites.
The main objective of this work is to characterise and describe this kind of plant community
using ecomorphology and phenomorphology and
to study the relationships with the ecological
parameters of the biotope where it develops. Another objective is to analyse the standardised data
to discuss the originality or similarity of Rhododendron forest to other kind of typically Mediterranean forest (Quercus suber). We also try to make
an approach to eco-phenomorphological grouping
of species following a combination of their ecomorphological and phenophasic patterns. Finally,
we try to find characters that support the relict
origin of Rhododendron forests and its eco-phenomorphological relations to present lauroid vegetation.
Methods
Vegetation
The studied vegetation type corresponds to riparian forest dominated by Rhododendron ponticum
(Scrophulario laxiflorae–Rhododendretum pontici
Pérez Latorre et al. 2000). Called ‘ojaranzal’ in
Spanish, this community is uniquely found in SW
Iberia within western Europe (Spain and Portugal)
(Pérez Latorre et al. 1996) and belongs to the
Western Mediterranean relict lauroid vegetation
(Rhododendretalia pontici Pérez Latorre et al.
2000, Pruno-Lauretea azoricae Oberdorfer ex
Sunding 1972) (Cabezudo and Pérez Latorre
2001). In Spain, its most important representative
is found in Andalusia (Los Alcornocales Natural
Park, Cadiz Province; Pérez Latorre et al. 1999)
while much smaller populations are found in the
Sierras de Monchique and Vouzela in Portugal
(Pereira Dı́as and Barros de Sa Nogueira 1973;
Malato Beliz 1982). The most unusual floristic
characteristic of this community is the brio-pteridophytic stratum with species which have a
Macaronesian-Atlantic optimum or paleomediterranean origin (such as Lepidopilum virens Card.,
Tetrastichium fontanum (Mitt.) Card., Neckera
intermedia var. laevifolia (Schiffn.) Renauld and
Cardot, Homalia webbiana Mont., Isopterygium
bottini (Breidl.) Broth-Bryophyta-. Culcita macrocarpa C. Presl., Diplazium caudatum (Cav.) JermyPteris incompleta Cav., Vandenboschia speciosa
(Willd.) Kunkel -Pteridophyta-) (Salvo 1990; Guerra et al. 2003).
Study site
The study site was chosen taking into consideration
the presence of most of the species belonging to the
community and the state of conservation. The selected R. ponticum stand (Figure 1) lies within a
protected area (Los Alcornocales Natural Park,
Los Barrios municipality, Dehesa de Ojén) in the
province of Cadiz (Spain). The riparian site occupies a 10 m wide, 20 m long stretch at 350 m a.s.l.
(536¢ W/367¢ N). A permanent stream flows on
siliceous sands, the bed of which contains large
rocky blocks. The soil data (Table 1) point to a low
pH and the absence of carbonates, while climatic
data (Figure 2, Table 2) reveal a typically Mediterranean warm climate (thermomediterranean
bioclimatic belt with an average annual temperature of 17.6 C) and much rain (humid ombrotype,
average annual rainfall of 1078 mm) although with
a dry period between July and August.
229
Figure 1. Little rectangle: distribution of Rhododendron ponticum L. in Spain and general view of the study area. Black dot: location of
the selected plot in the Natural Park of ‘Los Alcornocales’ (Sierra de Ojén, Cádiz province). White dots correspond to other
R. ponticum forests (localities taken from (Pérez Latorre et al. 1999, 2000).
230
Table 1. Main soil characteristics in the area of the study site.
Parameter
Type/data
Soil type
Rock type
pH horizon A (H2O)
C/N horizon A
CO3 horizon A
pH horizon B (H2O)
C/N horizon B
CO3 horizon B
Alfisol
Siliceous sandstones
5.6
15.7
0
4.7
11.2
0
Table 2. Weather station at Los Barrios (Polvorilla), Cádiz
(534¢ W/3615¢ N).
J
120
Mean annual rainfall
(mm)/2
100
Mean annual
temperature (ºC)
80
60
40
20
0
J
F M A M J
J A
S O N D
Figure 2. Climatic diagram of the study area.
Ecomorphology
For this ecomorphological study, we used the
method standardised by Orshan (1982, 1986),
while following the proposal by Orshan (1989) for
the phenological study. A selective plant inventory
was made in the locality, according to the presence
of the most representative species within the
community. The plot measured 200 m2, enough to
include the whole diversity of species. The inventory was made following Braun-Blanquet (1979),
including environmental data and plant cover of
the species, which was divided into persistent and
non-persistent (Table 3). Following the recommendations of Orshan (1986) we only took into
account the persistent or arid-active species (Evenari et al. 1975), that is to say, those that bear
aerial active shoots throughout the year and which
are therefore adapted to the Mediterranean
drought season. Arid-passive plants such as
F
M
A
M
J
J
A
S
O
N
D
Mean annual rainfall (mm)
163.5 169.5 104.7 84.7 50.5 14.8 0.9 2.9 27.7 101.8 191.5 165.8
Mean annual temperature (C)
13.2 14.6 15.2 15.9 18.2 21.3 23 22.4 20.8 16.9 15.3 14.3
It (Thermicity index) = 425, lower thermomediterranean bioclimatic belt with lower humid ombrotype (Rivas Martı́nez
classification 1987). Mean annual temperature = 17.6 C.
Absolute minimum temperature = 2 C. Absolute maximum
temperature = 40 C. Mean annual rainfall = 1078 mm. Days
of rainfall = 61.5.
ephemerals (terophytes) and ephemeroids (some
geophytes and hemicryptophytes) were not included in the studies, because their shoots disappear during the unfavourable season (Evenari
et al. 1975). The ecomorphological characters
(growth forms) were determined for each species in
the field. Twenty-eight characters of those proposed by Orshan (1986) were studied as well as
fruit type (fleshy, dry), as proposed in (Pérez Latorre et al. 1995). Afterwards a species/character
data matrix was made and the percentage of
presence of each character expression was calculated on the basis of number of species showing
that character (see Appendix A). For the ecomorphological description of the communities and
the subsequent comparison, we used the characters
and indices proposed by Pérez Latorre and Cabezudo (2002) including Estimated Biomass of the
Species (EBS) = plant height (m) · crown diameter (m) · canopy or branch density (%), and
Estimated
Biomass
of
the
Community
(EBC) = sum of EBS’s of all the species. Figure 3
shows the relative flat form of each species and its
positioning into the community structure.
Phenology and phenophasic indices
Data concerning the different reproductive phenological phases (flower bud formation, flowering,
fruit setting, seed dispersal) and vegetative phenological phases (vegetative growth and leaf
shedding of dolichoblasts and brachyblasts) were
recorded through monthly visits during a complete
annual cycle (2002–2003), for each arid-active or
persistent species (Evenari et al. 1975; Orshan
1989). Pteridophytes were excluded of calculations
231
Table 3. A: relevé in the study site, total community cover 100%, north facing slope 15, area: 10 · 25 m. Relative cover based on
Braun-Blanquet index (4 = 61–80%, 2 = 21–40%, 1 = 11–20%, + = 1–10%). B: percentage of presence of species in 16 localities
of the distribution area of R. ponticum forests (taken from Pérez Latorre et al. 1999, 2000); * = characteristic species of the community.
Persistent arid-active plants
*Rhododendron ponticum (Ericaceae)
*Ilex perado var. iberica (Aquifoliaceae)
*Hedera maderensis subsp. iberica (Araliaceae)
Quercus canariensis (Fagaceae)
Smilax aspera (Smilacaceae)
Ruscus hypophyllum (Ruscaceae)
*Frangula alnus subsp. baetica (Rhamnaceae)
Ruscus aculeatus (Ruscaceae)
*Laurus nobilis (Lauraceae)
Alnus glutinosa (Betulaceae)
Viburnum tinus (Caprifoliaceae)
Lonicera periclymenum subsp. hispanica (Caprifoliaceae)
Phyllirea latifolia (Oleaceae)
*Diplazium caudatum (Athyriaceae)
*Scrophularia laxiflora (Scrophulariaceae)
*Pteris incompleta (Pteridaceae)
Ephemerals arid-passive plants
Arisarum proboscideum (Araceae)
Sibthorpia europaea (Scrophulariaceae)
Ranunculus ficaria (Ranunculaceae)
Brio-pteridophytic synusial species
Athyrium filix-femina (Athyriaceae)
Vandenboschia speciosa (Hymenophyllaceae)
Davallia canariensis (Davalliaceae)
because of the lack of standard reproductive phenophases. A minimum of 30 individuals of each
species were selected and/or marked when possible, while a phenomorphological herbarium
(MGC) with representative samples of the phenological phases was collected. For each species a
phenological calendar was drawn up (see Appendix B), excluding uncommon events (Castro Dı́ez
and Montserrat Martı́ 1998). The frequency of
each phenophase taking place in each month was
calculated for the set of species. The phenological
calendar of each community was constructed as a
function of the seasonality of the following phenological phases: flower bud formation, flowering,
fruit setting, seed dispersal, vegetative growth and
leaf shedding. The descriptions of the species and
vegetation using phenophases and phenophasic
indices (Active Phenophasic Period of the Species
APS, Active Phenophasic Period of the Community APC and Index of reproductive/vegetative
Activity of the Species RVA), were made following
the model of (Pérez Latorre and Cabezudo 2002).
A
B
Relative cover
Presence (%)
4
2
1
1
1
1
+
+
+
+
+
+
+
+
+
+
100
70
90
75
60
55
85
55
50
45
40
30
30
15
15
10
2
+
+
35
35
10
1
1
+
75
10
25
Phenophasic patterns of the species were taken
from the models described by Montenegro et al.
(1989: 385) (patterns A, B, C, D, and E), although
a new phenophasic pattern (F) is here proposed to
describe the almost total coincidence of vegetative
growth with those of flower bud formation and
flowering (Figure 4).
Nomenclature
Valdés et al. (1987) and Castroviejo et al. (1986)
were used for this work (Valcárcel 2002, for Hedera; Corley and Crundwell 1991, for Bryophyta).
Results
The main ecomorphological and phenomorphological data are indicated in Appendices A and B.
As a result, we made the ecomorphological characterisation and the phenophasic calendar.
232
meters
30
25
Quercus
canariensis
Quercus
canariensis
20
15
Hedera
maderensis
Alnus
glutinosa
Hedera
maderensis
Frangula
baetica
10
Viburnum
tinus
5
2.5
Scrophularia
laxiflora
Ruscus
aculeatus
s lop e
Lonicera
his p an ic a
Ilex
perado
Laurus
nobilis
Rhododendron
ponticum +
Smilax aspera
Rhododendron
ponticum +
Smilax aspera
Phillyrea
angustifolia
Ruscus hypophyllum
0
Pteris incompleta
Diplazium caudatum
stream
Figure 3. Community structure and biotope. The species are represented to scale in two-dimensional forms, according to the ecomorphological characters of plant height x crown diameter and placed in their most common position in the biotope. Hedera
maderensis climbs on Quercus canariensis trunks whereas Smilax aspera grows into the canopy of Rhododendron-like tall shrubs and
Lonicera hispanica lies on the canopy of Phillyrea-like tall shrubs.
Ecomorphological characterisation
Phenophasic calendar
Rhododendron ponticum forest: phanerophytic,
scarcely spinescent, holoxyle, multi-layered and
evergreen community, with a maximum height of
22 m. Leaves are shed periodically on average
every 18 months; the leaves are predominantly
sclerophyll, with a 7% degree of tomentosity, and
mostly of about 38 cm2 (micro-mesophyll). The
average duration of the plants is 34.8 years; most
are adapted to after-fire regeneration by below
ground buds pattern. Vegetative growth and
flowering mainly occur in spring and fleshy fruits
are predominant.
For phenophasic calendar see, Figures 2, 5 and 6,
and Table 2. Flower buds formation from autumn
to spring, thus avoiding maximum temperatures
and the summer rainfall minima; peak flowering in
spring with a secondary maximum in autumn,
both warm, rainy seasons with many hours of
daylight; peak of fruit setting in summer, coinciding with maximal temperatures and rainfall
minima, lasting into autumn with its abundant
rainfall and moderate temperatures; abundant
seed dispersal in autumn, reaching a maximum in
winter, when temperatures are at their lowest
233
Discussion
The discussion is made following paragraphs
dealing with ecomorphological characters, phenological phases, phenophasic indices, phenophasic
patterns and eco-phenomorphological groups.
Ecomorphological characters
For ecomorphological characters see Tables 4 and 5.
Figure 4. Phenophasic patterns of the species that represent the
phenophasic sequence and overlapping of growth and flowering. (A) first: growth, second: growth and buds overlapping,
third: flowering. (B) first: growth, second: buds, third: flowering, no overlapping. (C) first: growth, second: buds and growth
overlapping, third: flowering and growth overlapping. (D) first
buds, second: flowering, third: growth, no overlapping. (E) first:
buds, second: flowering and growth overlapping, third: growth.
(F) first: buds and growth overlapping, second: flowering and
growth overlapping. White rectangle = vegetative growth, grey
rectangle = flower bud formation, black rectangle = flowering. Patterns ‘‘A’’ to ‘‘E’’ from Montenegro et al. (1989). Pattern ‘‘F’’ proposed here.
(though mild), the days are the shortest and rainfall plentiful; dolichoblast vegetative growth
maximal in spring, coinciding with an increase in
temperatures, continued rainfall and moist soils;
dolichoblast leaf shedding maximal in summer,
coinciding with the highest temperatures and
scant, if any, rainfall, although the second peak in
winter coincides with the lowest temperatures and
abundant rainfall.
Structure of the community and renewal buds
Biotype distribution (based on renewal bud position) shows that this is a forest dominated by
phanerophytes, as occurs in other types of Mediterranean forests studied (Oberdorfer 1960; Romane 1987; Floret et al. 1990; Danin and Orshan
1990; Pérez Latorre and Cabezudo 2002). However, the slight difference in stratification of this
lauroid type is quite well reflected in the scaled
distribution of biotypes according to plant heights
(Figure 3). The first layer of trees (‘roof’) is constituted by mesophanerophyte (5 species) below
which there is a dense layer of microphanerophytes (3 species). Three species of vines (one
phanerophyte, one amphiphyte and one chamaephyte) climb between these two layers. At ground
level and in areas of low luminosity grow cryptophytes (two species), hemicryptophytes (two
species) and one of the two amphiphytes, adapted
Vegetative Phenological Phases
sp %
sp %
Reproductive Phenological Phases
100
90
80
70
60
50
40
30
20
10
0
100
90
80
70
60
50
40
30
20
10
0
E
E
F
M
A
M
J
J
A
S
O
N
F
M
A
DVG
FBF
FWL
FS
M
J
J
A
S
O
N
D
D
LSD
BVG
LSB
SD
Figure 5. Time curse of the reproductive phenological phases in
the community expressed as the monthly percentage of species
that show each phenophase. Flower bud formation (FBF),
flowering (F), fruit setting (FS) and seed dispersal (SD).
Figure 6. Time curse of the vegetative phenological phases in
the community expressed as the monthly percentage of species
that show each phenophase. Dolichoblast vegetative growth
(DVG), leaf shedding dolichoblast (LSD), brachyblast vegetative growth (BVG) and leaf shedding brachyblast.
234
Table 4. Some important ecomorphological characters in the studied species.
Ecomorphological character
RW
OS
BC
SP
LS
LT
LC
LD
SO
FT
sB
Alnus glutinosa
Diplazium caudatum
Frangula alnus baetica
Hedera maderensis iberica
Ilex perado iberica
Laurus nobilis
Lonicera peryclimenum hispanica
Phyllirea latifolia
Pteris incompleta
Quercus canariensis
Rhododendron ponticum
Ruscus aculeatus
Ruscus hypophyllum
Scrophularia laxiflora
Smilax aspera altissima
Viburnum tinus
mePh
H
mePh
ePh
mePh
mePh
Am
miPh
H
mePh
miPh
Cr
Cr
Am
eCh
miPh
L
L
L
L
L
L
Bb/L
L
L
L
L
Sh
Sh
Bb/Sh
Bb
L
corky
no
smooth
papery
smooth
smooth
flaky
smooth
no
corky
flaky
smooth
smooth
no
smooth
smooth
no
no
no
no
leaves
no
no
no
no
no
no
leaves*
leaves*
no
stems
no
20–56
>1640
20–56
20–56
20–56
20–56
20–56
12–20
>1640
56–180
20–56
2–12
12–20
20–56
20–56
20–56
no
no
no
no
no
no
yes
no
no
no
no
no
no
no
no
no
Ma
sE
Ma
sE
S
S
Ma
sE
sE
sE
sE
S
Ma
Ma
S
sE
6–14
26–38
6–14
26–38
14–26
26–38
<6
14–26
14–26
6–14
6–14
14–26*
14–26*
<6
14–26
14–26
D
E
D
E
E
E
D
E
E
D
E
E
E
E
E
E
d
–
f
f
f
f
f
f
–
f
d
f
f
d
f
f
50
2
43
18
22
25
3
8
2
206
35
0.2
1
0.2
21
5
RW = renewal buds position (mePh = mesophanerophyte, miPh = microphanerophyte, ePh = escandent phanerophyte,
Am = amphiphyte, H = hemicryptophyte, Cr = cryptophyte, eCh = escandent chamaephyte). OS = organs shed rhythmically
(Bb = basipetal branch shedders, L = leaves, Sh = shoots). BC = bark consistency. SP = spinescence. LS = dolichoblast leaf
size (cm2). LT = leaf tomentosity. LC = leaf consistency (Ma = malacophyll, sE = semisclerophyll, S = sclerophyll,).
LD = dolichoblast leaf duration (months). SO = seasonality of assimilating organs (E = evergreen, D = deciduous). FT = fruit
type (d = dry, f = fleshy). sB = estimated biomass. * = phylloclades.
Table 5. Comparison among communities using ecomorphological characters and phenophases. Bold characters are mean differences
(data for Quercus community from Pérez Latorre and Cabezudo 2002).
Ecomorphological characters
Renewal buds position
Spinescence
Stratification
Maximum height
Organs shed
Bark consistency
Leaf consistency
Tomentosity (%)
Leaf size average (cm2)
Photosynthetic stems (species %)
Life duration leaves
Life duration plants
After fire vegetative regeneration
Main season of shoot growth
Main flowering season
Fruit type predominant
Biomass estimated
Phenological phases
Flower bud formation
Flowering
Fruit setting
Seed dispersal
Dolichoblast vegetative growth
Leaf shedding dolichoblast
Brachyblast vegetative growth
Leaf shedding brachyblast
Phytosociological class
Rhododendron forest
Quercus forest
diverse
almost absent
multi-strata
20–30 m
leaves
smooth > flaky
sclerophylly predominant
7%
micromesophyll (38)
44%
average 18 months
average 35 years
present 75% species
spring
spring-(bi-multiseasonal)
fleshy
28
micro-mesophanerophytic
absent
multi-strata
20–30 m
leaves and branches
flaky > smooth
sclerophylly predominant
33%
micromesophyll (28)
19%
average 19 months
average 40 years
present
bi-multiseasonal
spring
fleshy
18.3
autumn–winter–spring
spring–autumn
summer–autumn
autumn
spring
summer
throughout the year
summer
Pruno-Lauretea
winter–spring
spring
summer–autumn
autumn–winter
spring–summer
summer
brachyblast absent
brachyblast absent
Quercetea ilicis
235
to the light limiting factor. The existence of this
‘‘roof’’, which creates a favourable microclimate,
is perhaps responsible for the persistence of the
rest of the substrates. Regardless of the height of
the phanerophytes, canopy diameters of 2–5 m
predominate, suggesting strong competition for
horizontal space, although the height of Quercus
canariensis Willd. permits it to reach diameters of
15 m, while the multi-trunks growth of Rhododendron permits a canopy in excess of 5 m. Canopy density is generally very high (>75%), which
gradually diminishes the amount of light reaching
successive layers, meaning that the understorey
must be composed of shade-tolerant species, with
large shoots and leaves, maintained by the good
humidity of the biotope (Smith and Huston 1989).
The estimated biomass for this community is 27.6,
placing it well above the shrublands of Cistus (0.4)
and even Quercus suber forests (18.3) (Pérez Latorre and Cabezudo 2002).
Shoots, barks, post-fire regeneration
The community as a whole shows lignified shoots,
only the ephemeroids (not included in the study)
are axyle (Arisarum Miller, Ranunculus L., etc.).
Smooth barks predominate (8 species), unlike in
typical Mediterranean woods of Q. suber where
the predominant type is flaky (Pérez Latorre and
Cabezudo 2002). Only two species (Lonicera hispanica and Rhododendron ponticum) show shedding barks (which are also flaky) leading to an
accumulation of this inflammable dry material on
the floor. Two species with protective corky bark
(Alnus glutinosa and Quercus canariensis) are
capable of post-fire regeneration while most (12
species) regenerate by epicormic or rhizome buds
below ground level. Only ferns, the vine Lonicera
periclymenum subsp. . hispanica (Boiss. and Reuter) Nyman and the amphiphyte Scrophularia
laxiflora Lange die after fire. In this respect, Rhododendron forests closely resemble typical Mediterranean Q. suber forests, although no species
with aerial regenerating epicormic buds are to be
found (Pérez Latorre and Cabezudo 2002) for
which fire would lead to temporary destruction of
community stratification and loss for several years
of low light conditions necessary for the maintenance of relict pteridophytes and bryophytes (Salvo
1990; Cabezudo et al. 1995; Guerra et al. 2003).
Photosynthetic organs
One of the most important aspects of the forest is
the leaves area. Large leaves predominate in the
community (micro-mesophyll, 20–56 cm2), with
the presence of a very large-leafed species (Quercus
canariensis) with 56–180 cm2 (mesophyll) and two
ferns with megaphyll leaves. Such characteristics
differentiate this wood even further from the
Mediterranean Q. suber forest with its predominant leaf size of 2–20 cm2 and where it is also
possible to find nanophyll-leaved species
(<2 cm2). This predominance of large leaves and
high canopy densities provides strong shade to the
lower strata and the ground, almost totally
inhibiting the presence of herbaceous undergrowth, but preserves the relict ferns, which would
otherwise disappear through the photo-destruction
of its pigments (Ratcliffe et al. 1993). Only in autumn and winter does the intensity of the shade
diminish due to the loss of leaves of the three
deciduous species (Q. canariensis, Frangula alnus
ssp. baetica (Reverchon and Willk.) Rivas Goday
ex Devesa and Alnus glutinosa (L.) Gaertner). The
biomass represented by these large leaves is
maintained by the favourable environmental conditions of the area with a mean temperature above
the minimum for vegetative activity, abundant
rainfall, mists and high soil humidity levels (Werger and Ellenbroek 1978; Keshet et al. 1990), unlike those found in other Mediterranean woods of
average humidity, such as Q. suber forests (Pérez
Latorre and Cabezudo 2002). The predominance
of large leaves may also be due to the low level of
nutrients in the soil (Givinish 1987) and lent
weight by the presence of Alnus, a known nitrogenfixer. The dense shade may explain the high percentage of species with photosynthetic stems
(44%), three small species (Ruscus L. spp. and
Scrophularia) which grows in the understorey,
Smilax aspera L., which only climbs to intermediate height levels, and Hedera maderensis ssp.
iberica McAllister, whose stems remain below tree
canopies. The most interesting case is that
regarding Ilex perado var. iberica Loes. and Laurus
nobilis L., both small trees, with their long-lasting
photosynthetic branches (3–5 years) and height of
half a metre or more, which have low light and
water requirements corresponding to stress-tolerant plants (long-lasting aerial parts, sclerophyll
leaves, photosynthetic stems) (Grime 1979; Chapin
236
et al. 1993) but which grow in a biotope which
theoretically provides good year-round conditions.
(Danin and Orshan 1990) points to the total
inhibition of the latter due to the conditions imposed by the dense forest, except for the climber
chamaephyte Smilax.
Leaves morphology
The combination of green and glabrous leaves (15
species), without defence against dry conditions
such as hairs and resins would provide (Oppenheimer 1960), and horizontally inserted leaves (12
species) predominates, which, together with the
large areas of the leaves, produces a very attenuated
light in intermediate and low layers of the vegetation
and produces a humid biotope (Keshet et al. 1990).
The glabrous leaves with some sclerophyll degree
are in contrast with the malacophyll-glabrous
combination typical of Mediterranean shrublands
(Pérez Latorre and Cabezudo 2002). A group of
malacophyll-glabrous plants (Alnus and Frangula)
corresponds to winter-deciduous species whose
leaves last less than 1 year. Sclerophyll leaf implies
adaptation to water stress (Parsons 1976; Campbell
and Cowling 1985; Givinish 1987) and is therefore
somewhat surprising in these conditions where there
is no lack of water (Grieve 1953). However, it is
probably the result of the pre-Mediterranean origin
of the concerned species and their relict status
(Axelrod 1975; Herrera 1984). Even it may reflect
‘ghost’ paleoclimatic conditions different from the
present day (Went 1971) or simply is the result of the
nutrient-poor soils (Table 1) (Mooney et al. 1983)
(with Alnus as a nitrogen-fixer) typical of tropical
woods (Larcher 1977) with which R. ponticum forest
may be ecomorphologically related.
Spinescence is unusual in the forest, as it is in
other humid Mediterranean woods (Pérez Latorre
and Cabezudo 2002) and, curiously, only appears
on leaves or phylloclades (Smilax, not always, Ilex,
not always, and Ruscus spp.) and not on stems.
Long-lasting (2 or 3 years) leaves predominate, a
characteristic positively associated with increased
rainfall (Keshet et al. 1990) in Mediterranean
vegetation (Orshan 1982), conferring an evergreen
appearance to the community, except in the case of
the small group of winter-deciduous and malacophyll species (Alnus, Frangula, Quercus, Lonicera)
whose leaves last less than a year and which
probably have an origin, as floristic contingent,
different to the sclerophyll evergreen species
(which, nevertheless, partially lose their leaves in
summer). The phanerophytes/chamaephytes index
Seasonality, fruits
Rhododendron forest is an evergreen community,
reflecting mature Mediterranean formations of
sucesional stages. However, there is a deciduous
contingent of three species (Alnus glutinosa, Frangula alnus subsp. baetica, Quercus canariensis)
which points to a certain degree of coldness and
humidity (Keshet et al. 1990), the first of which no
longer applies, so that Alnus and Frangula, at least,
probably come from a floristic contingent (EuroSiberian), a relict of cold periglacial eras. The
community is characterised by plants (10 species)
with a predominantly spring growth pattern and
which include the deciduous plants, and another
six species showing multi-seasonal growth (evergreens). Growth stops from mid-summer to midwinter, which does not reflect the good prevailing
hydric and climatic characteristics (Table 2). This
disagreement reinforces the possibility of an origin
in differently-adapted floras. The flowering season
underlines this contradiction, since although typical Mediterranean spring flowering predominates
(6 species), two species (Alnus, Viburnum tinus L.)
flower in winter and two (Hedera, Smilax, climbers) in autumn (Herrera 1984). The species showing multi-seasonal flowering or spores dispersal
are, curiously, relict ferns and Ruscus ssp. The
fleshy fruits are characteristic of other Mediterranean woods (Pérez Latorre and Cabezudo 2002)
and point to the summer availability of ground
water, which disagrees with the typical and almost
total absence of summer rains.
Phenological phases
For phenological phases see Figure 5, 6 and 7 and
Table 5.
Flower buds formation
This phenophase is maintained throughout the
year, with minimum levels in summer and maximum in September, unlike in woods of Quercus
237
suber, when the maximum occurs in February–
March. Such long periods (lasting more than
5 months) are frequent, sometimes preceding a
short flowering period (Alnus, Rhododendron,
Viburnum) and sometimes lasting the whole year
(Ruscus); while in the case of Laurus nobilis it is
due to two flowerings, one in spring and one in
autumn. The shortest period (1 month) occurs in
Quercus canariensis.
Flowering
Maximum flowering occurs in spring, reducing to
almost zero in summer, the least favourable season, and with a significant secondary peak in autumn. Spring flowering species include Frangula
baetica, Quercus canariensis, Rhododendron ponticum and Ilex perado. Those flowering in autumn
are Hedera maderensis and Smilax aspera, while
Laurus nobilis flowers in both spring and autumn,
although no fruits appear in the latter. Alnus glutinosa and Viburnum tinus flower in winter, while
both species of Ruscus flower practically the whole
year, except in the middle of summer. Phillyrea
latifolia L. did not flower during the study period,
as occurred in the study of a subhumid Mediterranean wood of Quercus suber (Pérez Latorre and
Cabezudo 2002), perhaps due to an excess of shade
(Sack et al. 2003). The combination of different
flowering periods and a secondary peak in autumn
are similar to those detected by Pérez Latorre and
Cabezudo (op. cit.). The pteridophytes (Diplazium,
Pteris) sporulated throughout the year.
Fruit setting
Fructification begins in spring and is at a height
throughout the summer, with a secondary peak in
autumn and a minimum in winter, as similar to the
pattern of Quercus suber woods observed by Pérez
Latorre and Cabezudo (2002). Since most are fleshy fruit, it is not surprising that fructification lasts
until autumn. The longest fruiting period is that of
Ilex perado and Laurus nobilis (8 months) and the
shortest occurs in Frangula, Smilax and Lonicera
(2 months, despite the fleshy nature of the fruit).
Both species of Ruscus bear fruit throughout the
year, the result of several flowerings and supported
by the permanent shaded conditions (Sack et al.
2003), although few individuals of the respective
populations do so at a time.
Seed dispersal
Dispersion shows two maxima, in autumn and
winter, with a minimum as spring turns to summer, at which time other antagonist phenophase
(flowering) is at its height, as occurs in Mediterranean woods of Quercus suber (Pérez Latorre and
Cabezudo 2002). Summer dispersing species are
Scrophularia laxiflora (dry fruit/nuts) and Lonicera
hispanica, while Ruscus spp. disperses its fruits
throughout the year, although always in isolated
individuals. The longest seed dispersal period is
that shown by Alnus, Ilex and Viburnum, which
maintain mature fruit on their branches for up to
6 months.
Leaf shedding
Leaf shedding is maximal in summer, coinciding
with the dry period and decrease in the water level
of the stream, which is typical of evergreen species
suffering a facultative and partial loss of dolichoblast leaves. A second maximum occurs at the
beginning of winter due to the presence of deciduous species (Frangula, Quercus, Alnus) the first
continuing to shed leaves until the following
spring. Neither Ruscus spp. nor Smilax shed their
leaves (phylloclades in the former) since the
branches are completely renewed. Quercus canariensis does not shed all of its leaves but maintains a
small percentage even at the beginning of spring,
when the new branches are formed, behaviour
similar to that recorded for Quercus faginea ssp.
broteroi (Cout.) A. Camus in Quercus suber woods
(Pérez Latorre and Cabezudo, 2002). This may be
the result of the favourable climatic conditions of
the study area.
Vegetative growth
Vegetative growth is maximal during the spring
and is zero in August, September and October,
unlike in other Mediterranean woods (Pérez Latorre and Cabezudo 2002) where the growth, although low, continues in these months. No species
238
Phenophasic indices
sp %
For phenophasic indices see Appendix B. The
Active Phenophasic Period of Species (APS),
which indicates the number of months with
favourable conditions for reproductive activity
and growth, points to three phenomorphological patterns in the community. A very heterogeneous group making up the majority of
species and representative of the community
shows activity practically throughout the year.
These are Alnus, Hedera, Ilex, Laurus, Lonicera,
Scrophularia, Rhododendron, Ruscus, Viburnum
and the pteridophytes Diplazium and Pteris.
Another group shows activity during 8–
9 months (Quercus, Smilax), while Frangula alnus concentrates its activity into 4 months. Of
special interest is Phillyrea, in whose adult
population no reproductive phenophasic activity
was detected, probably because of the aboveDead Matter
100
90
80
70
60
50
40
30
20
10
0
DM
E
F
M
A
M
J
J
A
S
O
N
D
Figure 7. Time curse of the presence of dead matter on the
plants shoots.
Active Phenophasic Period of Community (APC)
sp %
continues growing throughout the year, which is
perhaps surprising, given that the climatic (see
Figure 2 and Table 2) and soil (permanent riparian
water) conditions are suitable for a year-round
growth. However, there are partial flower buds
formation and fructification peaks, which presumably compete for the resources necessary for
growth (Castro Dı́ez et al. 2003). Some species do
keep growing for several months (7–9 months in
the case of Scrophularia, Hedera and Pteris), while
Diplazium is the only species that continues to grow
throughout the year. The shortest growth period is
that of Frangula, Quercus, Rhododendron, Ilex and
Phillyrea, which all last for 2 months or less. The
accumulation of dead matter on the plants (branches, inflorescences, etc.) is greatest in spring (80%)
and least in autumn (35%) (Figure 7).
100
90
80
70
60
50
40
30
20
10
0
E
F
M
A
M
J
Rhododendron forest
J
A
S
O
N
D
Quercus suber forest
Figure 8. Time curse of the APC index (monthly percentage of
species that show phenophasic activity). Comparison between
Rhododendron and Quercus suber communities.
mentioned reasons. (Pérez Latorre and Cabezudo 2002) also identified three groups in
Mediterranean Q. suber woods, although less
heterogeneous in their case.
As regards the Active Phenophasic Period of
the Community (APC) (Figure 8), which indicates the percentage of active species (in the sense
of APS) for each month during the year, the
community as a whole shows phenophasic activity between 60 and 100% of the year, but with a
spring maximum and winter minimum, with a
small burst during autumn. This pattern shows
that, despite the good conditions of the biotope
as regards temperature and humidity, winter
slows down the biomass-forming and reproductive activity. In comparison, Mediterranean
woods of Quercus suber showed a slightly higher
activity in winter, summer and autumn than
Rhododendron forest although both winter and
summer conditions are worse than in our study
area. The community under study only showed a
slightly higher APS in spring. However, the
overall curve obtained for the APS is almost
identical for both the Rhododendron forest and
Quercus suber wood.
The Index of reproductive/vegetative Activity of
the Species (RVA) (Table 6) gives an idea of the
different strategies with respect to the time and
resources spent in a balance between reproductive
and vegetative phenological phases. Most of the
plants (12 species), including all the trees and
shrubs, are above 1, indicating the predominance
of reproductive over vegetative phenophases. The
maximum was obtained by Rhododendron with 6
and Ilex with 5.5. Four species (all climbers) score
less than 1, the minimum (0.33) belonging to
the amphiphyte of the understorey Scrophularia
239
Species
RVA Phenophasic patterns
Scrophularia laxiflora
Lonicera peryclimenum hispanica
Smilax aspera altissima
Hedera maderensis iberica
Frangula alnus baetica
Ruscus hypophyllum
Viburnum tinus
Ruscus aculeatus
Alnus glutinosa
Laurus nobilis
Quercus canariensis
Ilex perado iberica
Rhododendron ponticum
Phyllirea latifolia
0,33
0,5
0,8
0,86
2
2
2,2
2,4
3
4
4
5,5
6
–
C
B
B
B
F
E
E
E
E
E
F
E
D
D
laxiflora, which shows a predominance of vegetative over reproductive phenophases. Similar patterns were obtained by (Pérez Latorre and
Cabezudo 2002) in Quercus suber woods, where all
the trees and shrubs scored above 1, except some
shrubs (Cytisus L., Genista L.) and Erica arborea L.
Phenophasic patterns
The species of the community can be grouped as
follows (see Figures 4 and 9): no species have
pattern A; pattern B is showed by the three
climbers of the community (Hedera, Lonicera and
Smilax); pattern C (flower buds formation and
flowering coinciding) corresponds to the only
amphiphyte (Scrophularia); pattern D (Phillyrea
and Rhododendron); pattern E, major species (Alnus, Ilex, Laurus, Ruscus spp. and Viburnum);
pattern F is for deciduous trees (Frangula and
Quercus). Grouping these patterns into two basic
types, the species can be divided into: (a) the five
species that grow first and then flower
(A + B + D) and (b) those that grow and flower
at the same time (C + E + F) (9 species). Similar
phenophasic patterns were seen in the Mediterranean wood of Quercus suber (Pérez Latorre and
Cabezudo 2002). There were no significant differences between the communities as regards patterns
B, D, E and F, although the first three parameters
were slightly higher in the Rhododendron forest
and F was higher in the Q. suber wood. Pattern C
is much less common in the Rhododendron forest.
Eco-phenomorphological groups
It is possible to create a series of eco-phenomorphological groups in communities by combining the
phenophasic behaviour of the species and given ecomorphological characters. In this way, groups with
different adaptations and behaviour can be identified in the same biotope and within the same community (Pérez Latorre and Cabezudo 2002). Seven
such groups appear in the Rhododendron forest (see
Table 7) as a function of the main flowering/spores
dispersal season, phenophasic pattern and biological type (plant size) plus seasonality and leaf consistency. The result is somewhat surprising since the
response of the vegetation to such a homogeneous
biotope with so few species shows very different
phenophasic and ecomorphological adaptations,
which might suggest different contingents of plants
adapted to the biotope over a long period of time
(Herrera 1984). The only strictly winter-flowering
species are Alnus and Viburnum, which show a very
similar phenophasic calendar and identical pattern,
but very different biological types and leaves. The
spring-flowering species, Frangula and Quercus,
show almost identical calendar and pattern, differentiating themselves from the following group in
Phenophasic patterns
45
40
35
species %
Table 6. RVA index and phenophasic patterns of the species of
the community.
30
25
20
15
10
5
0
A
B
C
Rhododendron forest
D
E
F
Quercus suber forest
Figure 9 Percentage of species presenting each phenophasic
pattern (A–F). Comparison between R. ponticum forest and
Quercus suber forest.
240
Table 7. Grouping of species according to similarity of phenology and selected ecomorphological characters (renewal buds position,
type of fruit, seasonality, spinescence and presence of leaves).
Species
FBF
F
FS
SD
DVG LSD PPT
Alnus glutinosa
Viburnum tinus
Frangula alnus baetica
Quercus canariensis
Ilex perado iberica
Laurus nobilis
Rhododendron ponticum
Phillyrea latifolia
Hedera helix
Smilax mauritanica altissima
Lonicera peryclimenum hispanica
Scrophularia laxiflora
Ruscus aculeatus
Ruscus hypophyllum
Diplazium caudatum
Pteris incompleta
UA
AW
S
S
S
AWS
AWS
–
U
UA
SU
S
AWS
UAW
–
–
W
W
S
S
S
S
S
–
A
A
U
SU
AWS
AWS
year
year
SU
SU
U
UA
UA
SUA
UA
–
A
A
U
U
year
year
–
–
AW
AW
UA
AW
AW
A
AW
–
W
W
U
UA
year
year
–
–
S
WS
S
S
S
S
SU
S
WSU
SU
S
WS
S
SU
year
WS
AW
U
AWS
W
SU
U
U
S
U
–
U
–
–
–
–
–
E
E
F
F
E
E
D
D
B
B
B
C
E
E
–
–
Mesophanerophyte malacophyll deciduous tree
Microphanerophyte semisclerophyll evergreen tall shrub
Mesophanerophyte malacophyll deciduous tree
Mesophanerophyte semisclerophyll deciduous tree
Mesophanerophyte sclerophyll evergreen tree
Mesophanerophyte sclerophyll evergreen tree
Microphanerophyte semisclerophyll evergreen tall shrub
Microphanerophyte semisclerophyll evergreen tall shrub
Scandent phanerophyte semisclerophyll evergreen
Scandent chamaephyte sclerophyll evergreen
Scandent phanero-chamaephyte malacophyll deciduous
Hemi-chamaephyte malacophyll evergreen herb
Cryptophyte sclerophyll evergreen small shrub
Cryptophyte malacophyll evergreen small shrub
Hemicryptophyte pteridophyte fern
Hemicryptophyte pteridophyte fern
FBF: flower bud formation, F: flowering, FS: fruit setting, SD: seed dispersal, DVG: dolichoblast vegetative growth, LSD: leaf
shedding dolichoblast, PPT: phenophasic pattern. Bold letters indicate common or unique differential characters of the groups.
Seasons: W = winter, S = spring, U = summer and A = autumn.
that they are deciduous. Ilex, Laurus and Rhododendron are also very similar, including their
eco-morphological traits (especially of the leaves),
although the first show an E pattern and the third a
D type. With autumnal flowering, Hedera and
Smilax show almost identical calendars and phenophasic patterns but differ in their architecture and
ecomorphology. The only species that flower in
summer are Lonicera and Scrophularia, with very
similar calendar but different patterns and biological types. The two species of Ruscus are, unsurprisingly, very similar, with multi-seasonal
reproductive phenophases. Phillyrea distances itself
from all the plants of the forest because of its lack of
a reproductive phenophase, but is included in the
group of Rhododendron because of its vegetative
phenophases, phenophasic pattern and biological
type. Diplazium and Pteris are grouped together, as
is to be expected, because of the level of their
pteridophytic form and multi-seasonal spores dispersal (reproduction).
Conclusions
Similarities between Rhododendron forests and
Quercus suber woods are phenologically reflected
(Table 5) in fruit setting, seed dispersal, APS and
phenophasic patterns. As regards common ecomorphological traits, the most important aspects
are the almost complete absence of spinescence,
maximum trees height, sclerophylly, leaf and plant
duration, after-fire vegetative regeneration and fleshy fruits.
Among the differential features (Table 5) are the
greater number of layers in Rhododendron forest,
the greater amount of estimated biomass, the
predominance of smooth over flaky barks, the
greater leaf size (micro-mesophyll), the higher
percentage of photosynthetic stems and less
marked tomentosity. Phenologically, a differentiating character is the maximum flower buds formation that is reached at the beginning of autumn
and which lasts throughout the year, the four
species (25% of the total) that flower in autumn/
winter and the low prevalence of the C-type phenophasic pattern (flower bud formation and
flowering during the last stage of growth).
The relict status of Rhododendron forest can be
attributed to several factors that contrast with the
prevailing Mediterranean macroclimate, such as
the good conditions that prevail in riparian biotopes throughout the year. Stress tolerance characteristics, such as sclerophylly, are to be found,
which may have three possible explanations: (a) the
poor quality and low pH of the soils, (b) adaptation
to water stress and (c) the possibility that the
community originated from an ancient subtropical
vegetation during the Tertiary. The phenology
follows seasonal Mediterranean rhythms and does
not reflect the good conditions which last
241
throughout the year, since, given these, vegetative
and flowering activity might be expected throughout the year, which is not the case (zero DVG in
August, September and October; minimum APC in
winter). It is possible that this discordance between
adaptation and biotope conditions is due to an
adaptation to longer cycles (several years) of
drought, during which the stream would dry up
completely, creating conditions similar to those
that would arise from the absence of rainfall.
Lastly, the combination of evergreen-sclerophyll
leaves, average to large seeds, zoochory dispersion
(fleshy fruit) and late sucesional stages points to a
flora derived from tertiary-type paleotropical conditions (Herrera 1982, 1984; Axelrod 1975).
The paleotropical relict origin of this community may be supported by the discordance between the adaptive significance of the characters
studied and the biotope where they occur, constituting a good criterion (besides floristic singularity in the bryo-pteridophytic stratum) for
ecosystem conservation under EEC Directive 92/
43 referring to EU ‘‘habitats’’ and as Special
Conservation Area of the future European Nature Network 2000.
In the Pontic area (eastern Mediterranean)
Rhododendron occurs as understorey of temperate
deciduous forests of Fagus orientalis (Filibeck
et al. 2004) while in the Iberian Peninsula Rhododendron grows in Mediterranean relict lauroid
forests as this work points out. This is a thoughtprovoking point for future investigations, but we
will not get out of the impasse until we gain palaeobotanical information, almost completely
lacking at the present day.
Other paths for further investigation may be
proposed such as the eco-physiology of relic ferns
linked exclusively to the extreme ambient of shade
and phenological calendar of this kind of forest.
Studies on functional and taxonomical relationships of the mediterranean species of the genera
Frangula, Ilex and Laurus to their Macaronesian
related species will add information about the
origin of this Rhododendron forests. Remains a
palaeobotanical mystery, up to date, the absence
of R. ponticum in the north of Morocco, few
kilometres far from the Spanish Rhododendron
forests. Effects of climate change in this fragile
water-dependant ecosystems is other avenue of
future research. Finally, phytosociological comparisons between the Rhododendron Iberian forests
and those of Bulgaria (Strandzja) may clarify the
common origin of this kind of relic paleomediterranean vegetation.
Acknowledgements
Project REN 2000-1155 GLO (C.A.I.C.Y.T.,
Spain) ‘‘Diversidad vegetal, ecologı́a y estructura
de los bosques lauroides relı́cticos del sur de la
Penı́nsula Ibérica: fitocenosis, especies crı́ticas,
variabilidad genética de poblaciones y conservación’’ has supported the studies. Dr. J. Carrión
from the University of Murcia (Spain) and anonymous referees have made some valuable suggestions on the manuscript.
Appendix A.
Appendix A.
Plants
Renewal bud:
mesophanerophyte
Renewal bud:
microphanerophyte
Renewal bud:
escandent
phanerophyte
Renewal bud:
hemicryptophyte
Renewal bud:
amphiphyte
Renewal bud:
cryptophyte
Renewal bud:
chamaephyte
Organs shed:
leaves
Organs shed:
amphiphyte
Organs shed:
shoots
Organs shed:
branches basipetal
Plant height:
50–100 cm
Plant height:
senseless
Plant height: 5–10 m
Plant height: 1–2 m
Plant height: 2–5 m
Plant height: 10–20 m
% of
plants
5
31
3
19
1
6
2
13
2
13
2
13
1
6
11
69
2
13
2
13
1
6
3
19
3
19
3
2
2
2
19
13
13
13
242
Appendix A. Continued.
Appendix A. Continued.
Plants % of
plants
Plant height: 20–30 m
1
Crown diameter: 2–5 m
6
Crown diameter: 1–2 m
3
Crown diameter: senseless
3
Crown diameter: 50–100 cm
1
Crown diameter: 25–50 cm
1
Crown diameter: 5–10 m
1
Crown diameter: >10 m
1
Canopy density: 75–90%
5
Canopy density: >90 %
4
Canopy density: 50–75%
4
Canopy density: 25–50%
3
Stem consistency: holoxyle
15
Stem consistency: hemixyle
1
Bark consistency: smooth
7
Bark consistency: none
4
Bark consistency: flaky
2
Bark consistency: corky
2
Bark consistency: papery
1
Bark thickness <2
14
Bark thickness 10–20 mm
1
Bark thickness 20–50 cm
1
Bark shedding rhythm none
14
Bark shedding rhythm 2–5 years
1
Bark shedding rhythm >5 years
1
Spinescence absent
13
Spinescence leaves
2
Spinescence stems
1
Size larger leaves 20–56 cm2
10
Size larger leaves >1640 cm2
2
Size larger leaves 12–20 cm2
2
1
Size larger leaves 56–180 cm2
Size larger leaves 2–12 cm2
1
Size smaller leaves: no leaves
15
Size smaller leaves <0.2–2 cm2
1
Length of larger leaves 5–10
10
Length of larger leaves >50 cm
2
Length of larger leaves 2–5
2
Length of larger leaves 10–20
2
Length of smaller leaves: no sm. leaves
15
Length of smaller leaves 2–5
1
Length of photosynthetic stems:
9
no phot. stems
Length of photosynthetic stems
5
>50 cm
Length of photosynthetic stems 20–50
2
Width of larger leaves 20–50
11
Width of larger leaves: >50 cm
5
Width of smaller leaves: no sm. leaves
15
Width of smaller leaves 5–10 mm
1
Width of photosynthetic stems: no phot. stems 9
Width of photosynthetic stems 2–3 mm
4
Width of photosynthetic stems 3–5 mm
2
Width of photosynthetic stems 5–10
1
Leaf colour: all green
15
Leaf colour green and glaucous
1
6
38
19
19
6
6
6
6
31
25
25
19
94
6
44
25
13
13
6
88
6
6
88
6
6
81
13
6
63
13
13
6
6
94
6
63
13
13
13
94
6
56
31
13
69
31
94
6
56
25
13
6
94
6
Leaf angle mainly horizontal
Leaf angle mainly vertical
Leaf angle all transitions
Leaf tomentosity: non tomentose
Leaf tomentosity lower side
Leaf consistency: semi–sclerophyll
Leaf consistency malacophyll
Leaf consistency sclerophyll
Surface resins absent
Ratio leaves/assimilating stems: all assim.
leaves
Ratio leaves/assimilating stems: leaves >
stems
Ratio leaves/assimilating stems: leaves
aprox. = stems
Life duration of plant 2–5 years
Life duration of plant 5–25 years
Life duration of plant 25–50 years
Life duration of plant 50–100 years
Life duration of plant >100 years
Life duration larger leaves 14–26 months
Life duration larger leaves 6–14 months
Life duration larger leaves <6 months
Life duration larger leaves 26–38 months
Life duration smaller leaves: no sm. leaves
Life duration smaller leaves: <6 months
Life duration assimilating stems: no ass. stems
Life duration assimilating stems 1–2 years
Life duration assimilating stems 2–3 years
Life duration assimilating stems 3–5 years
Seasonality of assimilating organs evergreen
Seasonality of assimilating organs winter
deciduous
Seasonality of assimilating organs summer
deciduous
Main season of shoot growth spring
Main season of shoot growth bi-multiseasonal
Main flowering season spring
Main flowering season bi-multiseasonal
Main flowering season autumn
Main flowering season winter
Main flowering season summer
Main flowering season: no flowering
Vegetative regeneration after fire plant killed
Vegetative regeneration after fire below
ground buds
Vegetative regeneration below ground non
epicormic buds
Trophic types autotrophic only
Trophic types N fixing
Fruit type: fleshy or fleshy cotyledons
Fruit type: dry
Fruit type: no fruits
Plants
% of
plants
12
2
2
15
1
7
5
4
16
9
75
13
13
94
6
44
31
25
100
56
6
38
1
6
2
5
6
2
1
7
4
2
3
15
1
9
2
2
3
12
3
13
31
38
13
6
44
25
13
19
94
6
56
13
13
19
75
19
1
6
10
6
63
38
6
4
2
2
1
1
4
8
38
25
13
13
6
6
25
50
4
25
15
1
11
3
2
94
6
69
19
13
243
Appendix B.
244
Appendix B. Continued.
245
Appendix B. Continued.
246
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