This work is dedicated to Prof. G. Orshan.

Use of monocharacteristic growth forms and phenological phases todescribe and differentiate plant communities in Mediterranean-typeecosystems

A.V. Pérez Latorre* and B. CabezudoDepartamento de Biología Vegetal. Facultad de Ciencias, Universidad de Málaga, Apartado 59, E-29080Málaga, Spain; *Author for correspondence (e-mail: avperez@uma.es; fax +34 952131944)

Received 10 April 2001; accepted in revised form 31 October 2001

Key words: Cistus shrublands, Mediterranean vegetation, Phenological indexes, Plant functional types, Quercussuber forests

Abstract

The ecomorphological and phenological study was carried out within a Mediterranean vegetation context, inQuercus suber forests, which have been substituted by shrublands of Cistus spp within two Natural Parks in thesouth of the Iberian Peninsula. The ecomorphological characters that show meaningful differences between bothtypes of vegetation are: location of renewal buds, spinescence, stratification, maximum height of the vegetation,organs periodically shed, leaf consistency, leaf tomentosity, leaf size, and life duration of leaves, plant duration,vegetative regeneration after fire, main vegetative growth season, main flowering season and fruit type. The phe-nological phases also help to discern between forest and shrubland, specially flower bud formation, fruiting, seeddispersal, and the existence of brachyblast vegetative growth and brachyblast leaf shedding. We propose threenew indexes based on phenological phases: “active period of the species” (APS), “active period of the commu-nity” (APC) and “reproductive/vegetative activity of the species” (RVA). The results of their application, in com-bination with the ecomorphological characters, have proved promising in describing vegetation and in clearlydifferentiating communities. The results also show the existence of different ecomorphological groups of plantsat community level, with consequent ecological, historical, phytocoenological and adaptive implications

Introduction

Methods that study terrestrial vegetation according toplant functional types (Box 1987) are considered toshow a good correlation with atmospheric models(Box 1996), climatic change models (Chapin et al.1996) and predictive models based on the dynamicsof vegetation at a global level (Noble and Gitay1996). Such methods are used as an alternative to thetraditional systems of phytogeographical classifica-tion (Nemani and Running 1996). Within these mor-phofunctional systems, Orshan (1982, 1983, 1986,1989) proposed the use of monocharacteristic growthforms (ecomorphological characters) and phenomor-phology to study Mediterranean flora. This method-ology can be used in small areas and makes it possi-

ble to determine the functional patterns ofMediterranean ecosystems and to describe themthrough their ecomorphological and phenomorpho-logical attributes (Floret et al. (1987, 1990)). Thesepatterns are related to the functionalism and adaptivecharacters of the components (species) of the ecosys-tems (Mooney 1974 Le Roux et al. 1984 Pierce 1984)and permit us to detect groups of species adapted topast conditions that were different from those exist-ing today (Herrera (1984, 1987)). Such studies havebeen carried out in Australia (Pate et al. 1984), Chile(Orshan et al. 1984), France (Floret et al. (1987,1990) Romane 1987), Israel (Danin and Orshan 1990Keshet et al. 1990) and in the Iberian Peninsula,where they were centred on the typically Mediterra-nean vegetation of Quercus suber and Quercus rotun-

231Plant Ecology 161: 231–249, 2002.© 2002 Kluwer Academic Publishers. Printed in the Netherlands.

difolia forests, Cistus shrublands and other shrub veg-etation. From this ecomorphological andphenomorphological point of view, the Quercus suberforests have been studied by Cabezudo et al. (1993)and Pérez Latorre et al. (1995, 2001) and, as part of awide study, by Caritat et al. (1997), while the Cistusshrublands have been studied by Cabezudo et al.(1992) and Navarro and Cabezudo (1998). These re-sults complement the floristic and phytocoenologicalstudies on these communities in the south of the Ibe-rian Peninsula (Pérez Latorre 1993 Pérez Latorre etal. (1993, 1994, 1996, 1997)).

The objective of this work was to improve and testa methodology based on the phenological and eco-morphological behaviour (phenological phases andgrowth forms) of the communities as an instrumentto characterise and differentiate both vegetationstages (forest and shrubland). We also tried to deter-mine if the observed structural and phenological dif-ferences are due to adaptation to the different newecological conditions as a result of the successionprocesses. The results obtained are compared withthose obtained using the same methodology in othercommunities of the Western Mediterranean, such asthose studied by Floret et al. (1987, 1990) and Na-varro et al. (1994), Navarro and Cabezudo (1998),Castro Díez and Montserrat Martí (1998).

Material and methods

Study sites

The studied plant communities are distributedthroughout the south of the Iberian Peninsula (Málagaprovince, Andalusia) (Figure 1). The forest corre-sponds to the phytosociological association Teucriobaetici-Quercetum suberis Rivas-Martínez in DíezGarretas, Cuenca and Asensi 1988. The substitutionshrubland corresponds to Lavandulo caesiae-Genistetum equisetiformis Rivas Goday and RivasMartínez 1968 (Nieto Caldera et al. 1991 PérezLatorre et al. (1993, 1994)). Both communities growfrom sea level to 1000 m in the thermo-mediterraneanand meso-mediterranean bioclimatic belts (RivasMartínez 1987) on acid soils with an annual rainfallabove 600 mm. The successional dynamism betweenboth communities is caused by recurrent fires andovergrazing, with the attendant consequences of soilerosion and floristic, physiognomic, phytosociologi-

cal and structural changes in the communities (PérezLatorre et al. 1994) (Figure 2).

The representative study areas were selected ac-cording to the presence of the most characteristic spe-cies of both communities (Table 2). The Quercussuber forest is found in the “Sierra de las Nieves”Natural Park (Málaga province) 700 m above sealevel, developing on micaschists and well-structuredeutric cambisols. The climatic data from the NationalMeteorology Institute can be observed in Figure 3 andTable 1. The area is included in the meso-mediterra-

Figure 1. Distribution area of the communities studied and loca-tion of study sites (Málaga province, Spain). A: Quercus suber for-est, B: Cistus shrubland.

Figure 2. Physiognomy, soil depth and dynamism of the studiedplant communities. (mePh: mesophanerophyte, miPh: microphan-erophyte, naPh: nanophanerophyte, Ch: chamaephyte).

232

nean bioclimatic belt (mean annual temperature 16°),with a humid ombro-type (1000 mm average annualrainfall) with a dry period from June to mid Septem-ber. The Cistus shrubland is found in the Natural Parkof “Los Montes” (Málaga province) at 800 m abovesea level, developing on schists and severely erodedeutric cambisols. The climatic data can be observedin Figure 3 and Table 1. The area is included in themeso-mediterranean bioclimatic belt (mean annualtemperature 15 °) with a subhumid ombro-type (800mm average annual rainfall) with a dry period fromthe end of May to about the end of September.

Inventories were made to study the floristic com-position of the communities in their distribution area(Table 2). The structure of the forest and of the shru-bland as regards the species they contain can be ob-served in Table 2 and the structure of the communitiesin Figure 2. In the distribution area of the Quercussuber forests, the species with the greatest presencewere Quercus suber, Erica arborea, Arbutus unedo,Quercus broteroi, Pistacia lentiscus and Daphnegnidium. In the study plot, the species with the great-est cover were Quercus suber, Viburnum tinus andArbutus unedo. In the distribution area of the shru-bland, the species with the greatest presence are: La-vandula stoechas, Genista umbellata, Ulex parviflo-rus, Cistus monspeliensis, Cistus albidus and Cistusladanifer, while in the study plot, the species withgreatest coverage are Cistus ladanifer, Cistus mon-speliensis and Cistus albidus.

Methods

EcomorphologyFor the ecomorphological study the method standar-dised by Orshan (1982, 1986) was used, while wefollowed the proposal in Orshan (1989) for the phe-nological study. A selective inventory was made foreach locality, according to the presence of the mostrepresentative species of each community. The plotsmeasured 500 m2, enough to include the whole diver-

Table 1. Climatic data in the study areas. P = annual average rainfall (mm), T = annual average temperature (°C), It = Thermicity index(Rivas Martínez), Ia = Aridity index (De Martonne), ETP = Potential evapotranspiration (mm), BA = Annual balance (P-ETP), EST = rainfallseasonality (seasons from greater to lesser P; W = winter, S = spring, U = summer, A = autumn), CP = cold period, DP = dry period, Ic =continentality index (Gorezynski).

Study site P T It Ia ETP BA EST CP DP Ic

Forest Las Nieves Natural Park 962 16 398 33.9 954 +8 WSAU 1–2 3.5 28.0

Shrubland Los Montes Natural Park 788 15 281 33.1 719 +69 WASU 3–4 3.5 22.0

Figure 3. Climatic diagrams of the study areas: A = forest (LasNieves, Natural Park); B = shrubland, (Los Montes, Natural Park).C = Daylight hours in Málaga.

233

sity of species. The inventories were made followingBraun-Blanquet (1979), with the inclusion of environ-mental data and plant cover of the species, which wasdivided into persistent and non-persistent. Followingthe recommendations of Orshan (1986) we only tookinto account the persistent or arid-active species (Eve-nari et al. 1975), that is to say, those that bear aerialactive shoots throughout the year and which are there-fore adapted to the Mediterranean drought season. Ar-id-passive plants such as ephemerals (therophytes)and ephemeroids (geophytes and some hemicrypto-phytes) were not included in the studies, because theyshed their shoots during the unfavourable season(Evenari et al. 1975). The ecomorphological charac-ters (growth forms) were determined for each speciesin the field. Twenty-eight characters of those pro-

posed by Orshan (1986) were studied as well as fruittype (fleshy, dry), as proposed in Pérez Latorre et al.(1995). Afterwards a species/character data matrixwas made for each studied community and, for eachcommunity, the percentage of presence of each char-acter expression was calculated on the basis of num-ber of species showing that character. For the ecomor-phological description of the communities and thesubsequent comparison, we used the following char-acters: seasonality of assimilating organs, renewalbud location, spinescence, stem consistency, plantheight (stratification), organs periodically shed, leafconsistency, leaf tomentosity (%), average leaf size,life duration of leaves, fruit type, mean life durationof plant, vegetative regeneration after fire, main sea-son of shoot growth and flowering. An index sug-gested by Orshan, the “estimated biomass of the com-munity” (EBC), was only used to compare thecommunities. This is a non-dimensional number,based on the product of the quantitative characters,plant height, crown diameter and canopy density, foreach species of the community. [Estimated biomassof the species (EBS) = plant height (m or cm) ×crown diameter (m or cm) × canopy density (%)],[Estimated biomass of the community (EBC) = sumof EBS’s of all the species].

Phenology

Data concerning the different reproductive phenolog-ical phases (flower bud formation, flowering, fruit set-ting, seed dispersal) and vegetative phenologicalphases (vegetative growth and leaf shedding of doli-choblasts and brachyblasts) were recorded throughmonthly visits during a complete annual cycle, foreach arid-active or persistent species of every com-munity studied (Orshan 1989). Data were based on aminimum of 30 individuals of each species, while aphenomorphological herbarium (MGC) with repre-sentative samples of the phenological phases was col-lected. For each species a phenological calendar wasdrawn up (see Appendix), excluding uncommonevents (Castro Díez and Montserrat Martí 1998). Thefrequency of each phenophase taking place in eachmonth was calculated for the set of species of eachcommunity. The phenological calendar of each com-munity was constructed as a function of the season-ality of the following phenological phases: flower budformation, flowering, fruit setting, seed dispersal,vegetative growth and leaf shedding.

Table 2. A = percentage of presence of the species based on theinventories taken in the distribution area of the communities. B =species cover in the selected inventories.

A B

Q. suber forest presence % cover %

Abies pinsapo 6 1

Arbutus unedo 41 30

Calluna vulgaris 3 1

Cytisus grandiflorus 26 1

Cytisus villosus 26 1

Daphne gnidium 82 1

Erica arborea 71 5

Genista linifolia 18 1

Phyllirea angustifolia 47 5

Phyllirea latifolia 29 1

Pistacia lentiscus 32 1

Pistacia therebintus 3 1

Quercus broteroi 38 10

Quercus suber 100 75

Rhamnus alaternus 15 1

Viburnum tinus 29 50

Cistus shrubland

Cistus albidus 71 10

Cistus ladanifer 50 30

Cistus monspeliensis 79 20

Daphne gnidium 64 5

Genista umbellata 100 5

Inula viscosa 36 5

Lavandula stoechas 100 5

Phlomis purpurea 79 5

Ptilostemon hispanicus 57 5

Retama sphaerocarpa 64 1

Ulex parviflorus 100 5

234

Phenophasic indexes

To characterise and compare the communities andspecies studied, the PSI (phenophasic sequence) in-dex (Castro Díez and Montserrat Martí 1998) wasused. With the objective of improving the methodol-ogy and for comparing the results, we propose threenew phenophasic indexes.

1. “Active Phenophasic Period of the Species”(APS), defined as the number of months per yearin which each species shows activity with respectto the phenological phases that indicate favourableconditions: flower bud formation (FBF), flowering(F), fruit setting (FS) and vegetative growth ofdolichoblasts (DVG) and brachyblasts (BVG), dis-carding those which are exclusively mechanical(seed dispersal of dry fruits) due to external vec-tors (seed dispersal of fleshy fruits) or those whichcause the reduction in the body of the plant (leafshedding). This index can vary from a maximumof 12, in those species shown to be active all theyear round, to a minimum of 1, as in the case ofephemeral therophytes in arid zones.

2. “Active Phenophasic Period of the Community”(APC), which represents the number of speciesthat show activity in each month with respect tothe phenological phases that need favourable con-ditions for their development, as was explained forAPS. The variation of this index through the wholeyear permits us to compare the adaptation of thephenological strategies of the communities to theenvironment in which they develop. This is amonthly index that may vary from the total num-ber of species (when all of species are found to beactive) to zero (when none is found to be active).

3. “Index of reproductive/vegetative Activity of theSpecies” (RVA), which represents the relationshipbetween the sum of months with reproductive phe-nological phases (flower bud formation, flowering,fruit setting) and the number of months with veg-etative phenological phases (vegetative growth).This index gives an idea of the different strategiesof the plants with respect to the time and resourcesspent in the two types of phenological phases. Spe-cies with a low RVA spend a lot of time and re-sources on vegetative functions to the detriment ofreproductive functions, and conversely, a highRVA indicates a predominance of the reproductivefunctions to the detriment of the vegetative. Thisindex is tested to delimit groups of species with

similar adaptive strategies to Mediterranean envi-ronments.

Nomenclature

Valdés et al. (1987) and Castroviejo et al. (1986–2000) have been used for this work.

Results

Ecomorphology

Quercus suber forest: ecomorphologicalcharacterisationThe forest is characterised by phanerophytes (Ta-ble 3), as Floret et al. (1990) and Danin and Orshan(1990) found in France and Israel, respectively. Thedegree of stratification is well illustrated by the exist-ence of mesophanerophytes, microphanerophytes andnanophanerophytes, with the intermediate stratumshowing the greatest degree of diversity. Such a strat-ification bears close resemblance to the forest ofQuercus ilex studied in France by Romane (1987).Species with crowns of 2 to 5 metres in diameter pre-dominate, although the crowns of some species canreach 10 metres. The predominant canopy density ofthe different species is 50–90%, which indicates theexistence of a considerable biomass. The estimatedbiomass (EBC) is 18.3.

All the species of the community (Table 3) showholoxyle stems, with predominantly flaky barks andcontinuous shedding, which contributes to the accu-mulation of dead, dry matter on the soil. Two species(Quercus suber and Quercus broteroi) show corkybarks which means that they have adapted to firethrough vegetative regeneration. With the exceptionof Abies pinsapo, all the species show vegetative re-generation after fire (Cabezudo et al. 1995), the “be-low ground buds” regeneration model prevailing. Thedominant tree species in the forest (Quercus suberand Quercus broteroi) reflect the “above ground epi-cormic buds” model of regeneration, which permitsthem a quick regeneration of their canopies and lim-its the light reaching the plant strata below.

Species with a dolichoblast leaf size between 2 and20 cm2 (Table 3) predominate, although four specieshave leaves measuring between 20 and 180 cm2; therest show leaves of less than 2 cm2. The predomi-nance of large leaves (micromesophyll) is due to thehigh rainfall and to the decreased light conditions in

235

the below-trees strata (Werger and Ellenbroek 1978Givinish 1987 Keshet et al. 1990 Pérez Latorre 1993).Green leaves with horizontal insert angle are domi-nant, which means that there is a dense shadow underthe crowns. The moderate summer drought leads to aprevalence of semisclerophyll non-tomentose leaves,compared with the combination malacophyll-tomen-tose (Parsons 1976 Oppenheimer 1960 Campbell andCowling 1985 Givinish 1987). Spinescence is not afeature of the forest. Most leaves last more than 14months, a character positively correlated with an in-crease in rainfall (Keshet et al. 1990) and the Medi-terranean influence (Orshan 1982), which confers an

evergreen seasonality character to the community,though with a partial maximum shedding of leaves insummer. Other groups with an average leaf-life of 6to 14 months are generally malacophyll or winter de-ciduous. The phanerophytes/chamaephytes index(Danin and Orshan 1990) points to the total inhibi-tion of chamaephytes by the phanerophytes and theclimax optimum of the forest.

The forest is characterised by plants showing amulti-seasonal vegetative growth pattern (Table 4),which is more or less constant throughout the wholeannual cycle. Multi-season species, which grow fun-damentally in spring and summer, predominate. This

Table 3. Most representative ecomorphological characters in the studied species. RW = renewal buds position (mePh = mesophanerophyte,miPh = microphanerophyte, nPh = nanophanerophyte, Ch = chamaephyte, Am = amphiphyte), OS = organs shed rhythmically (Bb = basi-petal branch shedders, Ba = acropetal branch shedders, L/S = leaves/shoots, L = leaves), BC = bark consistency, S = 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), CO = relative cover.

Ecomorphological character RW OS BC S LS LT LC LD SO FT CO

Q. suber forest

Abies pinsapo mePh Ba corky no 0,1–0,2 no S > 62 E – 1

Arbutus unedo miPh L flaky no 56–180 no sE 14–26 E f 30

Calluna vulgaris nPh Ba flaky no < 0,1 no S 6–14 E d 1

Cytisus grandijlorus nPh Ba/L flaky no 0,2–2,2 yes Ma 6–14 D/E d 1

Cytisus villosus miPh Ba smooth no 2–12 yes Ma 6–14 E d 1

Daphne gnidium Am L/S smooth no 2–12 no Ma 6–14 E f 1

Erica arborea miPh Ba flaky no < 0,1 no S 26–38 E d 5

Genista linifolia miPh Ba flaky no 0,2–2 yes Ma 6–14 E d 1

Phyllirea angustifolia miPh L smooth no 2–12 no sE 26–38 E f 5

Phyllirea latifolia miPh L smooth no 12–20 no sE 14–26 E f 1

Pistacia lentiscus miPh L flaky no 2–12 no sE 26–38 E f 1

Pistacia therebintus miPh L flaky no 2–12 no Ma 6–14 D f 1

Quercus broteroi mePh L corky no 20–56 yes sE 6–14 D/E f 5

Quercus suber mePh L corky no 20–56 yes sE 14–26 D/E f 70

Rhamnus alaternus miPh L smooth no 12–20 no sE 14–26 E f 1

Viburnum tinus miPh L smooth no 56–180 no Ma 26–38 E f 50

Cistus shrubland

Cistus albidus nPh Ba flaky no 12–20 yes Ma < 6 E d 10

Cistus ladanifer nPh Ba papery no 12–20 yes Ma 6–14 E d 30

Cistus monspeliensis nPh Bb smooth no 2–12 yes Ma < 6 E d 20

Daphne gnidium Am L/S smooth no 2–12 no Ma < 6 E f 5

Genista umbellata Ch Ba papery no < 2 yes Ma < 6 E d 5

Inula viscosa Ch Bb smooth no 2–12 yes Ma < 6 E d 5

Lavandula stoechas Ch Bb flaky no < 2 yes Ma 6–14 E d 5

Phlomis purpurea Ch Bb flaky no 20–56 yes Ma < 6 E d 5

Ptilostemon hispanicus Ch Bb flaky leaves 20–56 no sE 14–26 E d 5

Retama sphaerocarpa nPh Ba smooth no – – – – E d 1

Ulex parviflorus nPh Ba flaky stems – – – – E d 5

236

multi-seasonality and the high percentage of specieswith summer growth (dry season) indicate optimumconditions for vegetal development in the study area.Though spring flowering, a characteristic of the Med-iterranean (Orshan 1989), is predominant, there aresome interesting exceptions which flower in autumn(Arbutus unedo, Daphne gnidium and Calluna vul-garis), which perhaps represent a set of plantsadapted to different conditions from those which pre-vail today (Herrera 1984).

The forest (Table 4) is a phanerophytic, non-spinescent, holoxyle, multi-stratum evergreen com-munity, with a maximum height of 23 metres. Leavesand branches are shed periodically; the leaves aresemisclerophyll, with a 33% degree of tomentosity,and micromesophyll (28 cm2) that are shed on aver-age every 19 months. The average life of the plants is40 years; the predominant regeneration after fire pat-tern is by below ground epicormic buds; vegetativegrowth is bi-multiseasonal, maximum flowering is inspring and fleshy fruits are predominant.

Cistus shrubland: ecomorphologicalcharacterisation

The shrubland is characterised (Table 3) by the pre-dominance of chamaephytes and nanophanerophytes,and an average plant height of 1–2 m. The commu-nity shows only one stratum and great homogeneityas regard stem consistency, since all the species areholoxyle. The estimated biomass (EBC) is 0.4.

The shrubland is a pyrophytic community (Ta-ble 3), which is mostly regenerated through seeds,with a scarce representation of vegetative regenera-tion, typical of the forest (Cabezudo et al. 1995). Theflaky and papery barks and their periodic sheddingmay play an important role in adaptation to fire dueto the accumulation of dead, dry matter which igniteseasily. Furthermore, the absence in the shrubland ofcorky barks that protect the body of the plant duringfires (Cabezudo et al. 1995) forces many species intoafter-fire regeneration through seeds.

With respect to renewal bud location (Table 3), thephanerophyte:chamaephyte ratio is low (0.66) and ex-

Table 4. Comparison between forest and shrubland concerning the most representative ecomorphological characters and phenological phases.

Ecomorphological characters Shrubland Forest

Renewal buds position chamaephytic-nanophanerophytic micro-mesophanerophytic

Spinescence scarce absent

Stratification mono-stratum multi-strata

Maximum height 2–5 m 20–30 m

Organs shed branches and barks leaves and branches

Leaf consistency malacophyll predominant sclerophyll predominant

Tomentosity (%) 78% 33%

Leaf size microphyll (average 14 cm2) micromesophyll (average 28 cm2)

Life duration leaves average 6.5 months average 19 months

Life duration plants average 16 years average 40 years

After fire vegetative regeneration absent present

Main season of shoot growth multi-seasonal bi-multiseasonal

Main Flowering season spring-summer spring

Fruit type predominant dry predominant fleshy

Biomass estimated 0.4 18.3

Phenological phases Shrubland Forest

Flower bud formation spring-summer winter-spring

Flowering spring-summer spring

Fruit setting summer summer-autumn

Seed dispersal summer-autumn autumn-winter

Dolichoblast vegetative growth spring-summer spring-summer

Leaf shedding dolichoblast summer summer

Brachyblast vegetative growth throughout the year absent

Leaf shedding brachyblast throughout the year absent

237

presses the adaptation strategy to the summer drought(Danin and Orshan 1990). The shrubland and the“maquis” studied by Orshan et al. (1984) in Israelshow a similar percentage of phanerophytes andchamaephytes. The periodical shedding of branchesalso constitutes an adaptation to the unfavourablesummer season, when the above species reduce theirbody size to avoid transpiration. The canopy densityis low, probably as a response to the scarcity of water(reduction of leaf area) and to the excess of light. Thealmost complete absence of spinescence might indi-cate that other types of adaptation to drought andgrazing pressure may be common in this community.The predominance of microphyll leaves indicates anadaptation to water stress due to the low rainfall andscarce retention of water in the soil (Werger and El-lenbroek 1978 Givinish 1987) or an adaptation to thehigh luminosity resulting from a low degree of veg-etation cover (Keshet et al. 1990). The presence ofphotosynthetic stems, the above-mentioned characterand the absence of leaves in some species also pointsto the adaptation to drought. The predominance of aglaucous leaf colour is due to the tomentosity and tothe presence of waxes and resins in the leaves, an ad-aptation to solar radiation and drought, as mentionedby Keshet et al. (1990) in Israel. The high degree oftomentosity, is reflected in the percentage of tomen-tose:total species (78%) and gives a good idea of thedegree of adaptation to minimise the loss of water.The dominance of malacophyll species, with unligni-fied leaves which last a year or less, falling at leastpartially in summer, also constitutes an adaptation tosummer water stress. The shrubland species shedsdolichoblast leaves on an annual or six monthly ba-sis. This fact may constitute an adaptation to the sum-mer drought and to the low annual rainfall (Keshet etal. 1990). The phanerophyte:chamaephyte ratio (Da-nin and Orshan 1990) is 5/5, which indicates this ad-aptation to the summer drought (abundance ofchamaephytes). The presence of species with a leafsurface covered by resins is related to other charac-ters associated with adaptation to summer droughtthrough decreased transpiration. This character alsoconstitutes a defence mechanism against grazing byimparting a bad taste and decreasing digestibility, fur-ther it inhibits seed germination (Herrera 1984) andcontributes to a quick spread of fires in the commu-nity.

The community is characterised (Table 4) by theshort average lifespan of its plants compared withthose of other Mediterranean ecosystems such as

Quercus suber forests (Pérez Latorre et al. 1995). Thecommunity is evergreen, one of the main characteris-tics of Mediterranean vegetation. This multi-season-ality may be indicative of the fact that the ecologicalparameters are mild enough to permit vegetativegrowth during more than one season. Although sum-mer is, ecologically, the most unfavourable season,the bi-seasonality of flowering (spring-summer) couldreflect a slight moderation of the climatic conditions,at least at the beginning of summer.

The shrubland (Table 4) is a chamaephytic-nanophanerophytic, scarcely spinescent, holoxyle,monostratum and evergreen community, with a maxi-mum plant height of 2.5 metres. Branches and barksare shed periodically; there is predominance of mala-cophyll leaves, with 78% tomentosity accompaniedby microphyll leaves (average of 14 cm2) that areshed on average after 6 and a half months. The aver-age life of plants is 16 years; the absence of after firevegetative regeneration is a predominant characteris-tic; vegetative growth is multi-seasonal, maximumflowering occurs in spring-summer and dry fruits arepredominant.

Phenological phases

Quercus suber forest: phenophasic characterisationFlower bud formation shows two peaks in winter andspring (Figure 4; Appendix). May shows the smallestnumber of species in this phenophase while the great-est number of species begin this phase in January.Viburnum tinus and Abies pinsapo show the longestperiod of flower bud formation (8 and 7 months re-spectively), while the deciduous species (Quercusfaginea, Quercus suber and Pistacia terebinthus)show the shortest period (1 month). Calluna vulgarisand Arbutus unedo show flowering buds at the begin-ning of summer and flower in autumn. Maximumflowering occurs in spring. The most unfavourablemonth for flowering initiation is July, and the mostfavourable are February and March. Arbutus unedo,Calluna vulgaris and Daphne gnidium flower in au-tumn and Cytisus villosus in winter. Phyllirea latifo-lia flowers unevenly and fruit setting (affected nor-mally by insect bites) is sporadic and scarce. Quercussuber shows very uneven flowering and fruit settingwithin the populations, which could indicate wide ge-netic variability in the study site. Fruit setting is at itsmaximum period in summer – autumn, and beginslargely in April and May. Arbutus unedo has a ripen-ing period lasting one year while the fruits of Vibur-

238

num tinus, Phyllirea latifolia and Quercus suber ripenin slightly more than six months (fleshy fruit or seedswith fleshy cotyledons). Cytisus villosus, Cytisusgrandiflorus, Daphne gnidium and Erica arboreahave the shortest ripening period (dry fruits). Seeddispersal is concentrated in autumn and winter. Thisphenophase, which is absent during the months ofApril and May, begins in October – November. Ericaarborea, Rhamnus alaternus and Cytisus villosus dis-perse in summer. Viburnum tinus and Pistacia lentis-cus show the longest period of seed dispersal (6–7months). Abies pinsapo needs a whole year to com-plete its reproductive phenological cycle, since it setsits flowering buds at the beginning of autumn, flow-ers in spring, shows vegetative growth in summer,while seed dispersal occurs in autumn.

There is a partial shedding of dolichoblast leavesduring the summer (Figure 5; Appendix), probably asan adaptation to water stress. The months of Septem-ber and October see very few species undergoing thisphenophase, which starts in April and May. Callunavulgaris sheds leaves all year round. Cytisus villosusand Pistacia lentiscus do not have any particular pe-riod for leaf shedding. Pistacia terebinthus loses allits leaves in winter while Quercus broteroi and Quer-cus suber, though shedding leaves primarily in winterand spring, respectively, never lose them totally.Quercus broteroi does not completely lose its leavesduring the winter, so that old and new leaves appeartogether in spring, giving the impression that it is anevergreen species, a behaviour that can be explainedby the mild climatic conditions of the study area. Itseems that the case of Quercus suber is special,among all the Quercus of the Iberian Peninsula (Ruizde la Torre 1971), since it behaves as a deciduousspecies in spring, with a very high percentage of leaf

shedding, though, as in the case of Quercus broteroi,it does not lose all its old leaves and the appearanceof new leaves gives the impression of an evergreenspecies. Vegetative growth occurs largely in spring –summer, although February and March are the mostcommon months for it to begin. Calluna vulgaris andGenista linifolia show dolichoblast vegetative growthall year round, while Erica arborea and Pistacia len-tiscus do so for 9 and 8 months, respectively. Cytisusgrandiflorus and Cytisus villosus show typical wintergrowth.

The forest presents the following phenologicalcharacteristics (Figures 3, 4 and 5; Appendix): flowerbud formation around the end of winter and the be-ginning of spring, coinciding with the increased tem-peratures; flowering in spring, a temperate and rainyseason with longer daylight hours (Figure 3); fruitsetting in summer, coinciding with the maximumtemperatures and minimum rainfall; seed dispersal inautumn, a temperate and rainy season with ever-de-creasing daylight hours (Figure 3); dolichoblast veg-etative growth in spring, coinciding with an increasein temperatures and considerable rainfall; dolicho-blast leaf shedding in summer, coinciding with maxi-mum temperatures and minimum rainfall.

Cistus shrubland: phenophasic characterisation

Flower bud formation shows a maximum in springand the beginning of summer (Figure 6; Appendix),while the minimum occurs during autumn. The long-est period of flower bud formation is shown byDaphne gnidium (5 months) and Cistus ladanifer (4months), while Retama sphaerocarpa only shows

Figure 4. Time curse of the reproductive phenological phases inthe forest expressed as the monthly percentage of species that showeach phenophase. Flower bud formation (FBF), flowering (F), fruitsetting (FS) and seed dispersal (SD).

Figure 5. Time curse of the vegetative phenological phases in theforest expressed as the monthly percentage of species that showeach phenophase. Dolichoblast vegetative growth (DVG) and leafshedding dolichoblast (LSD).

239

flower buds during 1 month. Ulex parviflorus beginsflower bud formation in autumn. Flowering is at amaximum in spring and the beginning of summer.The minimum occurs in autumn – winter with onlyone species flowering. This pheno-phase usuallystarts in April and June, although Daphne gnidiumflowers in autumn and Ulex parviflorus in winter andspring (for 8 months) and though the winter flowersdo not produce fruits. Fruit setting is concentrated insummer but may last into the beginning of autumn,reducing until it disappears in winter and a large partof the spring. May and June are basically the monthswhen fruit setting starts. Most of the species show ashort ripening period (dry fruits), although the fruitsmay be retained without dispersal for up to 4 months,except in the case of Genista umbellata, which onlyretains its fruits for 2 months. The fruit setting ofDaphne gnidium occurs in autumn – beginning ofwinter. Seed dispersal occurs mainly at the end ofsummer and in autumn, after which it falls drasticallyto zero at the end of winter and spring. Seed dispersalbegins in July and August. The shortest period of seeddispersal occurs in Genista umbellata, that is 2months, while the most common is 4–5 months.

Dolichoblast vegetative growth occurs in spring –summer (Figure 7), with March and April being themost favourable months for growth to begin. Certainspecies show long growth periods such as Cistus al-bidus (9 months) and Lavandula stoechas (11months), although others such as Retama sphaero-carpa and Genista umbellata have a much shorterperiod (3 months). Dolichoblast leaf shedding occursbasically in summer, probably as an adaptation to wa-ter stress, since the transpirable surface of the plant isthus reduced. The beginning of this phenophase oc-curs in June and July and is practically absent in win-

ter and spring. Phlomis purpurea and Lavandula sto-echas only shed dolichoblast leaves during 2 monthsof the summer, while Ptilostemon hispanicus does soin a more gradual manner during the 6 months ofsummer and autumn. Daphne gnidium also shedsleaves during the autumn. Brachyblast vegetativegrowth is maintained practically throughout the year,although at a lower rate in summer. Brachyblast leafshedding also occurs throughout the year. The simul-taneous shedding of this type of leaf was not detectedin Inula viscosa, probably because all leaves trans-formed into dolichoblast leaves, which are subse-quently shed.

The shrubland presents the following phenologicalcharacteristics (Figures 3, 6 and 7; Appendix): flowerbud formation in spring and summer, coinciding withan increase in temperatures and longer hours of day-light; flowering in spring and the beginning of sum-mer, a temperate and rainy period with increased day-light hours (Figure 3); fruit setting in summer,coinciding with maximum temperatures and lighthours and minimum rainfall; seed dispersal at the endof summer and in autumn, as rainfall increases andtemperatures and light hours decrease (Figure 3);dolichoblast vegetative growth in spring – summer,coinciding with an increase in temperatures and a de-crease in rainfall; dolichoblast leaf shedding in sum-mer, a season with maximum temperatures and min-imal rainfall; brachyblast vegetative growth all yearround with a maximum in winter, the coldest andwettest season; brachyblast leaf shedding all yearround.

Figure 6. Time curse of the reproductive phenological phases inthe shrubland expressed as the monthly percentage of species thatshow each phenophase. Flower bud formation (FBF), flowering(F), fruit setting (FS) and seed dispersal (SD).

Figure 7. Time curse of the vegetative phenological phases in theshrubland expressed as the monthly percentage of species thatshow each phenophase. Dolichoblast vegetative growth (DVG),leaf shedding dolichoblast (LSD), brachyblast vegetative growth(BVG) and leaf shedding brachyblast (LSB).

240

Phenophasic indexes

Q. suber forestPhenophase Sequence Index (PSI) (Table 5a). Apply-ing the interval proposed by Castro Díez and Mont-serrat Martí (1998), only two groups can be clearlyidentified from the total number of species, one con-sisting of deciduous trees and shrubs (Quercus suber,Quercus broteroi and Pistacia terebinthus) (PSI <0.5) and the other composed of the rest of the species(PSI > 0.6). Cytisus villosus and Calluna vulgaris fallwithin the PSI interval that separates the two groups(0.5 < PSI < 0.6). The PSI separates the specieswhose phenological phases overlap due to their de-ciduous condition.

Active Phenophasic Period of the Species (APS)(see Appendix). This index points to the existence ofdifferent groups of species within the forest. Onegroup (Abies pinsapo, Arbutus unedo, Calluna vul-garis, Erica arborea, G. linifolia and Viburnum tinus)shows activity throughout the annual cycle with someof the species (Arbutus unedo and Calluna vulgaris)flowering in autumn. Another group shows activityduring fewer months (7 to 9) (Cytisus grandiflorus,Cytisus villosus, Pistacia terebinthus, Quercus brot-eroi and Quercus suber). These constitute a winter-deciduous group, although some species are summer-deciduous, and some of them have photosyntheticstems. An intermediate group (the rest of the species)shows a small decrease in phenophasic activity last-ing 1–2 months, the decrease in activity normally oc-curring in winter, when temperatures are lower.

Active Phenophasic Period of the Community(APC) (Figure 8). This index shows a minimum inNovember and February (with a small increase in De-cember and January), while the maximum occurs dur-ing spring and summer (from April to July). This dis-tribution indicates that winter is the season with theleast favourable environmental conditions for generalactivity and biomass creation, while spring and thebeginning of summer are the most favourable periods.In the community as a whole, the APC remains above75% of the species throughout the year, which can betaken as a characteristic of this type of Mediterraneanforest. A study of the APC in other types of commu-nity more restricted by prevailing environmental con-ditions (high mountain, arid zones), would probablyresult in much lower APC values.

Index of reproductive/vegetative Activity of theSpecies (RVA) (TABLE 5b). One group of speciesshows values of RVA < 1, representing a predomi-

nance of vegetative phenophases to the detriment ofreproductive. Genista linifolia and Erica arboreashow the lowest values. Another group shows RVA >1, representing a predominance of reproductive phe-nophases to the detriment of vegetative. Quercusbroteroi and Abies pinsapo show the highest values.

Cistus shrubland

Phenophase Sequence Index (PSI) (Table 5a). The in-terval proposed by Castro Díez and Montserrat Martí(1998) show all the species to be grouped around PSI> 0.6, the only exception being Inula viscosa with aPSI of 0.43.

Active Phenophasic Period of the Species (APS)(see Appendix). The most common values of this in-dex are 11–12 months, although there is a group ofspecies adapted to an inactivity period in autumn –winter (Genista umbellata and Retama sphaerocar-pa), which show values of 5 and 7 months, respec-tively. However, the absence of functional leaves inthese plants should be emphasized and it is their sim-ple anatomical – functional shoot structure whichgives them a low APS. Daphne gnidium with an APSof 9 months (end of winter and beginning of springwith no activity) and Inula viscosa with an APS of10 (no activity at the beginning of spring or mid au-tumn) show another type of adaptation, which is sea-sonal. The predominance of an 11–12 month APS inthe species may indicate sufficiently mild environ-mental conditions for the development of vegetationpractically all year round, except in winter, which af-fects some species.

Active Phenophasic Period of the Community(APC) (Figure 8). This index clearly shows a mini-mum during winter and autumn, while the maximumoccurs during spring and summer (from April toJuly). Despite the two unfavourable seasons, the APCis maintained above 75% of the species throughoutthe year so that the same remarks as those made forthe forest apply.

Index of reproductive/vegetative Activity of thespecies (RVA) (Table 5b). One group of speciesshows values of RVA < 1, representing a predomi-nance of vegetative phenophases to the detriment ofreproductive. Phlomis purpurea, Cistus albidus andPtilostemon hispanicus show the lowest values. An-other group shows RVA > 1, representing a predomi-nance of reproductive phenophases to the detrimentof vegetative. Ulex parviflorus, Daphne gnidium andRetama sphaerocarpa show the highest values.

241

Discussion

Ecomorphological comparison between Q. suberforest and Cistus shrubland

Table 4 compares the studied communities using themost important growth forms and the phenologicalphases. Substitution of the forest by shrubland impliesnot only floristic and phytosociological changes butalso, as is observed in the table, changes in the mor-phological and functional standards of the vegetation.The predominance of high phanerophytes up to 20 mand the multi-layer stratification that characterises theforest contrasts with the predominance of the chamae-phytes and small phanerophytes of 1–2 m, and theuni-layer stratification in the shrubland. The biomassthat constitutes the forest is supported by a greaterquantity of available resources (preserved and deepsoil) and the relative independence from general at-mospheric conditions unlike in the case of shrubland.The rhythmic production of dead matter, such asbranches, is common to both communities. However,the forest periodically renews its leaves, while theshrubland sheds leaves, in a way that is dependent onthe severity of the summer drought but, above all,shedding of bark. With respect to leaf characters, theshrubland is characterised by tomentosity (an adapta-tion to the water stress already mentioned), whilesuch a feature is scarce in the forest, (although it doesshow this character, being a Mediterranean commu-nity). The average leaf size reflects the light and xe-ric environmental conditions in the shrubland (micro-phyll adaptation) and milder conditions in the forest(micromesophyll adaptation). Finally, the life span ofphotosynthetic leaves is much greater in the forestthan in the shrubland, where leaves are renewed rela-

Table 5a. Studied species grouped according to the values of theirphenophasic indexes PSI (5a), and RVA (5b). Index values ofDaphne gnidium, f = in the forest, s = in the shrubland.

Species PSIPistacia lentiscus 0,91Abies pinsapo 0,9Phillyrea latifolia 0,83Cistus ladanifer 0,81Retama sphaerocarpa 0,8Rhamnus alaternus 0,79Phillyrea angustifolia 0,78Arbutus unedo 0,76Viburnum tinus 0,75Phlomis purpurea 0,71Erica arborea 0,7Genista linifolia 0,7Daphne gnidium (f) 0,69Ulex parviflorus 0,66Cistus monspeliensis 0,66Cistus albidus 0,64Lavandula stoechas 0,63Genista umbellata 0,63Daphne gnidium (s) 0,61Ptilostemon hispanicus 0,6Cytisus grandiflorus 0,6Calluna vulgaris 0,52Cytisus villosus 0,5Pistacia terebinthus 0,5Inula viscosa 0,43Quercus suber 0,43Quercus broteroi 0,33

Table 5b. Species RVA

Phlomis purpurea 0,4Ptilostemon hispanicus 0,5Cistus albidus 0,5Genista linifolia 0,5Erica arborea 0,5Lavandula stoechas 0,6Inula viscosa 0,7Cistus ladanifer 0,7Calluna vulgaris 0,8Cytisus villosus 0,9Rhamnus alaternus 0,9Cytisus grandiflorus 1,2Genista umbellata 1,3Retama sphaerocarpa 1,3Daphne gnidium (f) 1,4Daphne gnidium (s) 1,8Arbutus unedo 2Ulex parviflorus 2,2Viburnum tinus 2,4Quercus suber 3Phillyrea angustifolia 3,3Pistacia lentiscus 4Phillyrea latifolia 5,5Abies pinsapo 6Quercus broteroi 7

Figure 8. Time curse of the APC index in both studied communi-ties.

242

tively quickly. The idea that the forest represents aclimax community is supported by the duration ofplants, which is almost three times that observed inthe shrubland, which grows more rapidly and showsa colonising ability on many occasions. The fact thatthe forest biomass (18.3) is greater than that of theshrubland (0.4) also lends weight to this idea.

Phenological comparison of Q. suber forest andCistus shrubland

The phenological phases (Table 4) are clearly concen-trated in spring/summer in the case of the shrubland,its phenophasic activity being confined to the climati-cally most favourable months (abundant rainfall andmild temperatures). On the other hand, the forestshows a more rhythmic behaviour, each phenophasebeing set in a specific season, beginning in winterwith flower bud formation, continuing in spring withflowering, in autumn with fruit setting and ending inwinter with seed dispersal. The presence of brachy-blasts in the shrubland causes this community to showall year round vegetative activity. Flower bud forma-tion is earlier in the forest than in the shrubland,which has a slightly longer flowering period. How-ever, the other reproductive phenological phases (fruitsetting and seed dispersal) are longer and later in theforest due to the predominance of fleshy fruits. Thischaracter (type of fruit) separates the forest, with itsfleshy fruits and seeds with rich cotyledons (Quercus)and zoochory from the shrubland, with dry fruits andmechanical seed dispersal.

The PSI does not discern between functionalgroups of species in the two studied communities (Ta-ble 5a). Ecomorphologically different species (mala-cophyll chamaephytes, aphyll nanophanerophyteswith dry fruits and sclerophyll microphanerophyteswith fleshy fruits) are grouped in the same interval(PSI > 0,6). It is notable that both communities havea similar APC index value (Figure 8) despite the eco-morphological and phenophasic differences alreadydescribed. That is to say, two quite different plantgroups have adapted in a similar manner to varyingenvironmental conditions during their annual cycle.The Index of reproductive/vegetative Activity of theSpecies (RVA) (Table 5b) clearly separates the ele-ments of the shrubland and of the forest. Taking theintermediate score of Daphne gnidium (1.4 in the for-est and 1.8 in the shrubland) as a yardstick, the spe-cies with a lower RVA value largely belong to theshrubland, while those which have a higher RVA

largely belong to the forest. The exceptions for theshrubland group are Erica arborea (capable of devel-oping in Cistus shrublands and heathlands), Genistalinifolia and Cytisus villosus (coloniser species in for-est clearings) and Calluna vulgaris (a heathland spe-cies). Rhamnus alaternus, in theory, should belong tothe forest group, but shows a special intermediatefunctional position, as observed, too, by Aronne andWilcock (1997). In the forest group the only excep-tion is Ulex parviflorus, which basically belongs be-cause of its long flowering period.

Table 6 shows the grouping of the species of bothplant communities according to their phenophasic be-haviour and selected ecomorphological characters (re-newal buds position, type of fruit, seasonality, pres-ence/absence of leaves). The most striking feature isthe richness of growth form combinations and varietyof phenophasic behaviour. There is no evidence ofone single adaptive type common to each plant com-munity studied, but rather that within each commu-nity groups of species exist with similar adaptations.In the forest up to 8 groups can be identified and inthe shrubland 5 such groups. In the forest Cytisus vil-losus and Phyllirea latifolia are the only species thatbegin to flower in winter; Abies pinsapo shows itsown peculiarities because it is a gymnosperm anddoes not bear fruit; Cytisus grandiflorus, Erica ar-borea and Genista linifolia flower in spring, dispersetheir seed in early summer and have dry fruits, whiletwo of them show all-year-round growth; Phillyreaangustifolia, Rhamnus alaternus and Viburnum tinusbegin to flower in spring, generally show seed dis-persal in autumn and have fleshy fruits; Pistacia tere-binthus and Quercus broteroi are the only winter-de-ciduous species; Quercus suber is amesophanerophyte, spring-deciduous, and also hasvegetative growth in spring; Daphne gnidium flowerstowards the autumn, and is the only amphiphyte sensuOrshan; Arbutus unedo and Calluna vulgaris (bothEricaceae) flower in autumn – winter, while the lat-ter is the only species with spring seed dispersal. Inthe shrubland Cistus albidus, Cistus ladanifer andCistus monspeliensis flower only in spring and arenanophanerophytes; Lavandula stoechas and Phlomispurpurea flower between spring and summer and arechamaephytes; Genista umbellata, Ptilostemon his-panicus and Retama sphaerocarpa flower in summerand show xeromorphological adaptations in the formof aphyll or spinescent stems; Daphne gnidium andInula viscosa flower in summer – autumn; Ulex parvi-

243

florus shows flowers during three seasons, and is theonly aphyll and spinescent nanophanerophyte.

General discussion

Differences between the two communities studiedwith respect to physiognomy, floristic compositionand phytosociology were already known. To thisknowledge we can now add the findings of this work,

Table 6. Grouping of species according to similarity of phenology (FBF: flower bud formation, F: flowering, FS: fruit setting, SD: seeddispersal, DVG: dolichoblast vegetative growth and LSD: leaf shedding dolichoblast) and selected ecomorphological characters (renewalbuds position, type of fruit, seasonality, spinescence and presence of leaves). Bold letters indicate common or unique differential charactersof the groups. Seasons: W = winter, S = spring, U = summer and A = autumn.

Species groups

Forest FBF F FS SD DVG LSD

Cytisus villosus WS WS SU UA WSU – Microphanerophyte with dry fruits

Phillyrea latifolia W WS SUA AW SU SU Microphanerophyte with fleshy fruits

Abies pinsapo W S SUA AW SU U Mesophanerophyte with conesCytisus grandiflorus WS S SU UA WS U Microphanerophyte with dry fruits

Erica arborea WS S SU UA YEAR S Microphanerophyte with dry fruits

Genista linifolia WS S SU UAW YEAR U Microphanerophyte with dry fruits

Phillyrea angustifolia WS S SUA A SU U Microphanerophyte with fleshy fruitRhamnus alaternus WS S SU U SUA U Microphanerophyte with fleshy fruitViburnum tinus YEAR S SUA AW SU U Microphanerophyte with fleshy fruit

Pistacia terebinthus S S SUA A SU AW Microphanerophyte winter-deciduousQuercus broteroi S S SUA AW SU AWS Mesophanerophyte winter-deciduous

Quercus suber SU SU UAW AW SU SU Mesophanerophyte spring-deciduous

Daphne gnidium (forest) UA UA UAW AW SU WSU Amphiphyte with fleshy fruit

Arbutus unedo UA AW YEAR AW SU SU Microphanerophyte with fleshy fruit

Calluna vulgaris UAW AW AWS WS YEAR YEAR Nanophanerophyte with dry fruit

Shrubland FBF F FS SD DVG LSD

Cistus albidus S S SU UAW WSU U Nanophanerophyte with dry fruit

Cistus ladanifer WS S SU SUA SUA U Nanophanerophyte with dry fruit

Cistus monspeliensis S S SU UA SUA SU Nanophanerophyte with dry fruit

Lavandula stoechas WS SU U AW AWS U Chamaephyte with dry fruit

Phlomis purpurea S SU SU U AWS U Chamaephyte with dry fruit

Genista umbellata SU U U U S – Chamaephyte aphyll with dry fruit

Ptilostemon hispanicus SU U UA UA SU UAW Chamaephyte spinescent with dry fruit

Retama sphaerocarpa U U UA UAW S – Nanophanerophyte aphyll with dry fruit

Daphne gnidium UA UA AW AW SU SU Amphiphyte with fleshy fruit

Inula viscosa SU UA UA UA SUA UA Chamaephyte with dry fruit

Ulex parviflorus AWS AWS U U SUA – Nanophanerophyte aphyll spinescent

244

which concentrates on the adaptation of the two com-munities to the environment where they develop.

Based on ecomorphological characters, the Quer-cus suber forest and the Cistus shrubland remain de-fined and separated by characters related to the loca-tion of renewal buds, adaptation to fire, drought orlight. The greatest differences observed refer to bio-logical types, vegetation stratification, biomass esti-mation, renovation of vegetative organs, leaf charac-ters and average lifespan of plants.

Phenological phases, too, define and differentiateboth communities. The forest is characterised by agroup of species that flower in autumn and which,despite their evergreen nature, lose an important partof their leaf cover in summer, while vegetativegrowth occurs during a large part of the year. Thephenophasic indexes show that the forest is activethroughout the year, winter only affects some species;spring and the beginning of the summer are the mostfavourable periods. The shrubland is characterised byalmost universal spring flowering. Vegetative growthtakes place in spring – summer; there is a drastic re-duction in dolichoblast leaf cover in summer; brachy-blast leaves are shed throughout the year. The phe-nophasic indexes point to year round phenophasicactivity and to the existence of species that discrimi-nate between forest and shrubland (very low APC).The winter is the least favourable season for somespecies of the community, while spring and summerare the periods of maximum activity. The most out-standing phenophasic differences between the twocommunities can be explained by the fact that theshrubland concentrates practically all its phenophasicactivity to spring and summer, while the forest showsactivity all year round. This long activity period indi-cates a certain degree of independence from the cli-mate, since the forest develops in deeper soil with agreater capacity to conserve water and a microclimatethat is milder inside the forest than outside. Fruit typealso separates the communities, fleshy fruits prevail-ing in the forest and dry fruits in the shrubland; thisalso influences the duration of the phenologicalphases of fruit setting and seed dispersal, which arecorrespondingly longer in the forest. The phenologi-cal phases coincide with seasonal changes in the cli-matic conditions, typical of the Mediterranean, inboth shrubland and forest. A comparison of the APCindex in both communities shows that, in spite of thelarge floristic, ecomorphological and phenologicaldifferences, both communities behave in a very simi-lar manner with respect to their active phenological

phases, during the annual cycle. The proposed “repro-ductive/vegetative activity of the species” index sep-arates the species into those considered as character-istic of substitution stages (shrubs) and thosecharacteristic of climax communities (forests).

By combining the ecomorphological charactersand the phenological phases, it is possible to groupspecies according to their adaptive strategies. Thegreat variety of ecomorphological and phenophasicbehaviours detected within a plant community as aresponse to given environmental conditions means anincrease in the already high Mediterranean diversity(which takes into account the number of species aswell as communities or types of landscape). The dif-ferences between climax plant communities and sub-stitution stages (in our case forests and shrublands)are clearly defined by a comparison between thegrowth – forms and the phenological phases of bothcommunities, which also provides basic informationon their adaptation to the environments where theydevelop. The value of using growth forms and phe-nological phases to characterise and catalogue theMediterranean plant communities is made clear inthis work, even in communities linked by successionand that develop in the same phytogeographical area.Also such studies may be of great use when under-taking restoration of natural areas and in fauna-relatedstudies, especially as regards possible pasturage (bio-mass production) and the relationship between ani-mals and flowering (pollination) and fruit setting(feed).

A study of Mediterranean vegetation based ongrowth forms and phenological phases can, therefore,make a great contribution to existing classifications(ecological, physiognomic, phytosociological, etc.). Itmay be considered a suitable way of cataloguing thevegetation of the Mediterranean Basin, as Orshan(1982) proposed and as Floret et al. (1990), amongothers, began to do. These authors also indicated theneed to deepen our knowledge of this type of study.

Appendix

245

246

247

References

Aronne G. and Wilcock C. 1997. Reproductive phenology in med-iterranean macchia vegetation. Lagascalia 19: 445–454.

Braun-Blanquet J. 1979. Fitosociología. Bases para el estudio delas communidades vegetales. Ed. Blume, Madrid.

Box E.O. 1987. Plant life forms and Mediterranean environments.Annali di Botanica 45: 7–42.

Box E.O. 1996. Plant functional types and climate at the globalscale. Journal of Vegetation Science 7: 309–320.

Cabezudo B., Navarro T., Pérez Latorre A.V., Nieto Caldera J.M.and Orshan G. 1992. Estudios fenomorfológicos en la veget-ación del sur de España. I. Cistus. Acta Botanica Malacitana17: 229–237.

Cabezudo B., Pérez Latorre A.V., Navarro T. and Nieto CalderaJ.M. 1993. Estudios fenomorfológicos en la vegetación del surde España. II. Alcornocales mesomediterráneos (Montes deMálaga, Málaga). Acta Botanica Malacitana 18: 179–188.

Cabezudo B., Pérez Latorre A.V. and Nieto Caldera J.M. 1995. Re-generación de un alcornocal incendiado en el sur de España(Istán, Málaga). Acta Botanica Malacitana 20: 143–151.

Campbell B.M. and Cowling R.M. 1985. Relationships betweenvegetation structure and the environment in the Cape, SouthernAfrica: a review. Annali di Botanica 43: 245–255.

Caritat A., Pérez Latorre A.V. and Romo A. 1997. The distributionof ecomorphological types as related to altitude in the Mont-seny mountain range (NE Spain). A preliminary study. ScientiaGerundensis 23: 63–71.

Castro Díez P. and Montserrat Martí G. 1998. Phenological patternof fifteen Mediterranean phanaerophytes from Quercus ilexcommunities of NE Spain. Plant Ecology 139: 103–112.

Castroviejo S. et al. 1986–2000. Flora Iberica: plantas vascularesde la Península Ibérica e Islas Baleares. CSIC, Madrid.

Chapin F.S., Bret-Harte M.S., Hobbie S.E. and Zong H. 1996. Plantfunctional types as predictors of transient responses of arcticvegetation to global change. Journal of Vegetation Science 7:347–358.

Danin A. and Orshan G. 1990. The distribution of Raunkiaer lifeforms in Israel in relation to the environment. Journal of Veg-etation Science 1: 41–48.

Evenari M.E., Schulze D., Kappen L., Buschbom U. and LangeO.L. 1975. Adaptive mechanisms in desert plants. In: VernbergE.J. (ed.), Physiological adaptation to the environment. Ameri-can Institute of Biological Sciences, New York, pp. 111–129.

Floret C., Galan M.J., Le Floc’h E., Orshan G. and Romane F.1987. Growth form as a tool in characterising vegetation ofsmall areas. A trial in a Mediterranean environment. Vegetatio71: 3–11.

Floret C., Galan M.J., Le Floc’h E., Orshan G. and Romane F.1990. Growth forms and phenomorphology traits along an en-vironmental gradient: tools for studying vegetation? Journal ofVegetation Science 1: 71–80.

Givinish T.J. 1987. Comparative studies of leaf form: assessing therelative role of selective pressures and phylogenetic constraints.New Phytologists (Suppl.) 106: 131–160.

Herrera C. 1984. Patrones morfológicos y funcionales en plantasdel matorral mediterráneo del sur de España. Studia Oecologica5: 7–34.

Fig

ure

9.Ph

enol

ogic

alca

lend

arof

the

stud

ied

spec

ies.

FBF:

flow

erbu

dfo

rmat

ion,

F:flo

wer

ing,

FS:

frui

tse

tting

,SD

:se

eddi

sper

sal.

DV

G:

dolic

hobl

ast

vege

tativ

egr

owth

,L

SD:

leaf

shed

ding

,do

licob

last

,B

VG

:br

achy

blas

tve

geta

tive

grow

th,

LSB

:le

afsh

eddi

ngbr

achy

blas

t.A

PS:

Inde

xof

Act

ive

Peri

odof

the

Spec

ies.

Dar

kgr

ey:

repr

oduc

tive

phen

olog

ical

phas

es.

Cle

argr

ey:

vege

tativ

eph

enol

ogic

alph

ases

.

248

Herrera J. 1987. Flower and fruit biology in southern Spanish med-iterranean shrublands. Annals of Missouri Botanical Garden 74:69–78.

Keshet M.A., Danin A. and Orshan G. 1990. Distribution of eco-morphological types along environmental gradients in Israel: 1.Renewal bud location and leaf attributes. Ecologia Mediterra-nea 16: 151–161.

Le Roux A., Kyriacou X.L. and Orshan G. 1984. The phenomor-phology of selected plants in Mediterranean-type ecosystems ofSouth Africa. Bulletin de la Société Botanique de France (Ac-tualités Botaniques) 131: 441–450.

Mooney H.A. 1974. Plant form as related to environment. In:Tüxen R. (ed.), Handbook of vegetation science IV. W. Junk,Den Haag.

Navarro T., Nieto Caldera J.M., Pérez Latorre A.V. and CabezudoB. 1994. Estudios fenomorfológicos en la vegetación del sur deEspaña. III. Comportamiento estacional de una comunidad debadlands. Acta Botanica Malacitana 18: 189–198.

Navarro T. and Cabezudo B. 1998. Estrategias fenomorfológicasde especies de un matorral mediterráneo (Andalucía, España).Acta Botanica Malacitana 23: 133–148.

Nemani R. and Running S.W. 1996. Implementation of hierarchi-cal global vegetation classification in ecosystem function mod-els. Journal of Vegetation Science 7: 337–346.

Nieto Caldera J.M., Pérez Latorre A.V. and Cabezudo B. 1991.Biogeografía y series de vegetación de la provincia de Málaga(España). Acta Botanica Malacitana 16 : 417–436.

Noble I.R. and Gitay H. 1996. A functional classification for pre-dicting the dynamics of landscapes. Journal of Vegetation Sci-ence 7: 329–336.

Oppenheimer H.R. 1960. Adaptation to drought: xerophytism.UNESCO Arid Zone Research Service 15: 105–138.

Orshan G. 1982. Monocharacter growth-form types as a tool in ananalytic-synthetic study of growth forms in Mediterranean typeecosystems. A proposal for an inter-regional program. EcologiaMediterranea 8: 159–171.

Orshan G. 1983. Approaches to the definition of Mediterraneangrowth forms. In: Kruger F.J. (ed.), Mediterranean type ecosys-tems: The role of nutrients. Springer-Verlag, Berlín, New York,USA, pp. 86–100.

Orshan G., Leroux A. and Montenegro G. 1984. Distribution ofmonocharacter growth form types in mediterranean plant com-munities of Chile, South Africa and Israel. Bulletin de la So-ciété Botanique de France (Actualités Botaniques) 131: 427–439.

Orshan G. 1986. Plant form as describing vegetation and express-ing adaptation to environment. Annali di Botanica 54: 7–37.

Orshan G. (ed.) 1989. Plant phenomorphological studies in Medi-terranean type ecosystems. Kluwer Academic Publishers, Dor-drecht.

Parsons D.J. 1976. Vegetation structure in the mediterranean shrubcommunities of California and Chile. Journal of Ecology 64:435–447.

Pate J.S., Dixon K.W. and Orshan G. 1984. Growth and life formcharacteristics of Kwongan species. In: Pate J.S. and BuurdsJ.S. (eds), Kwongan-Plant life in the sand plain. University ofWestern Australia, Perth, pp. 84–100.

Pérez Latorre A.V. 1993. Estudio fitocenológico de los alcornocalesandaluces. Universidad de Málaga.

Pérez Latorre A.V., Nieto Caldera J.M. and Cabezudo B. 1993.Contribución al conocimiento de la vegetación de Andalucía.II. Los alcornocales. Acta Botanica Malacitana 18: 223–258.

Pérez Latorre A.V., Nieto Caldera J.M. and Cabezudo B. 1994.Contribución al conocimiento de la vegetación de Andalucía.III. Series de vegetación caracterizadas por Quercus suber L.Acta Botanica Malacitana 19: 169–183.

Pérez Latorre A.V., Cabezudo B., Navarro T. and Nieto J.M. 1995.Caracterización fenológica y eco-morfológica de alcornocalesandaluces (Málaga, España). Anales del Jardín Botánico deMadrid 54: 554–560.

Pérez Latorre A.V., Galán de Mera A., Deil U. and Cabezudo B.1996. Fitogeografía y vegetación del sector Aljíbico (Cádiz-Málaga, España). Acta Botanica Malacitana 21: 241–267.

Pérez Latorre A.V., Navas P., Cabezudo B. and Nieto Caldera J.M.1997. Los jarales de la clase Cisto-Lavanduletea en el sur de laPenínsula Ibérica (Andalucía, España). Acta Botanica Malaci-tana 22: 171–185.

Pérez Latorre A.V., Caritat A. and Vilar L. 2001. Use of growthform for describing and comparing Quercus suber forest in theIberian Peninsula. Phytocoenologia 31: 109–121.

Pierce S.M. 1984. A synthesis of plant phenology in the fynbosbiome. South African National Scientific Programmes report88. CSIR, Pretoria.

Rivas Martínez S. 1987. Memoria y Mapas de las Series de Veg-etación de España. MAPA-ICONA, Madrid.

Romane F. 1987. Efficacié de la distribution des formes de crois-sance pour l’analyse de la végétation á l’échelle régionale. Uni-versité de Marseille, Marseille.

Ruiz de la Torre J. 1971. Arboles y arbustos. Inst. Forestal de In-vestigaciones y Experiencias, Madrid.

Valdés B., Talavera S. and Galiano E.F. (eds) 1987. Flora vascularde Andalucía Occidental. Ketres, Barcelona.

Werger M.J. and Ellenbroek G.A. 1978. Leaf size and leaf consis-tence of a riverine forest formation along a climatic gradient.Oecologia 34: 297–308.

249