INIA Report. Project No. 1FD97-0911-C3-1 Causas del decaimiento y Seca de las masas de Quercus mediterrneas. Tcnicas de amortiguamiento.

2000 Ministerio de Ciencia y Tecnologa. Instituto Nacional de Investigacin y Tecnologa Agraria y Alimentaria (INIA).
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Oak decline and vegetation dynamics in southwestern Spain

Santiago Sardinero1*, ngel Fernndez-Cancio1, Ins Pereira1 & Emilio Manrique2
Instituto Nacional de Investigacin y Tecnologa Agraria y Alimentaria (INIA), Centro de Investigacin Forestal (CIFOR), Carretera de La Corua km 7, 28040 Madrid, Spain. 2 Escuela Universitaria de Ingeniera Tcnica Forestal, Universidad Politcnica de Madrid, Ciudad Universitaria, 28040 Madrid. * E-mail: sardiner@inia.es Key words: Drought, oak mortality, Querco-Oleion sylvestris, Pistacio-Rhamnetalia alaterni, Temperature raise, Vegetation shift.
1

Abstract We present vegetation data from SW Spain that has suffered oak decline and massive death, primarily Quercus rotundifolia Lam (roundleaved holm oak), and Quercus suber L (cork oak). Vegetation plots were recorded according to plant-forms and physiognomic-ecological plant formations. The sampling was done using the abundance/dominance values of the 6-grade scale of Braun-Blanquet. These values were transformed into a 0-9 ordinal scale. Multivariate data analysis was carried out, including Hierarchical Clustering and Correspondence Analysis. Oak decline has occurred fundamentally in the most thermal orientations (SW and S), steep slopes and thin soils, and also in soils with high clay content (luvisols, vertisols). This process is best observed in contacts between thermophyllous evergreen oak-wild olive woodlands (Querco-Oleion sylvestris) and semiarid evergreen preforestal shrubby vegetation (Pistacio-Rhamnetalia alaterni). The hypothesis of an increment of yearly water stress (progressive increment of temperatures plus increment of rainfall irregularity) is presented as the trigger mechanism for physiological and pathogenic processes leading to oak decline.

Introduction Humans have recognized vegetation as one of the characteristics that better define landscapes. Phytoclimatology studies the relationships between plants and climate, making possible to directly correlate their geographic distribution, and also that of the vegetation formations they constitute, with climatic parameters. Temperature, precipitation, and evapotranspiration are the three factors that better determine the basic conditions of existence for terrestrial plants. Plants adapt to environmental conditions adopting especial plant-forms, physiological processes, and seasonal habits. The adoption of particular combinations of plant-form, and leaf and plant size within a particular climatic interval results in positive water and energy balances. Limits for this climatic interval act as the most important mechanism of environmental limitation (Box 1981). The usefulness of world and regional-scale vegetation models has also been well recognized in plant geography since the early 19th Century. Such models have generally been based on environmental limitations, correlating distributional limits of vegetation types with macroclimatic factors (see Tuhkanen 1980). Presently, the existence of climatic databases, vegetation and flora databases, and large computers, has made possible to obtain bioclimatic models at world and regional scale (Box 1991).

Among global and regional models that relate climate and vegetation, and phytoclimatic classifications can be mentioned those of Schimper (1898), Martonne (1926), Kppen (1930-1936), Rbel (1930), Holdridge (1947), Thornthwaite (1948), Emberger (1954), Bagnouls & Gaussen (1957), Walter & Lieth (19601967), Troll & Paffen (1964), Mather & Yoshioka (1966), Hueck & Seibert (1972), Walter (1973), Ozenda (1975), Walter & Box (1976), Box (1981). In Spain phytoclimatic classifications of Rivas-Martnez (1987, 1997) and Allu Andrade (1990, 1997) are widely used. The Earth is involved in a process of global warming (Overpeck et al. 1997, Jones et al. 1998, Mann et al. 1998, 1999, Huang et al. 2000), correlated with a substantial and sustained raise in carbon dioxide atmospheric concentration during the last two centuries (Keeling & Whorf 1998, Etheridge et al. 1998). In the last century, average temperature of the Earth surface has risen 0.6 0.2 C, and the four warmest years on record since 1860 have all occurred since 1990 (Houghton et al. 1996). Calibrated and verified models for Iberia and the Balearic Islands until year 2050 (Ribalaygua et al. 1998, submitted; Born et al. submitted) estimate that average temperature could rise 1.5 C more. Also, during the last decades extreme precipitation events have occurred (Fernndez Cancio & Manrique 1998, Manrique & Fernndez Cancio 2000), alternating long dry periods and dramatic rainy years. In this context, Houghton et al. (1996) have determined that 1

INIA Report. Project No. 1FD97-0911-C3-1 Causas del decaimiento y Seca de las masas de Quercus mediterrneas. Tcnicas de amortiguamiento. 2000 Ministerio de Ciencia y Tecnologa. Instituto Nacional de Investigacin y Tecnologa Agraria y Alimentaria (INIA).
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vegetation shifts is one of the processes that it is probably to occur. Oak mortality in southwestern Spain has been detected (Fernndez Melndez & Montero 1993, Montoya Oliver & Mesn 1994, Allu Andrade 1994, 1995, 1997, Montoya 1995, Fernndez-Cancio 1999, Sardinero et al. 1999). This mortality attributed to biotic agents (Brasier 1992, 1996, Brasier et al. 1993, 1994, Tusset et al 1997), or abiotic (Fernndez-Cancio & Manrique 1998, Gil Pelegrn 1997) seems to respond to a climatic perturbation affecting Spain during the 20th century and it is especially active since 1980 (Candela 2000). A systematic field sampling of areas presenting generalized oak decline has been carried out in order to detect their distribution patterns, and also the distribution of several bioindicators (taxa and plant communities) which could help to recognize where this process is taking place, and where is most likely to occur in the present climate change scenario.

rotundifolia; the latter presenting advanced decline and death.

Methods Data sampling The study is based on a data matrix composed of 25 relevs and 85 taxa. The sampling was done during 1999. Plot locations were subjectively chosen where generalized oak decline and death had occurred, recording vegetation according to plant-forms and physiognomic-ecological plant formations (Rbel 1930; Raunkiaer 1937; Braun-Blanquet 1964; Ellenberg & Mueller-Dombois 1967a,b; Whittaker 1975; Box 1981). The data was recorded in the 6grade scale of Braun-Blanquet (1964). Data analysis The abundance/dominance values of the 6-grade scale of Braun-Blanquet were transformed into a 0-9 ordinal scale according to van der Maarel (1979), obtaining the data matrix to be analyzed. Similarity Ratio was used to calculate the distance among every pair of relevs in the resemblance matrix. A Complete Linkage clustering was carried out with the program NCLAS (Podani 1993, 1994). A Correspondence Analysis (CA) was carried out with the program PRINCOMP (Podani 1993, 1994).

Figure 1. Hierarchical clustering using similarity ratio and complete linkage.

Results Classification Classification results are shown in Figure 1. Four groups have been distinguished: Group 1. Includes Olea europaea var. sylvestris (wild olive) formations, with minor amounts of Quercus 2

The preforestal layer is characterized by thermophyllous bioindicators like Pistacia lentiscus, Rhamnus oleoides, and Asparagus albus. Retama sphaerocarpa dominates the leguminous nanophanerophytic layer. Genista hirsuta or Cistus albidus dominate tall shrub formations. If there is Cistus ladanifer it generally shows decline and death. Lavandula sampaiana dominates the short shrub formation, accompanied by Urginea maritima and Asphodelus aestivus. Hyparrhenia hirta constitutes hemicryptophytic grass layers. These formations are located in the most thermal orientations (S and SW), steep slopes and thin soils, on Guadalquivir, Guadiana, and Tajo Basins. Samples belonging to this group also can be found on vertisols. Group 2. Includes open woodlands dominated by tall shrubs as Cistus albidus and/or Cistus ladanifer, accompanied by Cistus salvifolius, Cistus crispus, Genista hirsuta, and Phlomis purpurea, with spread Quercus rotundifolia and Olea sylvestris codominating. Pistacia lentiscus, Quercus coccifera and Rhamnus alaternus dominate the preforestal

INIA Report. Project No. 1FD97-0911-C3-1 Causas del decaimiento y Seca de las masas de Quercus mediterrneas. Tcnicas de amortiguamiento. 2000 Ministerio de Ciencia y Tecnologa. Instituto Nacional de Investigacin y Tecnologa Agraria y Alimentaria (INIA).
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nanophanerophytic layer. Asparagus albus is less frequent than it is in Group 1. Retama sphaerocarpa dominates the leguminous nanophanerophytic layer. Lavandula sampaiana dominates the short shrub layer, accompanied by Urginea maritima and Asphodelus aestivus. Hyparrhenia hirta constitutes a hemicryptophytic grass layer. Nerium oleander and Rubus ulmifolius might be found dominating in depressions. Oak defoliation is generalized in this group. These formations are located in thermal orientations (S and SW), on less pronounced slopes than Group 1. Soils can be granitic, the less is soil development the much severe is oak defoliation. On red soils with high clay content oak defoliation is also generalized. Group 3. Includes open woodlands dominated by tall shrubs as Cistus ladanifer, who generally shows decline and death, accompanied by Cistus albidus, Cistus monspelliensis, Rosmarinus officinalis, Genista hirsuta and Phlomis purpurea, with spread Quercus rotundifolia dominating the phanerophytic layer, and less amounts of Olea sylvestris. Pistacia lentiscus and Quercus coccifera dominate the preforestal nanophanerophytic layer. Retama sphaerocarpa dominates the leguminous nanophanerophytic layer. Lavandula sampaiana dominates the short shrub layer. The presence of Phyllirea angustifolia, Lavandula luisieri, Quercus suber (cork oak), and Astragalus lusitanicus, indicates higher precipitation. Orientation and soils are similar to those of Group 2. This vegetation contact in N orientations with roundleaved holm oak-cork oak closed mixed forests that generally present several spread dead individuals. Hyparrhenia hirta is less frequent than it is in Group 2. Nerium oleander and Rubus ulmifolius might be found dominating in depressions. Oak decline is generalized in this group. Group 4. Tree layer is dominated by Quercus rotundifolia with low or null amounts of Olea sylvestris, growing on heavy granitic substrates. The preforestal nanophanerophytic layer lacks of thermophyllous bioindicators present in Groups 1, 2, and 3. Retama sphaerocarpa dominates the leguminous nanophanerophytic layer, with high frequency of Cytisus scoparius. Cistus ladanifer dominates the tall shrub layer, and Lavandula sampaiana the short one. Hyparrhenia hirta might establish hemicryptophytic grass layers. Agrostis castellana lawns grow in depressions. These formations are located in thermal orientations (S and SW). These samples are located from the Tajo Basin to the north. Oak decline decreases in the same way. Ordination Ordination results are shown in Figures 2 and 3. Classification results were superimposed on the 3

ordination diagram (Figure 2; Group 1 = circles, Group 2 = squares, Group 3 = triangles, Group 4 = diamonds).
3 CA for samples

1 2 Axis 2

3 4

Figure 2 -3 -3 Axis 1 2

Figure 2. CA ordination for samples. Numbers correspond to vegetation groups described in the text.

6 CA for plants
AcaMol

Ar u

Ita

AriBae
F ru Jas

StiGig

ue Su

om MyrC im AriS euFru T RhaAla Asp Tam Com Acu O leSyl ri C PisT n is is er Mo cH C Ci s BryDio GenHir Da ni pG RubPer Ur CisAlb Da nn gM RetS PhlPur MerA ph ar up Ci NerOle Um bR sS AriPau ot a m al HelSer eR vS OsyAlb ng CytSco Q u La Ph y A La v Lu i L ad Cis RosO ff AgrCas nIm p EriArb Lo SciHol DigTha AstLus VibTin HalVis JunOxy ArbUne SecTin
Rh As
r

AspAph

aO

lHi

pA

Lu

Ba

le

Axis 2

lb

r Lu

s Di a

ir

yp

Th

yM

as

Ca

rH

is

Q Pi ue s L Co en c

Pa

Figure 3 -4 -4 Axis 1 3

Figure 3. CA ordination for taxa.

The central part of Figure 3 is occupied by plants characteristic to infrailicine stages: Olea sylvestris, Rhamnus alaternus, Asparagus albus, Rhamnus oleoides, Ballota hirsuta, very well represented in Group 1. Displacement in Figure 2 from Group1 toward Groups 2 and 3 means that tall shrubs, as Cistus crispus, Cistus albidus, Genista hirsuta, Cistus salvifolius, Phlomis purpurea, Cistus ladanifer, acquire more relevance (Figure 3); Nerium oleander is getting progressive importance in depressions.

INIA Report. Project No. 1FD97-0911-C3-1 Causas del decaimiento y Seca de las masas de Quercus mediterrneas. Tcnicas de amortiguamiento. 2000 Ministerio de Ciencia y Tecnologa. Instituto Nacional de Investigacin y Tecnologa Agraria y Alimentaria (INIA).
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Quercus coccifera, Pistacia lentiscus, Cistus monspelliensis and Rubia peregrina are located within Groups 1, 2, and 3. Phyllirea angustifolia, Lavandula luisieri, Lonicera implexa, Rosmarinus officinalis, Erica arborea, Astragalus lusitanicus, Quercus suber, Viburnum tinus and Arbutus unedo are located in the bottom-right part of Figure 3. This is related to higher precipitation in Group 3. Displacement in Figure 2 from Group 1 toward Group 4 is related to higher frequency of Cytisus scoparius, Mercurialis annua, Digitalis thapsi, Juniperus oxycedrus, Stipa gigantea, and Agrostis castellana lawns (Figure 3). This is related to displacement from southwestern toward central Spain. Acanthus mollis, Arum italicum, and Aristolochia baetica are located in the right-upper part of Figure 3. They differentiate relev no. 1, a thermophyllous wild olive formation in Guadalquivir River on vertisols.

dynamics, oak decline and death means that wild olive formations are moving out of the river basins. 4. In other orientations it can be detected oak decline and death in several individuals, but in all cases the proportion is much less, constituting what we know as spread deaths (few deaths among many trees that form a closed canopy). 5. Massive oak decline also occurs in depressions, on soils with vertic properties, i.e., with clay that hydrates in rainy seasons (winter, spring, fall), and dehydrates in the dry season (summer), helping to the continuous mixture of surface horizons (FAOUNESCO 1991). 6. Chromic luvisols in southern Badajoz (Tierra de Barros = Mudland) have high productivity rates, and they are cultivated with olives (Olea europaea var. europaea) and vines (Vitis sylvestris). Nevertheless, in marginal situations natural vegetation can be recognized. Quercus rotundifolia and Olea sylvestris dominate the tree layer. Pistacia lentiscus and Quercus coccifera dominate the nanophanerophytic layer. Cistus albidus, Cistus monspelliensis, Cistus ladanifer, and Genista hirsuta dominate the tall shrub layer, accompanied by Phlomis purpurea and Teucrium fruticans. Lavandula sampaiana dominates the short shrub layer. Data on this vegetation might be found in Rivas Goday (1964), Santos et al. (1989), and Ladero et al. (1990). These soils are characterized by the progressive migration of materials from the surface toward deeper horizons during humid seasons (winter, spring, fall), beginning by soluble salts and carbonates, and following by a gradual translocation of clay into an intermediate horizon (argilic B horizon; FAO-UNESCO 1991). On these soils a generalized oak decline and death has occurred, most frequently in thermal orientations. Pistacia lentiscus, Quercus coccifera, Cistus albidus, Phlomis purpurea, and Teucrium fruticans are thermophyllous bioindicators frequently present. At the same latitude they are not present on soils without clay (acidic substrates in general). 7. The phytosociological alliance Querco-Oleion sylvestris (Barbro et al. 1981; Rivas-Martnez et al. 1986; Benabid & Fennane 1994) in SW Iberia gather evergreen-sclerophyll oak and wild olive forests and woodlands. Its optimal thermotype is Thermomediterranean, although it can reach the Lower Mesomediterranean bioclimatic belt. Its ombrotype is dry or wetter. Plants belonging to the Order Pistacio-Rhamnetalia alaterni (Rivas-Martnez 1975) dominate the preforestal nanophanerophytic layer, and also constitute permanent plant communities on very dry soils. Significant plants belonging to this order are Pistacia lentiscus, Rhamnus oleoides ssp. oleoides, Quercus coccifera, Rhamnus oleoides ssp. angustifolia, Myrtus 4

Conclusion and discussion 1. Natural vegetation on the most thermal orientations (S and SW), steep slopes, and thin soils in the Tajo, Guadiana, and Guadalquivir Basins consists of wild olive formations. They are dominated by Olea sylvestris, accompanied by several thermophyllous taxa as Rhamnus oleoides, Asparagus albus, Hyparrhenia hirta, and in Guadiana and Guadalquivir Basins, Pistacia lentiscus (Rivas-Martnez 1975, 1987; Prez Chiscano 1976; Santos et al. 1989; Ladero et al. 1990; Cano & Valle 1990; Melendo 1998). This vegetation locally reaches the Duero Basin (Navarro et al. 1987). 2. Natural vegetation on vertisols of the Guadalquivir Basin and NW Morocco consists also in wild olive formations. They are accompanied by a preforestal nanophanerophytic layer dominated by Pistacia lentiscus with Chamaerops humilis, Daphne gnidium, Rhamnus oleoides, Rhamnus alaternus, Quercus coccifera, Myrtus communis. Vines as Clematis cirrhosa, Aristolochia baetica, Bryonia dioica, Tamus communis, Vinca difformis, Smilax altissima, Asparagus albus, and Asparagus aphyllus. A herb layer having Arum italicum, Arisarum subexertum, Asphodelus ramosus. Several mediterranean shrubs as Phlomis purpurea and Teucrium fruticans (Benabid 1984; Rivas-Martnez 1987; Prez Latorre et al. 1993; Galn de Mera 1993). 3. Massive oak decline occurs in the most thermal orientations (S and SW), thin soils (granitic and basic rocks), and steep slopes. These situations can be observed with highest frequency in the thermal gradient that goes from the warmest vertical walls in the Tajo, Guadiana, and Guadalquivir rivers, toward the external river basin. In terms of vegetation

INIA Report. Project No. 1FD97-0911-C3-1 Causas del decaimiento y Seca de las masas de Quercus mediterrneas. Tcnicas de amortiguamiento. 2000 Ministerio de Ciencia y Tecnologa. Instituto Nacional de Investigacin y Tecnologa Agraria y Alimentaria (INIA).
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communis, Chamaerops humilis, Osyris alba, Osyris quadripartita, Asparagus albus, Teucrium fruticans, etc. In territories with semiarid ombrotype they represent the climatic climax. They configure mediterranean preforests and shrubs, primarily evergreen-sclerophyll (see Barbro et al. 1992). All these authors agree in that it is the water stress and primarily the summer water stress what moves vegetation from Querco-Oleion sylvestris toward Pistacio-Rhamnetalia alaterni. 7. The confirmation that massive oak decline and death is occurring primarily in the most thermal orientations (S and SW), steepest slopes, and tiniest soils in vegetation contacts between wild olive and oak formations seems to support this model. Furthermore, observations made in points 2, 5, and 6, i.e., total absence, massive death, or generalized oak defoliation on soils having high clay content, lead us to hypothesize that during the dry season (summer), clay, with its high capacity for water retention, drives oaks to be water stressed. During exceptional long dry periods and/or with exceptional high temperatures water stress could be the limiting factor for oaks. Ecophysiological studies are needed on all the important plants in the transect from QuercoOleion sylvestris toward Pistacio-Rhamnetalia alaterni, especially studies on how these plants face water stress. All these observations point to water stress as the limiting factor for oaks, leading to physiological and pathogenic processes that have been related to oak decline (Brasier 1992; Brasier & Scott 1994; Brasier 1996; Tuset et al. 1996, Montoya & Lpez Arias 1998).

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INIA Report. Project No. 1FD97-0911-C3-1 Causas del decaimiento y Seca de las masas de Quercus mediterrneas. Tcnicas de amortiguamiento. 2000 Ministerio de Ciencia y Tecnologa. Instituto Nacional de Investigacin y Tecnologa Agraria y Alimentaria (INIA).
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