Changes in the summit flora of a Mediterranean mountain (Sierra Nevada, Spain) as a possible effect of climate change/Cambios en la flora de alta montana de una montana mediterranea (Sierra Nevada, Espana) como un posible efecto del cambio climatico.
The average annual surface temperature has increased by 0.8[degrees]C in Europe during the past century (ALCAMO & al., 2007) and two to three fold greater rates of warming are projected for the 21st century (NOGUES-BRAVO & ARAUJO, 2006). Specifically, on the Iberian Peninsula climate change projections for the current century predict an increase of the average temperature by 0.4[degrees]C/decade in winter and 0.7 [degrees]C/decade in summer, for the least favourable scenario (A2 of the IPCC), and by 0.4[degrees]C and 0.6[degrees]C/decade, for the most favourable scenario (B2 of the IPCC) (FERNANDEZ-GONZALEZ & al., 2005).
Mountain ecosystems are especially sensitive to climate change because they are limited by low temperatures (CHAPIN & KORNER, 1994; PRICE & BARRY, 1997; KORNER, 2003; PAULI & al., 2005) being its flora very particular with many endemics (DEL EGIDO & PUENTE, 2011; GAVILAN & al., 2012). Therefore, climate warming is expected to cause significant changes in high mountain plant diversity such as upward shifts from lower altitudes, extinctions or changes in the competitive relations among plants (GUISAN & al., 1995; BENISTON & al., 1997; KORNER, 2003; Grabherr & al., 2010).
Most of the revisitation studies in high mountain vegetation reported an increase in the number of species in the Alps (GRABHERR & al., 1994, 2001; BAHN & KORNER, 2003; WALTHER, 2005) and the Scandes (KLANDERUD & BIRKS, 2003). More recent evidences confirm the previously observed increases in species numbers in the Alps (HOLZINGER & al., 2008; VITTOZ & al., 2008; ERSCHBAMER & al., 2008; KULLMAN, 2010; WIPF & al., 2012, in press). A recent range contraction of subnival to nival species at their lower range margin, however, has been observed in the Alps (PAULI & al., 2007).
On this framework, the first high-altitude permanent plots in southern Spain were established in 2001 as part of the Global Observation Research Initiative in Alpine Environments (GLORIA, http://www.gloria.ac.at), whose main aim is to provide long-term observation series on the state of alpine biota. Permanent plots were established along a standardised sampling design on mountain summits along an elevation gradient, where vegetation data and time series of temperatures were recorded (PAULI & al., 2004).
The recent multi-region report of the first resurvey seven years after the establishment of the GLORIA sites in Europe (PAULI & al., 2012) has clearly confirmed that changes in vascular plant species exist. Therefore, the aims of the present study are to analyze the detailed changes in the Sierra Nevada (Spain, ES-SNE) related to slope exposure, summit elevation and the altitudinal distribution range of the species.
MATERIALS AND METHODS
The Sierra Nevada is located in the south-east of the Iberian Peninsula (37[degrees]N, 3[degrees]W), within the Baetic Range of Mountains (Figure 1). It contains numerous summits that exceed 3000 m asl, including the highest peak of the Peninsula (Mulhacen, 3482 m asl).
Within its upper part (above 2600 m asl) four sampling summits were permanently marked in 2001 according to the criteria specified in the GLORIA field manual (PAULI & al., 2004). These summits are situated in the western central zone of the range: Machos (MAC) 3327 m asl; Tosal Cartujo (TCA) 3150 m asl; Cupula (CUP) 2968 m asl and Pulpitito (PUL) 2778 m asl (Figure 1).
All the summits share a Mediterranean bioclimate in its pluviseasonal oceanic variant (RIVAS-MARTINEZ & al., 2007). Summer drought is pronounced and the 700-1500 mm/year rainfall occurs almost exclusively as winter snow at altitudes above 2500 m. All summits are formed by siliceous bedrocks except one (Pulpitito), having a less acidic substrate. The lower summits are dominated by dwarf shrub communities, while the higher ones are composed by grasses, hemicryptophytes and cushion chamaephytes scattered on open psycroxerophylous soils.
The standardized GLORIA Multi-Summit Approach (PAULI & al., 2004) was used to establish four permanent sampling summits in 2001. Each summit was divided into eight summit area sections (SASs), aligned along the four cardinal directions (N, S, E and W). The four upper sections cover the area from the highest summit point (HSP) to the 5m contour line, while the four lower ones from the 5-m to the 10 m contour line. All vascular plant species were recorded in each one of the sections.
Four quadrat clusters (3 m x 3 m) were established in each cardinal direction (N, E, S, W) at 5 m below the highest summit point. In each of the four 1 [m.sup.2] corner-plots of each cluster, a complete list of plant species and an estimation of their percentage cover and frequency (i.e. their presence in 100 divisions of 10 x 10 cm) was recorded in the 1 m x 1 m plots. Further, a data-logger (StowAway TidbiT, Onset Corporation, Massachusetts, uSA) was installed at 10 cm below the soil surface in each central quadrat of the clusters to measure soil temperatures at hourly intervals. In 2008, the resurvey involved the same procedure used in 2001.
We used as sources for taxonomy, distribution and altitudinal ranges of the species Flora Europaea (TUTIN & al., 1964-1980), Flora Iberica (CASTROVIEJO & al., 1986-2009) and regional floras or vegetation studies (MARTINEZ-PARRAS & al., 1985; MOLERO MESA & PEREZ-RAYA 1987; MOLERO MESA & al., 1996; GIMENEZ & GOMEZ, 2002).
To investigate whether the species richness of vascular plants in the summit area sections (SASs) changed between 2001 and 2008, we used a 3-way ANOVA using summit, slope exposure (N, S, E, W) and year as factors, and accounting for their interactions. A 3-way ANOVA was also used to investigate whether vascular plant species richness and Shannon Index (DEL RIO & al., 2003) changed in the 1 [m.sup.2] plots between 2001 and 2008. We verify whether species frequency changed in relation to slope exposure and/or summit elevation by a Chi-Square test for heterogeneity or independence ([chi square]-Test). In the cases where the results of ANOVA were significant, we tested for differences between groups within a factor though a Tukey and Bonferroni test. Statistical analyses were performed using SPSS 15.0.
We analyzed mean annual soil temperature data from each cardinal direction of each summit (n = 16) from January 2002 to December 2008. The trends of these temperatures, in each of the 16 time series, were studied fitting a straight line model. Temperature data were also used to calculate the growing season period, defined as the number of days with mean daily soil temperatures > 2[degrees]C (ERSCHBAMER & al., 2008; Vittoz & al., 2010). Thus, the first day was that with a mean temperature > 2[degrees]C, as long as this temperature was maintained for at least 6 days, while the end of the period was defined as the first day with a mean temperature < 2[degrees]C also maintained for at least 6 days. Then, we calculated the mean growing season length along the main compass direction at the four Sierra Nevada summits.
TRENDS IN SPECIES RICHNESS
Considering all summits together, species richness changed from 79 to 78 taxa during the seven year period. Overall, seven species disappeared from all summits areas, four of them showing a narrow distribution area (Poa minor subsp. nevadensis, Vitaliana primuliflora subsp. assoana, Plantago nivalis and Coincya monensis subsp. nevadensis) and three having a wider distribution (Galium rosellum, Luzula hispanica and Rhamnus pumilus). Six species were newly found: four annual species with widespread distribution (Erophila verna, Cuscuta sp. in PUL-E10, Viola sp. in TCA-S11 and Veronica sp. in CUP-S11) and two restricted to the Sierra Nevada massif (Senecio nevadensis and Linaria glacialis) (Appendix 1).
At the summit level (Table 1), absolute and mean species richness values showed a decreasing trend between 2001 and 2008, although these changes were not statistically significant. A detailed overview of locally new and locally lost taxa is shown in Table 2. At the plot scale (Table 1), the total species richness in the 1 [m.sup.2] plots decreased on two summits (PUL and TCA), stagnated on CUP and increased on MAC.
Species richness and Shannon Weaver Index did not significantly change at the 1 [m.sup.2] scale. In spite of this, changes in the sub-plot species frequency within the 1 [m.sup.2] plots were significantly ([chi square]-; p = 0.040) influenced by slope exposure: the number of species with decreasing frequency was larger than those with increasing frequency in every cardinal direction. Consistently, the number of disappearing species was larger than the number of appearing ones (Table 3).
Percentage cover data showed changes in the majority of the recorded species, but they were not significant. Despite that, we have believed an interesting item to show in Table 4 those cases where either increases or decreases of [greater than or equal to] 1% in absolute terms (referred to 1 [m.sup.2] plots) in more than two plots from 2001 to 2008 were observed.
Mean annual soil temperatures from the four cardinal directions of the four summits from 2002 to 2008 are shown in Figure 2. There was no clear warming trend within this short period. Differences among the main cardinal directions were, however, discernible. Southern slopes were the warmest, with the exception of the TCA summit, where higher temperatures occurred in the eastern direction in some years.
The mean duration of the growing season for each summit is shown in Table 5. The longest growing seasons were at the southern and eastern expositions, meanwhile the shortest growing season period was inconsistent among the cardinal directions, although it was least common at southern slopes.
This study provides detailed information about the changes occurred in the GLORIA summits of Sierra Nevada, a crucial mountain region to understand the impact of climate change in the Mediterranean regions (PAULI & al., 2012).
In contrast to the majority of changes observed in European alpine sites (Grabherr & al., 1994, 2010; WALTHER & al., 2005; KULLMAN, 2010), species richness was stagnating or decreasing on all summit sites. This is also indicated by the plot-level data (abundance in the 1 [m.sup.2] plots), where the number of species with decreasing abundance exceeded the increasers and the number of species not found again was larger than that of newly appearing ones in all cardinal directions.
Among the few studies which provide these kind of evidences, a decline of high-elevation species, are KLANDERUD & BIRKS (2003) from the Scandes and PAULI & al. (2007) from the Alps. Adding to these, a recent Europe-wide GLORIA study (PAULI & al., 2012), which includes Sierra Nevada GLORIA sites, showed similar observations in other Mediterranean mountains (Corsica/France, Lefka Ori-Crete /Greece), where an average decrease in species number was recorded opposed to an average increase for the boreal and temperate summits, even though species predominantly showed an upward shift across all three biomes.
PAULI & al. (2012) hypothesized that the observed species declines could indicate range retractions through a combination of rising summer temperature and stable to decreasing rainfall, facts also reported by the comprehensive governmental assessment report on the effects of climatic change in Spain (FERNANDEZ GONZALEZ & al., 2005). In the Sierra Nevada, as in other Mediterranean high mountains, the stagnation or decrease of species numbers and the local disappearance of species is particularly worrying because its flora has a high percentage of endemic and relict taxa, where further declines can result in irretrievable losses on the phylogenetical level (SANZ ELORZA & al., 2003; FERNANDEZ CALZADO & al., 2012).
Changes in species cover (in the 1 [m.sup.2] plots), despite are not significant or marginally significant, may partly be related to the local habitat situation, but in some cases seem to reflect directional changes with respect to the altitudinal range of species and to moisture requirements of species, trend that will need confirmation in the future. A related pan-European paper, which included the Sierra Nevada GLORIA sites (GOTTFRIED & al., 2012), consistently showed that species of lower elevations were immigrating to or expanding within higher-elevation sites. This 'thermophilisation' signal was significant across the entire European data set, but also for some single GLORIA sites such as for Sierra Nevada. The cover decrease of Genista baetica, Erodium cheilanthifolium and Festuca indigesta, common species on the lowest summit (PUL), might be related to the abundant unstable substrate which could impede their establishment. The cover increase of some chamaephytes on the other summits, such as Reseda complicata, Thymus serpylloides subsp. serpylloides and particularly of Alyssum spinosum, as well as the new appearance of Senecio nevadensis on two summits, however, could likely have been boosted by climate change, as their upper distribution ranges were observed to be expanding in the upper zone of the Sierra Nevada (FERNANDEZ CALZADO & MOLERO MESA, 2011a, b; and personal observations of the authors). A continued increase of chamaephytes such as Alyssum spinosum may further facilitate the arrival of other lower elevation species by acting as nurse plants (CALLAWAY & al., 2002; CAVIERES & al., 2006; KAMMER & al., 2007; GRABHERR & al., 2010). Festuca indigesta, a common graminoid of the oromediterranean belt that reaches its upper limit on CUP, was increasing in cover on this summit, whereas Festuca clementei, a common endemic restricted to the uppermost belt, was decreasing on TCA.
Several other high-elevation endemics, such as Saxifraga nevadensis, Artemisia granatensis, Vitaliana primuliflora subsp. assoana, Poa minor subsp. nevadensis and Coincya monensis subsp. nevadensis were not found in 2008. The same accounts for Luzula hispanica on CUP and Plantago nivalis on TCA, and Ranunculus demissus was decreasing in cover on CUP. According to the knowledge about the ecology of the studied species, it is likely that observed changes are associated with the reduction of water availability.
Species richness in the summit areas was stagnating or decreasing and, at the plot scale, species abundance were more commonly declining and the numbers of local disappearances were larger than new appearances.
Both changes in presence and absence of species as well as of species cover appear to reflect in several cases shifts along a moisture gradient.
Declines in species richness as well as in species cover are surprising in comparison to similar studies in the Alps and Scandes, but are in accordance with the results of other Mediterranean GLORIA sites reported in the pan-European GLORIA studies: a thermophilisation of the species composition of high mountain plant communities and a predominant upward-shift of species across Europe's mayor biomes, but declines in species numbers in the Mediterranean region, which could result from a combined effect of rising temperatures and restricted water availability.
Ongoing climate change impacts on the high-elevation flora of Sierra Nevada are expected to continue in the view of model predictions of further warming and decreasing precipitation (FERNANDEZ GONZALEZ & al., 2005, Christensen & al., 2007) and are worrisome insofar as a large proportion of the vascular plant flora is highly endemic and restricted to the uppermost elevation zone.
Appendix 1 List of plant species in GLORIA summits (2001 and 2008) (Distribution: Ne, Nevadense, Be, Baetican, Ib, Iberian, Ib-N, Iberian-northern African, Eu, European, Eu-N, European-northern African, Others, widely distributed) Taxa name Family Distribution Acinos alpinus subsp. meridionalis Lamiaceae Eu-N Aethionema saxatile subsp. Brassicaceae Eu-N marginatum Agrostis nevadensis Poaceae Ne Alyssum purpureum Brassicaceae Ne Alyssum spinosum Brassicaceae Eu-N Andryala agardhii Asteraceae Be Anthyllis vulneraria subsp. Fabaceae Ne pseudoarundana Arenaria armerina Caryophyllaceae Ib-N Arenaria pungens Caryophyllaceae Be Arenaria tetraquetra subsp. Caryophyllaceae Ne amabilis Artemisia granatensis Asteraceae Ne Asperula aristata subsp. scabra Rubiaceae Eu-N Biscutella glacialis Brassicaceae Ne Campanula willkommii Campanulaceae Eu Carduus carlinoides subsp. Asteraceae Ne hispanicus Cerastium ramosissimum Caryophyllaceae Others Chaenorhinum glareosum Scrophulariaceae Ne Cirsium gregarium Asteraceae Be Coincya monensis subsp. nevadensis Brassicaceae Ne Crepis oporinoides Asteraceae Be Cuscuta sp. Cuscutaceae -- Cystopteris fragilis subsp. Athyriaceae Others fragilis Dactylis juncinella Poaceae Ne Deschampsia flexuosa subsp. iberica Poaceae Ib Dianthuspungens subsp. brachyanthus Caryophyllaceae Ib-N Draba hispanica subsp. laderoi Brassicaceae Ne Erigeron frigidus Asteraceae Ne Erigeron major Asteraceae Be Erodium cheilanthifolium Geraniaceae Be Erophyla verna Brassicaceae Others Erysimum nevadense Brassicaceae Be Eryngium glaciale Apiaceae Ib-N Euphorbia nevadensis Euphorbiaceae Ib Euphrasia willkommii Scrophulariaceae Ib-N Festuca clementei Poaceae Ne Festuca indigesta Poaceae Ib-N Festuca pseudeskia Poaceae Ne Galium pyrenaicum Rubiaceae Ib Galium rosellum Rubiaceae Be Genista baetica Fabaceae Be Herniaria boissieri Caryophyllaceae Be Hieracium castellanum Asteraceae Eu Holcus caespitosus Poaceae Ne Iberis carnosa subsp. embergeri Brassicaceae Ne Jasione crispa subsp. amethystina Campanulaceae Ne Juniperus communis subsp. Cupressaceae Others hemisphaerica Juniperus sabina Cupressaceae Others Jurinea humilis Asteraceae Eu-N Lactuca perennis subsp. granatensis Asteraceae Be Leontodon boryi Asteraceae Be Lepidium stylatum Brassicaceae Ne Leucanthemopsis pectinata Asteraceae Ne Linaria aeruginea subsp. nevadensis Scrophulariaceae Ne Linaria glacialis Scrophulariaceae Ne Lotus corniculatus subsp. glacialis Fabaceae Ne Luzula hispanica Juncaceae Ib Myosotis minutiflora Borraginaceae Others Nepeta amethystina subsp. laciniata Lamiaceae Ne Plantago nivalis Plantaginaceae Ne Plantago radicata subsp. Plantaginaceae Eu granatensis Paronychia polygonifolia Caryophyllaceae Others Pimpinella procumbens Apiaceae Ne Poa ligulata Poaceae Ib-N Poa minor subsp. nevadensis Poaceae Ne Poa nemoralis Poaceae Eu Prunus prostrata Rosaceae Others Rhamnus pumila Rhamnaceae Eu-N Ranunculus demissus Ranunculaceae Others Reseda complicata Resedaceae Ne Saxifraga nevadensis Saxifragaceace Ne JO Sedum amplexicaule subsp. Crassulaceae Others tenuifolium Sedum dasyphyllum Crassulaceae Eu-N Sempervivum nevadense Crassulaceae Be Senecio boissieri Asteraceae Ib Senecio nevadensis Asteraceae Ne Senecio pyrenaicus subsp. Asteraceae Be granatensis Sideritis glacialis Lamiaceae Ne Silene boryi Caryophyllaceae Ib Teucrium lerrouxii Lamiaceae Ib Thymus serpylloides subsp. Lamiaceae Ne serpylloides Trisetum glaciale Poaceae Ne Veronica sp. Scrophulariaceae -- Viola sp. Violaceae -- Viola crassiuscula Violaceae Ne Vitaliana primuliflora subsp. Primulaceae Ne assoana
The setup of the permanent plots and data collection (2000-2003) was supported by the FP-5 project GLORIA-Europe (EVK2-CT-2000-0006) of the European Commission. Resurvey (2008) was supported by the Swiss MAVA Foundation for Nature Conservation and by a number of national funding agencies. We thank A. San Miguel Ayanz for his valuable comments and suggestions.
Received: 11 March 2013
Accepted: 21 November 2013
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Maria Rosa Fernandez Calzado & Joaquin Molero Mesa (*)
* Department of Botany. Faculty of Pharmacy. University of Granada. Campus de Cartuja. E-18071 Granada, Spain. E-mail: email@example.com
Table 1 Total species richness in the summit area (pooled from eight SASs) and the 16-[m.sup.2] plots (pooled from 16 single 1[m.sup.2] plots) of each of the four study summits in 2001 and 2008 (mean number for SASs and 1[m.sup.2] plots [+ or -] standard deviation). Summit area sections (SAS) Summit 2001 2008 Pulpitito 47 (30 [+ or -] 7.2) 45 (24.4 [+ or -] 6.4) Cupula 52 (29.3 [+ or -] 4.5) 50 (26.6 [+ or -] 5.1) Tosal Cartujo 40 (20 [+ or -] 4.8) 39 (18.4 [+ or -] 4.4) Machos 18 (8.5 [+ or -] 3.2) 16 (7 [+ or -] 2.7) 1 [m.sup.2] plots Summit 2001 2008 Pulpitito 31 (9.3 [+ or -] 3.9) 27 (8.1 [+ or -] 2.9) Cupula 32 (11.6 [+ or -] 2.9) 32 (10.8 [+ or -] 2.3) Tosal Cartujo 20 (5.8 [+ or -] 1.7) 18 (5.3 [+ or -] 2.1) Machos 13 (0.8 [+ or -] 1.1) 14 (0.9 [+ or -] 1.3) Table 2 New and lost taxa at the Sierra Nevada summits from 2001 to 2008. (Distribution: Ne, Nevadense, Be, Baetican, Ib, Iberian, Ib-N, Iberian-northern African, Eu, European, Eu-N, European-northern African, Others, widely distributed) Summit New species Lost species PUL Cuscuta sp. Erysimum nevadense (Be) Erophila verna (Others) Hieracium castellanum (Eu) Galium rosellum (Be) Rhamnus pumilus (Eu-N) CUP Cystopteris fragilis (Others) Artemisia granatensis (Ne) Erophila verna (Others) Carduus carlinoides subsp. Erysimum nevadense (Be) hispanicus (Ne) Senecio nevadensis (Ne) Erigeron major (Be) Veronica sp. Hieracium castellanum (Eu) Luzula hispanica (Ib) Saxifraga nevadensis (Ne) Vitaliana primuliflora subsp. assoana (Ne) TCA Cirsium acaule subsp. Arenaria armerina (Ib-N) gregarium (Be) Biscutella glacialis (Ne) Erigeron frigidus (Ne) Coincya monensis subsp. Euphorbia nevadensis (Ib) nevadensis (Ne) Linaria glacialis (Ne) Cystopteris fragilis (Others) Viola sp. Galium pyrenaicum (Ib) Plantago nivalis (Ne) MAC Crepis oporinoides (Be) Cystopteris fragilis (Others) Linaria glacialis (Ne) Herniaria boissieri (Be) Senecio nevadensis (Ne) Leontodon boryi (Be) Linaria aeruginea subsp. nevadensis (Ne) Poa minor subsp. Nevadensis (Ne) Table 3 Total number of species with frequency changes within the 1-[m.sup.2] plots (hundred 0.1 x 0.1 m subplots) in each of the four cardinal directions of the four Sierra Nevada summits combined. Species Increasing/ Appearing/ response Decreasing Disappearing East 36/42 17/30 North 26/46 3/14 South 41/49 15/21 West 31/41 13/16 Table 4 Species with cover changes (2001 versus 2008) in the 1-[m.sup.2] plots of [greater than or equal to] 1% (in absolute terms, referred to 1[m.sup.2] plots) in more than two quadrates on the four Sierra Nevada summits. Change Summit Species in cover PUL Genista baetica [down arrow] Erodium cheilanthifolium [down arrow] Festuca indigesta [down arrow] CUP Alyssum spinosum [up arrow] Thymus serpylloides subsp. serpylloides [up arrow] Festuca indigesta [up arrow] Ranunculus demissus [down arrow] Crepis oporinoides [down arrow] Deschampsia flexuosa subsp. iberica [down arrow] TCA Arenaria tetraquetra subsp. amabilis [up arrow] Reseda complicata [up arrow] Jasione crispa subsp. amethystina [down arrow] Festuca clementei [down arrow] MAC Alyssum spinosum [up arrow] Table 5 Mean growing season length (2002 to 2008), in days, along the main compass directions at the four Sierra Nevada summits. MAC TCA CUP PUL N 128.86 164.71 192.57 200.71 S 167.86 156.57 200.00 218.29 E 149.29 168.43 165.14 188.57 W 128.29 165.71 192.14 204.43
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|Title Annotation:||articulo en ingles|
|Author:||Calzado, Maria Rosa Fernandez; Mesa, Joaquin Molero|
|Date:||Jan 1, 2013|
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