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M. Cueto et al. Malacitana Vascular flora the Western Mediterranean (Andalusia, Spain) 81 Acta Botanica 39.in81-97 Málaga, 2014

DIVERSITY AND ECOLOGICAL CHARACTERISTICS OF THE VASCULAR FLORA IN THE WESTERN MEDITERRANEAN (EASTERN ANDALUSIA, SPAIN) Miguel CUETO1*, Gabriel BLANCA2, Carlos SALAZAR3 & Baltasar CABEZUDO4 Departamento de Biología y Geología, Universidad de Almería, 04120 Almería, Spain. 2 Departamento de Botánica, Universidad de Granada, 18071 Granada, Spain. 3 Departamento de Biología Animal, Biología Vegetal y Ecología, Universidad de Jaén, 23071 Jaén, Spain. 4 Departamento de Biología Vegetal (Botánica y Fisiología Vegetal), Universidad de Málaga, 29071 Málaga, Spain. *Author for correspondence: [email protected] 1

Recibido el 9 de septiembre de 2014, aceptado para su publicación el 1 de octubre de 2014

ABSTRACT . Diversity and ecological characteristics of the vascular flora in the western mediterranean (Eastern Andalusia, Spain). This work highlights the importance of the vascular flora of eastern Andalusia within the hotspot of the Mediterranean Basin, analysing 3726 taxa, which represent 42.0% of the Iberian Peninsula and 29.9% of European floras. Notably, 10.3% of the flora is endemic (350 taxa), constituting 34.3% of the endemism of the Iberian Peninsula and Balearic Islands, and 2.6% of those of the Mediterranean Basin, which include 6 of the 17 endemic genera of the Iberian Peninsula and Balearic Islands, 3 of these being exclusive of the territory analysed. Chamaephytes and hemicryptophytes make up 45.5% of the flora, with decreasing values for the therophytes as altitudes rise (41.1-8.1%) and increasing values for hemicryptophytes (22.4%-63.2%). Mediterranean taxa comprise 39.2%, followed by Ibero-North African taxa (15.3%), Iberian (13.9%), European (11.3%) and local endemism (9.4%). Of these taxa, 12.5% are threatened, according to the criteria of the UICN. Tanacetum funkii, exclusive of this area, is considered extinct. Only 6 families account for 43.5% of the threatened taxa. The Mesomediterranean thermotype (600-1400 m a.s.l.) harbours 82.8% of the species, with a maximum of between 700-800 m a.s.l. and with two zones of minimums: from -100 to 0 m a.s.l. and from 3300 to 3400 m a.s.l. Key words. Floristic richness, endemic species, altitudinal distribution, life forms; conservation. RESUMEN. Diversidad y características ecológicas de la flora vascular en el Mediterráneo Occidental (Andalucía Oriental, España). Se resalta la importancia de la Flora vascular de Andalucía Oriental en el hotspot Cuenca Mediterránea; para ello se analizan los 3.726 táxones (especies y subespecies) detectados This work was undertaken within the framework of the contracts supported by Consejería de Medio Ambiente, Junta de Andalucía [Studies on Andalusian Flora, UGR 30C0200100], [Experimental design of indicators and methodology of the monitoring programme of the effects of global change in arid and semiarid zones of eastern Andalusia (Glocharid), 852/09/M/00], [Development of predictive models and a monitoring and alert system of the effects of global change on biodiversity and the functioning of the ecosystems in the SE Iberian Peninsula (Segalert), P09-RNM-5048] .

Acta Botanica Malacitana 39. 2014

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en dicha flora, que suponen más del 42 % de la flora de la Península Ibérica y el 29,9 % de la flora europea. Un 10,3 % de la flora es endémica (350 táxones), que representan el 34,3 % de los endemismos peninsulares y baleares y el 2,6 % de los de la Cuenca Mediterránea, entre los que se incluyen 6 de los 17 géneros endémicos de la Península Ibérica e Islas Baleares, 3 de ellos exclusivos del territorio analizado. Caméfitos y hemicriptófitos suponen el 45 % de la flora, con valores decrecientes para los terófitos al ascender en altitud (del 41,1 al 8,1 %) y creciente para los hemicriptófitos (del 22,4 al 63,2 %). Un 39,2 % de los táxones son mediterráneos, seguidos por los iberonorteafricanos (15,3 %), los ibéricos (13,9 %) y en quinto lugar los endemismos locales (9,4 %). El 12,5 % de los táxones están amenazados según los criterios de la UICN y de un 5,1 no se tienen datos suficientes. Tanacetum funkii, exclusivo de este área, se considera extinto. Solo 6 familias concentran el 43,5 % de los táxones amenazados. En el termotipo mesomediterráneo (600-1.400 m) se desarrollan el 82,8 % de las especies presentes, con un máximo entre los 700-800 m, y mínimos en el ámbito marino (-100-0 m) y, por el contrario, en las cotas más elevadas (3.300-3.400 m). Palabras clave. Riqueza florística, endemismos, distribución altitudinal, formas de vida, conservación.

INTRODUCTION The declaration of legally protected areas is the starting point for the conservation of biodiversity. In this sense, it is desirable to identify the areas having the greatest biodiversity (Stanley 1987). In 2011, the IUCN had 130,709 protected areas registered at the national level and 27,188 at the international (http://www.wdpa.org/Statistics.aspx), with a surface area of 24,236,478.69 km2 (some 10.9 % of the land surface of the earth and 2.2% of the water surface). Due to the needs of natural resources and cultivation by humans, it is probable that these figures will never reach values high enough to cover a significant percentage of biological diversity, and it will be necessary furthermore to evaluate whether the areas that are listed are effective at protecting the biodiversity on the world scale. With the aim of establishing conservation priorities, different organizations (World Conservation Monitoring Centre [WCMC], Birdlife International, Conservation International, etc.) have pointed out different areas that have high levels of biodiversity and endemic species and that moreover are submitted to a high degree of threat, these areas being called hotspots (Myers 1988, 1990; Primack and Ros 2002). After the work of Médail and Quézel (1997), Myers et al. (2000) listed the Mediterranean

Basin as one of the 25 biodiversity hotspots at the world level, stressing the importance of areas of the Mediterranean type in five of such areas. In a later work, Médail and Quézel (1999) underscored the importance of Andalusia in the Mediterranean Basin, together with Morocco, as part of one of the main biodiversity centres existing in the basin. With this work, the objective is to analyse and highlight the importance of vascular flora of eastern Andalusia, as it is an exceptional focal point within the hotspot of the Mediterranean Basin, in good part due to its strategic position of bridging the Iberian Peninsula and North Africa at the end of the Miocene. As well as serving as a refuge for the flora of continental Europe during the period of the glaciations (Blanca 1993) and bearing diverse substrates with nutritional imbalances for plants (volcanic, peridotitic, dolomitic, gypsiferous, saline, etc.), giving rise to in a multitude, at the world level, of territories rich in endemic taxa (Johnston 1941; Rivas Goday and Esteve Chueca 1972; Rivas Goday 1973, 1974; Esteve and Varo 1975; Rivas Goday and López González 1979; Kruckerberg 1986, 1992; Cabezudo et al. 1989; Mota et al. 1993, 2008; Cowling et al. 1995; Stevanovič et al. 2003; Reeves and Adigüzel 2004; Safford et al. 2005; Figueroa 2006; Selvi 2007; Fuente et al. 2007; Heads 2008; Lendínez et al. 2011; Mota et al. 2011; García-Barriuso et al. 2012; Pérez-Latorre et al. 2013), as well

M. Cueto et al. Vascular flora in the Western Mediterranean (Andalusia, Spain) 83

as being due to the climatic and pedological aridity (Mota et al. 2003; Ozenda 2008), climatic and orographic variety, in addition to geographical isolation and abrupt ecological gradients (Blanca 1997). MATERIAL AND METHODS Eastern Andalusia is situated in the south-east of the Iberian Peninsula, composed of the administrative provinces Almería, Granada, Jaén, and Málaga (fig. 1). This territory covers an area of 42,079 km2, of which 9301 (22.1 %) are currently protected, a figure that rose to 15,160.6 km2 (36 %) after new spaces were proposed as Sites of Community Importance (SCI). Also, 10 spaces are included on the list of RAMSAR (Salinas del Cabo de Gata; Albufera de Adra; Punta Entinas-Sabinar; Humedales y turberas de Padul; Laguna Honda; Laguna del Chinche; Laguna Grande; Laguna de Fuente de Piedra; Lagunas de Campillos; Lagunas de Archidona)

Figure 1. Geographic position of eastern Andalusia and environmental units defined in that zone (according to Blanca et al. 2009). Administrative provinces: A (Almería); J (Jaén); G (Granada) and M (Málaga). Environmental units: 1 (Aljibe); 2 (Almería); 3 (Alpujarras); 4 (Axarquía); 5 (Cazorla); 6 (Granada); 7 (Guadalquivir); 8 (Guadiana Menor); 9 (Mágina); 10 (Nevada-Filabres); 11 (Ronda); 12 (Sierra Morena); 13 (Trevenque-Almijara) and 14 (Vélez-Baza).

and 5 Biosphere Reserves (Intercontinental del Mediterráneo Andalucía (España)-Marruecos; Cabo de Gata-Níjar; Sierras de Cazorla, Segura y Las Villas; Sierra Nevada; and Sierra de las Nieves y su entorno). This is an abrupt terrain, with an average slope of 22% and an altitudinal range from 0 m on the coast to 3480 m of Mulhacén Peak in Sierra Nevada (the highest peak of the Iberian Peninsula), these extremes lying only 33.7 km from each other in a straight line. In such an extensive territory, and given the high environmental heterogeneity (altitude, lithology, soil, climate), Blanca et al. (2009) identified 14 environmental units: Aljibe, Almería, Alpujarras, Axarquía, Cazorla, Granada, Guadalquivir, Guadiana Menor, Mágina, Nevada-Filabres, Ronda, Sierra Morena, Trevenque-Almijara, and Vélez-Baza (fig. 1). The physical and climatic characteristics of these units are summarized in tables 1 and 2, respectively. The data base used was compiled during the preparation of the Flora Vascular de Andalucía Oriental (Blanca et al. 2009, 2011), resulting from the review of the herbaria located in the territory (ALME, GDA, HUAL, JAEN, and MGC), as well as in others in Spain (mainly BC, BCF, MA, MAF, and SEV) and afterwards others in different countries (COI, G, MPU, among others). Also, collections were made in areas without prior records, which are deposited in the aforementioned herbaria (initials according to the Index Herbariorum). The systematic treatment and nomenclature followed is the same as in Blanca et al. (2009, 2011). The chorological elements used were those proposed by Takhtajan (1986), with modifications used by Blanca et al. (2009) to enable comparisons and groupings of the floristic catalogue. The life forms correspond to those of Raunkiaer (1934): phanerophytes, chamaephytes, hemicryptophytes, geophytes, therophytes, helophytes, and hydrophytes.

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Figure 2. Number of genus and species in the 10 families with the greatest representation of the vascular flora of eastern Andalusia. Unit

Surface area (km2)

Range

Altitude (m) Average

Average slope (%)

Predominant Substrates

Aljibe

641.5

0-1120

346.1

26.4

Siliceous

Almería

4533.3

0-1380

377.9

16.3

Basic

Alpujarras

2060.8

0-2240

793.2

34.3

Basic and Siliceous

Axarquía

2046.3

0-1217

313.8

26.1

Siliceous

Cazorla

3040.3

367-2380

1225.6

33.2

Basic

Granada

3375.5

360-1660

813.8

18

Basic

Guadalquivir

7074.1

13-1920

495.8

11.5

Basic

Guadiana Menor

3469.5

410-1731

913.7

13.9

Basic

Mágina

2313.9

451-2160

1085.3

23.7

Basic

Nevada-Filabres

2683.1

220-3480

1613.1

34.4

Siliceous

Ronda

3195.3

0-1900

714.1

30.1

Basic

Sierra Morena

3129.9

160-1280

599.3

21

Siliceous

Trevenque-Almijara

1521.9

0-2220

1015.5

36.4

Basic

Vélez-Baza

3111.7

479-2260

1126.6

18.4

Basic

Table 1. Physical characteristics of the environmental units in eastern Andalusia.

M. Cueto et al. Vascular flora in the Western Mediterranean (Andalusia, Spain)

Unit

Temperature (º C)

85

Rainfall (mm)

Range

Average

Range

Average

Aljibe

13.1-18.2

16.6

595.1-1760.3

959.1

Almería

12.7-20.3

17.6

152.8-413.1

266.7

Alpujarras

9.3-19.2

15.5

176.1-680.2

407.2

Axarquía

12.8-19.4

16.8

410.1-1009

548.4

Cazorla

7.8-18.1

12.8

289.4-1145.8

725.6

Granada

10.6-17.6

14.6

326.4-802.9

502.5

Guadalquivir

9.5-19.2

16.7

393.5-859.2

511.1

Guadiana Menor

10.8-17.6

14.3

270.9-718.5

361

9-18.1

13.8

335-802.9

543.9

Nevada-Filabres

4.6-18.8

11.7

211.6-746.1

439.5

Ronda

8.1-19.1

14.9

347.3-1360.5

710.4

Sierra Morena

12.6-19

16

443.9-832.2

558.8

Trevenque-Almijara

9.2-18.4

14

375.5-1003

590.3

7.6-17

13.7

212.1-510.8

351.9

Mágina

Vélez-Baza

Table 2. Climatic characteristics of the environmental units in eastern Andalusia (prepared from the data of Rediam, 2013)



Families

Genera

Taxa



%



%



%

Pteridophytes

20

12.1

28

2.8

63

1.7

Gymnosperms

5

3

14

1.4

36

1

Liliidae

31

18.8

208

21.2

677

18.2

Ranunculiidae

104

63

727

74

2941

78.9

Magnoliidae

4

2.4

4

0.4

8

0.2

Ceratophylliidae

1

0.6

1

0.1

1

0.02

140

84.8

940

95.7

3627

97.3

Angiosperms

Total Angiosperms Total Vascular Flora

165

982

3726

Table 3. Representation of the large groups of vascular plants in the vascular flora of eastern Andalusia.

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In the analysis of the state of conservation of plants, the reference works used were Blanca et al. (1999, 2000), Cabezudo et al. (2005), Bañares et al. (2004, 2007, 2009, 2011), and Moreno (2008), using the threat categories of the IUCN (2001). The belts of vegetation used in the altitudinal distribution correspond to the thermotypes of Rivas Martínez (2007): Thermomediterranean, Mesomediterranean, Supramediterranean, Oromediterranean, and Cryoromediterranean. RESULTS AND DISCUSSION Taxonomical composition The vascular flora of eastern Andalusia is composed of 165 families, including 982 genera and 3462 species, or 3726 taxa (species + subspecies). In relation to other figures provided for the entire Iberian Peninsula by Aedo et al. (2013), the flora studied here represents more than 42% of the peninsular flora on a surface area of less than 10% of the overall peninsula, and 29.9% of European flora. Table 3 lists each of the large plant groups studied. The distribution of species in large taxonomical groups is similar to that of overall Spanish flora: pteridophytes, 1.7% vs. 1.9% for Spain; gymnosperms 1% vs. 0.6% for Spain; angiosperms 97.8% vs. 97.6% for Spain. The majority group, angiosperms, represents 97.3% of the flora. Notable are the Ranunculiidae, with 78.9%. In eastern Andalusia, the families with the greatest representation are Asteraceae (126 genera, 415 species), Fabaceae (48 genera, 330 species), and Poaceae (115 genera, 301 species); also significant are the families Caryophyllaceae (31 genera, 174 species), Brassicaceae (65 genera, 171 species), Lamiaceae (33 genera, 159 species), and Apiaceae (62 genera, 137 species). The 10 families richest in species of eastern Andalusian flora constitute 54.2% of the genera and 55.8% of the species present in

the area (fig. 2). The genera with the greatest representation are Centaurea (48 species/18 locally endemic), Silene (45/3), Teucrium (40/14), Trifolium (39/0), Carex (33/2), Ononis (33/0), Vicia (32/1), Galium (32/8), Ranunculus (31/3), and Astragalus (30/3). The endemic flora of eastern Andalusia is made up of 350 taxa, i.e. 10.3% of the total, representing 34.3% of the total endemic taxa of the Iberian Peninsula and Balearic Islands and 2.6% of the Mediterranean Basin. Of the 12 richest areas in endemic taxa of the Iberian Peninsula and Balearic Islands identified by Moreno et al. (2013), 4 are found in eastern Andalusia (Subbetic System; Málaga and Granada; Sierras of Grazalema and Ronda; and Sierra Nevada). The families with the highest number of endemic taxa (fig. 3) are Asteraceae (60 taxa, 17.1%), Brassicaceae (34 taxa, 9.7%), and Lamiaceae (32 taxa, 9.1%). Of the 17 endemic genera of the Iberian Peninsula and Balearic Islands (Aedo et al. 2013), 6 are present in eastern Andalusia, with 3 being exclusive of this area (Castrilanthemum, Cazorla unit; Euzomodendron, Almería unit, and Rothmaleria, Trevenque-Almijara unit). Life forms The analysis of the distribution of taxa according to life forms (fig. 4) reflect a predominance of therophytes (1283; 33%), followed by hemicryptophytes (1163; 30%), chamaephytes (594; 15%), phanerophytes (396; 10%), geophytes (347; 9%), hydrophytes (55; 2%), and helophytes (33; 1%). In table 4 appears the proportion of taxa according to the life forms by thermoclimatic belts. The results of the distribution of life forms according to altitude coincide with the trends expressed by other authors (e.g. Voliotis 1982; Cueto et al. 1991). The proportion of therophytes reaches its maximum in the Thermomediterranean (41.1%), close to the value of 56% indicated by Braun-Blanquet

M. Cueto et al. Vascular flora in the Western Mediterranean (Andalusia, Spain) 87

(1979) for the El Golea desert in the Sahara and far higher than the 15% cited by Cowling et al. (1996) for local flora of all the regions with a Mediterranean climate, progressively diminishing with altitude (with a lowering of temperature and dryness). Meanwhile, the hemicryptophytes increase with altitude until reaching their maximum in the Cryoromediterranean (63.2%), a value similar to the 68% indicated by Braun-Blanquet (l.c.) for the zone between 2000-3000 m of the Alps. Chorological spectrum The predominant floristic element in peninsular Spain is Mediterranean in the broad sense, followed by the Euro-Siberian and, third, endemic elements (Moreno 2012). In eastern Andalusia (fig. 5) the Mediterranean taxa also predominate (1461; 39.2%), followed by Iberian-North African (569; 15.3%), Iberian (453; 139%), European (423; 11.3%), and in fifth place of endemic taxa of the study area, eastern Andalusia (350; 9.4%). One of the main causes of the uniqueness of the flora of this territory is due above all to the percentage of Iberian-North African species, the floristic element that, on the contrary, represents a minority group for the overall Iberian Peninsula. The Iberian-North African elements penetrate the south and east and differentiate beginning from a xerophytic and heterogeneous stock of ancestral plants, resulting in numerous endemic taxa having affinities with Saharan-Arab taxa (Thompson 2005). Some of the genera of the most genuinely Mediterranean flora, and which have a diversification centre in the Iberian Peninsula, are: Genista, Narcissus, Linaria, Helianthemum, Thymus, Teucrium, etc. (Thompson l.c.). In eastern Andalusia, all these genera are represented by more than 10 species, with the genus Teucrium reaching 40 species. These are autochthonous taxa that developed and differentiated during the Tertiary

due to isolation processes of the microplates and climatic change during this period (Zohary 1973). The number of taxa present in the environmental units (fig. 6) ranges from a maximum of 1894 taxa (50.8%) in the Ronda unit to a minimum of 1129 taxa (30.3%) in the Guadiana Menor. It was noted that the units with the greatest number of taxa are those having substrates that are predominantly basic (Ronda, Trevenque-Almijara, Cazorla, Vélez-Baza, etc.), with the exception of Nevada-Filabres, in fourth place, on siliceous substrates, followed by the Alpujarras unit, where basic and siliceous substrates coexist. The singularity of each environmental unit can be evaluated by the presence of elements that differ from the others. The data compiled support the territorial division established on the basis of environmental characteristics and expert knowledge. Notable are the units Nevada-Filabres, Almería, and Cazorla, with 181 differential elements (72 local endemic taxa), 128 (28), and 126 (27), respectively (tab. 5). The Sierra Nevada, from the geological standpoint, including the central siliceous core (within the Nevada-Filabres unit) and the north-western calcareous part (within the Trevenque-Almijara unit) is the most important centre of plant diversity in the western Mediterranean region (Molero-Mesa and Pérez Raya 1987; Blanca 1996; Heywood 1996; Blanca et al. 1998; Fernández 2012). The data reveal its floristic importance, since, of the 350 endemic taxa of eastern Andalusia, 174 belong to Sierra Nevada, 90 being exclusive of this massif. On the other hand, 215 taxa of eastern Andalusian flora are exclusive to Sierra Nevada, some being: Alpine, such as Artemisia umbelliformis Lam. and Gentiana alpina Vill.; Arctic-Alpine, such as Ranunculus glacialis L. and Saxifraga oppositifolia L. subsp. oppositifolia; Circumboreal, such as Silene rupestris L. and

88

Acta Botanica Malacitana 39. 2014

Figure 3. Number of local endemic taxa and number of genera with local endemic taxa by family of the vascular flora of eastern Andalusia.

Figure 4. Distribution of the vascular flora of eastern Andalusia according to the life forms of Raunkiaer (1934).  

M. Cueto et al. Vascular flora in the Western Mediterranean (Andalusia, Spain) 89

Figure 5. Chorological spectrum of the vascular flora of eastern Andalusia.

Figure 6. Number of taxa of the vascular flora of eastern Andalusia in each environmental unit of the territory studied.

90

Sibbaldia procumbens L.; Holarctic, such as Asplenium septentrionale (L.) Hoffm. subsp. septentrionale and Gymnocarpium robertianum (Hoffm.) Newman; or Iberian-North African, such as Murbeckiella boryi (Boiss.) Rothm., Epilobium atlanticum Litard. & Maire, and Saxifraga trabutiana Engl. & Irmsch. State of conservation Figure 7 summarizes the state of conservation of the flora analysed. Of the taxa evaluated (2600), 69.8% are included in the category LC (less concern), 10.4% (389) in NT (nearly threatened), 7.7% (286) in VU (vulnerable), 2.7% (102) in EN (endangered), 2% (76) in CR (critically endangered), 0.2% (8) EX (extinct), and 5.2% (192) in DD (deficient data). From the figures, it can be deduced that 1.25% of the taxa (464) are threatened (VU, EN, and CR), while 5.2% do not have sufficient data, which at least in part could increase the number of threatened taxa. In Spain, 25 taxa are considered extinct (Aedo et al. 2012); of these16 survive in the wild of other countries; 4 more are conserved ex situ; and 5 are known only from herbarium specimens: Carduncellus matritensis, Kunkeliella psilotoclada, Normania nava, Pharbitis preauxii, and Tanacetum funkii (Moreno 2008). Of this latter, only the original sheet collected on Sierra Nevada (Granada) is known, and thus can be considered the only Andalusian species extinct in the strict sense, although some authors have pointed out that doubts remain on whether it is an independent species or simply a deviant form of another species (Aedo et al. l.c.). According to the Spanish Red List (Moreno 2008), Andalusia is the second territory with the greatest number of threatened taxa (227) after the Canary Islands, with 247 taxa. In particular, the Betic Sierras (especially the province of Granada) appear as a critical area for the conservation of flora (Moreno 2008). The five families with the greatest presence

Acta Botanica Malacitana 39. 2014

on the Spanish Red List are: Asteraceae (279 taxa), Plumbaginaceae (111), Lamiaceae (109), Fabaceae (108), and Brassicaceae (96). In terms of genera, these are Limonium (84 species and subspecies), Sideritis (36), Centaurea (31), Argyranthemum (27), and Armeria (27). In eastern Andalusia, the families with the most threatened taxa being (tab. 6): Asteraceae (72), Brassicaceae (37), and Fabaceae (35). Distribution by altitude By altitudinal distribution, the taxa follow a normal distribution (fig. 8): a maximum in the band at 700 to 800 m, harbouring 2.252 taxa; and minimums from 0 to -100 m with 4 taxa, restricted to the marine ambit, and from 3.300 to 3.400 m on land, 32 taxa. The Mesomediterranean and Thermomediterranean thermotypes occupy 93.2% of the territory, followed by the Supramediterranean (5.7%), Oromediterranean (1%) and Cryoromediterranean (0,1%) (Table 7). However, although the proportion of taxa present in the thermotypes with the greatest surface area (Meso- and Thermomediterranean) is 87.1%, in the upper 3 thermotypes (with a surface area of only 6.8%), the proportion is 49.4%, indicating that the thermotypes with the smallest surface area nevertheless have a substantial number of taxa. In comparison with the latitudinal gradient, the altitudinal one implies a sharp variation in environmental conditions over short distances (Odland 2009), characterized generally by a fall in temperatures and evapotranspiration, a rise in moisture and solar radiation, and a decline in competition (e.g. Barry 1992; Körner 1999; Löffler 2002). The variation in species richness over an altitudinal gradient has two main models: a regular decline in richness with an ascent in altitude, and a normal distribution with the highest value of species richness near the middle of the gradient. The two options have been related to different environmental

M. Cueto et al. Vascular flora in the Western Mediterranean (Andalusia, Spain) 91

 

Figure 7. Percentages of taxa of the vascular flora of eastern Andalusia according to the categories of the IUCN (2001). NE: not evaluated; DD: deficient data; LC: least concern; NT: nearly threatened; VU: vulnerable; EN: endangered; CR: critically threatened; EX: extinct.

Figure 8. Number of taxa within the vascular flora of eastern Andalusia according to altitude (in bands of 100 m).

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Thermotypes

G

T

Hyd

Hel

Hemi

C

Ph

Thermomediterranean

8.7

41.1

1.6

0.9

22.4

14.2

11.1

Mesomediterranean

9.9

34.9

1.2

0.9

28.6

14.2

10.3

Supramediterranean

9.1

28

0.8

0.5

36.6

17

8

Oromediterranean

8

16

0.5

0.7

48.8

20.3

5.6

Cryoromediterranean

7

8.1

1.6

0.6

63.2

18.4

1.1

Table 4. Distribution, by thermotypes, of the plant taxa (in %) according to life forms. G: geophytes; T: therophytes; Hyd: hydrophytes; Hel: helophytes; Hemi: hemicryptophytes; C: chamaephytes; Ph: phanerophytes

Unit

Nº Families

Nº Genera

Nº Taxa

End. unit

End. Eastern Andalusia

Aljibe

22

48

51

1

9

Almería

41

92

128

28

44

Alpujarras

19

26

27

8

59

Axarquía

11

16

16

1

20

Cazorla

36

96

126

27

88

Granada

9

12

13

1

17

Guadalquivir

13

20

24

0

3

Guadiana Menor

16

26

29

5

21

Mágina

9

12

13

4

49

Nevada-Filabres

45

111

181

72

110

Ronda

25

47

65

32

77

Sierra Morena

29

53

70

4

4

Trevenque-Almijara

23

35

47

30

100

Vélez-Baza

15

23

24

6

61

Table 5. Differential elements (exclusively present in a unit) in the environmental units established for the flora of eastern Andalusia. Nº of Families: number of families to which the differential elements belong; Nº Genera: number of genera to which the differential elements belong; Nº of Taxa: number of differential taxa (species and subspecies) of each unit; End. Unit: number of exclusive endemic taxa in each unit; End. Eastern Andalusia: number of endemic taxa of the vascular flora of eastern Andalusia present in each unit.

M. Cueto et al. Vascular flora in the Western Mediterranean (Andalusia, Spain) 93

 

Asteraceae Brassicaceae Fabaceae Poaceae Caryophyllaceae Veronicaceae Apiaceae Lamiaceae Plumbaginaceae Ranunculaceae Orobanchaceae Rubiaceae Geraniaceae Cistaceae Rosaceae Saxifragaceae Amaryllidaceae Campanulaceae Fumariaceae Gentianaceae Zannichelliaceae Boraginaceae Chenopodiaceae Lentibulariaceae

VU 43 26 18 14 13 13 9 11 4 8 6 7 6 5 3 5 4 4 4 4 6 3 2 5

EN 16 6 7 5 4 2 3 3 9 3 3 4 0 1 2 1 2 2 2 1 0 1 0 0

CR 13 5 10 1 2 4 4 2 3 2 3 1 2 1 2 1 0 0 0 1 0 1 3 0

Threatened 72 37 35 20 19 19 16 16 16 13 12 12 8 7 7 7 6 6 6 6 6 5 5 5

Table 6. Families with more than 5 taxa in the threatened category. VU: Vulnerable; EN: Endangered; CR: Critically threatened; Threatened: sum of VU, EN, and CR. Thermotypes

Altitude (m)

Area

Taxa

Surface (km2) and (%)

Number and (%)

Taxa/km2

<600

19586.5 (46.4)

2411 (69.6)

0.12

Mesomediterranean

600–1400

19729.1 (46.8)

2866 (82.8)

0.14

Supramediterranean

1400–2000

2400.4 (5.7)

1735 (50.1)

0.72

Oromediterranean

2000–2800

438.5 (1)

583 (16.8)

1.32

Thermomediterranean

Cryoromediterranean

>2800

42.5 (0.1)

180 (5.2)

4.23

Thermo and Mesomediterranean

0-1400

39315.6 (93.2)

3245 (87.1)

0.08

Supra, Oro and Cryoromediterranean

>1400

2881.4 (6.8)

1841 (49.4)

0.64

Table 7. Altitudinal range, surface area, and percentage of the total of eastern Andalusia, number of plant species, and percentage of the total of eastern Andalusia, and number of species per area for each thermotype.

Acta Botanica Malacitana 39. 2014

94

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