The mycobiota - Universidad de Granada

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Nova Hedwigia

83

1—2

233—247

Stuttgart, August 2006

The mycobiota (Agaricales, Boletales, Russulales) from Andalusia (southern Spain): chorological and biogeographical analysis by

A. Ortega(1)* and F.B. Navarro(2) (1)

Dpto. de Botánica. Facultad de Ciencias. Universidad de Granada. C/Severo Ochoa s/no. 18071 Granada (Spain). E-mail: [email protected]

(2) Dpto. Forestal. Área de Recursos Naturales. Centro de Investigación y Formación Agraria. Instituto Andaluz de Investigación y Formación Agraria, Pesquera, Alimentaria y de la Producción Ecológica (CICE, Junta de Andalucía). Camino de Purchil s/no. Aptdo. 2027. 18080 Granada (Spain). E-mail: [email protected]

With 4 figures and 3 tables

Ortega, A. & F.B. Navarro (2006): The mycobiota (Agaricales, Boletales, Russulales) from Andalusia (southern Spain): chorological and biogeographical analysis. - Nova Hedwigia 83: 233-247. Abstract: This study analyses the Andalusian mycobiota (Agaricales, Boletales, Russulales) from a biogeographical standpoint. For this, the best-represented groups of macrofungi have been selected, including ectomycorrhizal and saprotrophic species, to discover whether their chorological behaviour is different or similar. Locations in a number of important chorological sectors in Andalusia have been sampled, namely the Aljíbico, Gaditano-Onubense, Guadiciano-Bacense, Malacitano-Almijarense, Mariánico-Monchiquense, Nevadense, Rondeño and Subbético sectors. A broad, current bibliography on Andalusian mycoflora has been used, as well as material on deposit in the GDA-GDAC (University of Granada) herbarium. Of all the references reviewed in this bibliography, those selected cover 15 genera in the orders of Agaricales, Boletales and Russulales. Correspondence Analyses (CA) have been made to establish the relationships between the different chorological sectors. The chorological sectors Aljíbico and Mariánico-Monchiquense have a greater representation of mycorrhizal than saprotrophic taxa, and this situation is the opposite in the Rondeño, Nevadense, Guadiciano-Bacense, Subbético and Malacitano-Almijarense sectors. The Gaditano-Onubense chorological sector is clearly differentiated from the rest. The Aljíbico and Mariánico-Monchiquense chorological sectors are closely related. With regard to the other areas, there is a certain relation between the Rondeño, Guadiciano-Bacense and Malacitano-Almijarense sectors. The chorological sector Nevadense is differentiated from the other eastern Andalusian sectors. The macrofungi species of any one area or region constitute a good parameter to perform biogeographical studies. The ectomycorrhizal species respond, due to their distribution, above all to the biogeographical characteristics of a region, whilst the saprophytic species respond to the local pluviometric regime. Key words: Macrofungi, biogeography, chorological sectors, CA.

*Corresponding author: [email protected] DOI: 10.1127/0029-5035/2006/0083-0233

0029-5035/06/0083-0233 $ 3.75 © 2006 J. Cramer in der Gebrüder Borntraeger Verlagsbuchhandlung, D-14129 Berlin · D-70176 Stuttgart

233

Fig. 1. Geopolitical map of Andalusia (Spain).

Introduction Andalusia, located the southern Iberian Peninsula (Fig. 1), occupies an 87,597 km² and, geographically, is characterised by its diversity, which provides ecological wealth expressed by extraordinary floral and mycological variability, revealing this territory as a hot spot of biodiversity (Akeroyd & Heywood, 1994; Comes, 2004). In terms of mycoflora, our data (Table 1), 1955 macrofungi species constitute the current Andalusian mycological catalogue, of which 1003 belong to the Agaricales, 64 to the Boletales and 118 to the Russulales. This information has been compiled from an abundant bibliography used as a starting point to examine (1) the diverse taxonomic groups comprising the macrofungi of the region of Andalusia (e.g. Ortega & Galán, 1981; Moreno et al., 1984; Ortega & Aguilera, 1987; Ortega & Gª Buendía, 1989; Ortega & Vizoso, 1991; Ortega, 1992; Ortega, 2000; Ortega & Esteve-Raventós, 1996; Calonge, 1997; Esteve-Raventós, 1999; Moreno-Arroyo et al., 1999; Ortega & Tellería, 2000; Dueñas et al., 2001), (2) the mycological component of the flora in some of the most representative nature parks of Andalusia (e.g. Calonge & Tellería, 1980; Gómez et al., 1993, 1995, 1999; Moreno et al., 1994; Ortega, 1994; Ortega et al., 1986, 1994, 1996, 1997a, 1997b, 2002; Romero de la Osa, 1991, 1992, 1993, 1996, 2002), (3) the mycological flora in the provinces of Córdoba (Martínez Macarro, 1996), Jaén (Jiménez Antonio, 1994; Jiménez Antonio & Reyes, 1999) and Málaga (Jiménez Gross & Ayala, 1992) and (4) a recent compilation of Andalusian macrofungi published by Moreno-Arroyo (2004). Remarkable biogeographical variability also typifies Andalusia (Fig. 2), given that, according to Rivas-Martínez (1987), Rivas-Martínez et al. (1997) and Valle et al. 234

Table 1. Distribution (no. species) by taxonomic groups and geopolitical provinces of the Andalusian Mycobiotaa GROUP

AL

CA

CO

GR

HU

JA

MA

SE TOTAL

Pyrenomycetes Leotiales Pezizales Heterobasidiomycetidae Aphyllophorales Agaricales Boletales Russulales Gasteromycetes

n.c. 2 11 n.c. 45 52 4 2 34

20 21 33 14 97 276 21 61 40

4 10 61 8 68 265 19 40 79

5 16 94 14 179 556 33 34 58

11 12 50 8 120 417 51 69 55

10 9 67 19 196 503 34 48 59

19 14 20 5 108 263 10 14 45

n.c. n.c. 9 n.c. 15 135 5 24 28

53 54 157 31 354 1003 64 118 121

TOTAL

150

583

554

989

793

945

498

216

1955

a AL: Almería, CA: Cádiz, CO: Córdoba, GR: Granada, HU: Huelva, JA: Jaén, MA: Málaga, SE: Sevilla. n.c. = not collected.

(2003, 2004), five chorological provinces and 13 biogeographical sectors can be differentiated, the fundamental characteristics of which are summarized in Table 2. Regarding the chorological aspects of Andalusian mycobiota, Ortega & Linares (2003) established a distribution by geopolitical provinces of 1876 macrofungi, while also delineating an ecological distribution of these taxa and comparing the mycoflora of western and eastern Andalusia. Similarly, Moreno-Arroyo et al. (2005: 163) reported new data, increasing the number of Andalusian macrofungi to 2343 taxa, which are distributed by geopolitical provinces.

Fig. 2. Biogeographical map of Andalusia (Spain) taken from Rivas Martínez et al. (1997). 1: Mariánico-Monchiquense Sector, 2: Gaditano-Onubense Sector, 3: Aljíbico Sector, 4: Algarviense Sector, 5: Hispalense Sector, 6: Rondeño Sector, 7: Malacitano-Almijarense Sector, 8: Subbético Sector, 9: Alpujarreño-Gadorense Sector, 10: Nevadense Sector, 11: Guadiciano-Bacense Sector, 12: Murciano-Almeriense Province, 13: Manchego Sector.

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236

MurcianoAlmeriense CastellanoMaestrazgoManchega

LusoExtremadurense Gaditano-OnuboAlgarviense

Bética

Provinces

0 2 0 3 1

Manchego (MN)

2

C

S S S Y-C

S

C C-S

S

2 2 2

C-Y

4

Hispalense (HI) AlpujarreñoGadorense (ALP) MariánicoMonchiquense (MM) Aljíbico (ALJ) Gaditano-Onubense (GO) Algarviense (ALG) (ALM)

C

2

MalacitanoAlmijarense (MA) GuadicianoBacense (GU) Nevadense (NE)

C C

Sustrata

2 3

no. distrits

Subbético (SU) Rondeño (RO)

Sectors

Table 2. Biogeographical characteristics of Andalusiab

M, (S)

T, (M) T T T, M

T, M, (S)

T, M T, M, (S)

M, S, O, C

M, S, O

T, M, S, (O)

M, S, O T, M, S, (O)

Thermotypes

PcQr, BhQr, RlQr, DoPs, RmJp (DlAg, AdQr, JpPs) AdQr, AdQp, GbJn, EfFc

PcQr, BhQr, DlAg, DoPs, JpPs, JpPh PcQr, BhQr, SmQr (DlAg, DoPs, PbAp, BmAp) SmQr, PcQr, BhQr, RmJp, DlAg, DoPs

Principal series of vegetation

(Sem), Sec

Su, Hu Sec, Su Sec, Su Sem, (Sec)

TbQs, McQs (RhQc, TcOs, Psa) OQs, Psa OQs, Psa MePa, Zl, MZl, BgPl, ChRl, Mt, Yc, Yt, Ys (PcQr, AdQm, RlQr) BrQr, RlJp

Sec, Su, Hu, (Hi) Sec, Su SmQr, PcQr, TcOs Sem, Sec, (Su) PcQr, BhQr, SmQr, AdQr, BgPl (DlAg, DoPs, McQs) Sec, Su, (Hu) McQr, PbQr, McQs, SaQs (AuQp, SaQp)

Sec, Su, Hu Sec, Su, Hu, (Hi) (Sem), Sec, Su, (Hu) Sem,Sec,(Su)

Ombrotypes

Explanations Table 2: bSubstrata (predominant): C = carbonated, S = siliceous, Y = gypsum. Thermotypes: T = thermomediterranean, M = mesomediterranean, S = supramediterranean, O = oromediterranean, C = crioromediterranean. Ombrotypes: Sem = semiarid, Sec = dry, Su = subhumid, Hu = humid, Hi = hyperhumid. Series of vegetation: Ad-Qp: Adenocarpo decorticantis-Querceto pyrenaicae S., Ad-Qr: Adenocarpo decorticantis-Querceto rotundifoliae S., AuQp: Arbuto unedonisQuerceto pyrenaicae S., BgPl: Bupleuro gibraltarici-Pistacieto lentisci S., BhQr: Berberido hispanicae-Querceto rotundifoliae S., BmAp: Bunio macucae-Abieteto pinsapo S., BrQr: Bupleuro rigidi-Querceto rotundifoliae S., ChRl: Chamaeropo humilis-Rhamneto lycioidis S., DlAg: Daphno latifoliae-Acereto granatensis S., DoPs: Daphno oleoidis-Pineto sylvestris S., EfFc: Erigeronto frigidi-Festuceto clementei S., GbJn: Genisto baeticae-Junipereto nanae S., McQr: Myrto communisQuerceto rotundifoliae S., McQs: Myrto communis-Querceto suberis S., MePa: Mayteno europaeiPeriploceto angustifoliae S., MZl: Mayteno europaei-Zizipheto loti S., OQs: Oleo-Querceto suberis S., PbAp: Paeonio broteroi-Abieteto pinsapo S., PbQr: Pyro bourgaeanae-Querceto rotundifoliae S., PcQr: Paeonio coriaceae-Querceto rotundifoliae S., RhQc: Rusco hypophylli-Querceto canariensis S., RlQc: Rhamno lycioidis-Querceto cocciferae S., RlQr: Rubio longifoliae-Querceto rotundifoliae S., SaQs: Sanguisorbo agrimonioidis-Querceto suberis S., SmQr: Smilaco mauritanicae-Querceto rotundifoliae S., TbQs: Teucrio baetici-Querceto suberis S., Zl: Zizipheto loti S., JpPh: Junipero phoeniceae-Pineto halepensis S., JpPs: Junipero phoeniceae-Pineto salzmannii S., Mt: Broom formation of edaphoxileropholous tabernense vegetation upon subsaline loam, Psa: Geoseries edaphoxileropholous coastal psammophilous thermomediterranean mediterranean-iberoatlantic, RlJp: Rhamno lycioidis-Junipereto phoeniceae S., RmJp: Rhamno myrtifoliae-Junipereto phoeniceae S., TcOs: Tamo communis-Oleeto sylvestris S., Yc, Ys, Yt: edaphoxileropholous vegetation formations on gypsum. The date within parenthesis indicates puntual presence. Data obtained from Rivas Martínez et al. (1997) and Valle et al. (2003, 2004).

Finally, with reference to the chapter on the ecology of the Andalusian macrofungi, we should point out that currently, general data is available on the biological behaviour of numerous species (Ortega & Linares, 2003). The group Aphyllophorales sensu lato lignicoles has been studied in greater depth (Ortega & Lorite, 2000a, b; Ortega & Navarro, 2004), whereas only one study has been made on Agaricales sensu lato (Ortega & Linares, 2002). This latter study focuses on the Pinus ssp. forests in the province of Granada, while another study treats the phenological behaviour of the macrofungi in the evergreen oak forests of Granada province (Galán et al., 1983). The above facts highlight the value of these kind of studies, and the present work in particular establishes a distribution, by chorological sectors, of an important group of macrofungal species belonging to the Agaricales, Boletales and Russulales, while analysing the current relationship (similarities and differences) between the chorological sectors represented in Andalusia, based on the presence or absence of a series of selected taxa which belong to the genera Agaricus, Amanita, Aureoboletus, Clitocybe, Cortinarius, Entoloma, Inocybe, Mycena (incl. Roridella), Psathyrella, Tricholoma, Boletus, Suillus, Xerocomus, Lactarius and Russula. The distributions of the saprotrophic and mycorrhizal species are also analysed in the chorological sectors studied. Materials and methods To study the biogeographical (chorological) distribution of the Andalusian mycobiota (macrofungi), we used the database compiled by Ortega (unpublished) as a starting point, and selected (Table 1) 606 taxa belonging to 15 genera, and to the Agaricales, Boletales and Russulales, which are best

237

represented of Andalusian mycoflora (see Appendix S1). Of these taxa, 457 are included in Agaricus, Amanita, Clitocybe, Cortinarius, Entoloma, Inocybe, Mycena, Psathyrella and Tricholoma (which represent 45% of the Andalusian species belonging to the Agaricales), 40 taxa belonging to the Aureoboletus, Boletus, Suillus and Xerocomus genera (which represent 64% of the Andalusian species belonging to the Boletales) and 109 taxa belonging to Lactarius and Russula (which represent 93% of the Andalusian species of the Russulales). In Table 3, compiled from this data, the saprophytic and mycorrhizal species have been separated in order to establish the distribution of each of these biological groups. Only the chorological sectors in which more than 60 species belonging to the chosen groups (10% of the species) were considered. After the length of gradient was tested by DCA (Detrended Correspondence Analysis), a unimodal method (Correspondence Analyses, CA) was also used (see Ter Braak, 1985; Ter Braak & Šmilauer, 1998) with the computer programme CANOCO 4.0 and STATISTICA 6.0. The aim was to determine similarities and differences between the chorological sectors based on their mycological flora (Figs. 3 and 4). The database (Ortega, unpublished) was compiled (Table 1) using abundant bibliography on Andalusian mycology, as well as material on deposit in the Herbarium of the University of Granada (GDAGDAC). In classifying the chorological sectors, we followed the work of Rivas-Martínez et al. (1987), RivasMartínez et al. (1997) and Valle et al. (2003, 2004).

Results 1. Mycofloristic aspects: Table 3 illustrates the total absolute number and the percentage (relative number) of taxa in each of the selected genera for each chorological sector analysed. It also includes the total number of saprotrophic (185) and ectomycorrhizal (421) species, in addition to the absolute and relative number of saprotrophic and mycorrhizal species in each chorological sector. The following data are noteworthy: No relative data is available for the chorological sectors Hispalense, AlpujarreñoGadorense, Algarviense, or Manchego, and there is a paucity of data on the MurcianoAlmeriense province, and therefore none of these chorological units will be used for our analysis. As shown in Table 3, the relationship between the saprophytic and mycorrhizal species (the quotient between the number of saprotrophic and mycorrhizal species) is less than 1 in two of the chorological sectors studied, Aljíbico (0.79) and Mariánico-Monchiquense (0.96), revealing the predominance of the mycorrhizal over saprophytic taxa. This is because of the forest formation in two chorological sectors of western Andalusia, mainly Quercus suber forests, which are richer in mycorrhizal species (56.2%; unpublished data) than is the forest consisting mainly of Quercus ilex subsp. ballota, which has 38% percentage of mycorrhizal species (unpublished data) and which grows in the chorological sectors preferably located in eastern Andalusia: Nevadense, Rondeño, Guadiciano-Bacense, Subbético and Malacitano-Almijarense, (where the ratios between the saprophytic and mycorrhizal species are 1.79, 1.78, 1.48, 1.14 and 1.13, respectively). The chorological sector of Gaditano-Onubense, despite its location in western Andalusia, has a ratio of 1.09, which denotes an equilibrium between saprophytic and mycorrhizal species, given that the psammophilous communities are quite restrictive for the growth of ectomycorrhizal species. 238

Table 3. Relative number of species of each one of the selected genera in each chorological sector in Andalusia and relative number of saprotrophic and mycorrhizical species by biogeographical sectorsc GENERA

RO

MA

SU

NE

GU

ALJ

GO ALG MM ALM Tot. sp.

Agaricus Clitocybe Entoloma Mycena Psathyrella Absolute number Relative number

20.0 53.1 14.6 54.7 44.8 71 38.4

53.3 53.1 39.0 58.5 31.0 89 48.1

83.3 59.4 48.8 62.3 48.3 111 60.0

30.0 53.1 65.9 45.3 44.8 90 48.6

13.3 37.5 7.3 11.3 10.3 28 15.1

23.3 46.9 31.7 49.1 20.7 67 36.2

30.0 6.3 9.8 7.5 13.8 23 12.4

6.7 n.c. n.c. n.c. n.c. 2 1.1

80.0 30.0 90.6 3.1 65.9 2.4 64.2 1.9 48.3 6.9 128 14 69.2 7.6

30 32 41 53 29 185 100%

Amanita Cortinarius Inocybe Tricholoma Boletus Aureoboletus Suillus Xerocomus Lactarius Russula Absolute number Relative number TOTAL Absolute number Relative number

16.3 10.6 31.9 47.7 31.8 100 71.4 30.0 23.5 5.3 91 21.6

25.6 61.1 51.4 54.5 31.8 n.c. 71.4 40.0 38.2 12.0 179 42.5

51.2 51.3 63.9 45.5 45.5 n.c. 100 30.0 52.9 49.3 221 52.5

20.9 31.9 22.2 38.6 31.8 100 71.4 30.0 29.4 13.3 114 27.1

4.7 5.3 19.4 25.0 n.c. n.c. 42.9 n.c. 14.7 2.7 43 10.2

51.2 31.0 44.4 38.6 45.5 100 42.9 60.0 55.9 62.7 192 45.6

37.2 2.7 18.1 6.8 n.c. n.c. 28.6 10 5.9 10.7 48 11.4

n.c. n.c. n.c. n.c. n.c. n.c. n.c. n.c. n.c. n.c. n.c. n.c.

90.7 n.c. 56.6 n.c. 54.2 n.c. 86.4 4.5 90.9 n.c. 100 n.c. 71.4 28.6 100 n.c. 85.3 2.9 76.0 n.c. 302 5 71.7 1.2

43 113 72 44 22 1 7 10 34 75 421 100%

162 26.7

268 44.2

332 54.8

204 33.7

71 11.7

259 42.7

71 11.7

2 0.3

430 71.0

606 100%

RO 38.4 21.6

MA 48.1 42.5

SU 60.0 52.5

NE 48.6 27.1

GU 15.1 10.2

ALJ 36.2 45.6

GO ALG MM ALM Tot. sp. 12.4 1.1 69.2 7.6 185 11.4 n.c. 71.7 1.2 421

Saprótrofas Micorrizógenas

19 3.1

c MM: Mariánico-Monchiquense Sector, GO: Gaditano-Onubense Sector, ALJ: Aljíbico Sector, ALG: Algarviense Sector, RO: Rondeño Sector, MA: Malacitano-Almijarense Sector, SU: Subbético Sector, NE: Nevadense Sector, GU: Guadiciano-Bacense Sector, ALM: Murciano-Almeriense Province. n.c. = not collected.

Another noteworthy fact is the poverty of species belonging to the genera selected in the Guadiciano-Bacense (71 species) and Gaditano-Onubense (71 species) sectors, despite that the sampling level was similar to that of the chorological sectors (Moreno-Arroyo, 2004), which perhaps implies certain ecological factors that limit the growth of macrofungi to some extent. The chorological sector having the greatest species richness was Mariánico-Monchiquense (430 species, 71%), and, with numerous localities sampled in the provinces of Córdoba, Huelva and Jaén, had ample representation of mycorrhizal (302 species, 71.7%). The Subbético chorological sector also registered notable mycological richness, registering 332 species (54.8%) , with a slightly lower percentage of saprotrophic taxa (60%) than in the chorological sector Mariánico-Monchiquense (69.2%) and a clearly inferior representation of mycorrhizal taxa (221 species, 52.5%). Another two chorological sectors which included a considerable number of macrofungal species were: the Malacitano-Almijarense (268 species, 44.2%), which, extending throughout the 239

Fig. 3. CA of the chorological sectors having the greatest wealth of Macrofungi species. MM: Mariánico-Monchiquense Sector, ALJ: Aljíbico Sector, GO: Gaditano-Onubense Sector, SU: Subbético Sector, RO: Rondeño Sector, NE: Nevadense Sector, GU: Guadiciano-Bacense Sector, MA: Malacitano-Almijarense Sector.

provinces of Granada and Málaga, has similar representation in saprotrophic (48.1%) and mycorrhizal (42.5%) species is quite similar; the Aljíbico sector, which has a limited geographical area, has surprising mycoflora richness, with 259 species identified (42.7%), and a clearly greater representation of mycorrhizal (45.6%) than saprotrophic (36.2%) taxa. This also occurs in the MariánicoMonchiquense chorological sector, reflecting the similar biogeographical characteristics in the two sectors. In the Nevadense sector (204 species, 33.7%) saproptrophic taxa clearly dominate (48.6%), due to the ample and exclusive representation of these taxa in the wet mountain pastures of the oromediterranean and cryoromediterranean belts in Sierra Nevada massif, ecosystems which are not found in any other chorological sector in Andalusia and are similar to the mountain meadowlands in Alpine and Pyrenean zones. Finally, in the Rondeño chorological sector (162 species, 26.7%) saprotrophic (38.4%) predominate over mycorrhizal (21.6%) species, just as in the previous case, due to the fact the ecosystems sampled (e.g. Quercus ilex subsp. ballota and Abies pinsapo forests)have abundant decomposing plant litter over the soil. 2. Chorological aspects: The data compiled in Table 3 may be useful to establish the degree of similarity or separation between the different chorological sectors, and, in order to do so, we have made some Correspondence Analyses (CA) (Figs. 3 and 4). The most significant results are shown below. 240

Fig. 4. CA of the chorological sectors having the greatest richness of macrofungi (with the exception of the Gaditano-Onubense Sector). MM: Mariánico-Monchiquense Sector, ALJ: Aljíbico Sector, SU: Subbético Sector, RO: Rondeño Sector, NE: Nevadense Sector, GU: Guadiciano-Bacense Sector, MA: Malacitano-Almijarense Sector.

The clear division established between the Gaditano-Onubense chorological sector (Doñana National Park and the zones bordering the provinces of Cádiz, Sevilla and Huelva) and the remaining chorological sectors (Fig. 3), is based on differences in the presence of macrofungi which are exclusive, or strongly linked, to continental or coastal dune formations. Therefore, the Gaditano-Onubense sector was eliminated from the graph in Fig. 4, so that we could establish relationships between the remaining chorological sectors. As shown in Fig. 4, the relationship between the Aljíbico and Mariánico-Monchiquense sectors is very close, and we can consider their mycological flora quite similar, highlighted by a predominance of symbiotic species belonging to the Amanita and Boletus sensu lato, Russulales, etc.. The plant communities analysed in these sectors prefer cork oak (Q. suber) forests growing on an acidic substrate, which is determined by a considerable level of oceanity. Fig. 4 also demonstrates that (1) the Subbético chorological sector tends to approach the Mariánico-Monchiquense sector, which is not surprising because some enclaves of the provinces of Córdoba and Jaén form part of this last sector, so that many widespread macrofungal species which grow in the Mariánico-Monchiquense sector (Córdoba and Jaén) may also be present in the bordering geographic areas of the Subbético sector. Also patent is its relationship with the Malacitano-Almijarense and Rondeño sectors on the basis of the types of plant communities there as well as the basic pH of the substrate. (2) The clear relationship between the rest of the chorological 241

sectors (except Nevadense), which expected because all of them share a predominance of basic soil and the presence of macrofungal species thrive in plant communities such as Quercus ilex subsp. ballota forests, Pinus ssp. reforestation stands. Within this group of chorological sectors, another situation to be expected is the close relationship between the Malacitano-Almijarense and Rondeño chorological sectors, given that the two sectors are geographically close and therefore share widely distributed macrofungal species; moreover, the zones sampled within these sectors share the same mountainous environment, plant communities (evergreen oak and pine forests) and a predominant carbonate substrate (limestones and dolomites). However, the Rondeño chorological sector differs from all of these sectors for the notable presence of Abies pinsapo forests. The Guadiciano-Bacense sector has similar ecological characteristics to those of Malacitano-Almijarense, sharing a geographical border, but it has less rainfall and therefore a lower number of species collected (71 taxa). Finally, isolation of the Nevadense chorological sector is noteworthy, as it shares middle-mountain forest formations with the Malacitano-Almijarense and Guadiciano-Bacense chorological sectors: evergreen oak forests, pine forests and riparian communities. However, these differ somewhat, in addition to the fact that the mycoflora which grows in the oromediterranean and cryoromediterranean belts of the Sierra Nevada is completely different. 3. Comparative mycofloristic analysis of the chorological sectors represented in Andalusia: As stated above, Table 3 indicates the percentage of representation (relative number) of each of the macrofungi genera studied, which include ectomycorrhizal and saprophytic species, represented in each of the Andalusian chorological sectors. MYCORRHIZAL SPECIES: Some groups of chorological sectors can be differentiated according to their mycological flora: (1) Chorological sectors of the Rondeño, Malacitano-Almijarense, Nevadense and Guadiciano-Bacense, are all characterised by: (i) a relatively scarce representation of the Amanita and, above all, Russula species; (ii) a significant representation of the Cortinarius (MA and NE sectors) and Tricholoma species (RO, MA and NE sectors); (iii) a moderate fructification of Inocybe species (RO, MA and GU sectors), which show considerable presence in evergreen oak and pine forests; (iv) and an excellent representation of the Suillus species, symbiotically associated with diverse reforested Pinus species which are frequent in these chorological sectors. (2) The chorological sectors Aljíbico and Mariánico-Monchiquense, in which the representation of the Amanita, Boletales and Russulales species is considerable, and the rarer presence of species belonging to the Suillus, in relation to other genera, due to the scant presence of Pinus afforestation The rarest species are Cortinarius, Inocybe and Tricholoma (ALJ sector), which are appreciably represented in the broad-leaved forests. (3) The Gaditano-Onubense chorological sectors manifest some peculiarities: (i) the common presence of sandy soils (some taxa belonging to the Inocybe genus), (ii) an oceanic character (notable presence of the Amanita and Russula species), and (iii) a good representation of Pinus pinea forests (presence of Lactarius deliciosus (L.) Gray and species belonging to Inocybe and Suillus). 242

SAPROTROPHIC

SPECIES:

The most important data are:

(1) The dissimilarities between the different sectors are less obvious than in the case of the ectomycorrhizal macrofungi, explaining why the saprotrophic species grow more directly in response to the rainfall regime in any one geographic zone, and also to the level of conservation of plant formations and the abundance of decomposing plant matter, in preference to the nature of substrate. (2) The Nevadense chorological sector shows the most appreciable differences with reference to the rest, since the most represented genus is Entoloma (subgenus Leptonia) and the least represented genera are Mycena, Agaricus, and Psathyrella. (3) The species of the selected genera which grow in the Gaditano-Onubense sector are also somewhat special, as the least represented genera are Clitocybe and Mycena, this being explained by the presence of fungi which grow on sandy and ruderalized substrate (e.g. Agaricus spp., Psathyrella spp.) or those which form part of heliophile plant communities (Entoloma cistophilum Trimbach, E. philocistus Hauskn. & Noordel., etc.). (4) The rest of chorological sectors have a significant presence of the Clitocybe and Mycena species, and least generalized Agaricus (MA, SU and MM sectors) or Entoloma species (SU and MM sectors). Discussion In this study, we examine the following. 1. Mycofloristic aspects. As shown in Table 3, the Aljíbico and MariánicoMonchiquense sectors show a greater representation of the mycorrhizal macrofungi taxa, since most were collected from cork oak forests. These forests are distinguished by their greater oceanity, the acidic pH of the substrate, an appropriate conservation level, and less decomposing organic matter on the soil, all this encouraging the fructifying of species of Amanita, Boletus, Russula, etc. The contrary situation arises particularly in the chorological sectors Nevadense, Rondeño and Guadiciano-Bacense, as well as in other forests located in eastern Andalusia, where the mainly basic pH of the soil, the greater continentality, the worse conservation level, and the greater amount of decomposing plant litter on the substrate favour the development of saprophytic macrofungi. The low number of macrofungi species found in the chorological sectors Gaditano-Onubense and Gaditano-Bacense can be explained, respectively, by the predominance of the psammophilous communities, these being highly restrictive for the development of macrofungi in the Gaditano-Onubense sector (GO) as well as the very low rainfall in the Gaditano-Bacense sector (GU). The Malacitano-Almijarense and Subbético chorological sectors are situated in an intermediate position due especially to a more favourable rainfall regime. The chorological sectors which have the highest number of species are MariánicoMonchiquense, Subbético, Malacitano-Almijarense and Aljíbico. This is explained in the three first cases by the large geographical surface area and quite favourable 243

climatic conditions for the development of macrofungi (above all in the MariánicoMonchiquense sector). The Aljíbico sector constitutes one of Mediterranean zones with the most biodiversity, due to the exceptional biogeographical particularities that characterize the natural park of Alcornocales (Cádiz and Málaga provinces), such as the excellent degree of conservation of the cork oak forests, and the favourable climate. 2. Chorological aspects. The macrofungi of a any particular zone constitute a good parameter for biogeographical analysis, since they may be used to distinguish the different chorological sectors which are present in a specific region or geographic zone. On the basis of their microflora, the chorological sectors analysed can be classified as follows. 2.a. Gaditano-Onubense sector is characterized by the presence of a number of species adapted to living in psammophilous communities, notably: Amanita boudieri Barla, Entoloma cistophilum Trimbach, Entoloma philocistus Hauskn. & Noordel., Inocybe amblyspora Kühner, Inocybe brunneorufa Stangl & J.Veselský, Inocybe pruinosa R.Heim, Mycena dunicola Esteve-Rav. et al., Psathyrella ammophila (Durieu & Lév.) P.D.Orton. 2.b. The Mariánico-Monchiquense, Aljíbico and Subbético chorological sectors are related especially by the considerable representation of ectomycorrhizal taxa, which express the state of maturity of the forests sampled (evergreen oak and cork oak forests), as well as a propitious climate for macrofungi to fructify. In addition, the Mariánico-Monchiquense and Subbético sectors, for their geographic proximity, share many species, mainly those with a wide ecological spectrum. 2.c. The Malacitano-Almijarense, Rondeño and Guadiciano-Bacense chorological sectors were related for sharing the same type of forest (Quercus ilex subsp. ballota and Pinus spp. forests) and the carbonate substrate. 2.d. The Nevadense sector is distinctly separated from the rest, since, although it shares with other sectors (MA and GU) the types of forests sampled (evergreen oak woodlands, pine forests, and riparian communities), these develop on an acidic substrate. Furthermore, the mycoflora that grows in the oromediterranean and cryoromediterranean belts of the Sierra Nevada is completely different, where the macrofungal species (e.g. Entoloma, subgenus Leptonia spp.) are very common. These fungal species can also be found in the high Alpine and Pyrenean mountain pastures, which are not present in any other chorological sector in Andalusia. 3. Comparative mycofloristic analysis. Three factors most influential the distribution of mycorrhizal species: (i) bioclimatic characteristics of the study area (oceanity and/or continentality), (ii) the substrate pH, and (iii) the plant-community type and its conservation level . With regard to the saprophytic species, the factors most govern their development are: (i) the local pluviometric regime and (ii) the amount of decomposing organic matter. With respect to the degree of representation (relative number, Table 3) of the selected macrofungi genera in each of the chorological sectors, there are certain mycofloristic differences between sectors located in western Andalusia (sectors ALJ, GO and MM) 244

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