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Chorology and habitat characterization of Fritillaria caballeroi an endemic and threatened species in the Central System (Iberian Peninsula)/Corologia y caracterizacion del habitat de la liliacea endemica y amenazadaritillaria caballeroi en las montanas del Sistema Central (Peninsula Iberica).


The genus Fritillaria L. (Liliaceae) is distributed across the Northern Hemisphere and, in particular, in temperate zones of Asia, Europe, North Africa and the West of North America (Grove, 1931). The centre of diversification of the taxon could be considered to be in Iran, in the middle of Asia or in the Caucasus (Guemes, in press). The number of recognized species in the genus varies, depending on the author, ranging from 50 (Krause, 1930) to almost 100 (Melchior, 1964). In Spain, traditionally five taxa have been addressed (Fernandez-Arias & Devesa, 1991a; Guemes, in press), although with different taxonomic ranges. One such taxon is the Spanish endemism (F. legionensis Llamas & J. Andres); two species are Iberian endemisms (F. caballeroi F.M. Vazquez and F. stenophylla Boiss. & Reuter), one en demism is Ibero-N. African (F. lusitanica Wikstr.) and the remaining species is Franco-Iberian (F. pyrenaica L.).

Caballero (1948) described Fritillaria falcata in the Sierra de Francia (Salamanca), pointing out that it was a very different species from other known species of the genus on the basis of its small size, its falcate leaves and the presence of a single flower. Later Rico (1985) proposed that it would be a form of F. lusitanica, indicating that the diagnostic characters are not always present within the same population. In their monographic work of the genus on the Iberian Peninsula Fernandez-Arias & Devesa (1991a) subordinate it to a subspecies of F. nervosa Willd., above all on the basis of its falcate, longitudinally folded leaves. More recently, Vazquez (2009) designated it a new name, F. caballeroi, owing to the existence of a previous heterotypic synonym for a Californian species (Beetle, 1944). Finally, in Flora Iberica Guemes (in press) maintains that denomination of F. caballeroi, together with the specific rank for the plant.

Fritillaria caballeroi is considered to be a vulnerable plant (VU), both in the Red Lists of the Flora Vascular Espanola (VV.AA., 2000; Moreno, 2008), and in the Atlas and Red Book (Amich & Bernardos, 2010), as a result of its reduced geographic distribution and of the continuing decrease in its number of mature plants. This taxon has not yet been assessed by the IUCN Red List (Iucn, 2012). Similarly, the species is catalogued as "Preferential Attention" in the catalogue of Flora Protegida de Castilla y Leon (Anonymous, 2007). An index record of the plant has been made for Castilla y Leon and a proposal for a micro-reserve for the plant has been put forwards (Amich & Bernardos, unpub. data), in concordance with the provisions of the Protection Decree for this Community (Anonymous, 2007).

The aims of the present work were: (1) to address in detail the distribution of this endemism in the Sistema Central, and (2) to characterize the habitats of the populations of F. caballeroi in the areas where it grows in the Sistema Central and compare the habitats of the different sierras.



Fritillaria caballeroi flowers towards the end of spring (May-June) and fruits in summer (July- August); insect pollinated; seeds dispersed by wind. Its chromosome number is 2n = 24 (FERNANDEZ- ARIAS & DEVESA, 1991b). In the central western part of the Iberian Peninsula, the species grows on siliceous rocks, either granites (Sierras de Bejar, Tormantos and la Estrella) or Armorican quartzes from the Silurian and Devonian (Sierra de Francia).

During the period 2008-2011 we visited most of the localities in the Iberian Peninsula in which Fritillaria caballeroi has been reported (Table 1) and we also surveyed other areas thought suitable for this taxon along the sierras of the Sistema Central, including the Portuguese territories of the Serra da Estrella.

Habitat characterization of fritillaria caballeroi

This study involved 57 species and 8 releves, and was performed following the method of Braun-Blanquet (1979). The syntaxonomic nomenclature followed was that suggested by Rivas-Martinez & al. (2002) and Rivas-Martinez (2011) and Costa & al. (2012). In the syntaxonomic synopsis of the syntaxa mentioned we also followed the ordering proposals of RivasMartinez & al. (2002) and Rivas-Martinez (2011), except for the association Minuartio juresii-Festucetum summilusitanae (in Costa & al. 2012), described after works of Rivas-Martinez.

The nomenclature used for the taxa of syntaxonomic interest was that of Flora Europaea (Tutin & al., 1964-1980) and Flora Iberica (Castroviejo, 1986-2012), except for Festuca summilusitana subsp. gredensis (Fuente & Ortunez) Rivas-Mart., Fuente & Ortunez in Itinera Geobotanica 18: 485 (2011), Jasiones crispa subsp. centralis (Rivas-Mart.) Rivas-Mart. in Anales Inst. Bot. Cavanilles 27:154 (1970), Minuartia recurva subsp. juresii (Willd. ex Schltdl.) Mattf. in Bot. Jahrb. Syst. 57, Beibl. 126: 31 (1921), and Silene ciliata subsp. elegans Link ex Brot. in Fl Lusit. 2: 185 (1804).

Fritillaria caballeroi flowers towards the end of spring (May-June) and fruits in summer (JulyAugust); insect pollinated; seeds dispersed by wind. Its chromosome number is 2n = 24 (Fernandez-Arias & Devesa, 1991b). In the central western part of the Iberian Peninsula, the species grows on siliceous rocks, either granites (Sierras de Bejar, Tormantos and la Estrella) or Armorican quartzes from the Silurian and Devonian (Sierra de Francia).

During the period 2008-2011 we visited most of the localities in the Iberian Peninsula in which Fritillaria caballeroi has been reported (Table 1) and we also surveyed other areas thought suitable



We first reviewed the published data on the distribution of this species and additionally revised some of the main herbaria pertaining to central western Iberia (HVR, COI and SALA). The total number of known localities are shown in Figure 1 and Table 1. In the explorations carried out in these territories we did not note new population nuclei. However, their presence in them cannot be ruled out, owing their extraordinary geomorphological complexity, the early phenology of the plant and the existence of favorable ecological conditions for growth.

The citation of Fritillaria caballeroi for the Montes de Leon (Paz & al., 2011) was later confirmed by Guemes (in press).

Thus, Fritillaria caballeroi was found to be a narrow endemic of the Orocantabric and Carpetan Leonese subprovinces (Atlantic European and Mediterranean West Iberian provinces, respectively, according to Rivas-Martinez & al., 2002; Rivas-Martinez, 2005, del Egido & Puente, 2011). In the Central System (Carpetan Leonese subprovince) we detected it in the following phytogeographic units: the Tormantino and Bejarano districts (Bejaran Gredensean biogeographical sector), the Altosalmantino district (Salmanticensean sector), and the Altoestrelensean district (Estrelensean sector). In the Sistema Central it has two small populations in the sierra de Tormantos; another two small populations in the sierra de Bejar; a single population with small subpopulations scarcely separated by a few tens of metres in the Sierra de Francia, and a single population in the Portuguese Serra da Estrela. Despite the different toponimies used in the citations referring to the Sierra de Francia, all of them correspond to only three municipalities and 3 squares of 10 x 10 km, which are contiguous (Figure 1, Table 1). The species' altitude range was from 1500 m for the populations in the Sierra de Francia up to 2250 m for the populations of the Sierra de Bejar. The southern and eastern limits of the populations are at 40[degrees] 13.009' and 05[degrees] 34.418' respectively (Sierra de Tormantos, Avila), while the northern and western limits are respectively in the Sierra de Francia (40[degrees]31.595') and the Serra da Estrella (07[degrees]35.199').


We elaborated a series of 8 releves (Table 2) in all locales in which Fritillaria caballeroi has been cited and at which we had the possibility of confirming its presence.

In the Sierras de Bejar and Tormantos (Table 2, rels. 1-4) Fritillaria caballeroi is found in vigorous silicolous, psychrophilous and cryoromediterranean meadows, which prosper on the summit zones of high mountains of the Sistema Central ("altioreina" communities) on crests, spurs and rock ledges that receive little snow. These are phytocoenoses with a predominance of caespitoses chamaephytes and are rich in Central Iberian palaeoendemisms, such as Armeria bigerrensis subsp. bigerrensis, Jasione crispa subsp. crispa, Minuartia recurva subsp. juresii, and Silene ciliata subsp. elegans.

These grasslands belong to the association Agrostio rupestris-Armerietum bigerrensis, endemic to the Bejaran-Gredensean, from the Picos de Los Galayos to the Sierra de Bejar (Sardinero, 1994, 2004; Rivas-Martinez, 2011), which is integrated within the optimum Carpetan alliance Minuartio-Festucion curvifoliae. This vegetation is representative of a mature stage of a series of climatophilous vegetation or of a permanent community of a xerophyte site.

By contrast, in the Sierra de Francia (Table 2, rels. 5-7,) Fritillaria caballeroi is integrated in open pioneer-like grasslands that prosper on shallow, developed siliceous soils, subjected to strong phenomena of cryoturbation, on the Oromediterranean ("oreina" communities), generally on topographic scenarios of crests, steep slopes, or rocky spurs where the accumulation of snow is of little magnitude. This chyonophobic nature means that F. caballeroi often shows a typically subrupicolous behaviour. In these grasslands there is a greater predominance of short, and caespitoses hemicryptophytes although the grasslands are also rich in chamaephytes and therophytes. They also have different Iberian or Central Iberian endemisms such as Arenaria querioides, Armeria salmantica, Festuca summilusitana subsp. summilusitana, F. summilusitana subsp. gredensis, and Jasione sessiliflora.

These communities lie within the association Arenario querioidis-Festucetum gredensis, endemic to the Bejarano-Gredense (Sanchez-Mata, 1989; Sardinero, 1994, 2004; Sardinero & Rivas-Martinez, 1999), although also reaching the Altosalmantino (Amich & Bernardos, 2010), integrated within Hieracio castellani-Plantaginion radicatae, an optimum Carpetan phytocoenosis (although also present in the oroiberian and orocantabric areas) like Minuartio-Festucion.

In most cases, these grasslands appear in clear areas close to oromediterranean Cytiso-Echinospartetum pulviniformis broom, forming a regressive replacement stage (herbaceous serial stage) of the sierra broom communities with pulviniform "cambriones", which form the natural potential vegetation of the Oromediterranean belt of the whole studied territory (Rivas-Martinez & al., 1987; Gavilan & al., 2011).

For the Serra da Estrella, Meireles (2010) cites Fritillaria nervosa subsp. nervosa (= F. pyrenaica L.), in agreement with the citation of Franco & RochaAfonso (in Franco 1994), but the Lynnean taxon is not present in Portugal (Guemes, in press). F. caballeroi (Guemes, in press) is found, however, in the Beira Alta (Serra da Estrella). Meireles (2010) indicates that Fritillaria lives in communities of Hieracio castellani-Plantaginion radicatae.

In the Serra da Estrella (Table 2, rel. 8) we have found F. caballeroi in communities referable to the recently described association Minuartio juresii-Festucetum summilusitanae (Costa & al., 2012), which replaces the association Arenario querioidis-Festucetum gredensis at the western end of the Central System, communities also belonging to the alliance Hieracio castellani-Plantaginion radicatae.

In the Montes de Leon, Paz & al. (2011) also report it flourishing in silicolous grasslands of Hieracio-Plantaginion, and in clearings of heather clumps and groups of Cytision oromediterranei.

The habitats of Fritillaria caballeroi are clearly different from those occupied by other species of the genus on the Iberian Peninsula. Thus, F. legionensis preferentially lives in hay meadows, semi-flooded meadows and intensely grazed acidophilous grasslands ("nardetas") in the Supra- and Orotemperate belts, above all in the alliances Arrhenatherion and Nardion (Llamas & Andres, 1983; Rodriguez-Rojo & al., 2012; Paz & al., 2011). Fritillaria pyrenaica is a taxon that is indifferent to the substrate, has a broad ecological distribution, and lives in areas ranging from grasslands and meadows to forest clearings, above all in the supra- and orotemperate storeys (Fernandez-Arias & Devesa, 1991a; Paz & al., 2011). Finally, F. lusitanica lives in scrub communities of Cytisetea (Gavilan & al., 2011), shrubland and forest clearings, ranging from the thermo- to the supramediterranean storey, whereas F. stenophylla occurs on palaeodunes and sandy soils in thermomediterranean shrub patches of the alliance Coremation albi (FernandezArias & Devesa, 1991a).

doi: 10.5209/rev_LAZA.2013.v34.n1.41685


We thank Dr C. Meireles for granting us access and permission to consult her PhD Thesis on the Flora and Vegetation of the Portuguese Serra da Estrella and Dr J. Rocha for the cartography.


Communities cited in the text:

FESTUCETEA INDIGESTAE Rivas Goday & Rivas-Martinez 1971

Festucetalia curvifoliae Rivas Goday & Rivas-Martinez ex Rivas-Martinez 1964 corr. Izco & Pulgar 2009

Minuartio bigerrensis-Festucion curvifoliae Rivas-Martinez 1964 corr. Rivas-Martinez, Fernandez-Gonzalez & Loidi 1999

Agrostio rupestris-Armerietum bigerrensis Rivas-Martinez in Rivas-Martinez, Fernandez-Gonzalez & Sanchez-Mata 1986

Jasiono sessiliflorae-Koelerietalia crassipedis Rivas-Martinez & Canto 1987

Hieracio castellani-Plantaginion radicatae Rivas-Martinez & Canto 1987

Arenario querioidis-Festucetum gredensis Rivas-Martinez, Sanchez-Mata & Source in Rivas-Martinez, Fernandez-Gonzalez & Sanchez-Mata 1986 corr. Rivas-Martinez & Sanchez-Mata 2002


Arrhenatheretalia elatioris Tuxen 1931

Arrhenatherion elatioris Koch 1926

NARDETEA STRICTAE Rivas Goday in Rivas Goday & Rivas- Martinez 1963

Nardetalia strictae Oberdorfer ex Preising 1950

Nardion strictae Br.-Bl. 1926

CISTO-LAVANDULETEA STOECHADIS Br.-Bl. in Br.-Bl., Molinier & Wagner 1940

Stauracantho genistoidis-Halimietalia calycini Rivas-Martinez, Lousa, T.E. Diaz, Fernandez-Gonzalez & J.C. Costa 1990

Coremation albi Rothmaler 1943


Juniperetalia hemisphaericae Rivas-Martinez & J.A. Molina in Rivas-Martinez, Fernandez-Gonzalez & Loidi 1999

Cytision oromediterranei Tuxen in Tuxen & Oberdorfer 1958 corr. Rivas-Martinez 1987

Cytiso oromediterranei-Echinospartetum pulviniformis Rivas-Martinez, Belmonte, Canto, Fernandez-Gonzalez, Fuente, J.M. Moreno, Sanchez-Mata & Sancho 1987


Amich, F. & Bernardos, S.--2010--Fritillaria caballeroi F.M. Vazquez--In: Banares, A., Blanca, G., Guemes, J., Moreno, J.C. & Ortiz, S. (Eds.). Atlas y Libro Rojo de la Flora Vascular Amenazada de Espana. Addenda 2010. Pp. 114-115. Dir. Gen. Med. Nat. Pol. For., Madrid.

Anonymous--2007--Decreto 63/2007, de 14 de Junio, por el que se crean el Catalogo de Flora Protegida de Castilla y Leon y la figura de proteccion denominada Microrreserva de Flora. Bol. Ofic. Castilla y Leon 119: 1319713204

Beetle, D.E.--1944--A monograph of the North American species of Fritillaria.--Madrono 7: 133-158.

Braun-Blanquet, J.--1979--Fitosociologia. Bases para el estudio de las comunidades vegetales--Ed. Blume. Madrid. 820 pp.

Caballero, A.--1947--. Dos excursiones botanicas a los alrededores de La Alberca (Salamanca, Caceres)--An. Inst. Bot. Cavanilles 7: 645-653.

Castroviejo, S. & al. (Coords.)--1986-2012--Flora iberica. Plantas vasculares de la Peninsula Iberica e Islas Baleares. Vols. 1-8, 10-15, 17-18, 21. R. Jard. Bot., C.S.I.C. Madrid.

Costa J.C., Neto C., Aguiar C., Capelo J., Espirito Santo M.D., Honrado J., Pinto-Gomes C., Monteiro-Henriques T., Sequeira M., Lousa, M.--2012--Vascular plant communities in Portugal (Continental, the Azores and Madeira)--Global Geobot. 2: 1-180.

Del Egido, F. & Puente, E.--2011--Valeriano apulae-Potentilletum nivalis y Festuco eskiae-Cytisetum oromediterranei: dos nuevas comunidades vegetales de la alta montana cantabrica--Lazaroa 32: 91-99.

Fernandez-Arias, M.I. & Devesa, J.A.--1991a--Revision del genero Fritillaria L. (Liliaceae) en la Peninsula Iberica --Stud. Bot. 9: 49-84.

Fernandez-Arias, M.I. & Devesa, J.A.--1991b--Estudio cariologico del genero Fritillaria L. (Liliaceae) en la Peninsula Iberica.--Stud. Bot. 9: 85-107.

Franco, A.--1994--Nova Flora de Portugal (continente e Acores). Vol. 3, Fasc. 1. Escolar Editora, Lisboa. 181 pp.

Gavilan, R.G., Vilches de la Serna, B. & Fernandez-Gonzalez, F.--2011--Syntaxonomical review of Cytisetea scopario-striati communities in central Spain--Lazaroa 32: 29-72.

Grove, A.--1931--The genus Fritillaria--The New Flora and Silva 3: 147-153.

Guemes, J.--In Press--Fritillaria L.--In: Rico, E. (Ed.). Flora Iberica, vol. 20. R. Jard. Bot., CSIC, Madrid.

Krause, K.--1930--Liliaceae--In: Engler, A. (Ed.). Die Naturlichen Planzenfamilien. Pp. 332-335. Leipzig.

IUCN--2012--IUCN Red List of Threatened Species. Version 2012.2. <>. Downloaded on 04 December 2012.

Llamas, F., & Andres, J.--1983--Fritillaria legionensis Llamas & Andres, sp. nov.--Lagascalia 11: 67-70.

Meireles, C.I.--2010--Flora e Vegetacao da Serra da Estrela: Aproximacao Fitossociologica da Vertente Meridional. Mem. Doc. (ined.). Univ. Jaen. 397 pp.

Melchior, H.--1964--Reihe Liliiflorae--In: Engler, A. (Ed.). Syllabus der Planzenfamilien, 2: 513-543. Berlin.

Moreno, J.C. (Coord.)--2008--Lista Roja 2008 de la Flora Vascular espanola.--Dir. Gen. Med. Nat. Pol. For. Madrid. 86 pp.

Moreno, J.C.--2002--Asientos para un atlas corologico de la flora occidental, 25. Mapa 0930--Cavanillesia Altera 2: 593-594.

Paz, E., Alonso, R., Ruiz, A. & Garcia-Gonzalez, M.E.--2011--El genero Fritillaria L. (Liliaceae) en la Cordillera Cantabrica (Espana)--Candollea 66: 383-395.

Rivas-Martinez, S.--2005--Avances en Geobotanica. R. Acad. Farm. Madrid. 128 pp.

Rivas-Martinez, S.--2011--Mapa de series, geoseries y geopermaseries de Vegetacion de Espana--Itinera Ge obot. 18: 5-800.

Rivas-Martinez, S., Diaz, T.E., Fernandez-Gonzalez, F., Izco, J., Loidi, J., Lousa, M. & Penas, A.--2002--Vascular plant communities of Spain and Portugal--Itinera Gebobot. 15: 5-922.

Rivas-Martinez, S., Belmonte, D., Canto, P., Fernandez-Gonzalez, F., de la Fuente, V., Moreno, J.M., SanchezMata, D. & Sancho, L.G.--1987--Piornales, enebrales y pinares oromediterraneos (Pino-Cytision oromediterranei) en el Sistema Central--Lazaroa 7: 93-124.

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Sardinero, S. & Rivas-Martinez, S.--1999--La vegetacion herbacea vivaz climatofila supra- y oromediterranea (Koelerio-Corynephoretea)en Gredos occidental, Sistema Central, Espana--Lazaroa 20: 55-69.

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Vazquez, F.M.--2009--Anotaciones taxonomicas y nomenclaturales a la flora de Extremadura: Fritillaria caballeroi F.M. Vazquez nom. nov.--Folia Bot. Extremad. 4: 122.

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Received: 8 April 2013

Accepted: 5 July 2013

Francisco Amich & Sonia Bernardos (*)

(*) Department of Botany. University of Salamanca. E-37007 Salamanca, Spain. E-mail:

Table 1
Known localities of Fritillaria caballeroi in the Central System

Country    Province     Locality         Site

Portugal   Beira Alta   Manteigas        Covao d'Ametade
Spain      Avila        Navalguijo       Sierra de Tormantos
Spain      Avila        Nava del Barco   Sierra de Tormantos, entre
                                         los circos de La Nava y
                                         los Caballeros
Spain      CAceres      Tornavacas       Sierra de Tormantos,
                                         Garganta La SerrA
Spain      CAceres      La Garganta      Sierra de Bejar, Hoya de
Spain      Salamanca    Candelario       Sierra de Bejar, El Calvitero
Spain      Salamanca    El Cabaco        Sierra de Francia, Pena de
Spain      Salamanca    El Cabaco        Sierra de Francia, Paso de
                                         los Lobos
Spain      Salamanca    El Cabaco        Sierra de Francia, cercanias
                                         del Collado de Pena Cabra
Spain      Salamanca    El Mallo         Sierra de Francia, Penas
                                         del Corzo
Spain      Salamanca    Monsagro         Sierra de Francia, cercanias
                                         del Collado de Pena Cabra

Country    Altitude   UTM           Geographical
                      coordinates   coordinates

Portugal   1575       29TPE2065     40[degrees] 19.735' N
                                    07[degrees] 35.199' W
Spain      2120       30TTK8155     40[degrees] 13.009' N
                                    05[degrees] 34.418' W
Spain      2180       30TTK8155     40[degrees] 13.011' N
                                    05[degrees] 34.420' W
Spain      1600       30TTK76       40[degrees] 15.533' N
                                    05[degrees] 42.273' W
Spain      2000       30TTK66       40[degrees] 15.364' N
                                    05[degrees] 49.321' W
Spain      2250       30TTK6967     40[degrees] 19.296' N
                                    05[degrees] 43.130' W
Spain      1675       29TQE4088     40[degrees] 30.481' N
                                    06[degrees] 10.047' W
Spain      1500       29TQE3988     40[degrees] 30.498' N
                                    06[degrees] 10.754' W
Spain      1575       29TQE3989     40[degrees] 31.039' N
                                    06[degrees] 10.733' W
Spain      1500       29TQE3890     40[degrees] 31.595' N
                                    06[degrees] 11.416' W
Spain      1575       29TQE3989     40[degrees] 31.038' N
                                    06[degrees] 10.732' W

Country    Voucher          Cited by

Portugal   MA 784248        GUEMES (in press)
Spain      MAF 135686       SANCHEZ-MATA & al. (1991)
                            AMICH & BERNARDOS (2010)
Spain      MAF 135687       SANCHEZ-MATA & al. (1991)
                            AMICH & BERNARDOS (2010)
Spain      HSS 25160        VAZQUEZ (2009)
Spain      HSS 9909/9290B   VAZQUEZ (2009)
Spain      MAF 131481       FERNANDEZ-ARIAS & DEVESA (1991a)
                            MORENO (2002)
Spain      MA 21354,        CABALLERO (1947)
           SALA 79649       FERNANDEZ-ARIAS & DEVESA (1991a)
Spain      SALA 77857       FERNANDEZ-ARIAS & DEVESA (1991a)
Spain      --               AMICH & BERNARDOS (2010)
Spain      SALA 13628,      RICO (1985)
Spain      --               AMICH & BERNARDOS (2010)

Table 2
Agrostio rupestris-Armerietum bigerrensis (1-4)
(Minuartio-Festucion curvifoliae, Festucetalia indigestae,
Festucetea indigestae)
Arenario querioidis-Festucetum gredensis (5-7)
Minuartio juresii-Festucetum summilusitanae (8)
(Hieracio-Plantaginion radicatae, Jasiono sessiliflorae-Koelerietalia
crassipedis, Festucetea indigestae)

Altitude (m)                             2100    2110    2150    2175
Area ([m.sup.2])                          40      50      45      45
Coverage (%)                              50      50      60      55
Exposure                                   N       N      NE      NE
Inclination (%)                           25      20      15      20
Releve number                              1       2       3       4

Jasione crispa subsp. centralis            2       3       2       1
Armeria caespitosa                         1       1       1       2
Silene ciliata subsp. elegans              1       +       2       1
Leucanthemopsis alpina                     1       1       1       +
Plantago alpina                            1       1       .       +
Luzula hispanica                           .       +       .       +
Agrostis rupestris                         .       +       +       .
Fritillaria caballeroi                     +       .       +       +
Agrostis delicatula                        1       1       +       1
Sedum brevifolium                          1       1       .       +
Festuca rivas-martinezii                   .       .       .       .
Festuca summilusitana subsp. gredensis     .       .       .       .
Arenaria querioides                        .       .       .       .
Jasione sessiliflora                       .       .       .       .
Hieracium castellanum                      .       .       .       .
Leucanthemopsis flaveola                   .       .       .       .
Armeria salmantica                         .       .       .       .
Armeria transmontana                       .       .       .       .
Plantago holosteum                         .       .       .       .
Koeleria crassipes                         .       .       .       .
Corynephorus canescens                     .       .       .       .
Scleranthus perennis                       .       .       .       .
Companions of Salicetea and Nardetea
Mucizonia sedoides                         1       +       .       +
Nardus stricta                             +       +       .       +
Dianthus gredensis                         1       1       .       .
Companions of Helianthemetea
Rumex angiocarpus                          .       .       .       .
Logfia minima                              .       .       .       .
Micropyrum tenellum                        .       .       .       .
Spergula arvensis                          .       .       .       .
Linaria elegans                            .       .       .       .
Arnoseris minima                           .       .       .       .
Cerastium ramosissimum                     .       .       .       .
Other companions
Senecio carpetanus                         +       +       +       .
Dianthus lusitanus                         .       .       .       .
Arrhenatherum carpetanum                   .       .       .       .
Luzula lactea                              .       .       .       .
Avenula pratensis                          .       .       .       .
Linaria nivea                              .       .       .       .
Poa bulbosa                                .       .       .       .
Ornithogalum concinnum                     .       .       .       .
Gagea lutea                                .       .       .       .
Teedalia nudicaulis                        .       .       .       .
Sedum hirsutum                             .       .       .       .
Jasione montana                            .       .       .       .
Cytisus oromediterraneus                   +       +       .       .
Hyacinthoides non-scripta                  .       .       .       .
Arenaria montana                           .       .       .       .
Conopodium majus                           +       .       +       .
Tulipa sylvestris                          .       .       .       .

Altitude (m)                             1575    1580    1585    1575
Area ([m.sup.2])                          100     50      75      50
Coverage (%)                              70      60      50      50
Exposure                                   E       E      NE      NE
Inclination (%)                           10      10      15      10
Releve number                              5       6       7       8

Jasione crispa subsp. centralis            .       .       .       .
Armeria caespitosa                         .       .       .       .
Silene ciliata subsp. elegans              .       .       .       .
Leucanthemopsis alpina                     .       .       .       .
Plantago alpina                            .       .       .       .
Luzula hispanica                           .       .       .       .
Agrostis rupestris                         .       .       .       .
Fritillaria caballeroi                     1       2       1       1
Agrostis delicatula                        2       1       1       +
Sedum brevifolium                          1       1       .       1
Festuca rivas-martinezii                   1       +       1       .
Festuca summilusitana subsp. gredensis     .       .       .       1
Arenaria querioides                        1       2       1       1
Jasione sessiliflora                       1       2       1       1
Hieracium castellanum                      2       1       1       +
Leucanthemopsis flaveola                   2       1       1       .
Armeria salmantica                         1       1       2       .
Armeria transmontana                       .       .       .       1
Plantago holosteum                         1       1       1       .
Koeleria crassipes                         1       1       .       1
Corynephorus canescens                     +       1       1       .
Scleranthus perennis                       1       1       .       .
Companions of Salicetea and Nardetea
Mucizonia sedoides                         .       .       .       .
Nardus stricta                             .       .       .       .
Dianthus gredensis                         .       .       .       .
Companions of Helianthemetea
Rumex angiocarpus                          1       1       1       1
Logfia minima                              1       1       1       .
Micropyrum tenellum                        +       1       +       .
Spergula arvensis                          +       +       1       .
Linaria elegans                            1       1       .       .
Arnoseris minima                           1       .       +       .
Cerastium ramosissimum                     .       1       +       .
Other companions
Senecio carpetanus                         .       .       .       .
Dianthus lusitanus                         1       1       1       1
Arrhenatherum carpetanum                   1       2       1       .
Luzula lactea                              1       2       2       .
Avenula pratensis                          1       2       1       .
Linaria nivea                              1       2       1       .
Poa bulbosa                                1       1       1       .
Ornithogalum concinnum                     2       1       2       .
Gagea lutea                                +       1       1       .
Teedalia nudicaulis                        +       .       1       1
Sedum hirsutum                             1       1       .       .
Jasione montana                            1       +       .       .
Cytisus oromediterraneus                   3       .       .       +
Hyacinthoides non-scripta                  1       2       .       .
Arenaria montana                           .       1       1       .
Conopodium majus                           .       1       1       .
Tulipa sylvestris                          .       1       1       .

Other species: Pterospartum tridentatum 3 in 5; Echinospartum ibericum
and Halimium alysoides 2 in 5; Erica arborea and E. australis 1 in 5;
Orchis champagneuxii 1 in 6; Narcissus rupicola + in 8.

Localities: SPAIN: 1: Avila, Navalguijo, Sierra de Tormantos,
30TTK8155, 06.06.2008, Amich & Bernardos; 2: ibidem, 06.06.2008, Amich
& Bernardos; 3: Avila, Nava del Barco, Sierra de Tormantos, near Circo
de La Nava, 30TTK8155, 06.06.2008, Amich & Bernardos; 4: ibidem,
06.06.2008, Amich & Bernardos; 5: Salamanca, El Cabaco, Sierra de
Francia, near Collado de Pena Cabra, 29TQE3989, 15.05.2008, Amich &
Bernardos; 6: ibidem, 15.05.2008, Amich & Bernardos; 7: Salamanca,
Monsagro, Sierra de Francia, near Collado de Pena Cabra, 29TQE3989,
15.05.2008, Amich & Bernardos. PORTUGAL: 8: Beira Alta, Manteigas,
Serra d'Estrella, Covao d'Ametade, 29TPE2065, 25.05.2010, Amich &
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Title Annotation:articulo en ingles
Author:Amich, Francisco; Bernardos, Sonia
Date:Jan 1, 2013
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