Geobotanical Synopsis of Klasea and Serratula (Asteraceae) in the Iberian Peninsula

International Journal of Geobotanical Research, Vol. nº 2. 2012. pp. 21-33

Paloma CANTÓ

Departamento de Biología Vegetal II, Facultad de Farmacia, Universidad Complutense. 28040 Madrid, Spain.

Abstract

This contribution presents new data on geobotany of Klasea and Serratula (Asteraceae) on the Iberian Peninsula. For every recognized taxon on point out the habitat: sigmetum, secondary vegetation units, bioclimate, biogeography and distributions. Two new combinations are proposed in Klasea. From the syntaxonomical point of view, three new nomina correcta are also proposed. Conservation aspects are discussed and new taxa are suggested for the Red List of Spanish Vascular Flora.

Keywords: Klasea, Serratula, Geobotany, Iberian Peninsula

Introduction (2007) and CANTÓ (2009) in recognising the genera Serratula and Klasea. In the syntaxonomic scheme we The aim of this contribution is to carry out a geobo- have followed RIVAS-MARTÍNEZ & AL. (2002) and tanical synopsis of the Klasea and Serratula in the Ibe- RIVAS-MARTÍNEZ (2011); we have also taken account of rian Peninsula. other contributions by CAPELO & AL. (2002), COSTA & While the generic delimitation within Cardueae and AL. (2002), LOUSA & AL. (2002), PINTO-GOMES & PAIVA the phylogeny of Klasea and Serratula have been studied FERREIRA (2005), PINTO- GOMES & AL. (2007) and (SUSANNA & AL., 2006; MARTINS, 2006), the geobotany PINTO-GOMES & AL. (2010). We follow RIVAS- of this tribe are relatively little known. However, various MARTÍNEZ (2007) for the biogeographic nomenclature; authors highlight the importance of this type of contribu- and for the distribution of political provinces we have tion: for example, the contributions of DEIL (2006, 2007) followed the codes of CASTROVIEJO & AL. (1986). We on Ptilostemum and Phonus, which are particularly determined the phytocoenological behaviour through the worth noting, and constitute a step forward in this type of relevés of the communities of Serratula and Klasea research. species found. We have followed the methodology of the After our recent revision of the worldwide natural Zürich-Montpellier school (BRAUN-BLANQUET, 1979; biogeographic and bioclimatic distribution of Serratula GÉHU & RIVAS-MARTÍNEZ, 1981), also taking into and Klasea (CANTÓ, 2011), we now provide a more account data from other authors. detailed study of the phytosociological behaviour of these taxa on the Iberian Peninsula. According to Results MARTINS (2006), the origin of the genus Klasea may be located in western Asia; however, the proliferation of Serratula tinctoria species in the western Mediterranean subregion, and Serratula tinctoria L. (Figure 1) is a Eurosiberian specifically on the Iberian Peninsula, points to this area species [it extends as far as the Mediterranean region of as another possible genetic centre. On the Iberian Penin- the Iberian Peninsula, Sicily and according to QUÉZEL & sula the range of bioclimatic zones, thermotypes and SANTA (1963) to Algeria in northern Africa] which is ombrotypes, combined with the variability of substrates adapted to different habitat conditions. In the Iberian and soil, particularly favours the high number of ende- Peninsula, it prefers acid or neutral soils developed on mic species. siliceous rocks, but also grows on calcareous substrata with low levels of carbonates. It lives in deciduous fo- Methods rests (Quercetalia roboris) of Quercus petraea, Q. pyre- In the current taxonomic scheme we agree with naica, Q. pubescens, Q. robur, Fagus sylvatica and their GREUTER (2003), GREUTER & VON RAAB-STRAUBE substitution communities, such as Nardus stricta grass- (2005), MARTINS (2006), SUSANNA & GARCÍA JACAS lands, meadows and pastures of Molinion coeruleae, or

Corresponding author: Paloma Cantó. Departamento de Biología Vegetal II, Facultad de Farmacia, Universidad Complutense. 28040 Madrid, Spain. [email protected] ISSN: 2253-6302 (print)/ISSN: 2253-6515 (on line) ©Editaefa DOI: 10.5616/ijgr 120003 22 Paloma Cantó

Serratula tinctoria 50-2000 m Temperate (temperate submediterranean) Pulmonario longifoliae-Querco pyrenaicae sigmetum Genistion floridae Linario triornithophorae-Querco pyrenaicae sigmetum Genistenion polygaliphyllae Carici sylvaticae-Fago sylvaticae sigmetum Serratulo tinctoriae-Nardetum strictae (decalcifitation) Polysticho setiferi-Fraxino excelsioris sigmetum Daboecio-Ulicetum gallii

Sileno maritimae-Ulicetum humilis Genisto occidentales-Ulicetum maritimi Mediterranean Fraxino angustifoliae-Querco pyrenaicae sigmetum Vicio sepio-Centaureetum carpetanae (edaphic compensation) Cynosurion cristati Melico uniflorae-Betulo celtibericae sigmetum Juncion acutiflori Table 1: Phytosociological synopsis of Serratula tinctoria on the Iberian Peninsula heathlands and moorlands (“brezales”, “tojares” and “brezal-tojares”). It also occurs in mixed edaphohygrophilous ash and oak forests through soil compensation (and in their borders and substitution stages). FER- NÁNDEZ GONZÁLEZ & SÁNCHEZ MATA IN SÁNCHEZ MATA (1989) propose Serratula tinctoria as a characteristic species of the association Vicio sepium-Centau- reetum carpetanae, which corresponds to the mesophi- lous perennial herbaceous borders of the forest ecosys- tem in these ash groves. In the Sistema Central range, FERNÁNDEZ GONZÁLEZ & A. MOLINA (1988) described the subassociation Querco pyrenaicae-Fraxinetum angustifoliae serratuletosum tinctoriae in the bottoms and foothills of the Lozoya Valley. S. tinctoria also lives in birch groves (Betula celtiberica) with a somewhat edaphohygrophilous character. Both on the Iberian Pe- ninsula and outside, we agree with RIVAS-MARTÍNEZ Figure 1: Serratula tinctoria in Pulmonario longifoliae- (1979) and with OBERDORFER (1983), respectively, that Quercetum pyrenaicae this macrospecies may be considered to be an bioindica- and in the Pyrenees [Esp.: B L], in subalpine pastures, tor species of late-mown nutrient-poor Molinion grass- alpine grasslands and rocky slopes between 1000 and lands, although it is also known in other parts of central 2000 m. Finally, the var. seoanei corresponds to extreme Europe in more nutrient-rich flood-meadows of the forms at the westernmost limit of the species' distribu- alliance Cnidion (BURKART 1998; HÖLZEL 1999, tion, and is well represented in ultrabasic and peridotitic BISSELS & AL. 2004). S. tinctoria prefers the Temperate serpentinized rocks in the province of La Coruña from macrobioclimate, but can also live in the Mediterranean the mountains of Cabo Ortegal to El Ferrol; and found Pluviseasonal-Oceanic bioclimate. Eurosiberian: Atlantic more rarely, in coexistence with the var. tinctoria in European, Central European, Pannonio-Carpatian, Atla- other territories in Western Europe: northwest of the sian, Cevenno-Pyrenean, Apennino-Balkan, Sarmatian, Iberian Peninsula, southwest France and southern En- Mediterranean: Mediterranean West Iberian, Mediterra- gland [Esp.: B Bi Bu C Ge Gu Lu O S SS Vi. Port.: BL nean Central Iberian, Balearic-Catalonian-Provençal. DL Mi (TM)]. The distribution of the political provinces in the Iberian Peninsula is: Esp.: B Bi Bu C Ge Gu Hu L Le Lo Lu M Klasea nudicaulis Na O Or P Po S Sg So SS T Vi Z. Port.: BL DL Mi Klasea nudicaulis (L.) Fourn. grows on basophilous, (TM). 50-2000 m. VII-XI. (Table 1). orophilous and quionophobous pastures and cushion-like Serratula tinctoria is an extraordinarily polymor- thickets, living on shallow soils, often suffering cryotur- phous species in its habitat and leaf morphology, which has favoured the recognition of infraspecific taxa and bation processes during the cold period. It can also be even species, most of which can be considered to be found on windy crests with a minimal winter snow layer, phenetic responses to the environment and to the cha- for example on Pyrenean flysch with calcareous borders. racteristics of the soils on which they grow. However in This species is a bioindicator of basophilous grasslands the Iberian Peninsula we accept three varieties. The var. developed on soils with ephemeral snow cover, often tinctoria is well represented on the Iberian Peninsula in with cryoturbation and without gleyic character on Fes- all the species' environments, from coastal promontories tuco hystricis-Ononidetea striatae communities. to heights of 2000 m. The var. alpina lives in high cen- In the Castilian-Cantabrian sector, it has been found tral and southern European mountains, particularly do- in the series Spiraeo obovatae-Querco fagineae sigme lomitic (Austria, Croatia, France, Italy and Switzerland), tum, both in Plantagini discoloris-Thymion mastigophori Geobotanical Synopsis of Klasea and Serratula (Asteraceae) in the Iberian Peninsula 23

Klasea nudicaulis 500-2800 m Temperate and Mediterranean

Plantagini-Thymion mastigophori (on cryoturbated soils) Spiraeo obovatae-Querco fagineae sigmetum Arctostaphilo-Genistetum occidentalis (on deeper soils)

Echinosparto horridi-Pino pyrenaicae sigmetum Serratulo nudicaulis-Asperuletum pyrenaicae Mediterranean Junipero sabinae-Pino ibericae sigmetum Erodio celtibericae-Erinacetum anthyllidis Festucetum hystricis Junipero sabinae-Pino latisquamae s. Daphno hispanicae-Pino nevadensis sigmetum Seseli granatensis-Festucetum hystricis Lino differentis-Salvietum lavandulifoliae Cephalanthero rubrae-Querco fagineae sigmetum helianthemetosum cani Characteristic of Festuco hystricis-Ononidetea striatae Table 2: Phytosociological synopsis of Klasea nudicaulis on the Iberian Peninsula

In the Subbetic sector, K. nudicaulis is found in su- pra- and oromediterranean areas with its optimum in a cacuminal topographic position, integrated in the association Seseli granatensis-Festucetum hystricis (Seseli granatensis-Festucion hystricis), with plants such as Seseli granatensis, Helianthemum canum and Festuca hystrix begins to Junipero sabinae-Pino latisquamae sigmetum. Mediterranean and temperate submediterranean bio- climates. Eurosiberian region: Atlantic European, Alpi- ne, Cevennese-Pyrenean, Apennine-Balkan provinces; and Mediterranean region: Betic, Mediterranean West Iberian, Mediterranean Central Iberian, Rifean-Tangerie- ran, Atlasian [SW Europe (Spain, France, Italy) and NW Africa (Morocco)]. From the supramediterranean belt (eventually mesomediterranean) to oromediterranean and Figure 2: Klasea nudicaulis in grasslands of Guadarrama orotemperate submediterranean. Esp.: A Ab Al B Bu Cs Sierran sector Cu Gr Gu Hu J L Le Lo Lu M Na Or P S Sa Sg So T Te V Va Vi Z Za. 500-2800 m. (V) VI-VII (VIII). (Table 2). with cryoturbation, and in Arctostaphilo crassifoliae- Klasea legionensis Genistetum occidentalis (Genistion occidentalis) in deeper soils. In the Iberian-Maestracense sector it has been Klasea legionensis (Lacaita) Holub, is an Iberian en- identified in Festucetum hystricis (Sideritido fontqueria- demic at the limit of the Atlantic European and Medite- nae-Arenarion microphyllae, Festuco-Poetalia ligulatae, rranean western-Iberian provinces. According to LADERO Festuco-Ononidetea) in the series Junipero sabinae-Pino & AL. (1985), this species occurs in perennial herb com- ibericae sigmetum, where in some sites it can be inte- munities with low moisture (Linarion triornitophorae) grated in the dwarf shrublands of Erodio celtibericae- on the outer fringes of woodlands in siliceous territories Erinacetum anthyllidis (Festuco-Ononidetea). In the in the southern Atlantic European and Carpetan-Leonese central Pyrenees, according to MONTSERRAT & VILLAR areas, in the Su and Supratemperate Submediterranean, (1987) it forms part of the association Serratulo nudi- humid and subhumid belts. caulis-Asperuletum pyrenaicae, which is typical of Deciduous acidophilous forests and preforests, forest windy summits with minimal snow cover or on slopes borders and dwarf shrublands; ● Northwest Iberian Pe- with moderate cryoturbation, above 1900 m, in Pyrenean ninsula. Esp.: Or Za. Port.: (Mi) TM. 900-1400 m. VII- flysch alternating with hard limestone. VIII. (Table 3). In calcareous islands in the Guadarrama Sierran sector, Klasea nudicaulis grows on basophilous pastures and Klasea pinnatifida cushion-like thickets which can be ascribed to the sub- Klasea pinnatifida (Cav.) Cass., is an Iberian-North association described by FERNÁNDEZ GONZALEZ (1991): African element [Iberian Península and NW Africa (Al- Lino differentis-Salvietum lavandulifoliae heliantheme- geria and Tunisia)]. Mediterranean basophilous forests tosum cani, although enriched with bioindicator elements and preforests, preferably sclerophilous, forest borders, of Festuco hystricis-Ononidetea, such as Anthericum "lastonares" (grasslands dominated by Brachypodium liliago, Fumana procumbens, Ranunculus gramineus retusum), thyme thickets and dwarf shrublands. Charac- var. gramineus, Inula montana, Seseli montanum; in all cases in the series of Guadarrama Sierran gall oak teristic of basophilous communities of open shrubs, forests. (Figure 2). usually with a preponderance of chamaephytes and low- 24 Paloma Cantó

Klasea legionensis 900-1400 m Mediterranean and submediterranean Holco mollis-Querco pyrenaicae sigmetum Carduo platypi-Festucetum durandii Characteristic of Linarion triornithophorae Endangered species “EN” Table 3: Phytosociological synopsis of Klasea legionensis on the Iberian Peninsula

Klasea pinnatifida 30-2200m Mediterranean Rubio longifoliae-Querco rotundifoliae sigmetum Teucrio-Thymenion piperellae Paeonio coriaceae-Querco rotundifoliae sigmetum Lavandulo-Echinospartion boissieri Daphno hispanicae-Pino nevadensis sigmetum Xeroacantho-Erinaceion anthyllioidis Saturejo-Velletum spinosae Junipero sabinae-Pino latisquamae s. Querco rotundifoliae sigmetum Sideritido-Salvion Characteristic of Rosmarinetea officinalis Table 4: Phytosociological synopsis of Klasea pinnatifida on the Iberian Peninsula

Klasea baetica subsp. baetica 30-1000 m Mediterranean Halimio atriplicifolii-Digitaletum laciniatae Pino acutisquamae-Querco cocciferae sigmetum

Paeonio coriaceae-Querco rotundifoliae sigmetum Asperulo asperrimae-Staehelinetum baeticae Characteristic of Staehelino-Ulicion baetici EN “endangered” Table 5: Phytosociological synopsis of Klasea baetica subsp. baetica on the Iberian Peninsula

Klasea baetica subsp. alcalae 30-900 m Mediterranean Teucrio baetici-Querco suberis sigmetum Genisto tridentis-Stauracanthetum boivini Characteristic of Genisto-Stauracanthetum boivinii

Table 6: Phytosociological synopsis of Klasea baetica subsp. alcalae on the Iberian Peninsula growing nanophanerophytes, on carbonate soils (Rosma- Klasea baetica rinetea officinalis), found in most alliances: Rosmarino- Ericion (Teucrio-Thymenion piperellae) in the Xativan Klasea baetica (Boiss.) Holub sensu lato, occurs in sector, Lavandulo-Echinospartion boissierii in the Betic subhumid and humid, thermo-mesomediterranean envi- province, Sideritido-Salvion in the Central Iberian Me- ronments in the south-western Iberian Peninsula and North Africa (Morocco, Algeria and Tunisia). Forests diterranean province and Xeroacantho-Erinaceion (Eri- nacetalia) in the Betic province, Xativan sector and NW and dwarf shrublands on acid or basic substrates; Esp.: Africa. (Figure 3). It is found from the thermomediterra- Ca Gr Ma. Port.: AAl Ag BL E R. 30-1000 m. V-VII. Three subspecies can be distinguished, forming a nean to the lower oromediterranean belt, particularly from dry to subhumid, in the mediterranean pluviseaso- pattern of geographical and soil vicariant taxa, although nal bioclimate; it also grows in semiarid to dry areas, in their areas of distribution may overlap in a few plot lo- cations. which case it can be found either in the "fenalar” (grass- K. baetica subsp. baetica grows on ultramaphic, ser- land dominated by Brachypodium phoenicoides), or in pentinised peridotitic and dolomitic rocks in southern contact with it or nearby, in search of some soil compen- Spain (mountains of Malaga and Ronda, and Sierra Elvi- sation. Abundant in the eastern part of the Iberian Penin- ra, Granada): Betican province and northwest Morocco sula and scattered in the west. Esp.: A Ab Al Ba Bu Ca (Rif): Rif-Tangerine province. In the Bermejense district Co CR Cs Cu Gr Gu Hu J Le Lo M Ma Mu Na P (S) Sa it is integrated in the series Pino acutisquamae-Querco (Se) Sg So Te To V Va (Vi) Z Za. Port.: AAl. 30-2200 cocciferae sigmetum; according to RIVAS GODAY & m. V-VII. (Table 4). RIVAS-MARTÍNEZ (1969) it forms part of the clear shru- Geobotanical Synopsis of Klasea and Serratula (Asteraceae) in the Iberian Peninsula 25

na variety is found in decarbonated sandy soils. In encla- ves with clayey-sandy soils, it is common to find transi- tional plants between both varieties. Klasea baetica (Boiss. ex DC.) Holub subsp. lusitanica (Cantó) Cantó & Rivas Mart. var. lusitanica (Cantó) Cantó, comb. nova [hoc loco] Basion.: Serratula baetica Boiss. ex DC. subsp. lusitanica Cantó f. lusitanica Cantó in Lazaroa 6: 52. 1985. [Ind. Loc.: “Portugal: Serra da Arrabida, solo calcareo”] Common in forests of holm oaks, gall oaks, kermes oaks, thyme, dwarf shrublands and perennial grasslands "lastonares" and "fenalares", preferably on calcareous substrates ● Port.: Ag BL E. It is endemic in the calcareous mountains of Portuguese Divisorian (Serra da Arrabida -Arisaro clusii-Querco broteroi sigmetum-, Serra do Aire and the Algarve (Querco alpestris-broteroi sigmetum). This taxon has its optimum in dwarf scrub vegetation such as for example Salvio sclareoidis-Uli- cetum densi. It can also be found in caespitose perennial grasslands over deep limy substrata and, in the opinion

Figure 3: Klasea pinnatifida on the Central Iberian Mediterranean province blands under cover of Pinus, rich in endemisms: Halimio atriplicifolii-Digitaletum laciniatae (Staehelino-Ulicion baetici, Cisto-Lavanduletea), with plants such as Hali- mium atriplicifolium subsp. serpentinicola, Genista la- nuginosa, Ulex baeticus, Digitalis obscura subsp. laci- niata. In the Rondense district it forms part of the series Paeonio coriaceae-Querco rotundifoliae sigmetum and is integrated in Asperulo asperrimae-Staehelinetum baeticae (Staehelino-Ulicion baetici, Cisto-Lavandule- tea), open shrublands with numerous chamaephytes which cover the rocks and denuded soils of the Carratra- ca mountains and other serpentine outcrops, with plants such as Linum carratracensis, Staehelina baetica, Cen- taurea carratracencis and Teucrium reverchonii. Esp.: Ma Gr. V-VII. (Table 5). K. baetica subsp. alcalae (Coss.) Cantó & Rivas Mart., colonize soils derived from Aljibe sandstone (Coastal Lusitano-Andalusian Province) and similar environments in the Rif-Tangerine and Constantine- Tunisian provinces. It is found in the series Teucrio baetici-Querco suberis sigmetum, forests of cork oak and Andalusian gall oak, thyme dwarf shrublands, prefe- Figure 4: Klasea baetica subsp. lusitanica var. lusitanica rably on sandstones. According to CANTÓ (1985) it is on Portuguese Divisorian sector characteristic of Genisto tridentis-Stauracanthetum boivinii. Southern Iberian Peninsula (mountains of Cadiz of PINTO-GOMES & AL. (2007), it is characteristic of and western Malaga) and NW Africa (Morocco, Algeria Avenulo occidentalis-Celticetum giganteae. It has also and Tunisia). Esp.: Ca Ma. 30-900 m. V-VII. (Table 6). been assigned (PINTO GOMES & PAIVA FERREIRA, 2005) K. baetica subsp. lusitanica (Cantó) Cantó & Rivas to Galio concatenati-Brachypodietum phoenicoidis. Accor- Mart., lives in forests of holm oaks, gall oaks, kermes ding to LOUSA & AL. (2002) it also thrives in the series of oaks, thyme thickets, dwarf shrublands and perennial the holm oak forests in the Portuguese Divisorian sector: grasslands, on calcareous substrates, and also in rock Lonicero-Querco rotundifoliae sigmetum, and the Algar- rose heathlands on sandy soils. ● W & S of Portugal. viense sector: Rhamno oleoidis-Querco rotundifoliae Port.: Ag AAl BL E R. 30-700 m V-VI. . (Table 7). This sigmetum. (Figure 4). is a highly polymorphous subspecies with regard to its Klasea baetica (Boiss. ex DC.) Holub subsp. lusita- leaf shape, but a certain correlation observed with ecolo- nica (Cantó) Cantó & Rivas Mart. var. sampaiana gical factors allows two varieties to be recognised. The (Cantó) Cantó, comb. nova [hoc loco] lusitanica variety prefers basic soils, while the sampaia- 26 Paloma Cantó

Basion.: Serratula baetica Boiss. ex DC. subsp. lusi- Ribatagus-Sadoan sector. The characteristics of the sub- tanica Cantó f. sampaiana Cantó in Lazaroa 6: 52. 1985. strate are that it is rich in bases, in contrast with the sur- [Ind. Loc.: “Portugal, Estremadura, entre Eiría y Alco- face sands, and has its optimum in two fundamental entre”] types in the shrubby communities Thymo camphorati- S. baetica subsp. lusitanica var. sampaiana (Cantó) Stauracanthetum spectabilis (Cisto-Lavanduletea, Stau- Cantó in Lagascalia 15 (extra): 380 (1988) racantho genistoidis-Halimietalia commutati, Corema- It lives in forests of cork oaks, Portuguese gall oaks, tion albi), with psamophilous character, or else in the oaks, rock rose thickets and heathlands, growing on endemic association of San Vicente Cape district, grow- sandy soils or on acid substrates (with a predominance of ing on limy soils covered by windblown sands: Ulicetum granites) or slightly basic, but highly decarbonated and erinacei (Eryngio-Ulicion erinacei, Rosmarinetalia offici- with a degree of hydromorphy, humid thermo-mesome- nalis, Rosmarinetea officinalis), as indicated by RIVAS- diterranean ● Port.: Ag AAl BL E R. (Table 7). MARTÍNEZ & AL. (1990). It can also be found in other serial stages of Aro neglecti-Querco suberis sigmetum, Klasea integrifolia group such as in the caespitose perennial grasslands on sandy The Klasea integrifolia group occurs in the south- soils of varying depths. In this regard, other authors western Iberian Peninsula; the common denominator is (PINTO-GOMES & AL., 2007) have ascribed it to the the sandy soil in thermo- and mesomediterranean ther- grasslands Avenulo hackelii-Celticetum sterilis (sub motypes. This group comprises two species: Klasea Serratula monardii) or subsequently (PINTO-GOMES & algarbiensis and Klasea integrifolia diversified into two AL., 2010) to Armerio macrophyllae-Celticetum gigan- subspecies: subsp. integrifolia and subsp. monardii. teae (sub Serratula monardii); according to CAPELO & Klasea algarbiensis AL. (2002), it is also integrated in Centaureo crocatae- Quercetum lusitanicae; ● SW Iberian Peninsula. Port.: K. algarbiensis (Cantó) Cantó, lives on windy sands Ag E. 5-100 m. IV-V. (Table 8). deposited over Cretaceous and Jurassic substrata in the coastal area of the Algarve and the southern coast of the

Klasea baetica subsp. lusitanica var. lusitanica 30-700 m Mediterranean Melico arrectae-Quercetum cocciferae Arisaro clusii-Querco broteroi sigmetum Salvio sclareoidis-Ulicetum densi Oleo sylvestris-Querco broteroi sigmetum Avenulo occidentalis-Celticetum giganteae Lonicero implexae-Querco rotundifoliae sigmetum Teucrio capitati-Thymetum sylvestris Querco alpestris- broteroi sigmetum Thymo lotocephali-Thymbretum capitatae Rhamno oleoidis-Querco rotundifoliae sigmetum Phlomido purpureae-Cistetum albidi Characteristic of Klaseo-Thymenion Klasea baetica subsp. lusitanica var. sampaiana 30-700 m Mediterranean Erico scopariae-Quercetum lusitanicae Asparagp aphylli-Querco suberis sigmetum Junipero navicularis-Quercetum lusitanicae Lavandulo viridis-Querco suberis sigmetum Lavandulo luisieri-Ulicetum jussiaei

Thymo capitellati-Stauracanthetum genistoidis Table 7: Phytosociological synopsis of Klasea baetica subsp. lusitanica on the Iberian Peninsula

Klasea algarbiensis 5-100 m Mediterranean Aro neglecti-Querco suberis sigmetum Thymo camphorati-Stauracanthetum spectabilis Osyrio quadripartitae-Junipero turbinatae s. Ulicetum erinacei Table 8: Phytosociological synopsis of Klasea algarbiensis on the Iberian Peninsula

Klasea integrifolia subsp. integrifolia 30-1000 m Mediterranean Arbuto unedonis-Querco pyrenaicae sigmetum Halimio ocymoidis-Ericetum umbellatae Sanguisorbo hybridae-Querco suberis sigmetum Characteristic of Ericenion umbellatae Table 9: Phytosociological synopsis of Klasea integrifolia subsp. integrifolia on the Iberian Peninsula Geobotanical Synopsis of Klasea and Serratula (Asteraceae) in the Iberian Peninsula 27

Klasea integrifolia AL. (2002) and COSTA & AL. (2002) it thrives in Erico- Klasea integrifolia (Vahl) Greuter, is diversified into Quercetum lusitanicae, and it could be considered to be a two subspecies: subsp. integrifolia and subsp. monardii. bioindicator of these communities, or a differential ele- The first is the most continental taxon in this group, and ment. At the northern limit of its distribution it can be is an endemism in sabulicolous communities (“thickets found in the series Sanguisorbo hybridae-Querco suberis of Cistus heathlands) in C, SW and S Spain; K. integri- sigmetum. Other authors (PINTO-GOMES & AL., 2007) folia subsp. monardii is an endemism in C, W and SW have ascribed it to the caespitose perennial grasslands of Iberian Peninsula, and occurs on sandy soils of the Plio- plio-pleistocene sandy soils with Euphorbio transtag- cene but also on paleodunes, and ranges from sabulico- anae-Celticetum giganteae and Avenulo hackelli-Celti- lous communities on red limestone in Conil-Chiclana to cetum sterilis. ● C, W and SW of the Iberian Peninsula. centre-western Portugal. (Figure 5). Forests of cork oak, Esp.: Ca H Za. Port.: Ag BA BAl AAl BL E R TM. 20- oak, pine forests, forest borders, heathlands, thickets of 600 m. V-VI. (Table 10). rock rose and Cistus shrubs; ● C, W and SW of the Ibe- rian Peninsula. Esp.: Av Ba Ca Cc CR H J Sa To Za. Port.: Ag BA BAl AAl BL E R. 20-1000 m. V-VII. K. integrifolia subsp. integrifolia Forests of cork oak, oak, pine forests, forest borders, heathlands and rock rose thickets; integrated mainly in the series Arbuto-Querco pyrenaicae sigmetum or also in cork oak forests Sanguisorbo hybridae-Querco suberis sigmetum. (Figure 6). In Extremadura it has its optimum in the heathlands of Halimio ocymoidis-Ericetum umbellatae ericetosum australis; ● C, SW and Southern Spain. Esp.: Av Ba Cc CR H J Sa To. 30-1000 m. V-VII. (Table 9).

Figure 6: Klasea integrifolia subsp. integrifolia on Montitoledan-Marianican territory

Klasea flavescens Klasea flavescens (L.) Holub sensu lato, comprises a group of stenochorous taxa in semi-arid to arid steppe Figure 5: Klasea integrifolia subsp. monardii on Sandy environments in south-western Europe and northern soils of Conil-Chiclana Africa; however in their distribution limits, it is some- times difficult to separate these subspecies, and there is K. integrifolia subsp. monardii (Dufour) Cantó some overlap. Klasea flavescens subsp. flavescens is a Forests of cork oak, pine, forest borders, thickets of Manchan endemism (CR, M, To) which extends as far as rock rose and Cistus developing preferentially on sandy the Betic province (Gr, J, Se), and grows on clayey or or silty-sandy soils; it is found integrated in the series clayey and gypsum soils with nitrogen influence, and is Aro neglecti-Querco suberis sigmetum and southwest characteristic of the Onopordion castellani alliance. K. Iberian sandy sites with Halimium halimifolium and in flavescens subsp. mucronata develops in Triassic limes- Asparago aphylli-Querco suberis sigmetum. It is found tone, phillites and volcanic soils, and is a western Medi- above all in substitution rock rose thickets (Coremation terranean element: S Iberian Peninsula (Murcian-Alme- albi, Stauracantho-Halimietalia commutati, Cisto-La- rian and Betic Provinces, Al, Gr, Mu), N Africa (Mo- vanduletea), for example in Stauracantho-Corematetum rocco, Algeria and Tunisia: Muluyan-Cabilian Province), albi in Ribatagano-Sadense, or in thickets of Cistus on and Sicily (Italo-Thyrrhenian Province). K. flavescens paleodunes in the Huelva area: Halimio halimifolii-Stau- subsp. leucantha grows on soils developed on marls or racanthetum genistoidis, and also in the rock rose marley-gypsicolous materials, and is an eastern and thicket: Halimio commutati-Cistetum bourgaeani, typi- southern Iberian endemism (Bardenan-Monegrese, Xa- cal of the zones of contact between the sandy sites of the tivan, Valencian-Tarragonian and Betic). K. flavescens Guadalquivir river and the limy-reddish sediments of the subsp. neglecta is an endemism in the southern coastal mountains; it can also be found (COSTA & AL., 2002) in territories of the Iberian Peninsula: Spain (Ma, Gr) and the heathlands-moorlands: Halimio lasianthi-Ulicetum Portugal (Al); it colonises neutral soils developed on minoris (Ericion umbellatae). According to CAPELO & dolomitic rocks. 28 Paloma Cantó

Klasea integrifolia subsp. monardii 20-600 m Mediterranean Stauracantho genistoidis-Corematetum albi Aro neglecti-Querco suberis sigmetum Halimio halimifolii-Stauracanthetum genistoidis Halimio calycinii-Cistetum bourgaeani Characteristic of Coremation albi EN “endangered”

Table 10: Phytosociological synopsis of Klasea integrifolia subsp. monardii on the Iberian Peninsula

Klasea flavescens subsp. flavescens 350-1500 m Mediterranean Asparago acutifolii-Querco rotundifoliae sigmetum Onopordion castellani Characteristic of Onopordion castellani

Table 11: Phytosociological synopsis of Klasea flavescens subsp. flavescens on the Iberian Peninsula

Klasea flavescens subsp. mucronata 5-1850 m Mediterranean Sideritido marminorensis-Thymetum hyemalis Mayteno europaei-Periploco angustifoliae sigmetum Sideritido osteoxyllae-Teucrietum charidemi Phlomido almeriensis-Ulici canescentis Odontito purpurei-Thymetum baetici Minorisigmetum (permanent character) Characteristic of Thymo moroderi-Sideritidion leucanthae

Table 12: Phytosociological synopsis of Klasea flavescens subsp. mucronata on the Iberian Peninsula

Klasea flavescens subsp. leucantha 50-500(1100) m Mediterranean Teucrio verticillati-Thymetum pallentis Chamaeropo humilis-Rhamno lycioidis sigmetum Thymo ciliati-Teucrietum verticillati Helianthemo alypoidis-Gypsophiletum struthii Rhamno lycioidis-Querco cocciferae sigmetum Helianthemo thibaudi-Gypsophiletum hispanicae Characteristic of Gypsophiletalia Strictly protected (Catalonia, Decree 328/1992, DOGC of 1 March 1993 (sub S. flavescens)

Table 13: Phytosociological synopsis of Klasea flavescens subsp. leucantha on the Iberian Peninsula

Klasea flavescens subsp. neglecta 10-1000 m Mediterranean Cisto clusii-Ulicetum rivasgodayani Vinco difformis-Ceratonio siliquae sigmetum Thymo lotocephali-Thymbretum capitatae Rhamno oleoidis-Querco rotundifoliae sigmetum Phlomido purpureae-Cistetum albidi Ulicetum erinacei Characteristic of Eryngio-Ulicion erinacei

Table 14: Phytosociological synopsis of Klasea flavescens subsp. neglecta on the Iberian Peninsula

Geobotanical Synopsis of Klasea and Serratula (Asteraceae) in the Iberian Peninsula 29

Hawthorn thickets with dwarf palms, "cornicales" found in other territories of the East of the Iberian Penin- (shrubs with Periploca angustifolia), forests of kermes sula, for example in the semiarid mesomediterranean oaks, thyme dwarf shrublands, grasslands, etc., in soils Low Aragonian (Helianthemo thibaudii-Gypsophiletum with varying degrees of nitrification, neutral or basic, hispanicae, Gypsophilenion hispanicae). ● E and SE more rarely acid due to decarbonation; W of the Medite- Spain. Esp.: A Ab Al B Cs Cu Gr Hu L Lo Ma Mu Na T rranean region. Esp.: A Ab Al B CR Cs Cu Gr Hu J L Lo V Z. 50-500(1100) m. V-VII. (Table 13). M Ma Mu Na Se T To V Z. Port.: Ag. 5-1850 m. IV- VII. K. flavescens subsp. flavescens It forms part of the Manchan series: Asparago acutifolii-Querco rotundifoliae sigmetum on calcareous or loamy soils, but in degraded stages of the series: roadside grasslands with greater or lesser degree of nitrification and ruderal and nitrophilous communities growing on deep dry calcareous or loamy soils dominated by "car- dueas": Onopordion castellani. (Figure 7). ● C and S Spain. Esp.: CR Gr J M Se To. 350-1500 m. V-VII. (Table 11).

Figure 8: Klasea flavescens subsp. mucronata on Almerian sector

Figure 7: Klasea flavescens subsp. flavescens on Manchan sector K. flavescens subsp. mucronata (Desf.) Cantó & Ri- vas Mart. Hawthorn thickets with dwarf palms, "cornicales", thyme dwarf shrublands, and "lastonares" (grasslands with Brachypodium retusum), etc., in calcareous, clayey soils, often volcanic in character, with neutral or weakly basic pH, in the thermomediterranean murcian-almerian; for example in the mining soils of Portman and Calblanche in Cartagena or in the mountains of Cabo de Gata, Rodalquilar mines: Sideritido marminorensis- Thymetum hyemalis and Sideritido osteoxyllae-Teucrie- tum charidemi (Thymo moroderi-Sideritidion leucan- Figure 9: Klasea flavescens subsp. leucantha in Helian- thae). (Figure 8). It reaches the Alpujarra, in terra rosa- themo alypoidis-Gypsophiletum struthi type soils (chromic luvisol), on limestone in association with phyllites: Odontito purpurei-Thymetum baetici K. flavescens subsp neglecta (Iljin) Greuter & Wage- (Saturejo-Thymbrenion). South of the Iberian Peninsula, nitz N Africa (Morocco, Algeria and Tunisia), and Sicily. Forests of holm and kermes oaks, thyme dwarf Esp.: Al Gr Mu. 5-1850 m. IV-VI. (Table 12). shrublands, grasslands, etc., in neutral or basic soils, pre- K. flavescens subsp. leucantha (Cav.) Cantó & Rivas ferentially on limestones and dolomites, less frequently Mart. in acid soils due to decarbonation; it is found in Cisto Thyme dwarf shrublands, preferentially on loams and clusii-Ulicetum rivasgodayani, thermomediterranean gypsum; in the thermomediterranean almerian: (Helian- Almijara Sierran thyme dwarf shrublands typical of do- themo alypoidis-Gypsophiletum struthii, Gypsophilo- lomitic substrates and in Thymo lotocephali-Thymbretum Santolinenion viscosae) and semi-arid thermo-mesome- capitatae, thyme dwarf shrublands on calcareous wea- diterranean alicantese-murcian (Thymo ciliati-Teucrie- thered sites, in eroded terra rosa, sporadically in these tum verticillati, Teucrio verticillati-Thymetum pallentis, communities in the Algarve on dolomitic terra rosa soils, Thymo-Teucrienion verticillati). (Figure 9). It is also (Saturejo-Thymbrenion); according to PINTO-GOMES & 30 Paloma Cantó

Map 1: ● Serratula tintoria. ○ Klasea baetica subsp. baetica. ▲ Klasea baetica subsp. alcalae ∆ Klasea baetica subsp. lusitánica. Klasea algarbiensis. ■ Klasea integrifolia subsp. integrifolia. □ Klasea integrifolia subsp. monardii

Map 2: Klasea nudicaulis. Klasea legionensis. ○ Klasea pinnatifida. ● Klasea flavescens subsp. flavescens. ■ Klasea flavescens subsp. mucronata. □ Klasea flavescens subsp. leucantha. Klasea flavescens subsp. neglecta Geobotanical Synopsis of Klasea and Serratula (Asteraceae) in the Iberian Peninsula 31

PAIVA FERREIRA 2005, its behaviour is broader and it red, possibly disappeared in the region). We consider also thrives in the dwarf shrublands of Eryngio-Ulice- that S. tinctoria –at least in Madrid– should be conside- nion erinacei; it can thus be considered characteristic of red with the VU category at the regional level, as the the alliance Eryngio-Ulicion ereinacei, which comprises southern European limit of the species. In contrast, S. both suballiances. It is also found in Phlomido purpu- tinctoria in Madrid is linked to relic forests of beech and reae-Cistetum albidi and in Thymo lotocephali-Thym- oak, or else finds refuge in wetlands. bretum capitatae. Other studies by PINTO-GOMES & AL. Since Klasea legionensis was included on the 2000 (2010) transfer it to the perennial grasslands over deep Red List as a species with the category of UICN Vulner- limy substrata: Serratulo flavescentis-Celticetum gigan- able “VU” (criteria: B1+2b, D2), numerous authors have teae (sub Serratula flavescens s.l.). S of the Iberian Pe- defended their proposals for greater conservation in ninsula and NW Africa (Morocco). Esp.: Gr Ma. Port.: Galicia (ORTIZ & AL., 1998; ROMERO BUJÁN, 2007; Ag. 10-1000 m. V-VII. (Table 14). SILVA PANDO & AL. 2008). Currently Klasea legionensis is catalogued on the 2008 Red List of Spanish Vascular Conclusions Flora [MORENO SAIZ (COORD.)] as an endangered species Serratula and Klasea species are found in specific “EN” (criteria: B1ab (i, iii, iv) + 2ab (i, iii, iv). vegetation series, adapted to forest vegetation in the case K. baetica subsp. baetica is classified both on the of Serratula, and can be adapted to forest vegetation type 2000 Red List and on the 2008 Red List of Spanish Vas- or a non-forest vegetation type in the case of Klasea. cular Flora as EN “endangered” (criteria B1 + 2d) and D Serratula tinctoria has its optimum in deciduous forests respectively. We propose the consideration of endange- and in their substitution communities. red taxon EN for K. baetica subsp. alcalae, taking into Klasea nudicaulis (Klasea sect. Klasea) is constant in account its reduced distribution area: S of the Iberian orophilous and quionophobous pastures and cushion-like Peninsula (mountains of Cadiz and western Malaga) and dwarf shrublands, living on shallow soils, of the Festuco NW Africa (criteria B2). hystricis-Ononidetea striatae. K. integrifolia subsp. monardii is an endemism of the The Klasea species sect. Demetria are elements in C, W and SW the Iberian Peninsula. It appears (sub dwarf and open shrubland communities. Their ombro- Klasea monardii) on the Red List of Vascular Flora of climatic distribution ranges from the arid in SW Spain to Andalusia (CABEZUDO & TALAVERA, COORDS., 2005) the humid in the western Iberian Peninsula. These com- and it is listed (sub Serratula monardii) in the Catalogue plexes are not only phylogenetically close, but also have of Protected Flora of Castile-León (Decree 63/2007 of 14 certain characteristic life strategies and ecological requi- June) as a species for "preferential attention". This taxon rements in common. We often find grassland vegetation is considered in the 2008 Red List of Spanish Vascular in spatial contact with Klasea-communities, dominated Flora with the category of “endangered EN" [criteria by Stipa tenacissima in the east or Celtica gigantea in B1ab (i, ii, iii, iv, v)+2ab (i, ii, iii, iv, v)]. We agree with the west; or in some cases Quercus lusitanica communi- this consideration and propose, for the same reasons, the ties, presenting a more or less similar mosaic. Within same protection treatment for the endemism of the C, section Demetria, we concede two extremes of adapta- SW and S of Spain, K. integrifolia subsp. integrifolia, a tion: a) towards xeric conditions (aggr. flavescens); and taxon which is mentioned in the regional catalogue of b) towards subhumid-humid conditions: group baetica endangered plant species of Extremadura (PALACIOS & and group integrifolia. Their subspecies are stenochorous AL., 2010), as is Klasea monardii. It should also be and their areas do not overlap, or overlap only slightly. pointed out that the photograph shown in this work These groups form patterns of geographical vicariant (PALACIOS & AL. 2010: 272) does not in fact correspond taxa. In contrast, the endemism K. legionensis is found in to Klasea integrifolia subsp. monardii (K. monardii), but a very restricted area. K. pinnatífida is widespread to Klasea pinnatifida; This last species is not included in throughout the Peninsula, and can adapt to more or less the Red List of Spanish Flora, although it could be con- mesophytic conditions. sidered with the special category of “rare” at the regional From the point of view of distribution, if there are level in Extremadura, as Klasea pinnatifida is restricted two or more different taxa in Klasea in a territory with to limestone sites. similar environmental conditions (soil and macrobiocli- K. flavescens subsp. leucantha is protected at the matic characteristics), each one specializes in a biocli- level of the autonomous regions: strictly protected (Ca- matic level. For example: in a transect from the Betic talonia, Decree 328/1992, DOGC of 1 March 1993: Province (Sierra Nevada mountains) to the Murcian- yermos de Aitona (cf DEVESA & ORTEGA OLIVENZA, Almerian province (Cabo de Gata mountains): Klasea 2004). Furthermore, BARRENO & AL. (1985) assigned it nudicaulis (oromediterranean) --- Klasea pinnatifida the special consideration of “rare”. In both cases this (supramediterranean) --- Klasea flavescens subsp. leu- taxon was identified as Serratula flavescens, although cantha (mesomediterranean) --- Klasea flavescens subsp. once the specimens were studied, we confirmed that they neglecta (thermomediterranean) --- Klasea flavescens were not in fact Klasea flavescens subsp. flavescens but subsp. mucronata (thermomediterranean). Klasea flavescens subsp. leucantha. In our opinion this subspecies warrants a national consideration of VU (cri- Implications for conservation teria B and C). On the other hand, Klasea flavescens subsp. flavescens, an endemism of the central and sout- S. tinctoria, cited in Algeria as “rare” (QUEZEL & hern Iberian Peninsula, which has very few populations SANTA, 1963), is catalogued on the regional level in in its area of distribution, is also susceptible to protection France in Nord Pas de Calais as CR* (critically endange- and should appear on the Red List of Spanish Vascular 32 Paloma Cantó

Flora. Klasea flavescens subsp. mucronata (sub Serra- K. algarbiensis (Cantó) Cantó in Ann. Bot. Fenn. 46: tula mucronata) is considered as a taxon of special inter- 437. 2009. est in the Murcia Red Book (SÁNCHEZ & AL., 2002); K. flavescens (L.) Holub in Folia Geobot. Phytotax. 12: these three subspecies are either endemisms or stenocho- 305. 1977. rous, and with a very specific habitat, and should be subsp. flavescens classified as endangered EN or as vulnerable VU. subsp. mucronata (Desf.) Cantó & Rivas Mart. in Lazaroa 5: 319. 1984. Syntaxonomy and nomenclature: proposal of “nomi- subsp. leucantha (Cav.) Cantó & Rivas Mart. in Laza- na correcta” roa 5: 319. 1984. We propose the follow corrections: subsp. neglecta (Iljin) Greuter & Wagenitz in Willde- - Klaseo nudicaulis-Asperuletum pyrenaicae P. novia 33: 59. 2003. Montserrat & Villar 1987 nom. mut. propos. hoc loco Acknowledgements [Serratulo nudicaulis-Asperuletum pyrenaicae P. Mont- serrat & Villar 1987 (art. 45)]. Correct name: Serratula I wish to thank Salvador Rivas-Martínez for his kind nudicaulis (L.) DC. should be Klasea nudicaulis (L.) help. Fourn. - Klaseo lusitanicae-Thymenion sylvestris Capelo, References J.C. Costa, Espirito-Santo & Lousa 1993 nom. mut. Barreno, E., Bramwell, D., Cabezudo, B., Cardona, propos. hoc loco [Serratulo estremadurensis-Thymenion M.A., Costa, M., Fernández Casas, J., Fernández-Ga- sylvestris Capelo, J.C. Costa, Espirito-Santo & Lousa liano, E., Fernández-Prieto, J.A., Gómez Campo, C., 1993 (art. 45)]. Correct name: Serratula estremadurensis Hernández Bermejo, E., Heywood, V.H., Izco, J., Franco should be Klasea baetica subsp. lusitanica Llorens, L., Molero Mesa, J., Montserrat, P., Rivas- (Cantó) Cantó & Rivas-Martínez. Martínez, S., Sáenz Laín, C., Santos Guerra, A., Val- - Klaseo neglectae-Celticetum giganteae C. Pinto dés, B. & Wildpret, W. 1985. Listado de plantas endé- Gomes & R. 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Holub in Folia Geobot. Phy- Castroviejo, S., Laínz, M., López González, G., Montse- totax. 12: 305. 1977. rrat, P., Muñoz Garmendia, F., Paiva, J. & Villar, L. subsp. baetica 1986. Flora Iberica. Plantas vasculares de la Península subsp. alcalae (Cosson) Cantó & Rivas Mart. in Laza- Ibérica e Islas Baleares. Vol. I. Real Jardín Botánico de roa 5: 319. 1984. Madrid. Madrid. subsp. lusitanica (Cantó) Cantó & Rivas Mart. in La- Costa, J.C., Capelo, J., Lousa, M. & Espirito Santo, M.D. zaroa 5: 319. 1984. 2002. Os sobreirais do Sector Divisorio Portugues: As- var. lusitanica (Cantó) Cantó parago aphylli-Quercetum suberis. Quercetea 3: 81-98. var. sampaiana (Cantó) Cantó Deil, U. 2006. Distribution, ecology and phytosociology K. integrifolia (Vahl) Greuter in Willdenowia 35: 235. of the N Moroccan endemic Ptilostemon leptophyllus 2005. (Compositae). Willdenowia 36: 1-10. subsp. integrifolia Deil, U. 2007. Habitat shift as a successful strategy – an subsp. monardii (Dufour) Cantó in Ann. 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