zyxwvutsrqp zyxwvu zyxwv zy Botanical Journal of the Linnean Society (1990), 103: 325-349. With 3 figures Infrasectional systematics of the genus Sideritis L. section Sideritis (Lamiaceae) D. RIVERA NUREZ, C. OBON DE CASTRO Departamento de Biologia Vegetal, Facultad de Biologia, Universidad de Murcia, Spain F. TOMAS-LORENTE, F. FERRERES AND F. A. TOMAS BARBERAN zyxwvuts zyx zyx Laboratorio de Fitoquimica, C.E.B.A.S.-C.S.I.C., Apdo 195, Murcia, Spain Received January 1989, revised and accepted for publication M a y 1989 RIVERA NUmEZ, D., OBON DE CASTRO, C . , TOMAS-LORENTE, F., FERRERES, F. & TOMAS-BARBERAN, F. A,, 1990. Infrasectional systematics of the genus Sideritis L. section Sideritis (Lamiaceae).A new taxonomic division of the section Sideritis is proposed on the basis of morphological, cytological and chemical characters. T h e following subsections are recognized: GrandiJlora, Ouata, Camarae, Linearifolia, (Gmnocarpne, Stachydioides, Lacailae, Hirmta, Chamaedryfolia, Arborescens, Flavouirens, Leucantha, Angustifolia, Serrata and Scordioides. Possible evolutionary pathways are discussed. ADDITIONAL KEY WORDS: Chemotaxonomy phylogeny - systematics - taxonomy. biogeography - - ecology - Labiatae CONTENTS Introduction . . . . Results . . . . , Subsection GrandiJlora . Subsection Ovata . . Subsection Camarae . Subsection Linearifha . Subsection Gymnocarpae Subsection Stachydioides Subsectiou Lacailae . Subsection Hirsutn. . Subscction Chamaedyfolia Subsection Arborescens . Subsection Flauovirens . Subsection Leucantha . Subsection Anpslifolia . Subsection Serrnta . , Subsection Scordioides . Discussion, . . , . Acknowledgements , . References. . . . , 002&4074/90/080325 + 25 $03.00/0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . , , . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 325 . , , . . . . . . , . . . . , . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . , . . . 326 326 329 330 331 332 333 335 336 336 338 338 340 341 342 344 345 346 348 348 zy 0 1990 The Linnean Society of London 326 zyxwvut zyxwvu zyxwvutsrqpo D. RIVEKA NUREZ E T A L IN'I'RODUC'I'ION The genus Sideritis is generally divided into four sections of perennials and two others of annual herbs. The perennials are to some extent geographically separated, each section being more or less endemic to a restricted area. Sections Empedocleopsis Huynh and Marrubiastrum (Moench) Benth. are endemics of Macaronesia, Empedoclea (Raf.) Benth. occurs in Italy, the Balkans, Aegean Islands, Turkey and the Near East. The section Sideritis is endemic to the western Mediterranean Region, ranging from the northern French Jura to the Southern Atlas of Morocco and from Cab0 de San Vicente in Portugal to the north of Italy in Europe and northern Tunis in Africa. A first attempt a t infrasectional systematics in section Sideritis was proposed by Font Quer (1924). This was based on the presence or absence of a ring of hairs inside the calyx, the carpostegium. According to this criterion Font Quer recognized two unequal subsections: the subsection Carpostegiatae Font Quer, which contained over 90% of the taxa included in section Sideritis and the other subsection Gymnocarpae Font Quer with only S. incana L. and related species devoid of a carpostegium. Many problems arose using this division. For instance, it is inconsistent to place populations of S. hyssopiflia L. (sensu Lato) and S. chamaedryfootia Cav., which lack the ring of hairs, in the subsection Carpostegiatae, or the specimens of S. stachydioides Willk., with a carpostegium, in the Gymnocarpae and it is possible to see additional examples of how difficulties arise in the building of the infrasectional classification on the basis of a single character. The definition of subsections based on character syndromes should be more reliable, but there are many problems with this, due to overlapping boundaries between the different characters (chemical, morphological, cytological, etc.) . A highland landscape is a good graphical model to depict variation in Sideritis: the chains of mountains are the subsections and the peaks and summits the species. The real populations in the field can be represented by the shoulders or faces. Interspecific breeding, introgressions, clinal variations, allelic fluctuations and phenotypical plasticity have puzzled taxonomists since Linnaean times: "Interspecific hybrids within Sect. Sideritis have been recorded from many localities in Spain and it seems probable that much of the taxonomic difficulty is due to the occurrence of these hybrids and possibly hybrid swarms" (Heywood, 1972). The aim of this paper is to provide reference points in this complicated section which are easy to separate and recognize, using traditional characters. Some subsections are composed of a single species, these are the highly evolved taxa according to Fernandez Peralta (1981). zy RESULTS We summarize here our taxonomic work on the morphology and chemistry, of which only fragments have been published before (Rivera & Obon, 1988a, b; Tomiis-Lorente et al., 1988; Tomiis-Barberan et al., 1988). Additional information was obtained from the bibliographical review of Guy (1987), but with some difficulty. The main problem with using this data concerns the lack of zyxwvutsr zyxwvu zy SY S'IE MAT1C:S OF SIDEKITIS 327 C zyxwvu zyxw zyxw zyxw zyxwvu Figure 1. Hair covering on the base of the stems. Scale bar = I mm. A. Siderzlii borgiae. B. S. ouata. C . S.camarae. D. S.hyssopijolia var. aranensis. E. S. brachycalyx. F. S.cantabrica. C . S. caslellana. H. S.carbonellii. I . S. jaualambrensis. J . S. linearijolia. K. S.glauca. L. S. incana. M. S. stachydioides. N. S. lacaitae. 0. S.augustinii. P. S.hirsuta. information for many of the critical taxa and the problems imposed by incorrect determinations of voucher specimens; for instance, much of the literature concerning the chemical properties of S. mugronensis Borja was published under the name S.funkiana Willk., which was itself not studied at all, cf. Guy (1987). Many specimens of glabrescent forms belong to S. incana L. sensu lato, from Morocco, are labelled in the herbaria as S. ochroleuca Willk. causing confusion in the attribution of the chemical analysis (Tom&-Barberin et a/., 1988). The subsections are compiled on the basis of the macromorphology, hair covering, chromosome number, flavonoids, essential oils, etc. The morphology and hair covering (Figs 1-3) were studied on dried specimens collected in the field by the authors, or belonging to the following herbaria: BM, K, LINN, LY, MA, MPU, MUB and P. There is no real gap in the hairiness or glabrescence in the throat of the calyx and there is no clear distinction between the presence or absence of a single ring of hairs inside the calyx. Specimens having bundles of hairs between the lobes of calyx appear in some groups, but they are also 328 zyxwvutsrqpo zyxwv zyxw D. RIVERA NURE2 87 AL. zyxwvu zyxwv zyxwvut zyx Figure 2. Hair covering on the basr of t h r stems. Scale bar = 1 mm. A. Siderilis hirsula var. altilabra. B. S. gadilana. C . S. hlrsula. D. S.granalensis. E. S. mscinonensis. F. S. liirsuta var. nivalis. G. S. endresii subsp. laxispicata. H. S. scordioide~ proles fonlii. I. S. chamaedvfolia. J. S. arborascens subsp. paulil. K. S. arborescrns subsp. arborescens. L. S. maireana. M. S.foetens. N. S. luteola. 0. S.per~z-larae. considered ‘gymnocarpics’ since they lack a continuous ring of hairs. The nomenclature of Stearn (1973) and Vaczy (1980) for the hair covering was followed. According to their length, five categories of hairs were separated; very short (70-400 pm), short (400-700 pm), medium (700-1000 pm), long (1000-1400 pm) and very long (1400-2000 pm). T h e hair covering of the bases of the young stems is uniform between individuals belonging to the same taxon. The flavonoid pattern was studied in the same specimens used for the morphological analysis as far as possible and the results summarized by Tomas-Lorente et al. (1988) and Tomas-Barberan et al. (1989); references to the voucher specimens are given at the end of each subsection. Flavonoids are good taxonomic markers (Guy, 1987), and have been successfully used to discriminate between interspecific hybrids (Tomas-Lorente et al., 1989). Chromosome numbers were studied by Ob6n & Plantrose, (personal communication), or obtained from the literature (Fernandez Peralta, 1981). Much dispersed chemical information concerning Siderilis has been collected by Guy (1987). Taxa are listed for each subsection; nomenclature follows Heywood (1972), Greuter, Burdet & Long (19861, Briquet (1891),Font Quer (1924),Jahandiez & zyxwvut zyxwvu zyxwv SYSTEMATICS O F SIDERITIS M 329 N zyxwv Q”i zyx zyxwvut zyxwvu i- Figure 3 . Hair covering on the base of the sterns. Scale bar = 1 mm. A. S i d e r i h pusilla. var. mlina. B. S . osleuxylla. C. S. ibanyezii. D. S. p u d l a . var. pusilln. E. S . leucanlha var. leucantha. F. S. leucantha var. incana. G. leucantha var. bourgapnnn. H. S. tragori,qanum. I. S . saetnbmsis. ,J. S.,funkinna. K . S. reuerchonii. L. S..serrata. M. S. iliciJlin. N. S. spinu1o.m. 0. S. glacialis. 1’. S. mariae. Q. S. wrdioide5. K. S . cauanillesii. Maire ( 1934) and Malagarriga ( 1968). Synonymies and infrasubsectional taxa are subject to further studies, currently in progress. Geographical distribution is summarized from the localities recorded on the labels of almost 3000 specimens reviewed. Types of most of the taxa were studied morphologically, but detailed references are given only for the type species of each subsection. The number of specimens reviewed in the different herbaria is given after the reference of publication for each taxon. No type specimens were analysed for flavonoid patterns. zyxwvu 1. Subsection CrandifIora Rivera & Obon, subsect. nov. Caulibus herbaceis, basi lignosis, calicus elongatis (usque ad 15 mm long.), foliis inferioribus lanceolatis, obtusis, longe petiolatis (45-90 x 12-15 mm). Verticillastri 10-multiflori. Carpostegiatae. Pilis caulinaris holotrichis. S. grandzjlora Salzm. ex Benth., Lab. Gen. et S’.: 577 (1834): in collibus Tingitanis, 1825 Salzmann (Holotype: Herb. Lindley, CGE; isotype K!). TYPE: 330 zyxwvutsrqpo zyxwvu zyxwvu zy D. RIVERA NUmEZ E T .4L. S. grandzjora Salzm. ex Benth., Lab. Gen. el Sp.: 577 (1834). 7 (BM), 3 (K), 1 (P). DISTRIBUTION: Dispersed localities in sou th-west Spain and north-west Morocco. TAXA: The form and size of the leaves and bracts and the high number of flowers, 10-30 per whorl are very typical of this species. The hair covering is densely shaggy on the stems, bracts and flowers giving to the whole plant a silvery appearance. Both very long and glandular hairs are present. Without epicuticular flavonoids and with only traces of vacuolar flavonoids, accumulating 7-allosyl ( 1 -+ 2) glucosides of chrysoeriol, luteolin and apigenin, phloroglucinol type flavonoids unsubstituted on the &position. Chromosome number unknown. The high number of flowers per whorl is found only here and in the subsection Serrata. The flavonoid pattern is quite different from the rest of the section and is very similar to that of the section Hesiodia. It is probably the most primitive subsection in the section Sideritis. zyxwv SPECIMENS USED FOR FLAVONOID ANALYSIS: et al., 1988). Tetouan, Morocco (Tomis-Barberan 2. Subsection Ovata Rivera & Obon, subsect. nov. Caulibus herbaceis, basi lignosis, foliis inferioribus longe petiolatis, crenatis vel dentatis, verticillastris congestis in spicam densam. Carpostegiatae. Pilis caulinaris holotrichis. S. ovata Cav., Icon. Descripl. Plant., 1: 36 (1791): in Peruvia.. . i n Regio horto Matritensi et in Pharmaceutico vulgo de la Priora.. .se halla en Vizcaya (MA). Cavanilles used both specimens grown from seeds at Madrid and dried specimens collected in northern Spain. Probably the collector was J . A . Pavdn and the plants and/or seeds belonged to his botanical collections accounting for why Cavanilles supposed the plant came from Peru. Probably due to the inimity between Cavanilles and Pavon, the former did not quote the latter collector. It is not clear if Cavanilles examined the specimen now at BM “an in horto quidam collitur, vix indigenam?” Pavon, afterwards labelled “Habitat in montibus santanderensibus” by Lagasca, which could also be a type specimen. TYPE: zyxwvu zyxw S. ovata Cav., Icon. Descript. Plant., 1: 36 (1791). 4 (BM), 2 ( K ) , 24 (MA), 1 (P). S. borgiae Andrks in L6pez Pacheco et al., Dos ESP. Fl. Leonesa in Fac. Biol. Ledn, 1: 3 (1979). 2 (MA). TAXA: DISTRIBUTION: Mountains of north-west Spain. The rhizomatous stems show the adaptations to glareous moving soils. T h e hair covering is generally shaggy on the stems, calyx and axis of the inflorescence. The leaves have the same type of hairs but these are laxly dispersed, being more abundant on the abaxial surface. T h e bases of the stems are holotrichous (the hairs are distributed all round the stem) covered by long (1000-1400 pm) or medium length (700-400 pm) hairs. Without epicuticular flavonoids. Accumulating mainly isoscutellarein 7-allosyl-glucosides with only zyxwvu zyxwv zyxwv zyxwvutsr SYSTEMATICS OF SZDERZTZS 33 I zyxwvuts zyxwv zyx traces of hypolaetin 7-allosyl ( 1 2) glucoside. Chromosome number unknown. Closely related to the next group through the similar habit of S. lurida from which it differs only in the continuous hair covering. Hybridizes with subsection Scordioides (Font Quer, 1924a). --f SPECIMENS USED FOR FLAVONOID ANALYSIS: S. ovata Cav., Valdegoira, Alava (Tomiis-Lorente et al., 1988). 3. Subsection Cumurae Rivera & Obon, subsect. nov. Plantae suffruticosae, caulibus lignosis. Folia floralia inferiora et superiora dissimilia, foliis caulinaris ellipticis, oblongis, lanceolatis, linearibusve. Carpostegiatae, rare gymnocarpae, glabrescentis vel villosis. Pilis caulinaris goniotrichis. TYPE:S. camarae Sennen, Diagn. Nouv. PI. Espagne et Maroc, 1928-35 263 (1936): Hab.-Logroiio. Peiia Irasa, F. Camara. Sennen wrote at the end of the name “(Pau) Sennen, nov.”. We have seen in MA specimens collected by F. Camara on August 21 1933, labelled in C. Pau’s handwriting, with the following: Sideritis hyssopzfolia var. camarae Pau. This taxon was implicitly cited by Sennen but no references of any publication were given. We have not traced the publication of this variety by Pau. This is a common error in Pau’s work where names of taxa have been given as “in schedulae et ad amicos”, and the normal rules of valid publication have not been followed. We therefore think the type should be chosen from specimens in the herbarium Sennen at La Salle-Bonanova, Barcelona. ISOTYPES were labelled under number 9791 in the series of 1935 of the exsiccata “Plantes d’Espagne et d u Maroc”. Unfortunately, we suspect these plants were never distributed, probably due to the sudden start of the Spanish Civil War (1936-1939). Isotypes only exist as the specimens of the Pau herbarium (MA) and those of the botanical collection of the Instituto Miguel Servet at Zaragoza included in the herbarium of F. Ciimara Niiio. zyxwvu zyxwvutsrq TAXA:S. camarae Sennen, Diagn. Nouv.PI. Espagne et Maroc 1928-35 263 (1936). 6 (MA). S. brachycalyx Pau, Bol. SOC.ESP. Hist. Nut., 20: 141 (1920). 7 (BM), 16 (MA), 2 ( P ) . S. hzssoPzJolia var. aranensis Font Quer, Trab. Mus. Cienc. Nut. Barcelona, 5 ( 4 ) : 26 (1924). 2 (MA). S. lurida Gay, J . Bot., 67 151 (1929) 6 (BM), 6 (K), 6 ( M ) , 3 (P). S. lurida var. relegata (Font Quer) Font Quer in Font Quer & Rothmaler, Cavanillesia, 7 178 (1936). 1 (MA). S. scordioides var. glabrata Benth., Lab. Gen. el Sp.: 578 (1834). 1 ( K ) . Central and North Spain, South France, in the Pyrenees, Cantabrian mountains, the Picos d e Europa, Gredos and the Logroiio province. DISTRIBUTION: T h e hair covering is shaggy, with patent hairs on the axis of the inflorescence, showing some tendency to glabrescence. The margins of the leaves are covered by antrorsely curved hairs. The bases of the stems are goniotrichous, covered by retrorsely curved medium, short and very short hairs. Normally the calyx is carpostegiate, but S. brachycalyx Pau lacks carpostegium in some populations, 332 zyxwvut zyxwvu zyxwvutsrqpon zyxwvut D. RIVERA NUmEZ E T AL. although this character is discontinuous through the range of the species. Chromosome numbers, n = 15, 16, 17 (Fernandez Peralta, 1981). This subsection is relatively homogeneous with a marked tendency to glabrescence in the north-western border of its distribution. Transitional populations exist within subsections Linearzfolia or Scordioides, probably due to the gene exchange in close neighbourhoods, but sometimes translated into a clinal variation. The retrorse direction of the curvature in the hair covering is used as a differential character between this subsection and the subsection Iinearzfolia which has antrorsely curved hairs. We have no data concerning flavonoid patterns. zyxwvut 4. Subsection Linearij’olia Rivera & O b h , subsect. nov. Plantae suffruticosae, caulibus lignosis erectis vel decumbentibus, foliis integerrimis vel serrulatis, inermis. Corollis pallide luteis vel flavis. Carpostegiatae. Pilis antrorsis, holotricis vel goniotricis, vestita. S. linearifolia Lam., En&. Metk., 2: 168 (1786): “On trouve cette plante en Espagne, elle a ttt cultivte au Jardin du Roi”. Lamarck wrote at the end of the diagnosis “V.V.”, vidit in vivo. The possible relevant type specimens of this species are in the Lamarck herbarium at P. There are three sheets under the same cover labelled S. linearifolia Lam. The first one is labelled only “Herbier de Lamarck/Acquis en Novembre 1886”. This specimen has broader leaves than stated in the diagnosis but for the rest fits well with the description. The second sheet has one specimen belonging to S. montana L. The third sheet is labelled; “Herbier de Lamarck/Acquis en Novembre 1886/de M r AndrelSideritis hispanicalSideritis linearifolia em.”. There are four specimens, the two in the centre are S. k a n a L. sensu lato and the two marginal ones are probably hybrids between S. incana L. and the plant described by Lamarck under the name of S. linearifolia. We interpret these to refer to the Spanish populations living near the Ebro Basin. These plants fit very well with the diagnosis and likely these were seen by Lamarck living a t the Royal Botanic Garden of Paris. The specimens of the herbarium are probably gatherings of latter generations changed under the influence of culture and hybridization. TYPE: zyx zyxw zyxwv S. linearzfolia Lam., En& Meth., 2: 168 (1786). 7 (BM), 4 (K), 1 (MA), 7 (P). S. alpina Villars, Hist. PI. Dauph., 2: 373 (1787). 7 (BM), 2 (K),4 (MA), 1 (MPU), 1 (P). S. brevispica Sennen & Elias, Bol. Real. SOC.Zber., 31: 1 15 ( 1934). 1 (P). S. carbonellii Socorro, Stud. Bot. (Salamanca) , I : 2 1 ( 1982). 4 (BM), 2 (K), 1 (MA), 1 (MUB), 4 (P). S. cantabrica Sennen & Elias, Bol. Real SOC.Zber., 31: 114, 1934. 10 (MA). 1 (P). S. castellana Sennen & Elias, Bull. Acad. Int. Geogr. Bot.: 228 (1914). 1 (BM), 8 (MA), 1 (P). S eynensis Sennen, Bol. Real SOC.Zber.: 373 (1926). 1 (MA). S. giennensis Font Quer, Cavanillesia, 1: 40 (1928), nom. inval. 1 (P). S. hyssopzfolia L., S.’ PI.:575 (1753). 6 (BM), 31 (K), 1 (LINN), 10 (MA), 1 (MPU), 5 (P). S. hyssopzfolia subsp. guillonii (Timb.-Lagr.) Nyman, Consp. Fl. Eur. Suppl. 2: 253 (1890). 5 (BM), 4 (K), 4 (MA), 1 (P). S.java1ambrensis Pau, Not. Bot. Fl. EspaEola, 1: 26 (1887). 6 (BM), 2 (MA), 1 (MUB), 3 (P). S. maura De Noe in Bal., PI. Alge‘rie 1852. no 564 (1852) [in sched.]. 2 (BM), 3 (K), 2 (MPU), 8 ( P ) . S. pastoris Sennen, Bol. Soc. Arag., 15: 248 (1916) & Bol. Real SOC. Zber.: 375 (1926). 10 (BM), 1 ( K ) , 5 (MA), 1 (P). TAXA: zyxwvu zyxwvu zyxw zyxwvuts SYSTEMATICS OF SIDERITIS 333 S. peyrei Timb.-Lagr., Mlm. Acad. Toulouse, Slr. 7, 4: 383 (1872). 1 (BM), 1 (K), 1 (MA), 1 (P). S. pyrenaica Poiret. Diet. Suppl., 2: 383, 1811. 24 (BM), 14 (K), 18 (MA), 1 ( M P U ) , 1 (MUB), 1 (P). S. scordioides var. angustzfolia Benth., Lab. Gen. et Sp.: 578 (1834). 1 ( K ) . S. scordioides var. elongata Benth., Lab. Gen. et Sp.: 578 (1834). 4 (K). S. vidali Sennen, Bol. Sac. h a g . , 1 5 247 (1916). 2 (BM). DISTRIBUTION: North Africa, south-east Spain, Ebro Basin, Mountains of Castilla, North Spain, Pyrenees, East France, north-west Italy. The hair covering is shaggy on the axis, calyx, bracts, leaves and bases of the stems, with antrorsely curved hairs. O n the basis of the young stems, the hair covering is holotrichous, inclining to goniotrichous in some populations. The length of the hairs is variable from medium length (1000-700 pm) to short (700-400 pm) or more rarely very short (400-70 pm). The presence of epicuticular flavonoids is restricted to certain taxa as S. linearzfolia or S. javalambrensis which produce mainly cirsilineol or 8-methoxycirsilineol, S. maura produces 8-methoxycirsilineol and xanthomicrol. Sideritis linearfolia and S. javalambrensis accumulate as vacuolar flavonoid isoscutellarein 4’-methyl ether 7-allosyl ( 1 -+ 2) glucoside. Other taxa, such as S. maura and the S. hyssopfolia group accumulate derivates of isoscutellarein or hypolaetin. Chromosome numbers: n = 13, 14, 15 (Fernandez Peralta, 1981). This group shows the similarities with subsection AngustiJolia in their overlapping area, but differs from them mainly in chemical, i.e. flavonoid patterns, and in their chromosome numbers. T h e subsection is more or less sympatric with the subsections Camarae, Gymnocarpae, Scordioides and Hirsuta. Many transitional taxa could be of hybrid origin. Hybrids were described with subsections Scordioides (Font Quer, 192 1 ) ) Gymnocarpae and Hirsuta (Font Quer, 1924a). FLAVONOID ANALYSIS: S. carbonellii Socorro: L a Sagra, Granada, Alcaraz, Peinado €8 Martinez (MUB). S. javalambrensis Pau: Sierra de Javalambre, Teruel, Toma‘s-Barbera’n (MUB). S. hyssopfolia subsp. guillonii Timb.-Lagr.) Nyman: Picos de Europa, Asturias (Tomas-Lorente et al., 1988). S. maura De Noe: Dahra, Oran, Algeria (Tomis-Barberan et al., 1988). S. lineartfolia Lam.: Zaida, Zaragoza, Rivera €8 D e L a Tore (MUB). S. pyrenaica Poiret.: San Antonia de Urquiola, Guipuzcoa (Tomas-Lorente et al., 1988). SPECIMENS USED FOR zyxwvu 5. Subsection Gymnocarpae Font Quer, T r a b . Mus. Cienc. N a t . Barcelona, 5( 4) : 3 (1924). Carpostegio nullo, foliis integris vel paucidentatis, nunquam spinosis. Bracteis calycibus brevioribus vel eos subaequilongis, a base dentatis, vel raro, integris. Floribus luteis vel roseis, rare albis. Plantae tomentosae vel glaberrimae, parce odorata. S. incana L., Sp. P1. ed. 2: 802 (1763):Habitat in Hispania, Loejling, Alstroemer (LINN). TYPE: T h e specimen number 729.9 of LINN is labelled as follows on the back “Hispanica 424. a Loefl./[Span. list 1753 n. 424. a det. Loefl.]”. There is another specimen numbered 729.10 belonging also to S. incana L., but unlabelled. Gin& 334 zyxwvut zyxwvu zyxwvutsrqpo D. KIVERA NLJmEZ E T AI, L6pez informs us that the text of Loefling’s list is 1752 (not 1753) “424 a. Sideritis calycibus tomentosis acutis, foliis linearibus. Synonyma nescio. ex horto Queriano”. Probably the plant collected by Loefling in Quer’s botanic garden was sent to Linnaeus and is the specimen 729.9, which must be considered as a zyxwv zyxwvu zyxwvutsrq zyxw HOLOTYPE. S. incana L., Sp. P1. ed. 2: 802 (1763). 17 (BM), 28 (K), 2 (LINN), 8 (MA), 42 (MUB), 10 (P). S. atlanlica Pomel, Nouu. Mal. Fl. All.: 120 (1874). 1 (P). S. atlantica var. neruosa Pomel, Nouv. Mat. Fl. Atl.: 121 (1874). S. glauca Cav., Icon. Descr., 2: 68 (1794). 6 (BM), 1 1 ( K ) , 1 (MA), 6 (MUB), 3 (P). S. guyoniana Boiss. & Reuter, Pugill. PI.: 98 (1852). 8 (BM), 12 (K), 13 (P). S. incana var. albgora Maire in Battand., Contr.: 68 (1919), & in Font Quer, Bol. SOC.ESP. Hist. Nut., 25 465 (1925). 2 (BM), 3 (P). S. incana var. albzjora subvar. roseijlora Font Quer, Bol. Sac. ESP. Hist. Nat., 25 465 (1925). S. incana var. albzJora subvar. depauperata Font Quer, Bol. Sac. Esp. Hist. Nut., 25 465 (1925). S. incana var. altiatlantica Font Quer, Bol. SOC.Esp. Hisl. Nut., 25 464 (1925). S. incana var. altiallantica subvar. recessa Font Quer, Bol. Sac. ESP.Hist. Nut., 25 464-465 (1925). 1 (P). S. incana var. angustiflia Font Quer, Bol. Sac. ESP. Hist. Nat., 25 465 (1925). S. incana var. aurasiaca Battand., FI. Alg.: 698 (1890). 1 (P). S. incana var. edetana Pau ex Font Quer, Trab. Mus. Cienc. Nut. Barcelona, 5 ( 4 ) : 7 (1924). 3 (MA), 6 (MUB). S. incana var. edetana subvar. saxzjiraga Font Quer, Bol. Soc. Esp. Hist. N a t . , 25 466 (1925). S. incana var. Jauovirens Maire, Bull. SOC.Hist. Nut. Afr. du Nord 274 (1916). S. incana var. longzfolia Font Quer, Bol. SOC.Esp. Hist. Nal., 25 466 (1925). S. incana var. accidentalis Font Quer, Trab. Mus. Cienc. N u t . Barcelona, 5 ( 4 ) : 7 (1924). S. incana var. regimontana Maire, Bull. SOC.Hisl. N u t . Afr. du Nord 274 (1916). 1 (P). S. incana var. robusta Font Quer, Bol. Sac. ESP. Hist. Nut., 25: 464 (1925). S. incana var. socovensis Rivera & Ob6n, Al-Basit, 24: 229 (1988). 2 (MUB). S. incana subvar. spinulosa Font Quer, Trab. M u s . Cienc. Nat. Barcelona, 5 (4 ): 7 (1924).S. incana var. tomentosa Battand. et Pitard, Contr. Etude F1. du Maroc: 32 (1918). 2 (MA), 2 (P). S. incana subsp. tunetana Murb., Acta Uniu. Lund., ser. 2, 1/4: 65 (1905). 1 (P). S. incana var. uirgala subvar. arairae (Maire) Font Quer, Bol. Sac. Esp. Hist. Nat., 25 463 (1925). 1 (P). S. incana var. uiridzfolia C. Vicioso, Anales Jard. Bat. Madrid, 2: 224 (1942). 1 (MA). S. matris-jiliae Emb. & Maire, PI. Marocc. Nov., 2: 7 (1929). 1 ( P ) . S. ochroleuca aucl. non Willk., Bot. Zeit., 17 282 (1859). 2 (P). S. pycnostachy Pomel ex Battand. & Trabut, Fl. de I’Algirie, I : 699 (1890). S. sericea Pers., Syn. PI., 2: 118 (1806). 1 (K), 4 (MA), 1 ( P ) . S. virgata Desf., F1. Atl., 2: 15, tab. 125 (1798). 20 (BM), 10 ( K ) , 45 (MA), 9 (P). TAXA: DISTRIBUTION: Spain, Morocco, Algeria and Tunisia. The hair covering varies from woolly to hoary on the bracts, leaves and stems, but sometimes these are glabrescent. There is no continuous ring of hairs in the throat of the calyx, but in some specimens isolated groups of hairs appear between the calyx lobes. The bases of the stem show typical hairs, longer than 2000 pm but much twisted. Without epicuticular flavonoids. Accumulating vacuolar flavonoids of the hypolaetin group, but S. glauca accumulates mainly isoscutellarein derivates. Chromosome numbers: n = 13, 14, 15, 17 (Fernandez Peralta, 1981). zyxwvut zyxwvu zyxwvu zyxwvu zyxwvuts zyxwvu SYS'I'EMAL'ICS OF SIDEKlT1.S 335 This group is relatively homogeneous showing very interesting geographical patterns of clinal variation in the density of the hair covering, dimensions of the leaves and colour of the corollae. Hybrids are known with the subsections Lineariflia (Font Quer, 1924a), Hirsuta (Font Quer, 1922, 1924b), Arborescens (Malagarriga, 1968), Leucantha (Font Quer, 1924;) and AngusLz&olia (Font Quer, 1921). S. incana L.: La Toba, Cuenca, Rivera @ De L a Torre (MUB); Ouarzazate, Morocco and Ayachi, Grand Atlas, Morocco (Tomas-Barberan et nl., 1988). S. incana var. edelana Pau: Entre Quesa y Bicorp, Rivera €5' Obdn (MUB). S. glauca Cav.: Sierra de Orihuela, Alicante, Rivera (MUB). S. guyoniana Boiss. & Reuter: Oran, Algeria (Tomas-Barberan el al., 1980). S. sericea Pers.: Quesa, Valencia (Tomiis-Lorente et al., 1988). S. uirgata Desf.: Grazalema, Cadiz, Alcarar (MUB); Sierra de Almansa, Albacete, Obdn (MUB). SPECIMENS USED FOR FLAVONOID ANALYSIS: 6. Subsection Stachydioides Rivera & O b h , subsect. nov. Plantae suffruticosae, foliis tomentosis, integerrimis vel pauce dentatis. Calycibus carpostegiatis, rare gymnocarpis, corollis roseis. S. stachydioides Willk., Bol. Zeit., 8 78 (1850): Sierra de Maria, Almeria, Willkomm (1845), Funk (1848) (COI not seen). TYPE: ISOTYPES: We have seen in BM two relevant specimens. O u r register number 1389 refers to one specimen collected by Funk in July 1848 in the Sierrra de Maria belonging to Herb. Auerswald, received in BM in 187 1. This is the same exsiccata cited by Willkomm in the diagnosis. The second specimen in BM, registered by us with the number 1390, belongs to the exsiccata of Willkomm from the Sierra de Maria, in the year 1845, cited also by the author in the original description. S. stachydioides Willk., Bat. Zeit., 8 78 (1850). 8 (BM), 7 (K), 2 (MUB), 5 (PI. DISTRIBUTION: Mountains of a very reduced area in south-east Spain. TAXA: The hair covering is woolly or tomentose, very dense. The hairs are similar to those of the subsection Gymnocarpae. Carpostegium is present. Without epicuticular flavonoids and with only traces of vacuolar flavonoids, accumulating luteolin and apigenin 7-p-cumaroil glucosides and lesser amounts of hypolaetin 7-allosyl ( 1 + 2) glucoside. Chromosome number: n = 17 (Fernandez Peralta, 1981 ). This species is separated off into another subsection because it differs markedly in its flavonoid pattern from that of the subsection Gymnocarpae, being more closely related to the section Marrubiastrum of the Canary Islands or the genera Phlomis and Marrubium. The presence of a carpostegium supports this separation. Hybrids have been described with the subsections Leucantha (Socorro, 1981) and Angustzfolia (Fernandez-Casas, personal communication). SPECIMENS USED FOR FLAVONOID (Tomb-Lorente et al., 1988), ANALYSIS: Velez Blanco, Granada 336 zyxwvut zyxwvu zyxwvutsrqpon zyxwv zyxwv D. RIVERA NUNEZ E l AL 7. Subsection Lacaitae Rivera & O b h , subsect. nov. Plantae suffruticosae, foliis lanceolato-linearis, sinuatis, villosis. Calycibus gymnocarpis. Corollis luteis. S. lucaitae Font Quer, Bol. SOC.ESP. Hist. &at., 24: 208 (1924):. . . montium Marianorum, pr. Santa Elena, 7 vi 1923, Lacaita (BC). TYPE: This specimen was sent by Lacaita to Font Quer and no duplicate was conserved by Lacaita in his herbarium, now in BM (at least, if so, we have not traced it). The specimen used for the description was sent in return to Lacaita, with the letter of Lacaita cited by Font Quer. This specimen is then the HOLOTYPE, number 26243 of the European Herbarium (BM). Font Quer has distributed specimens collected “ e loco classico” on 8 June 1924 by Font Quer & Gros. Lacaita collected on 9 July 1926, other specimens from the classical locality, which in detail is “in dumetis supra El Arroyo de Oruga, loco dicto Cerro Castillejo”. zyxwv S. lacaitae Font Quer, Boi. SOL.ESP. Hist. Nut., 24: 208 (1924). 5 (BM), 4 (K), 1 (MA), 10 ( M W , 1 (PI. TAXA: DISTRIBUTION: Northern Andalusia and south of Castilla. O n siliceous soils. The hair covering is laxly villous, with very long hairs (2000-1400 pm). Carpostegium absent. With traces of epicuticular flavonoids accumulating cirsimaritin. With vacuolar flavonoids accumulating hypolaetin-7-allosyl ( 1 + 2) glucoside and 3’-methyl ether derivatives. Chromosome number: n = 1 7 (Fernandez Peral ta, 1981) . This taxon seems to be of hybrid origin between the subsections Gymnocarpae and Hirsuta. SPECIMENS USED FOR FLAVONOID ANALYSIS: Sierra del Relumbrar, Albacete, Herranr (MUB). 8. Subsection Hirsuta, Rivera & O b h , subsect. nov. Plantae suffruticosae, caulibus lignosis vel herbaceis, erectis vel decumbentis Folia floralia retrorsa vel decidua in maturitatem, folia caulina obtusa, serrata vel lobata. Carpostegiatae. Foliis caulibusque hirsutis. TYPE: S. hirsuta L., Sp. PI.: 575 (1753). Habitat in G[allia]. Narbonensi. According to C. E. Jarvis (personal communication): “There seem to be a number of possible syntypes. Sheet 729.15 (LINN) is one with a possible duplicate in the small Linnaean collection in UPS.. .). There are also two specimens in the small Linnaean collection in the Bergius Garden (SBT) which may be possible syntypes. In addition, material in Herb. Burser, vol. X I I I . 59 (UPS) is a syntype”. We have studied specimens in LINN. The specimen numbered 729.14 has no label or writing but it belongs to 5’. hirsuta sensu lato. A pin joins sheets number 729.15 and 729.16. The latter has written on it only the capitals “H. U.”. The sheet 729.15 is labelled: “8 hispanica hirsutn” on the face and “Siderit. hispanica crenata, procumbens, pore ulbo majore Tourng. inst. Monnier” on the back. This last specimen is morphologically similar to those living nearby south of Madrid. zyxwvu zyxwv zyx zyxwvu zyx sYsrELmwcsOF SIDERITIS 337 TAXA: S. hirsuta L., Sp. PI.: (1753). 38 (BM), 50 ( K ) , 54 (MA), 30 (MUB), 1 1 (P). S. aculeata (Bubani) Font Quer, Butll. Inst. Catal. Hist. Nat., 214): 31 (1924). S. augustinii Sennen & Pau, Bull. h a d . Int. Giogr. Bot., 21: 119 (191 1 ) . 2 (MA), 2 (P). S. briquetiana Font Quer & Pau, Cavanillesia, 3 60 (1930). 26 (BM), 6 (P). S. bubanii Font Quer, Butll. Inst. Catal. Hist. Nut., 20: 141 (1920). 1 (K), 2 (MA). S. catalaunica Sennen & Pau, Bull. Acad. Int. Giogr. Bot., 21: 119 (191 1 ) . 1 (P). S. crenata Lapeyr., Hist. Abr. de la Fl. des Pyrin.: 331 (1813). 1 (BM). S. endressii Willk., Bot. zeit., 17: 276 (1859). 26 (BM), 9 ( K ) , 7 (MA), 1 (P). S. endressii f. laxispicata Degen & Debeaux, Bull. Acad. Int. Gigr. Bot., 1 7 196 (1907). 8 (BM), 4 (MA), 6 (P). S. gaditana Rouy, Zll. PI. Eur. Rarior., 17 137, tab. 417 (1902). 2 (BM), 10 ( K ) , 3 (MA). S. gouani Timb.-Lagr., M e m . Acad. Toulouse, Sir. 7(4): 382 (1872). 1 (BM). S. hirsuta var. altilabra Pau in sched. [nomen nudum]. 2 (BM), 3 (MA), 1 (P). S. hirsuta var. bracteosa Willk., Bot. z e d . , 1 7 284 (1859). S. hirsuta subsp. emporitana Cadev. in Cadev. & Font Quer, Fl. Catalunya, 4: 397 (1932). S. hirsuta var. granatensis Pau, Contr. Est. Flora Granada: 223 (1916). 2 (BM), 3 (K), 7 ( M A ) . S. hirsuta var. maritima subvar. angustifolia Font Quer, Butll. Inst. Catal. Hist. Nut., Ser. 2(4): 32 (1924). S. hirsuta var. maroccana Cosson in Battand., Contr. F1. Atl.: 69 (1919). 1 (BM), 2 (K), 1 (MA), 2 (P). S. hirsuta var. nivalis Font Quer, Butll. Inst. Catal. Hist. Nut., Ser. 2 ( 4 ) : 32 (1924). 15 (BM), 1 ( K ) , 2 (MA). S. hirtula Brot., Fl. Lusit., I : 161 (1804). S. imbricata H. Lindb. fil., Acta SOC.Sci. Fenn., Ser. B, Opera Biol., 112): 132 (1932). 2 (K). S. kebdanensis Sennen, Campagn. Bot. Maroc Or. 1930-35: 113 (1936) [pro parte]. 7 (BM), 1 (P). S. littoralis Timb.-Lagr., Bull. SOC.Bot. France, 22: 309 (1875). 2 (BM), 2 (MA), 2 (P). S. provincialis Jordan & Fourr, in sched. S. ruscznonensis Timb-Lagr., Mim. h a d . Toulouse, Sir. 7(2): 380 (1872). 1 (MA), 1 ( P ) . S. tomentosa Pourret, Mim. h a d . Toulouse, S i r . l ( 3 ) : 328 (1788). 21 (BM), 2 ( K ) , 53 (MA), 1 ( P ) . zyxwv DISTRIBUTION: Portugal, Spain, South of France, North Italy, Morocco, Algeria? Hair covering shaggy, with patent hairs on bracts, flowers, leaves and stems. The bases of the stems show a mixed hair covering of short retrorsely curved hairs and long patent or erecto-patent hairs. T h e length of the longer hairs varies between populations, being long (1000 1500 pm) or of medium length (700-1000 pm), sometimes short (400-700 pm). Without epicuticular flavonoids in some populations, but many southern populations of the Iberian Peninsula and North Africa produce cirsiliol, sideritoflavone, xanthomicrol and 8-methoxycirsilineol. Accumulating the vacuolar flavonoids isoscutellarein or hypolaetin and their 3’ or 4’-methyl ethers, and more rarely chrysoeriol. Essential oil with a-pinene (7.5-42.476), 1,8-cineol (4.2-15.40/,), d-cadinene (4.7-7.9%) and lesser amounts of sabinene (0-4.6(>&),fenchone (0-2.40/,), thymol (0-6.6%), etc. Chromosome numbers: n = 12, 14, 15, 28 (Fernandez Peralta, 1981). The mixed hair covering seems to be characteristic of this group, but the boundaries between species are mostly diffuse. This subsection hybridizes with the following other subsections: Arborescens (Font Quer, 1930), Leucantha (Font Quer, 1924c), Angustzfolia, Serrata, Scordioides (Font Quer, 1921) , Linearzfolia (Font Quer, 1924) and Gymnocarpae (Font Quer, 1922). Sideritis gaditana is considered by many authors to be of hybrid origin. Wild populations of this taxon were studied near El Puerto de Santa Maria (Cadiz, Spain), and found to be very uniform in habit, shape and details of morphology; no alleged parents 338 zyxwvut zyxwvu zyxwvutsrqpo zyx zyx zyxwv D. RIVERA NUmEZ E T A L . were in the neighbourhood. Probably the origin is with one ancient hybridization between subsections Hirsuta and Arborescens. ANALYSIS: S. briquetiana Font Quer & Pau: Djebel Kerker, Morocco (Tomas-Barberan et al., 1988). S. endressii f. laxispicata Degen & Debeaux: Cazorla, Jain, Alcaraz et al. (MUB). S. hirsuta var. grantensis Pau: Nerja, Malaga, Rivera (MUB). S. hirsuta L.: Zaida, Zaragoza, Rivera tY De La Torre (MUB); Murcia, Spain; Asni, Grand Atlas, Morocco; Michlifen, Moyen, Moyen Atlas, Morocco (Tomis-Barberan et al., 1988). S. hirsuta var. altilabra Pau: Enguera, Valencia (Tomas-Lorente et al., 1988). S. imbricata H. Lind. fil.: Tirarine, Djebel Aunsitene, Morocco (Tomas-Barberan et al., 1988). SPECIMENS USED FOR FLAVONOID 9. Subsection Chamaedryfolia Rivera & Obon, subsect. nov. Plantae suffruticosae, caulibus lignosis, erectis, rare decumbentis. Folia caulina obtusa, lobata. Bracteis calycibus brevioribus vel eos subaequilongis. Gymnocarpae vel minime carpostegiatae. Foliis caulibusque pilosis hirsutisque. S. chaemaedryfolia Cav., Icon. Descript., 4: 1 , tab. 301 (1797): Habitat in regni Valentini tractu, vulgo Collado de San Antonio inter Bocayrente et Baiieres. TYPE: We have found one type specimen at MA. In BM there is one specimen of Cavanilles dated 1803 and relabelled “TYPE SPECIMEN”. We are not sure that this one should be named as type, because the only date given is six years after the publication date. Additional problems arose concerning the locus classicus. The plants commonly named S. chamaedryfolia are typically growing on sandy soils in the region compressed between Villena and Baiieres, but were never found in the locality cited by Cavanilles. TAXA:S. chamaedryfolia Cav., Icon. Descript., 4: 1, tab. 301 (1797). 5 (BM), 1 (MA), 12 (MUB), 2 (P). S . f o n t i i Sennen, Bol. Soc. Iber. Cienc. flat., 31: 60, 1934. 1 (BM), 2 (MA). DISTRIBUTION: South-east Spain and southern Catalonia near the Ebro Basin. Hair covering of very short (70-400 pm) patent hairs on the stems bases mixed with more or less adpressed, antrorsely curved short (400-700 pm) or medium length (700-1000 pm) hairs. Bracts, flowers and spike axis covered with antrorsely curved short hairs. Essential oil with d-cadinene (9.5y0), a-pinene (7.8%), fenchone (7.8%), 1,8-cineol (6.8%) and lower amounts of sabinene (2.3%). Chromosome number unknown. This group seems to belong in the gap between the subsections Scordioides and Hirsuta, being adapted to specialized habitats such as sandy and gypsaceous soils. 10. Subsection Arborescens, Rivera & Obon, subsect. nov. Plantae fruticosae vel suffruticosae, caulibus floralibus lignosis. Folia fioralia inter se conformia, folia caulina lobata vel integerrima. Carpostegiatae. TYPE:S. arborescens Salzm. ex Benth., Lab. Gen. et S’.: 579 (1835): Hab. in Hispaniae monte Gibraltarico, Broussonet!, Salzrnann! (h. s. sp. comm. a cl. Bouschet-Doumeng.) . zyxwvu zyxwv SYSI‘EMA’I‘ICS OF SIBERI’TIS 339 zyxwvut zy zyx zyxwv We have found the HOLOTYPE at K. O n the sheet with number 532 of our register there are labels of different exsiccatae; the following labels must refer to the specimen near the right border of the sheet: “Herbarium/l854/Benthamianum”, “Sideritis scordioides/Gibraltar (ex. Broussonet)/Bouschet Doumeng/(82)/Sideritis arborescens Salzm./Benth Lab. 579/S. foetens/. . . [illegible]/l2443”. T h e only specimen cited in the diagnosis is this one, but by citing Salrmann as collector, the exsiccatae of Salzmann ought to be seen as SYNTYPES. T h e specimen at K situated on the right half of the sheet is registered by us with the number 540, and labelled as follows: “Sideritis arborescens. Mihi./Salzmann misit/August. 1825/in monte Gibraltarico. majo.” “Herb. J. Gay./Presented by Dr Hooker/February 1868.”, “Sideritis arborescens Salzm! exsicc./Ann. 1825.-Benth. Lab. p. 579. (Aug. 1834).”, “S.foetens Lag.”, could be also relevant as it was probably seen by Bentham before publication. S. arborescens Salzm. ex Benth., Lab. Gen. et Sp.: 579 (1835). 9 (BM), 15 (K), 1 1 (MUB), 3 ( P ) . S. angustzfolia var. lusitanica Font Quer, Flora Iberica Selecta, Cent. 1, n”73 (1934). 8 (BM), 2 ( K ) , 3 (MA), 80 (MUB), 8 ( P ) . 5‘. arborescens var. africana Font Quer et Pau in Font Quer, Iter Maroc. 1927. n”532 (1928) [in sched]. S. arborescens var. kebdanensis f. laxispica Font Quer et Sennen in Sennen, Diagnoses Nouv. Plantes d‘Espagne et du M a r o c de 1928 a’ 1935. 252, n”9723 (1936). S. arborescens var. ortonedue Font Quer & Pau, Cavanillesia, 3 TAXA: 60 (1930) [sensu strictol]. 10 (BM), 1 (P). S. arborescens subsp. Perez-larae Borja, Anales Jard. Bot. Madrid, 40: 278 (1983). 1 (MA), 15 (MUB). S.foelens Clemente ex Lagasca, Gen. et Sp. PI. Nov.: 18 (1816). 4 (BM), 5 (K), 3 (MA), 4 (MUB), 1 (P). S. foetens var. rivas-godayi Fernandez Casas, Anales Jard. Rot. Madrid, 32(2): 306 (1975). 1 (MA). S. getula Battand., Bull. Sac. Bol. France, 53 79 (1907). 2 (P). S. lutea Font Quer, Flora Hispanica Cent., 4. n”376 (1948) [nom. illeg.]. S. luteola Font Quer, T r a b . M u s . Cienc. Nut. Barcelona, 5(4): 32 ( 1924). 1 (BM), 1 ( K ) , 2 ( M A ) , 1 1 (MUB). S. maireana Font Quer & Pau in Font Quer, Iter M a r o c . 1927, n”533 (1928). 2 (BM), 3 (MA), 5 (MUB), 1 ( P ) . S. paulii Pau, Bol. Soc. Esp. Hist. N u t . , 21: 151 (1921). 5 (MA). S. paulii var. castellana Font Quer & Pau, Cavanillesia, 7 83, 1935. S. subatlantica Battand., Contr. FZ. Atlant.: 69 (1919). 13 (BM), 3 (P). 5’. subatlantica fa. heterostachya Sennen, Diagn. Nouv. PI, Espagne: 113 (1936). S. subatlantica fa. laxispica Pau & Font Quer in Font Quer, Iter Maroc. 1929. n”380 (1930). S. Jubatlantica var. riphaea Font Quer & Pau in Font Quer, Iter Maroc. 1927. n”530 (1928). DISTRIBUTION: South of the Iberian Peninsula and Morocco The hair covering is laxly shaggy, with patent hairs, on the calyx, axis and leaves, becoming less glabrous in some taxa. T h e bases of the young stems are goniotrichous, with very short (70-400 pm) retrorsely curved hairs on two opposed faces, while the other two are glabrescent. Some populations show a continuous pattern of hair covering on the bases of the young stems composed of retrorsely curved short (400-700 pm) or very short hairs. With epicuticular flavonoids, cirsiliol or sideritoflavone in S. maireana and S. foetens, which also produces xanthomicrol and 8-methoxycirsilineol. Accumulating derivates of hypolaetin or isoscutellarein. Essential oil with thymol (20%), p-cymene (19.8%), sabinene (8.67”) and lesser amounts of a-pinene (5.5%), 1,8-cineol (5.574) or d-cadinene (2.50/,) (S.foetens). Chromosome numbers: n = 12, 13, 15 (Fernandez Peralta, 198 1) . 340 zyxwvu zyxwvu zyxwvutsrqpo D. RIVERA N U R E 2 E l AL. This group is broadly distributed in southern Spain where transitional forms occur with subsection Hirsuta. In North Africa hybrids have been described with subsections Gymnocarpae (Malagarriga, 1968) and Hirsuta (Font Quer, 1930). Closely related to the next subsection. SPECIMENS USED FOR FLAVONOID ANALYSIS: S. arborescens var. kebdanensis Font Quer & Sennen: Kebdana, Morocco (TomAs-Lorente et at., 1988). S. foetens Clemente ex Lagasca: Barranco del Caballar, Almeria, Rivera (MUB). S. maireana Font Quer & Pau: Bu-Merziat, Morocco (TomAs-BarberAn et al., 1988). S. subatlantica Battand.: Bocaya, Hoceima, Morocco (Tomas-BarberAn et al., 1988). S. subatlantica f. heterostachya Sennen: Hidum, Morocco ( Tomas-Barberan et al., 1988). zy 11. Subsection Ftavovirens Rivera & Obbn, subsect. nov. Plantae suffruticosae, caulibus lignosis, erectis. Folia caulina obtusa, dentata vel lobulata. Carpostegiatae. Caulibus glabrescentis, hirsutis, a base pilis incurvatis munitis. S. flavovirens (Rouy) Font Quer ex Sennen = S. leucantha var. jlauouirens Rouy, Rev. Sci. Nut. Sir. 3, 3(2): 246 (1883): Puerto de Lumbreras. TYPE: This locality, Puerto de Lumbreras, was given by Rouy in the previous publication of his collections list (Rev. Sci. Nut. Sir. 3, 2(2): 241 (1882).At LY we have found one sheet with three specimens and two labels. One label includes the names of varieties of S. leucantha recognized by Rouy; the next label reads as follows: “PLANTES D’ESPAGNE/Flore de la Province de MurcialSideritis leucantha Cav./var. flavouirens Rouy/Puerto de LumbreraslCabezo de la Jara/7 Juin 1882/Legi G. ROUY”. The three specimens are all types showing the same morphological characters. S. leucantha var. flavouirens Rouy, Rev. Sci. Nal. Sir. 3, 3(2): 246 (1883). 7 (BM), 6 (K), 1 (LY), 3 (MA), 140 (MUB), 3 (P).S. almeriensis Pau, Bol. Sac. Arag., 7: 78 (1908). 2 (BM), 1 (K), 10 (MUB). S. debeauxii Font Quer, Butll. Inst. Catal. Hist. Nut., Ser. Z(5): 198 (1925). 1 (K). S. hirsuta var. maritima Font Quer, Butll. Inst. Catal. Hist. Nut., Sir. Z(4): 32 (1924). 2 (MA), 1 (MUB). S. hirsuta var. maritima subvar. rotundzfolia Font Quer, 1.6. S. hirsuta var. maritima subvar. pinnatifda Font Quer, 1.c. S. hirsuta var. maritima subvar. oscilans Font Quer, 1.c. S. ibanyezii Pau, Bol. Sac. Arag., 2: 68 (1903) & in Munuera, Mem. Sac. ESP. Hist. Nut., 2: 103 (1904). 1 (K), 1 (MA). S. ilorcitani Sennen, Butll. Inst. Catal. Hist. Nast., 32(4): 16 (1932) & Diagn. Nouv. P1. Espagne et Maroc 1928-35: 72 (1 936). S. leucantha var. carthaginensis Font Quer, Butll. Inst. Catal. Hist. Nut., 2 Ser., 5(7): 184 (1925). 10 (MUB). S. leucantha var. microphylla Willk., Bat. Zeit., 17(34): 289 (1859). S. ochroleuca Willk., Bat. Zeit., 17 282 (1859). 2 (BM), 7 (K), 6 (P). 5’. ochroleuca var. antiatlantica Maire, Contr. 1316. 2 (P). S. ochroleuca var. breuibracteata Font Quer, Cavanillesia, I : 38 (1928). S. ochroleuca var. denticulata Font Quer, Cauanillesia, I : 38 (1928). S. ochroleuca var. eremophylla (Maire) Font Quer, Cauanillesia, 1: 38 (1928). 1 ( P ) . S. ochroleuca var. mairei Font Quer, Cauanillesia, I : 38 (1928). S. ochroleuca var. maroccana Font Quer, Cavanillesia, I : 38 (1928). S. osteoxylla Pau, Bull. Acad. Int. Giogr. Bat. 16(2): 77 ( 1906). 8 (BM), 6 (K), 5 (MA), 2 (MUB), 4 (P). S. pusilla TAXA: zyxwvuts zyx zyxwvut zyxwvu zyxwv zyxwvutsr SYSTEMATICS OF SIDERITIS 341 (Lange) Pau, Bull. Acad. Int. Giogr. Bot. 16(2): 77 (1906). 5 (BM), 3 (K), 5 (MA), 35 (MUB), 6 (P). S. pusilla var. carthaginensis Font Quer, Trab. M u s . Cienc. N u t . Barcelona, 5(4): 23 (1924). 1 (MA), 3 (MUB). S. pusilla var. gracillima Font Quer, Trab. M u s . Cienc. Nut. Barcelona, 5(4): 22, 1924. S. pusilla var. littoralis Font Quer, Trab. Mus. Cienc. Nut. Barcelona, 5 ( 4 ) : 21 (1924). S. pusilla var. salina Font Quer, Trab. M u s . Cienc. .Nut. Barcelona, 5 ( 4 ) : 23 (1924). DISTRIBUTION: South-east Spain and North Morocco. The hair covering is shaggy on the axis, with patent or retrorsely curved hairs. O n the calyx the covering is shaggy with antrorsely curved hairs. T h e leaves are sometimes glabrescent. The base of the young stems is covered with short (700-400 pm) or very short (400-70 pm) hairs retrorsely curved, of goniotrichous distribution or less commonly holotrichous. With epicuticular flavonoids: cirsiliol, sideritoflavone, cirsimaritine, xanthomicrol and 8-methoxycirsilineol. Accumulating isocutellarein and hypolaetin or their derivates. Essential oil with a-pinene (18.7-32.7%), fenchone ( 1 1.9-12.4%), 1,8-cineol ( 10.4-10.570), sabinene (6.3-8.80/) and lesser amounts of d-cadinene (1.4-2.40/,). Chromosome numbers: n = 1 I , 13, 15 (Fernandez Peralta, 1981). This subsection seems to be related to the Arborescens group. But it is also close to subsection Leucantha, composed of hybrids, transitions or clines all very difficult to identify in the field. The most useful differential character is the retrorse direction of the hairs at the base of the stems, which is typical of the subsection Flavovirens, compared with the antrorse direction in the subsection Leucantha. zy zyxwvu S. Jauouirens (Rouy) Font Quer: Puerto Lumbreras, Murcia, Rivera (MUB). S. ochroleuca var. anliatlantica Maire: Ouarzazate, Morocco (Tomas-Barberan et al., 1980). S. ochroleuca var. maroccana Font Quer: Guercif, Morocco (Tomis-Barberan et al., 1988). S. osteoxylla Pau: Cab0 de Gata, Almeria, Alcaraz (MUB). S. pusilla (Lange.) Pau: Adra, Almeria, Rivera (MUB). S. pusilla var. curlhaginensis Font Quer: Cartagena, Murcia, Alcaraz (MUB). SPECIMENS USED FOR FLAVONOID ANALYSIS: 12. Subsection Leucantha Rivera & O b h , subsect nov. Plantae suffruticosae, caulibus lignosis, erectis. Folia caulina dentata vel integerrima, spathulata. Bracteis calycibus brevioribus vel eos subaequilongis. Corolla alba vel luteomaculata. Carpostegiatae. Axis spica rufescens. S. leucantha Cav., Icon. Descript., 4: 2, tab. 304 (1797): Habitat in regni Valentini tractu Collado d e San Antonio ubi floridam observavi mense Augusto (MA: Herbarium Cavanilles). TYPE: Font Quer (1924a) discussed the authenticity of the collection place stated on the label of Cavanilles. I t seems probable that Cavanilles confused this locality in the north of Alicante with the southern place of Orihuela, where he also collected botanical specimens; it is now the only locality of this plant. Sideritis leucantha Cav. does not grow at Collado de San Antonio, being too far north from its distribution area. It is curious to find at MA the specimen of Lagasca, named 342 zyxwvu zyxwvu zyxwvutsrqpo D. RIVERA NUNIEZ E T A L zy zyx zy zyx S. foetida, which is a typical S. Leucantha and which was collected “in collibus siccis circa Oriolam” flowering from January till August. It seems that Lagasca was also puzzled by the confusing publication of Cavanilles, but both specimens seem similar. S. leucantha Cav., Icon. Descript., 4: 2, tab. 304 (1797). 10 (BM), 11 (K), 1 (MA), 35 (MUB), 3 (P). S. bzjlora Porta, ALLi Imp. Regia Accad. Rouerelo, ser. 2 , 9 59 (1892). 2 (BM), 2 (K), 2 ( P ) . S. bourgaeana Boiss., Diagn. P1. Orient., Ser. Z(4): 34-35 (1859). 4 (BM), 6 (K). 110 (MUB), 4 (P). S. leucanlha var. incana (Willk.) Font Quer, Trab. Mus. Cienc. Nut. Barcelona, 5(4): 9 (1924). 4 (BM), 1 (K), 25 (MUB). S. leucantha var. integrzfolia Cosson in Sched. S. leucantha var. intermedia Font Quer, Trab. Mus. Cienc. Nut. BarceLona, 5(4): 10 (1924) [nomen nudum]. S. Leucantha var. meridionalis Font Quer, Trab. Mus. Cienc. Nut. Barcelona, 5(4):9 (1924). S. leucantha var. oblongfolia Rouy, Rev. Sci. Nut. Sir. 3,3(2): 246 (1883). S. leucantha var. paucidentata Willk., Bot. Zeit., 17(34): 289 (1859). S. leucantha var. serratzfolia Willk. Bot. Zeit., 17(34): 289 (1859). 3 (MUB). S. tragoriganum Lagasca, Gen. Sp. PI.: 18 (1816j. 8 (BM), 4 (K), 30 (MUB). TAXA: DISTRIBUTION: South-east Spain. The hair covering is very typical on the base of the young stems, composed of antrorsely curved hairs of variable length (Font Quer, 1925), being short 400-700 pm) or very short (70-400 pm) . Typically, the hairs have longer apical cells. The cover is holotrichous containing the epicuticular flavonoids cirsiliol, sideritoflavone, xanthomicrol, 8-methoxycirsilineol and traces of cirsimaritine and cirsilineol. Accumulating vacuolar flavonoids hypolaetin 7-allosyl ( 1 + 2) glucoside and its 3’-methyl ether derivative, hypolaetin 8-glucoside, and isoscutellarein 7-allosyl ( 1 -+ 2) glucoside. Essential oils containing a-pinene (23.6-33.5%), sabinene (10.4-10.9%) and 1,8-cineol (5.8-10.5y0); d-cadinene is present in lesser amounts (0-3.5%). Chromosome numbers: n = 13, 14 (Fernandez Peralta, 1981). Spontaneous hybrids with subsection Serrata, have been detected (Rivera & Obon, 1988b, 1989). These hybrids bear a certain resemblance to subsection Angustijiolia. Intermediate populations of this subsection with subsections Angustzfolia and Flauouirens are currently in study. The presence of hypolaetin 8-glucoside point towards the close relation with subsection Angustijiolia. S. leucantha Cav.: Santomera, Murcia, ALcaraZ (MUB). S. bourgaena Boiss.: Hellin, Albacete, Riuera (MUB). S. Leucantha var. intermedia Font Quer: Cieza, Murcia, Alcaraz (MUB). S. leucantha var. incana (Willk.) Font Quer: Entre Puerto Lumbreras y Velez Rubio, Murcia, Rivera (MUB), S. tragoriganum Lagasca) Torreblanca, Castellon, Borja (MUB); La Hoya de Altea, Alicante (Tom&-Lorente et al., 1988). SPECIMENS USED FOR FLAVONOID ANALYSIS: 13. Subsection Angustifolia Rivera & O b h , subsect. nov. Plantae suffruticosae, caulibus lignosis, tomentellis. Folia caulina lanceolata, acuta, mucronata, folia floralia inferiora et superiora dissimilia. Corollis luteis. Carpostegiatae. Axis spica leviter rufescens. TYPE: S. angustiflia Lagasca, Gen. et Sp. Nov.:18 ( 1816), emmendavit Borja, Anales zyxwvu zyxwv zyxwvu zyxwv zyxwv zyx zyxwvut SYSTEMAI’ICS OF SIDERITIS 343 Jard. Bot. Madrid, 30(2): 145-150 (1975): Habitat in montibus Regni Valentini, et praesertim circa Canales oppidum. No relevant specimen was found at MA. A neotype has been recently proposed by Socorro, Cano & Espinar (1988) from “Cerros proximos a Canals, Valencia”, Socorro (GDA). Font Quer ( 1924a) discussed the problem of this typification underlining the great variability of this taxon, but proposed no type specimen. Borja (1975) proposed that the locus classicus was Canal de Navarrks arguing that this plant did not grow at Canals, but again no type specimen was proposed. In our opinion, the choice of neotype made by Socorro, Cano & Espinar (1988) is not satisfactory because it does not solve the basic problem of the original designation of the taxon; the specimen was chosen from recent collections made in a well-known locality. In P we have found one specimen which formerly belonged to Dufour’s herbarium labelled as follows: “Sideritis angustiflia Lag. gen. et sp. (ex $so) forte S. linearzfolia Lam.? M . Dufour hisp. regn. valent.”. According to the text, the specimen was collected by Dufour and determined by Lagasca as S. angustiflia “ex ipso”. We are not sure that Lagasca considered this specimen before his publication of S. angustzfolia, normally the collector should be cited, as is usually the rule in the Lagasca writings. As Dufour’s name was not quoted this specimen was probably seen after publication, but having been accepted by the author, this is the best specimen that can be chosen as the neotype. S. angustzfolia Lagasca, Gen. et Sp. Nov.: 18 (1816). 13 (BM), 10 (K), 1 (MA), 150 (MUB), 2 (P). S.funkiana Willk., Bot. zeit., 17(33): 282 (1859). 1 (BM), 1 (K), 10 (MUB). S.jahandiezzii Font Quer, Publ. Junta Ci. Nut. Barcelona, Sir. Bot., 6 25 (1924). 1 (BM), 2 ( M P U ) , 4 (P). S. lagascana Willk., Bot. zeit., 17(33): 282 (1859). 60 (MUB). S. mugronensis Borja, Anales Jard. Bat. Madrid, 3 8 357 (1982). 1 (BM), 1 (K), 1 (MA), 30 (MUB). S. reverchonii Willk., Suppf. Prodr. Fl. Hisp.: 156 (1893). 4 (BM), 6 (K), 6 (MA), 20 (MUB), 6 (P). S. suetabensis Rouy, Bull. Soc. Bot. France, 29: 125 (1882). 1 ( K ) , 1 (MA), 6 (MUB), 1 (P). TAXA: South and East Spain. Centres of diversity occur in the eastern border of the Meseta, the Guadix-Eaza area with a secondary centre in Malaga. Represented by one taxon in Northern Africa. DISTRIBUTION: Hair covering shaggy on the stems with patent or antrorsely curved hairs. Calyx with antrorsely curved hairs. Leaves with some disperse hairs which fall away. The hair covering on the base of the young stems is shaggy, holotrichous, with antrorsely curved, more or less robust medium, short and very short hairs. Possessing the epicu ticular flavonoids: sideritoflavone, cirsiliol, 8-methoxycirsilineol and traces of cirsimaritin, cirsilineol and xanthomicrol. Accumulating as vacuolar flavonoids hypolaetin 7-allosyl ( 1 .+ 2) glucoside and its 3’-methyl ether, hypolaetin 8-glucoside and isoscutellarein 7-allosyl ( 1 -+ 2) glucoside. Essential oil with a-pinene (10.7-23.6%) and 1,8-cineol (14.4-19.4O/”), reduced amounts (less of 5%) of d-cadinene and sabinene. Chromosome numbers: n = 10, 12 (Fernandez Peralta, 1981). The hair covering shows the gradual change from the western forms with short and robust hairs, from Malaga (S. reuerchonii Willk.), to the eastern populations of S. angustzfolia Lagasca or S. saetabensis Rouy with thinner, medium or short 344 zyxwvut zyxwvutsrqpo D. RIVERA NUmEZ E T AL. hairs. Similarities in hair covering and flavonoids suggest that the mountainous taxa S. glacialis Boiss. of the subsection Scordioides or S. jaualambrensis Pau, belonging to the subsection Linearifolia, are related to those of subsection Angustzfolia living on neighbouring plains. Morphological relationships, fundamentally in the hair covering, are shown by the eastern taxon S. angustifolia Lagasca with the subsection Scordioides. The presence of hypolaetin 8-glucoside is shared only with the subsection Leucantha; this flavonoid appears only in traces in two other subsections. This group has one endemic chemotype producing 5-desmethylnobiletin and the diterpene borjatriol in the province of Albacete. I n south-east Spain clinal variations were detected within the subsection Leucantha. Spontaneous hybrids are known with the subsections Hirsuta, Gymnocarpae and Scordioides (Font Quer, 192 1) . zyxwvu zyxwv zyxwvuts S i d e r i h angustzfolia Lagasca: Ayora, Valencia, Rivera & D e L a Torre (MUB); Sierra d e Mariola, Alicante, Obdn, Carreras & Rivera (MUB); Fuente la Higuera, Valencia, Obdn, Carreras @ Rivera (MUB); Alpera, Albacete, Rivera (MUB). Sideritis jahandiezii Font Quer: Almis de Gigou (Tomis-Barberan et al., 1988). Sideritis mugronensis Borja: Los Llanos, Albacete, Rivera, (MUB); Las Mariquillas, Albacete, Rivera, (MUB); Sierra del Mugrbn, Albacete, Obdn (MUB). Sideritis reverchonii Willk.: Ronda, Malaga (Tomas-Lorente el al., 1988). Sideritis saetabensis Rouy: JBtiva, Valencia, Obdn, Carreras & Rivera (MUB). SPECIMENS USED FOR FLAVONOID ANALYSIS: 14. Subsection Serrata Rivera & Obbn, subsect. nov. Plantae fruticosae vel suffruticosae, caulibus lignosis, tomentellis, pilis glandulosis munitis, foliis lanceolatis, acutis, mucronatis, serratis vel spinoso-dentatis. Corollis luteis. Carpostegiatae. S. serrata Lagasca, Gen. et. Sp. Nov.:18 (1816). Habitat in montibus ditionis Tobarra oppidi in Murciae Regno (MA). Lagasca quotes clearly the label in Cavanilles’ handwriting “Sider, spinosa Cav. Herb.” which is the same found on the sheet 101033 at MA. T h e specimen on the right side was designated the lectotype (Bellot gL Casaseca, 1975). TYPE: TAXA:S. serrata Lagasca, Gen. et Sp. Nov.: 18 ( 1816). 2 (BM), 1 (MA), 90 (MUB). S. fragrans Costa ex Rouy, Illustr. PI. Eur. Rar., 2 271 (1899). S. iliczfolia Willd., Enum. Pl. Hort. Berol.: 606 (1809). 2 (BM), 5 (K), 10 (MUB), 1 (P). DISTRIBUTION: East Spain, from Ebro Basin to Catalonia. One isolated locality in the Albacete province. Sideritis iliczfolia has up to 16 flowers per whorl. T h e hair covering is glandular and shaggy on the stems, with long erecto-patent hairs. Calyx with antrorsely curved hairs, with a similar covering on the leaves. Bases of the young stems densely covered with very short glandular hairs (70-150 pm) mixed with long, medium, short or very short patent hairs. Possessing epicuticular flavonoids as cirsiliol, sideritoflavone, cirsimaritin, cirsilineol, xanthomicrol, 8-methoxycirsilineol, 5-desmethylnobile’tin and gardenin B. Accumulating the vacuolar flavonoids hypolaetin 7-allosyl ( 1 + 2) glucoside and its 3’-methyl ether, isoscutellarein 7-allosyl ( 1 4 2) glucoside and its 4’-methyl ether. Essential oil zyxwvu zyxwvu zyx SYSTEMATICS OF SIDERI’TIS 345 with sabinene (20.6%), a-pinene (14.9%) and 1,8-cineol (8.6%), but no significant amounts of d-cadinene (S. serrata). Chromosome number: 2n = 28 (Obbn & Plantrose, unpublished count for S. serrata). It seems to be related to the subsection Scordioides, with transitional forms of possible hybrid origin in the Ebro Basin (Font Quer, 1920). Sideritis serrata hybridizes freely with S. bourgaeana of the subsection Leucantha in their overlapping areas near Tobarra (Rivera & Obon, 198810, 1989). Font Quer (1921) has published one hybrid of S. iliczfolia with the group S. hirsuta. zyxwvu zyxwvu zyx ANALYSIS: Sideritis serrata Lagasca: Tobarra, Albacete, Rivera (MUB). Sideritis ilicifolia Willd.: Barbastro, Huesca, Proudhomme (MUB). SPECIMENS USED FOR FLAVONOID 15. Subsection Scordioides Rivera & Obon, subsect. nov. Plantae suffruticosae, caulibus lignosis erectis vel decumbentibus, humiles. Folia caulina obtusa, ovato-lanceolata vel spathulato-cuneata, dentata vel spinulosa aut serrato-spinosa. Corollis pallide luteis. Carpostegiatae. Pilis mollibus adpresis vestita. S. scordioides L., Syst. Nut. ed. 10, 2: 1098 (1 759): a D.Sauvagesio, but no locality is given. TYPE: Linnaeus also cites Barrelier ( 1 7 1 7 ) icon 343. Linnaeus in Sp. PI. ed. 2, 2.803 (1 763), gave further information on the locality, “Habitat Monspelii”, citing Sauvage as sender. C. E. Jarvis (personal communication) gave his opinion on the specimen 729.12 in LINN, as being a syntype with two further possible syntypes in the Bergius Garden, Stockholm (SBT), and drew attention to the citation of Barrelier. Specimen 729.12 in LINN could hardly be included in the currently accepted sense for Sideritis scordioides L. Further research is needed in SBT for typification. TAXA:5’. scordioides L., Syst. N a t . ed. 10, 2: 1098 (1759). 12 (BM), 1 (K), 8 (MA), 2 ( M P U ) , 5 (P). S. cauanillesii Lagasca, Gen. Sp. PI.: 18 (1816). 7 (BM), 9 (K), 10 (MA), 3 (MUB), 5 (P). S. crispata Willd., Enum. PI. Hort. Berol.: 606 (1809). S. glacialis Boiss., Biblioth. Universelle Gen2ve st?. 2(13): 410 (1838). 15 (BM), 15 (K), 5 (MUB), 1 ( P ) . S. glacialis var. pulvinata Font Quer in sched. 1 (BM). S. mariae Sennen, Bol. Real SOC.Iber. Cienc. &at. 31: 120 (1934). 1 (MA). S. spinosa Lam., En&. M i t h . , 2: 169 (1786). 2 (P). S. spinulosa Barnades ex ASSO,Intr. Oryctogr. Arugdn: 1 7 1 ( 1 784). 8 (BM), 2 (K),6 (MUB). S. spinulosa f. intermedia Font Quer, Bol. Real Soc. Iber. Cienc. Nut.: 137 (1920). S. subspinosa Cav., Icon. Descript., 3 5 (1795). 3 (BM), 3 (K), 1 (MPU), 1 (MUB), 1 (P). DISTRIBUTION: North Spain and South France. Centre of diversity around Ebro Basin and South Pyrenees. Isolated locality in Sierra Nevada. The hair covering is shaggy or tomentose on the sterns, calyx, bracts, leaves and bases of the stems, with adpressed erect very long, long, medium and rarely short hairs. Possessing of the epicuticular flavonoids sideritoflavone, cirsimaritin, cirsilineol, xanthomicrol. Accumulating the vacuolar flavonoids hypolaetin 7-allosyl ( 1 +. 2) glucoside and its 3’-methyl ether derivative, isoscutellarein 7-allosyl ( 1 -+ 2) glucoside and its 4’-methyl ether derivative. Essential oil with 346 zyxwvut zyxwvutsrqpo zyxwv zyx D. KIVEKA NUREZ E T A L a-pinene (19.876) and 1,8-cineol (13.8%) but with little sabinene (5.8%) and no significant amounts of d-cadinene (5’. spinulosa). Chromosome numbers: n = 1 1, 13, 14, 15, 17 (Ferniindez Peralta, 1981). This group displays the gradation from unarmed to spinose leaves, with unarmed populations under the name S. scordioides L., intermediate ones under S. cavanillesii Lag. and the spinose under S. spinulosa Barnadts. Hybridizes with the subsections Lineariflia, Hirsuta, Serrata, Angustifilia (Font Quer, 192 1 ) or Ouata (Font Quer, 1924a), in their sympatric areas. SPECIMENS USED FOR FLAVONOID ANALYSIS: S. glacialis Boiss.: Cerro del Almirez, Almeria, Robledo (MUB). (S. spinulosa Barnadts ex Asso: El Cerrato, Palencia (Tomiis-Lorente et al., 1988). DISCUSSION Section Sideritis is located in the western border of the Mediterranean Basin, with a Mediterranean climate, although some species belonging to this group extend towards the north end of the southern European region of temperate climate. Other species, such as S. glacialis, live under conditions very similar to those of the alpine mountains and, finally others, e.g. S. maireana, invade the boundaries of the Sahara Desert. The presence of endemic sections of the genus Sideritis in the Canary Islands could be interpreted as a relict of the Tertiary taxa living in the Mediterranean Basin and North Africa, which reached the islands as late as 2.5 M.y. B.P., and have survived under the influence of an oceanic climate (cf. Raven & Axelrod, 1974). From a palynological viewpoint, the Canarian species belonging to the sections Marrubiastrum or Empedocleopsis must be considered as ancestors of the Mediterranean section Sideritis (Huynh, 1972). Fernandez Peralta (1981) presumed that evolution took place both in the Mediterranean and Canarian regions in a more or less parallel way, originating in one herbaceous perennial ancestor with a chromosome number n = 14, morphologically similar to the sections Burgsdorfia and Hesiodia. This opinion entails the primitiveness of taxa with n = 13, 14 and the higher degree of evolution in taxa with n = 10, 11, 12, 15, 16, 17. As far as can be determined, the first branching in the section separated subsections Hirsuta and Gymnocarpae. This follows the theories of Willis (1922) of the largest distribution areas belonging to the oldest taxa. According to Huynh (1972) and Fernandez Peralta (1981) the primitive states of section Sideritis should be summarized as follows: plants living on stony places, with predominantly herbaceous stems; leaves clearly petiolated, broad, with the margin crenate to lobulated; bracts similar to the leaves in size, form and colour, or gradually changing from the lower part of the inflorescence to the apex; large flowers, with a very long pedicel; calyx actinomorphic, with the throat broad and the lobes very long; corolla shorter than calyx, pale yellow coloured; large seeds; hair covering shaggy with very long patent hairs or tomentose, holotrichous. These characters are more or less represented in S. grandgora, whose chromosome number is still unknown. This species has more than ten flowers per whorl, sometimes up to 30; it is possible to assume that a greater number of flowers per whorl is primitive, the number being gradually reduced to six or even fewer in the highly evolved forms. zyxwvut zyxwvu SYSI'EMAIICS OF SIDERI'TIS 347 Throughout evolution, the hair covering seems to have remained relatively constant, the primitive state being slightly ambiguous as is shown by the subsection Hirsuta, which has evolved a predominant direction of the hairs on the stems, either downwards or upwards. T h e hairs are retrorse in the subsections Arborescens, Camarae and Flauouirens, and antrorse in the rest. Separation of groups with different hair coverings seems to have been very old and not reversible: no mutants have been discovered showing this reversibility. Accordingly, the subsection Flauouirens cannot have evolved from subsection Leucantha as proposed by Fernandez Peral ta ( 1981 ) , but probably evolved from subsection Arborescens through reduction of the length of hairs and leaf dimensions. This also seems to be the direction of evolution in subsection Camarae, seen in the glabrous S. brachycalyx. Twisted hairs are exclusive to subsections Stachydioides and Gymnocarpae and the eastern Mediterranean section Empedoclea. T h e centres of diversity of the subsections are situated around the Tirrenian Basin from East Spain to North Tunis, and they are probably vicariants of the eastern section Empedoclea. I t is difficult to accept that plants with straight hairs have directly evolved from plants with very twisted hairs, as proposed by Fernandez Peralta (1981) for S. lacaitae, which cannot be interpreted as being derived from S. incana. Hybridization could be claimed to explain the origin of S. lacaitae, but the known hybrids of section Gymnocarpae also have twisted hairs (Rivera & Obbn, 1988a). T h e spiny leaves associated with the presence of stipitate glands on the stems are exclusive to the subsection Serrala, this subsection probably being older than S. hirsuta. Apart from this species, glandular hairs appear only in S. grandgora. The primitive groups very often lack epicuticular flavonoids. The evolved states should be: plants with woody stems; long, narrow leaves with the margin more or less entire without differentiated petiole; bracts dissimilar to the leaves, and similar to the inflorescence; six or fewer flowers per whorl; small flowers, with short pedicels; calyx more or less zygomorphic, with the throat narrow and the lobes short; corolla normally longer than the calyx, bright yellow coloured; small seeds; hair covering hoary or glabrous, goniotrichous. T h e chromosome number n = 17, according to Fernandez Peralta (1981), belongs to highly evolved taxa more or less adapted to rocky places, such as S. lacaitae, S. stachydioides, S. glauca and S. glacialis, every one belonging to different subsections. These plants are taxonomically easy to define, they have relatively reduced morphological variability, small areas, and probably have reached the higher degree of speciation within the section. As they were evolving under moister climatic conditions, they did not produce epicuticular flavonoids. Only S. glacialis evolving under a very cold climate, which implies physiological dryness, has produced epicu ticular flavonoids. Other groups, such as subsections Flavouirens, Leucantha or Anguslijolia, are currently speciating; the epicuticular flavonoids play an actual role in this process under dry climatic conditions. Variation occurs at the level of populations, mutants concerning flower colour, size and form of leaves and density of hair covering are relatively frequent within these taxa. Human disturbance has influenced recent evolution by providing opportunities for hybridization. Frequently, it is possible to see two or more species living closely in dense clumps, conditions which are very favourable to steady hybridization (Rivera & Obbn, 1989). zyxw 348 zyxwvu zyxwvu zyxwvutsrqpon D. RIVERA NUREZ E l AL. ACKNOWLEDGEMENTS The authors are indebted to Dr Borja who communicated much unpublished original work concerning morphology. Dr C. Guy very kindly allowed us to use her Ph.D. dissertation. REFERENCES BARRELIER, J., 17 14. Plantae per Galliam, Hispaniam et ltaliam obseruatae iconibus aeneis exhibitae. Paris: Stephan Gaveau. BELLOT, F. & CASASECA, B., 1975. Specimens “Typus” de quelques especes proposes par Mariano Lagasca, se trouvant dans I’herhier du jardin hotanique de Madrid. 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