Figure 1.23 T. maroaanus (Bot. J. Linn. Soc., 16: pi. 27).
Figure 1.23 T. maroaanus (Bot. J. Linn. Soc., 16: pi. 27).
6. Subsect. Pseudomarginati
7. Subsect. Serpyllum
I. Sect. Micantes Velen., Bei. Bot. Centr. 19(B2): 278 (1906) Typus: T. caespititius Brot.
Erect plants (North-African species) or caespitose; stems holotrichous; leaves flat, glabrous, long oblong-obovate; inflorescence spiciform, sometimes dense; lateral upper teeth of calyx very short or reduced.
It comprises three species, two of them are North African woody species, that occur in Morocco: T. satureioides and T. riatarum. The former inhabits the High Atlas region, while T. riatarum is a prostrate plant and lives in the Rif mountains. The Ibero-Macaro-nesian species T. caespititius occurs in the northwest of the Iberian Peninsula and also in Madeira and Azores. If we take into account their plesiomorphic features, like flat, non-revolute and glabrous leaves, and their geographical distribution, this section seems to be very old.
II. Sect. Mastichina (Miller) Bentham, Lab. Gen. Sp.: 340 (1834) Masticbina Miller, Gard. Diet. ed. 4 (2) (1754).
Erect plants with holotrichous stems, leaves flat, lanceolate to obovate; inflorescence capituliform; calyx very hairy, teeth similar and subulate, with long cilia.
This section is endemic to the Iberian Peninsula, and comprises T. mastichina, with two subspecies, and T. albicans. T. mastichina subsp. mastichina is a very common plant in Spain and Portugal. The subspecies donyanae occurs only in the southwest of the Iberian Peninsula around the 'Coto de Donana' and in some locations in the 'Algarve'. The other species, T. albicans, is also living in the southwestern pinewoods of Pinus pinea. T. mastichina subsp. donyanae and T. albicans, with 2n = 30 chromosomes, are probably the origin of the tetraploid apoendemic T. mastichina subsp. mastichina, a modern taxon that has spread throughout the entire Iberian Peninsula.
III. Sect. Piperella Willk., Prodr. Fl. Hisp. 2: 404 (1868). Typus: T. piperella L.
Erect or decumbent plants, with holotrichous stems and leaves obovate, flat and glabrous; flowers growing in lax verticillasters.
T. piperella is found at Valencia province and surroundings, and it is the unique species of this monotypic section, endemic to this region.
IV. Sect. Teucrioides Jalas, Bot. J. Linn. Soc. 64(2): 201 (1971). Typus: T. teucrioides Boiss. and Spruner.
Plants usually decumbent with leaves revolute, ovate or triangular-ovate; flowers in verticillasters. Endemic to the Balkan Peninsula. It inhabits the mountains of Greece and Albania. Three species can be recognised within this section: T. teucrioides, T. hartvigi, and T. leucospermus, that have been studied by Hartvig (1987). Chromosome numbers of these species are not yet known.
V. Sect. Pseudothymbra Bentham, Lab. Gen. Sp.: 341 (1834). Typus: T. Otocephalus G. Lopez and R. Morales (T. cephalotos auct. non L.)
Erect plants with holotrichous stems and linear revolute leaves, usually hairy and with cilia at the base; inflorescence capituliform with broad bracts; corolla very long.
In this section are included nine Iberian-North-African species, usually with long corollas, up to 2 cm, and bracts rather different from the leaves and subglobose inflorescence, except in subsection Anomalae. The North-African species are T. munbyanus, common and very variable, extending from the Middle Atlas and the Rif Mountains as far as the Algerian mountains. A difficult species with two subspecies and hybrids with T. algeriensis and T. willdenowii. T. bleicherianus is only known from three locations, one in Algeria and two more in the north of Morocco. The other species of this section are all Iberian.
We recognize two subsections:
VI. Subsect. Pseudothymbra (Bentham) R. Morales, Ruizia 3: 146 (1986). Inflorescence capituliform and bracts are very different from the leaves.
V2. Subsect. Anomalae (Rouy) R. Morales, Ruizia 3: 146 (1986). T. sect. Anomalae Rouy, Bull. Soc. Bot. France 37: 166 (1890). Typus: T. antoninae Rouy and Coincy
Flowers in verticillasters, bracts with similar appearance as the leaves.
VI. Sect. Thymus
Erect or radicant plants with holotrichous stems, revolute leaves, usually hairy; flowers in spiciform or globose inflorescences.
Western Mediterranean section, with three most important species: T. vulgaris, T. zygis and T. willdenowii. The first usually occurs on basic soils and is distributed in northern Italy, south of France and east of Spain. T. zygis is a very common species in all the Iberian Peninsula and T. willdenowii is common in North Africa (Morocco and Algeria) and also grows only in Gibraltar area in the Iberian Peninsula. We recognize two subsections:
VII. Subsect. Thymastra (Nyman ex Velen.) R. Morales, Ruizia 3: 146 (1986). T. sect. Thymastra Velen., Bei. Bot. Centr. 19(B2): 276 (1906).
Typus: T. capitellatus Hoffmanns, and Link.
Erect plants with triangular-ovate or lanceolate-ovate leaves, without cilia at the base; inflorescence more or less globose with bracts different from the leaves. VI2. Subsect. Thymus
Erect or subtended plants with leaves usually hairy, and ciliate or not at the base, with revolute margins and more or less linear, bracts broader than the leaves, but not very different.
VII. Sect. Hyphodromi (A. Kerner) Halacsy, Denkschr. Akad. Wiss. Wien 61: 252 (1894). Typus: T. bracteosus Vis. ex Bentham.
Plants usually subtended and rooting; stems holotrichous; leaves flat or revolute, usually not hairy; inflorescence frequently capituliform with bracts different from the leaves.
This section extends throughout the Mediterranean area and comprises around 60 species. From the three subsections, Subbracteati is characterized by more or less revolute or convolute leaves and seems to be Oriental. Only one species occurs in North Africa, from Morocco to Libya: T. algeriensis. Another species occurs in Central Spain: T. mastigophorus. T. spinulosus occurs in Sicily and Italy, and T. striatus in the Italian and Balkan Peninsulas. Both species are very variable. T. argaeus, T. brachychilus, T. cappa-docicus, T. cherlerioides, T. convolutus, T. pulvinatus, and T. revolutus occur in Turkey; T. boissieri, T. comptus, T. dolopicus, and T. plasonii in the Balkan Peninsula; T. atticus, T. parnassicus, and T. leucotrichus inhabit Turkey and the Balkan Peninsula. The last species also grows in Syria and in the Lebanon. T. integer is only found on the island of Cyprus. This species is probably not different from T. leucotrichus. T. samius occurs in the Aegean islands. T. borysthenicus and T. pallasianus occur north of the Black Sea, T. persicus south of the Caucasus, but only one location for this species is known.
We recognize three subsections:
VIII. Subsect. Subbracteati (Klokov) Jalas, Bot. J. Linn. Soc. 64(2): 205 (1971), emend. T. sect. Subbracteati Klokov, Not. Syst. (Leningrad) 16: 315 (1954) pro parte. Typus: T. pallasianus H. Braun.
VII2. Subsect. Serpyllastrum Huguet del Villar, Cavanillesia 6: 124 (1934). Lectotypus: T. bracteosus Vis. ex Bentham.
VII3. Subsect. Thymbropsis Jalas ex R. Morales, Anales Jard. Bot. Madrid 45(2): 562 (1989).
Typus: T. maroccanus Ball.
Subsection Serpyllastrum is a group of species characterized by the presence of prostrate stems and flat leaves more or less wide. Five species from this section are living in Spain: T. bracteatus, T. leptophyllus, T. fontqueri, T. granatensis and T. lacaitae. It is also well represented in the East, but no species occur in Italy and North Africa. T. aznavourii and T. bracteosus occur in the Balkan Peninsula; T. canoviridis, T. haussknechtii, T. pectinatus and T. spathulifolius are found in Turkey. T. zygioides extends from the Balkan Peninsula as far as the Crimean Peninsula and also in Turkey. This species and the Spanish endemic T. lacaitae are morphologically very similar. There is also a group of species that occur only in the Caucasus: T. dagestanicus, T. hadzhievii, T. helendzhicus, T. karjagnii, T. ladjanuricus, T. lipskyi, T. majkopiensis, and T. sosnowskyi. Seven more species from Central Asia are considered inside this subsection: T. cuneatus, T. eremita, T. incertus, T. irtyscbensis, T. kirgisorum, T. nerczensis, T. petraeus.
Subsection Thymbropsis includes the North African T. broussonetii, T. maroccanus, T. lanceolatus, T. numidicus, T. pallescens, and the two endemic species from Greece T. laconicus and T. holosericeus. Five more species from this section are found in Turkey: T. cariensis, T. cilicicus, T. eigii, T. leucostomus, and T. sipyleus. T. syriacus occurs in Lebanon, Syria and a location in northern Irak; T. bovei lives in the Sinai Peninsula, Israel, Jordan, Irak and Saudi Arabia; and T. decussatus in Sinai and Saudi Arabia. This group has predominantly North-African and East-Asian species.
VIII. Sect. Serpyllum (Miller) Bentham, Lab. Gen. Sp.: 340 (1834). Serpyllum Miller, Gard. Diet. ed. 4 (3) (1754).
Woody plants or only woody at the base, but with herbaceous appearance, usually subtended and rooting, with holotrichous stems or hairy only in two opposite sides or in the angles (goniotrichous or alelotrichous), leaves flat and usually ciliate at the base, with distinct lateral veins; inflorescence spiciform or more or less globose.
In this section there are around 120 species. They occur throughout the area of the genus, except in Madeira and the Azores. We find in the species of Serpyllum the widest chromosomal variation. There are also woody species that grow in the mountains in arid areas like T. origanoides on Lanzarote (Canary Islands); T. serrulatus and T. schimperi in Ethiopia, T. laevigatus in the southwest of the Arabian Peninsula. Another group of species are more or less herbaceous and occur in the Mediterranean mountains, and all of Eurasia and also along the coasts of Greenland. The species of the last group seem to be younger in evolutionary terms and have probably been actively evolving since the last giaciation when this group colonized the new lands free of ice. This group is also very difficult taxonomically and corresponds to the last three subsections. Few species of these subsections are present in the Mediterranean area. According to Jalas (1971), we divided this section into seven subsections.
Subsection Insulares comprises T. willkommii, an endemic species that occurs in the mountains of the provinces of Castellón and Tarragona (eastern Spain); T. richardii, with three subspecies: subsp. richardii from Majorca and Yugoslavia, subsp. ebusitanus from Ibiza and subsp. nitidus from Marettimo island near Sicily; the North-African T. dreatensis and T. guyonii, the Canary Island endemic T. origanoides and the endemic species to northwest Turkey T. bornmuelleri.
Subsection Kotscbyani includes a lot of Asian species, but only T. fallax and T. transcaucasicus occur in Turkey. Other interesting species occurring outside the Mediterranean area are T. laevigatus from the mountains of Yemen or T. schimperi and T. serrulatus from the Ethiopian mountains.
Subsection Pseudopiperellae comprises T. herba-barona from Majorca, Corsica, and Sardinia (Mayol et al., 1990) and T. nitens from the south of France.
Five species inhabiting the Balkan Peninsula belong to the subsection Isolepides\
T. bulgaricus, T. glabrescens, T. longedentatus, T. pannonicusm, and T. sibthorpii.
Subsection Alternantes includes T. linearis from the Himalaya mountains; the European T. pulegioides, T. froelichianus, T. alpestris, T. oehmianus, T. bihoriensis, and T. comosus.
Subsection Pseudomarginati includes the species T. longicaulis and T. praecox, very common in Europe and also in Turkey; T. nervosus, an endemic of the Pyrenees and the French Massif Centra/; T. ocheus, T. stojanovii, and T. thracicus from the Balkan Peninsula and the East Mediterranean region.
Subsection Serpyllum includes T. quinquecostatus from Japan, the European T. serpyllum and T. talijevii and other Russian species.
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