Micromeria microphylla (d'Urville, 1822) Bentham, 1834

Bogdanović, Sandro, Ljubičić, Ivica, Sáez, Llorenç & Brullo, Salvatore, 2024, Micromeria silicii, a new species from Croatia and its relationships with taxa of the M. microphylla group (Lamiaceae), Phytotaxa 662 (1), pp. 1-18 : 7-15

Remarks on Micromeria microphylla complex

Apart from the description of the new species from Croatia, we provide the taxonomical, nomenclatural and chorological consideration on the species that according to the literature belong to M. microphylla complex. In the Table 1 the main comparative morphological features regarding these species are listed. The investigated taxa are:

1) Micromeria acropolitana Halácsy ex Maire & Petitmengin (1908: 421)

≡ Satureja acropolitana (Halácsy) Greuter & Burdet in Greuter & Raus (1984: 302).

Type (lectotype designated by Bräuchler in Bräuchler et al. 2008a):— GREECE. In Acropoli Athenarum, 30 August 1906, R. Maire & M. Petitmengin 1073 (lectotype WU0040407!).

Description:—Stems up to 30 cm tall, branched, erect, densely hairy, reflexed, 0.1–0.4 mm long; leaves subsessile, acute, 1.8–5 × 0.4–1.5 mm, basal ones often cordate-lanceolate, upper ones oblong-lanceolate, densely hairy on both sides, with hairs 0.2–0.5 mm long; verticillasters of 2 pedunculate fascicles, each 2–5(–7)-flowered; calyx 2.4–2.5 mm long, with teeth 0.6–0.8 mm long, linear-triangular, densely pubescent, patent, tube hairs 0.5–1 mm long, teeth hairs shorter (0.1–0.3 mm long); corolla 3 mm long, with tube included in the calyx; stamens exserted from the tube.

Habitat and ecology: —Rocky places, on calcareous substrata and also in crevices of archeological site.

Iconography: — Fig. 5a in Tan et al. (2010).

Chromosome number: —2n = 30, Siljak-Yakovlev et al. (2011).

Distribution: — Greece, Attica, Acropolis of Athens ( Fig. 2 ).

Notes: —This species was described by Maire & Petitmengin (1908) almost simultaneously with Halácsy (1908) and it is successively accepted by Chater & Guinea (1972), Greuter et al. (1986), Doroszenko (1986), Tan et al. (2010), Siljak-Yakovlev et al. (2011) and Dimopoulos et al. (2013), while Bräuchler et al. (2008a) treated it doubtfully as a synonym of M. microphylla .

Specimina visa:— GREECE. Attica, Athen, Akropolis und Philopappos, Fels und steinige Abhange, ca. 150 m, 14 May 1967, W. Lippert & B. Zollitsch s.n. (M0158796!).

2) Micromeria carpatha Rechinger f. (1949: 208)

Type (lectotype designated by Bräuchler in Bräuchler et al. 2008a):— GREECE. Karpathos, 5 June 1886, Forsyth Major 165a (lectotype G00087102!, isolectotype Z000039167!).

= Micromeria microphylla Rechinger f. (1943: 526) non Bentham (1834: 377), pro. parte.

Description:—Stems 15–25 cm tall, very branched, erect, subglabrous; leaves subsessile, acute, 3.3–6 × 0.7–2.4(–3) mm, basal ones often cordate-lanceolate, upper ones oblong-lanceolate, scabrid, with very short hairs (0.1 mm long) on both sides; verticillasters of 2 pedunculate fascicles, each 1–3-flowered; calyx 3–3.5 mm long, with upper teeth 0.8–0.9 mm long, linear-triangular, lower teeth 1.2–1.5 mm long, pubescent, hairs patent, 0.1–0.2 mm long; corolla 3 mm long, slightly exserted, with tube included in the calyx; stamens exserted from the tube; mericarps 0.6 mm long.

Habitat and ecology: —Calcareous rupestrian stands, as cliffs, rocks, walls, ravines and boulders.

Iconography: — Fig. 5d in Tan et al. (2010).

Chromosome number: —unknown.

Distribution: — Greece, Karpathos island ( Fig. 2 ).

Notes: —This species was described by Rechinger (1949) and it is accepted by Tan et al. (2010), Dimopoulos et al. (2013) and Strid (2016). Chater & Guinea (1972), Greuter et al. (1983, 1986) treated it as a synonym of M. sphaciotica , while Doroszenko (1986) and Bräuchler et al. (2008a) as a synonym of M. microphylla .

3) Micromeria cypria Kotschy in Unger & Kotschy (1865: 270), Fig. 4

≡ Satureja graeca subsp. cypria (Kotschy) Holmboe (1914: 159) .

≡ Satureja cypria (Kotschy) Prain (1921:233) .

≡ Micromeria graeca subsp. cypria (Kotschy) Chapman (1949: 430) .

≡ Micromeria graeca subsp. cypria (Kotschy) Rechinger f. (1950: 430) .

Type (lectotype designated by Bräuchler in Bräuchler et al. 2008a):— CYPRUS. In scopulorum fissures montis Pentadactylos et Buffavento, 1500’ elevation, April 1862, Kotschy 338 (lectotype W 0014293!, isolectotypes JE 00007501! but excluded specimens marked by X, K 000193721!, P 00714527!, P 00714528!, S 08-10884!, BP-TRA 00573276!).

= Micromeria cypria var. villosissima Lindberg (1946: 29) Type: lectotype designated by Väre (2012):— CYPRUS. distr. Kyrenia, Ayios Hilarion, in fiss. rup. sicc., 7 June 1939, H. Lindberg s.n. (lectotype H-1413162!).

= Satureja graeca Linnaeus subsp. graeca var. pauciflora (Visiani) Briquet (1895: 420) pro. parte (type not located).

Description:—Stems 10–30 cm tall, ascending to erect, simply branched, hairy, hairs patent, 0.05–0.5 mm long; leaves subsessile, ovate to lanceolate, acute, 3–9 × 1.5–5 mm, hairy, with appressed hairs, 0.2–0.8 mm long on both sides; verticillasters of 2 pedunculate fascicles, each 1–7-flowered; calyx 3–4 mm long, tube with hairs 0.1–0.2 mm long, upper teeth 0.5–0.7 mm long, triangular, lower teeth 0.9–1.2 mm long, triangular-subulate, hairy, hairs patent, 0.2–0.3 mm long; corolla 5–5.5 mm long, with tube long exserted from the calyx; stamens exserted from the tube; mericarps 0.7 × 0.4 mm.

Habitat and ecology: —Calcareous rupestrian stands, as cliffs, rocks, walls, ravines and boulders.

Chromosome number: —unknown.

Distribution: — Cyprus Island, Kyrenia Mt range ( Fig. 2 ). According to Hand (2011) specimens identified as M. microphylla were collected in the western part of the island near Androlikou gorge.

Notes: —It was described by Kotschy in Unger & Kotschy (1865) and later accepted by Lindberg (1946) and Greuter et al. (1986), while Holmboe (1914) consider it as a subspecies of M. graeca . Besides, Meikle (1985) believes that M. cypria may fall within the variability of M. microphylla , since the differential characters between the two species do not result very clear. This opinion is also shared by Hand et al. (2011) too. Based on our investigations, using herbarium materials coming from the type locality, the Cyprian populations are morphologically quite distinct from the typical ones of M. microphylla , as well as from other species belonging to this group. Therefore, we agree with Greuter et al. (1986) to keep them distinct at a specific level as proposed by Kotschy in Unger & Kotschy (1865).

Specimina visa:— CYPRUS. Insula Cyprus, in rupium fissuris Mt Xerobuno, prope St. Claysostomo , May 1889, Pichler s.n. ( LY0495763 , P03833405 !) ; Kornos (Kyrenia range), In and among rocks in limestone ravines above Larnaca — Lapithou , 306–457 m, 3 April 1941, P. H. Davis 2995 ( H1490680 !) ; distr. Kyrenia, Pentadactylos, in fiss. rup. sicciss. juxta Armeniae Monastery , 5 July 1939, H. Lindberg s.n. ( H1413163 !) ; distr. Kyrenia, Boghazi, in saxosis sicciss. in valle rivuli supra opp. Kyrenia, 5 June 1939, H. Lindberg s.n. ( H1413161 !), distr. Kyrenia, Agios Hilarion , in fiss. rup. sicc., 7 June 1939, H. Lindberg s.n. ( H1413164 !) ; distr. Kyrenia, Buffavento, Castle , 23 April 1973, P. Laukkonen 283 ( H1088576 !) ; Kytrea , 2 February 1966, O. Houvila 65 ( H1088742 !) ; In M. Pentadactylos , May 1880, P. E. E. Sintenis & G. Rigo 121 ( K000193722 !, PRC 462224!, P03833404 !, P02971198 !, P02971199 !, P02971200 !) ; Pentadactylo range, Buffavento, rupi calcaree, 6 July 2013, G. Giusso del Galdo & S. Brullo s.n. ( CAT!) .

4) Micromeria hispida Boissier & Heldreich ex Bentham (1848: 215)

≡ Satureja candica Greuter & Burdet (1985: 302) . ≡ Satureja hispida (Bentham) Nyman (1855: 102) . ≡ Clinopodium hispidum (Bentham) Kuntze (1891: 515) . Type (lectotype designated by Bräuchler in Bräuchler et al. 2008a):— CYPRUS. Creta in rupestribus Eparchia Mirabello, April 1846, T.

de Heldreich (lectotype G-BOIS00769782!, isolectotypes GH00001324!, GOET004294!, WAG0004454!, K001070122!, STU-PH-

2012-002616!).

Description:—Stems 5–15 cm tall, procumbent or ascending, simple or sparingly branched, sparsely villous, hairs patent, 0.4–0.7 mm long; leaves subsessile, ovate to lanceolate, acute, 2.4–5.6 × 1.2–2.4 mm, densely villous, with patent hairs, 0.3–0.7 mm long on both sides; verticillasters of 2 pedunculate fascicles, each 1–3-flowered; calyx 4.5–5 mm long, with upper teeth 1.2–1.5 mm long, linear-triangular, lower teeth 1.6–2.2 mm long, pubescent, hairs patent, 0.4–0.8 mm long; corolla 5–6.5 mm long, with tube long exserted from the calyx; stamens slightly exserted from the tube.

Habitat and ecology: —Calcareous rupestrian stands, as cliffs, rocks, walls, ravines and boulders, as well as in xeric Mediterranean phrygana and grasslands.

Iconography: — Fig. 5e in Tan et al. (2010).

Chromosome number: —unknown.

Distribution: — Crete Island, central-eastern part ( Fig. 2 ).

Notes: —It was described by Bentham (1848) and transfered by Greuter & Burdet (1985) to the genus Satureja with a new name S. candica . This species is usually accepted in the current literature ( Chater & Guinea 1972, Doroszenko 1986, Turland et al. 1993, Jahn & Schönfelder 1995, Bräuchler et al. 2008a, Tan et al. 2010, Dimopoulos et al. 2013, Strid 2016).

Specimina visa:— CRETE. Selinaris. Nomos Lasithiou, dans le gorge traversèe par la route Agios Nikolaos , sur une pente rocheuse, a proximite du monastere d’Agios Georgios, April 1997, G. v. Buggenhout 18344 ( BR0000026283869 V!, BR0000027466636 V!, B100086801 !, P03281976 !, H1725504 !) ; Insel Kreta , Eparchia Ierapetras , Afendis Kavousi, Gipfel, 1 July 1994, R. Jahn s.n. ( B100086802 !) ; Selinaris , Kreta , russen Heraklion (Malia en Vrahassion) Agios Nikolaos, 3 May 1982, G. v. Buggenhout s.n. ( BR000002669245 V!, P03281975 !) ; Agios Georgios Selinaris (Kriti), April 1976, G. v. Buggenhout s.n. ( BR0000022434845 !) ; Agios Georgios Selinaris , gelegen op de weg Iraklion-Agios Nikolaos (Kreta, nomos Lassithiou), tussen rotsblocken bij het kloos, 18 April 1976, G. v. Buggenhout s.n. ( BR0000026189918 V!) ; Distr. Kaenurio , Montes Psiloriti (Ida), in saxosis calc. supra Vorisia, ca. 1300 m, 5 July 1942, K. H. Rechinger 14338 ( US 02809551!) ; In saxosis infra Korfes et Kroussona, distr. Malevisi, 4 June 1899, A. Baldacci 90 ( P03599402 !, P03599403 !, P03599404 !) ; Creta : eparch. Mirabello , int. Malea et Neochori, 21 April 1846, T. de Heldreich s.n. ( LY0017464 !) ; in rupestribus provincae Mirabello Cretae rariss, April 1846, T. de Heldreich s.n. ( K000193592 !) ; In rupestribus prov. Mirabello, Creta, 21 April 1846, T. de Heldreich s.n. ( TCD0018162 !) ; in rupestribus submontosis de Maglia à Kainourio Korio (Eparchia de Mirabello), 21 April 1846, T. de Heldreich 1422 (G-BOIS00769782!) ; In rupestribus pr. Kainurio-Chorio (Eparchia Mirabello) Cretae, 21 April 1846, T. de Heldreich 1422 ( WU0074660 !) .

5) Micromeria microphylla (d’Urville) Bentham (1834: 377) , nom. cons. prop. Fig. 5

≡ Thymus microphyllus d’Urville (1822: 327) , nom. cons. prop. ( Del Guacchio et al. 2022).

≡ Satureja microphylla Gussone (1828: 120) .

≡ Satureja microphylla Gussone (1844: 91) nom. illeg.

≡ Clinopodium microphyllum (d’Urville) Kuntze (1891: 515) .

Type (lectotype designated by Bräuchler in Bräuchler et al. 2008a):— MALTA. Thymus melitensis , Malte, d’Urville (lectotype P 04231225!), typ. cons. prop. ( Del Guacchio et al. 2022).

= Thymus melitensis d’Urville ex Gussone (1844: 91) , pro syn. nom. illeg.

= Micromeria melitensis Tineo in Chodat (1924: 245) pro syn. nom. illeg.

= Micromeria microphylla var. glabriuscula Bentham (1834: 377) (type not located).

= Micromeria microphylla var. gussonei Lojacono in Chodat (1924: 15) pro syn. nom. illeg.

= Micromeria microphylla var. villosa Bentham (1834: 377) (type not located).

= Thymus micranthus Gussone (1826: 245) non Brotero (1804: 176) (type not located).

= Piperella filiformis C. Presl (1826 : XXXVII) non Thymus filiformis Aiton (1789: 313) (type not located).

Description:—Stems 5–15 cm tall, procumbent or ascending, simply branched, hairy, hairs patent or reflexed, 0.05– 0.4 mm long; leaves subsessile, basal ones ovate to cordate, acute, 3–5 × 2.5–4 mm, hairy, with appressed hairs, 0.3–0.8 mm long on both sides, upper ones lanceolate to linear-lanceolate, acute, 3–6 × 0.9–1.6 mm, hairy, with hairs 0.1–0.4 mm long; verticillasters of 2 pedunculate fascicles, each 1–3-flowered; calyx 2.5–3.2 mm long, with upper teeth 0.8–1.1 mm long, linear-triangular, lower teeth 1–1.5 mm long, triangular, hairy, hairs patent, 0.3–0.8 mm long; corolla 4.5–6 mm long, with tube long exserted from the calyx, with throat glabrous; stamens exserted from the tube, filaments of upper stamens 0.8–1.5 mm long; style 2.3–4.5 mm long; mericarps 0.6–0.8 × 0.3–0.4 mm.

Habitat and ecology: —It occurs on rocky walls and rocky outcrops of calcareous and volcanic substrata.

Iconography: —Fig. 3147 in Fiori (1933), Fig. 5b in Tan et al. (2010).

Chromosome number: —unknown.

Distribution: — Malta Archipelago ( Malta, Gozo and Comino), Sicily and neighbouring islands (Favignana, Marettimo, Lampedusa, Linosa and Lipari), Italy (Apulia) in Lazio is adventitious ( Lucchese 1988, Pignatti 2018, Fig. 2 ).

Notes: —This species, described by d’Urville (1822) from Malta, was erroneusly lectotypified by Bräuchler in Bräuchler et al. (2008a) and successively correctly retypified by Ferrer-Gallego et al. (2020). In order to mantain the name proposed by d’Urville (1822) applied to the Maltese plant, Del Guacchio et al. (2022) propose Thymus microphyllus as nomen conservandum and the lectotype proposed by Bräuchler in Bräuchler et al. (2008a) as type conservandum. Currently, this species is considered endemic to Malta, Sicily and Italy by Greuter et al. (1986), Casha (2015) and Pignatti (2018).

Specimina visa:— MALTA. Comino, 15 April, 1987, S. Brullo , P. Pavone & G. Ronsisvalle s.n. ( CAT060.885 !) ; Malta, Naxxar, 16 April, 1987, S. Brullo , P. Pavone & G. Ronsisvalle s.n. ( CAT060.875 ) ; Malta, Wadi Babu , 24 September 1985, S. Brullo s.n. ( CAT060.884 !) ; Malta, Saint Julien, 11April 1987, S. Brullo , P. Pavone & G. Ronsisvalle s.n. ( CAT060.886 !) ; Malta, Misraim, Ghonoa , 28 June 1973, S. Brullo & G. Ronsisvalle s.n. ( CAT060.877 !) ; Malta, Mistra Bay , 26 June 1973, S. Brullo & G. Ronsisvalle s.n. ( CAT060.876 !) ; Malta, Salina Bay , 26 November 1985, S. Brullo s.n. ( CAT060.883 !) ; Malta, Ghemieri , 12 April 1984, S. Brullo & G. Ronsisvalle s.n. ( CAT060.881 !) ; Malta, Santi Valley ( Birgemma ), 12 April 1984, S. Brullo & G. Ronsisvalle s.n. ( CAT060.879 !) ; Malta, Dingli, 12 April 1984, S. Brullo & G. Ronsisvalle s.n. ( CAT060.880 !) ; Malta, Melleha , soil overlying calcareous rock, 22 October 1974, L. Y. Th. Westra & J. v. Rooden 166 ( U1402465 !) ; Insula Gozo, Ta’Lunzjata, small valley between Kercem and Victoria , 60 m, 15 April 1969, K. U. Kramer & L. Y. Th. Westra 4405 ( U1402467 !) ; Insula Malta steep S. escarpment of Mellieha Ridge , S. of Melleha, 125 m, 30 Aprile 1968, K. U. Kramer & L. Y. Th. Westra 4263 ( U1402466 !) ; Malta, la Valetta, rochers, 14 April 1938, R. Maire & M. Weiller 1193 ( MPU369177 !) ; Flora Melitensis , November 1882, Ruhmer 95 ( PAL37846 !) ; Insula Malta, in collibus saxosis, rara, 4 April 1896, M. F. Spencer s.n. ( B!). SICILY. Monserrato ( Modica ), 15 March 1971, S. Brullo s.n. ( CAT060.868 !) ; Monti Climiti , 9 June 1988, G. Fichera & F. Scelsi s.n. ( CAT060.867 !) ; Fiume Tellaro , 7 March 1969, S. Brullo s.n. ( CAT060.869 !) ; Modicano rupi, 26 April 1971, S. Brullo s.n. ( CAT060.870 !) ; Modica , 23 April 1983, S. Brullo s.n. ( CAT060.871 !) ; Cava d’Ispica , April 1987, F. Scelsi s.n. ( CAT060.873 !) ; Cava d’Ispica, 10 May 1987, S. Brullo , P. Minissale & F. Scelsi s.n. ( CAT060.872 !) ; Cava d’Ispica , 28 May 1988, S. Brullo s.n. ( CAT060.874 !) ; Fontane Bianche ( SR), 5 March 1989, G. Fichera s.n. ( CAT060.863 !) ; M. Climiti, Siracusa , 17 April 1979, S. Brullo & F. Fagotto s.n. ( CAT060.862 !) ; Cassibile-Cava Grande , June 1975, S. Brullo s.n. ( CAT060.861 !) ; Lipari , M. S. Angelo, 26 Aprile 1982, S. Brullo s.n. ( CAT060.860 !) ; Lampedusa, Capo Grecale, 21 March 1986, S. Brullo , P. Minissale, P. Pavone & G. Spampinato s.n. ( CAT060.859 !) ; Lampedusa, 17 March 1985, S. Brullo , P. Minissale & G. Spampinato s.n. ( CAT060.858 !) ; Lampedusa , 23 April 1977, S. Brullo s.n. ( CAT060.857 !) ; Linosa , April 1977, S. Brullo s.n. ( CAT060.856 !) ; Insel Lampedusa, Albero sole, 18 April 1959, J. Kohlmeyer 648 ( B!) ; Monte Amara , 13Aprile 1980, M. Grillo s.n. ( CAT060.855 !) ; Campolato , 4 Aprile 1980, M. Grillo s.n. ( CAT060.864 !) ; Siracusa , locis aridis rupestribus collim, April 1898, G. Rigo 124 ( B!, BP-TRA-00573278!)) ; Siracusa , tussen stenen van Grieks amphitheater, 7 April 1959, J. Th. Koster 6700 ( L2791360 !) ; Siracusa , 21 July 1897, A. Palyi s.n. ( BP-TRA 00573285 !) ; Favignana , in locis saxosis aridis, April 1890, H. Ross 63 ( B!, O-V2248247!, L2791363 !, BP-TRA-00573279!, BP-TRA-00573280!) ; In aridis insulae Favignana , 3 May 1855, E. & A. Huet du Pavillon (O-V2248248!, L2791361 !, WAG1711191 !, VTA058866 !) ; Terranova, in collibus sterilibus, April , A. Todaro 853 ( U1402468 !, VTA058867 !, LY0491623 !, PAL17198 !) ; Insula Favignana , April 1888, H. Ross s.n. ( AMD81847 !) ; Comiso , V. Tineo s.n. ( LY0740067 !) ; In Sicilia meridionalis, May , M. Lojacono s.n. ( PAL17326 !) ; Castelvetrano , 1890, G. Reina s.n. ( PAL17199 !) . ITALY. Gallipoli , in collibus aridis calcareis, May 1917, M. Guadagno s.n. ( PI039567 !) ; Ostia Antica, sui muri aridi delle rovine e nei pavimenti marmorei, 20 June 1988, F. Lucchese s.n. ( FI, RO) .

6) Micromeria rodriguezii Freyn & Janka (1874: 16) , Fig. 6

≡ Micromeria nervosa Bentham subsp. rodriguezii (Freyn & Janka) Nyman (1881: 590) .

≡ Satureja rodriguezii (Freyn & Janka) Pau (1900: 68) .

≡ Micromeria biflora var. rodriguezii (Freyn & Janka) Knoche (1922: 346) .

≡ Satureja cordata subsp. rodriguezii (Freyn & Janka) O. Bolòs & Vigo (1983: 94) .

≡ Micromeria filiformis subsp. rodriguezii (Freyn & Janka) Bonafè (1980: 56) .

≡ Satureja microphylla subsp. rodriguezii (Freyn & Janka) O. Bolòs & Vigo (1996: 301) .

≡ Satureja rodriguezii (Freyn & Janka) Rivas-Mart., Costa & Loidi (1992: 217) .

≡ Micromeria microphylla subsp. rodriguezii (Freyn & Janka) Romo (1994: 261) .

Type (lectotype designated by Bräuchler in Bräuchler et al. 2008a):— SPAIN. Minorca, in valle barranco del Favaret prope Mahon, 29 March 1873, C. F. Hegelmaier s.n. (lectotype STU-PH-2011-072456!).

= Micromeria rodriguezii f. major Sennen (1917: 192) Type: (lectotype designated by Rosselló & Sáez 2000):— SPAIN. Balearic Islands, Mallorca. Baléares: Route de Valdemora, à 6 km de Palma, 13 July 1914, Bianor (lectotype BC-Sennen 836596!).

= Micromeria knochei Sennen & Bianor in Sennen (1917: 193) Type: (lectotype designated by Rosselló & Sáez 2000):— SPAIN. Balearic Islands, Mallorca. Collines près de Palma, May 1913, F. Bianor Pl. Espagne-F. Sennen n.º 2039 (lectotype BC-Sennen 836597!).

= Satureja microphylla var. condensata L. Chodat (1924: 245) Type: (lectotype designated by Rosselló & Sáez 2000):— SPAIN. Balearic Islands, Mallorca, Torre den Pau, March–April 1921 (lectotype G!).

= Satureja microphylla subsp. majoricensis L. Chodat (1924: 247) (type not located).

= Micromeria rodriguezii var. acuminifolia Sennen (1933: 62) Type: (lectotype designated by Rosselló & Sáez (2000):— SPAIN. Balearic Islands, Mallorca Fossé—Route de palma à Valldemossa. Soller (Mallorca Baléares), 26 December 1911, Bianor (lectotype BCSennen 836599!).

≡ Satureja rodriguezii var. acuminifolia (Sennen) Sennen (1933: 62) .

≡ Micromeria rodriguezii var. major (Sennen) Sennen (1933: 62) .

≡ Satureja knochei (Sennen & Bianor) Sennen (1933: 63) .

Description:—Stems 5–25(–36) cm long, procumbent to suberect (sometimes erect), simple or branched, densely hairy to tomentose, with retrorse hairs, 0.1–0.3(–0.4) mm long; leaves subsessile, ovate to ovate-lanceolate, sparsely hairy, with appressed hairs, 0.05–0.4 mm long on both surfaces, acute to subacute at the apex, (1.2–)2–4(–6.5) × (1–)1.5–3.5(–4.5) mm; verticillasters of 2 pedunculate fascicles, each 2–5-flowered; calyx 2.5–4(–4.5) mm long, tube with antrorse hairs, 0.05–0.35(–0.5) mm long, with subequal teeth, linear-lanceolate to subulate, 0.8–1.7 mm long, hairs antrorse, 0.2–0.5(–0.6) mm long; corolla 5–6.5 mm long, with tube slightly exserted from the calyx; stamens included in tube; mericarps 0.8–1.2 × 0.3–0.6 mm.

Habitat and ecology: —Open scrub, rocky places, stone-walls and roadsides on calcareous, sandy and marl substrata.

Iconography: —Tab. 129A in Willkomm (1889: 68); Fig. 22 in Morales Valverde (1991).

Chromosome number: —2n = 50 ( Morales 1994).

Distribution: —Endemic to the Balearic Islands: Mallorca, Menorca, Eivissa and for Formentera ( Gil & Llorens 2001); recently reported by Senar et al. (2020) from a single location in Eastern Spain, Castellón province ( Fig. 2 ).

Notes: —This species was described by Freyn & Janka (1874), but often treated as synonym of M. filiformis Bentham (1834: 378) by Chater & Guinea (1972), or as subspecies of the latter ( Bonafè 1980; Bräuchler et al. 2008), while other authors (Mus Amezquita & Rosselló 1987, Morales Valverde 1991, Morales 2010) consider it as synonym of M. microphylla . It is recognized as a distinct species by Willkomm (1889), Chodat (1924) and Senar et al. (2020), or as subspecies of M. nervosa ( Nyman 1881) , of M. cordata ( Bolòs & Vigo 1983) and of M. microphylla ( Bolòs & Vigo 1996) .

Specimina visa:— SPAIN. Baleares: Ins. Minorca : barranco del Favaret prope Mahon, 29 March 1873, I. et C. F. Hegelmaier s.n. ( JE00007503 !) ; Ins. Minorca in declivibus apricis vallis Son Blanc, 3 April 1873, F. Hegelmaier s.n. (STU-PH-2011-069575!) ; Ins. Minorca , barranco de Algender ad rupes calcareas, 1 April 1873, F. Hegelmaier s.n. (STU-PH-2011-075532!) ; Ins. Majorca , ad castellum Bellver prope Palma, 12 April 1873, F. Hegelmaier s.n. (STU-PH-2011-073100!) ; Ins. Majorca , ad muriracata urbis Palma, 11 April 1873, F. Hegelmaier s.n. (STU-PH-2011- 073760!) ; Mallorca, Fossé—Route de Palma à Valldemossa. Soller, 26 Dec 1911, Bianor (BC-Sennen 836599!) ; Mallorca, Cala Mandrago , 18 April 2001, R. Böcker SP-M 333 ( B!) ; Mallorca, Collines près de Palma, May 1913, F. Bianor, Pl. Espagne-F. Sennen n.º 2039 (BC-Sennen 836597!) ; Mallorca, Chemins et collines autour de Palma, 11 July 1914, Bianor, Pl. Espagne-F. Sennen n.º 2037 ( BC 49867 !) ; Mallorca, Route de Valldemora [Valldemossa], à 6 km de Palma, 13 July 1914, F. Bianor, Pl. Espagne-F. Sennen n.º 2036 ( BC 49864 !, MA104550 !) ; Mallorca, Artà , 24 May 1915, L. Garcias Font ( BC 600942 !) ; Palma de Mallorca a Génova, 20 March 1918, P. Font-Quer ( BC 49939 !) ; 12 July 1932, Cala Major, Palma , 12 May 1932, sine leg. ( BC 49863 !) ; Palma, à Génova , 12 Apr 1933, F. Sennen, Pl. Espagne-F. Sennen n.º 8622 ( BC 82427 !) ; Artà , 15 Apr 1936, Kennedy ( BC 103461 ) ; Mallorca, S’Aigo de Palma , 4 Sept 1947, P. Palau Ferrer ( BC 601110 !) ; Mallorca, Torrent Gros, Pont d’Inca , 2 May 1949, P. Palau-Ferrer 166 ( BC 112645 !), Mallorca, entre Felanitx i Porto Colom, 2 May 1971, F. Masclans ( BC 607564 !) ; Mallorca, castell de Santueri, Felanitx , 370 m, rocky places, 24 Apr 2000, L. Sáez LS-5346 ( L. Sáez, pers. herb.) ; Menorca, Caminos cerca de Mahón, Apr 1870, J. Rodriguez-Femenias ( BC 654515 !) ; Alayor , 16 March 1874, J. Rodriguez-Femenias ( BC 654512 !) ; Menorca, Binisarmenya , 13 June 1876 J. Rodriguez Femenias ( BC 49871 !) ; Menorca, Fonduco, Mahón , grietas peñascos, 19 Jan 1913, P. Font-Quer ( BC 49861 !) ; Cala San Esteban , 19 May 1913, P. Font-Quer ( BC 49870 !) ; Santa Ponsa de Alayor , 30 May 1913, P. Font Quer ( BC 49872 !) ; Cala Figuera, Mahón , 8 July 1913, P. Font-Quer ( BC 49868 !) ; Menorca, San Clemente , 24 Dec 1913, P. Font-Quer 917 ( BC 49860 !) ; Menorca, Mahón, camino de Trepucó, 6 March 1914, P. Font-Quer ( BC 49866 ). Eivissa , Cala Jondal , 16 May 1918, Gros ( BC 49937 !). SPAIN. Castellon : Santa Magdalena de Polpis , cami del Saltet , pr. Mas de Xarpa , 120 m, depressió de desugàs de l’aigua de l’AP-7, pastures d’anuals, sobre gleres calcaires, 26 August 2018, R. Senar s.n. ( VAL241026 !) ; ibidem, cami de la autopista (margen derecha en sentido Tarragona), 124 m, 6 May 2019, B. Plana, P. P. Ferrer-Gallego & E. Laguna s.n. ( VAL2422218 !) .

7) Micromeria sphaciotica Boissier & Heldreich ex Bentham (1848: 220)

≡ Satureja sphaciotica (Bentham) Greuter & Burdet (1985: 306) .

Type (lectotype here designated):— GREECE. Creta in rupibus faucis Aratana, Sphakia, July 1846, T. de Heldreich s.n. (lectotype G- DC00478886!, isolectotypes G 00418066!, G-BOIS00418066!, G-BOIS00330680!, GOET 004296!). The holotype designated by Bräuchler in Bräuchler et al. (2008a) has to be rejected, because he cited the sentence reported in the protologue and not that of the herbarium label.

Description:—Stems (5–) 10–20 cm tall, procumbent or ascending, simple or sparingly branched, villous, hairs patent, 0.1–0.5(–0.8) mm long; leaves subsessile, ovate to lanceolate, acute, 3.5–8 × 0.9–2.5 mm, hairy, with patent hairs, 0.1– 0.7 mm long on both sides; verticillasters of 2 pedunculate fascicles, each with 1–5-flowered; calyx 2.5–3(–3.5) mm long, with upper teeth 0.6–0.7 mm long, linear-triangular, lower teeth 0.9–1 mm long, pubescent, hairs patent, 0.2–0.5 mm long; corolla 5–6 mm long, with tube long exserted from the calyx; stamens slightly exserted from the tube.

Habitat and ecology: —Cliffs and rocky walls on calcareous substrata.

Iconography: — Fig. 5c in Tan et al. (2010).

Chromosome number: —unknown.

Distribution: — Crete, gorges of Aradena near Sfakia ( Fig. 2 ).

Notes: —This species described by Bentham (1848) is treated as a doubtful taxon by Chater & Guinea (1972), or as synomym of M. microphylla by Doroszenko (1986) and Bräuchler et al. (2008a). It is accepted as valid species by Greuter & Burdet (1985), Turland et al. (1993), Jahn & Schönfelder (1995), Tan et al. (2010), Dimopoulos et al. (2013) and Strid (2016).

Specimina visa:— CRETE. Ile de Crete , Region des plateaux (400m – 600m), roches d’Aradena Sphakia, 1845, V. Raulin 280 ( P03599405 !), sub. M. hispida ; Gole di Aradena , 24 April 2022, S. Cambria s.n. ( CAT!) ; Calamintha minima , annua, thymi-folio, Inst. rej. herb. 194. herbier de Vaillant (P-Vaillant 04049884!); Calamintha minima annua Thymifolio, herb. Tournefort no. 1312 B (P-Tournefort 00652340!) .

Figures

FIGURE 2. Distribution map of species belonging to Micromeria microphylla group based on herbarium materials: M. silicii (red dots), M. microphylla (black dots), M.rodriguezii (green dots), M. acropolitana (blue dot), M. sphaciotica (yellow dot), M. hispida (orange dots), M. carpatha (pink dot) and M. cypria (purple dots).

FIGURE 4. Micromeria cypria. A. Habit. B. Branch with inflorescences. C. Cyme. D. Flowers. E. Open calyx ventral view. F. Open calyx dorsal view. G. Open corolla ventral view. H. Cyme peduncle. I. Leaves abaxial face. J. Leaves adaxial face. K. Stem indumentum. Illustration by S. Brullo based on material coming from Cyprus (type locality).

FIGURE 5. Micromeria microphylla. A. Habit. B. Branch with inflorescences. C. Stem indumentum. D. Cymes. E. Cyme peduncle. F. Open calyx ventral view. G. Open calyx dorsal view. H. Open corolla ventral view. I. Stlyles and stigmas. J. Leaves abaxial face. K. Leaves adaxial face. L. Mericarps. Illustration by S. Brullo based on living material coming from Malta (type locality).

FIGURE 6. Micromeria rodriguezii. A. Habit. B. Branch with inflorescences. C. Cymes. D. Stem indumentum. E. Calyx. F. Leaves abaxial and abaxial face. Illustration by L. Sáez based on material coming from Balearic Islands, Mallorca, Palma, Génova (BC 49863).