A morphological re-evaluation of the taxonomic status of Satureja L. (Lamiaceae, Nepetoideae, Mentheae) from flora of Iran Author Bordbar, Firouzeh Author Mirtadzadini, Mansour Department of Biology, Faculty of Science, Shahid Bahonar University of Kerman, Kerman (Iran) bordbar @ uk. ac. ir, fbordbar 2000 @ yahoo. com (corresponding author) mirtadz @ uk. ac. ir bordbar@uk.ac.ir text Adansonia 2024 3 2024-04-23 46 7 45 63 https://sciencepress.mnhn.fr/sites/default/files/articles/pdf/adansonia2024v46a7.pdf journal article 10.5252/adansonia2024v46a7 1639-4798 11058077 Genus Satureja L. Species Plantarum 567 ( Linnaeus 1753 ). — Satureja sect. Tragoriganum Benth., Labiatarum genera et species : 354 ( Bentham 1834 ) . — Satureja sect. Annuae Boiss. , Flora Orientalis 4: 562 ( Boissier 1879 ) . — Satureja sect. Eusatureiae Boiss. , Flora Orientalis 4: 563 ( Boissier 1879 ) . — Satureja sect. Subbilabiatae Boiss. , Flora Orientalis 4: 565 ( Boissier 1879 ) . — Satureja sect. Zatarioideae Boiss. , Flora Orientalis 4: 567 ( Boissier 1879 ) . — Satureja sect. Sabbatia Briq. in Engler & Prantl, Die natürlichen Pflanzenfamilien , 4, 3a: 298 ( Briquet 1895 -1897). TYPE SPECIES . — Satureja montana L. (designated by Doroszenko 1986 ). PERSIAN VERNACULAR NAMES . — This plant is known by local people as ( S. khuzistanica in Khuzestan ), Azbua ( S. longiflora Boiss. & Hausskn. in Kermanshah ), Ezgen and Ezghand ( S. kermanica in Kerman ), Oushan and Ourishan ( S. bachtiarica Bunge in Bakhtiari), Maraza ( S. bachtiarica in Kermanshah ), Hazbeh ( S. bachtiarica in Kordestan ) and the common name Marzeh for S. hortensis . DISTRIBUTION . — N, NW, W, SW and SE of Iran . ECOLOGY AND PHYTOGEOGRAPHY . — The distribution pattern of most of Satureja species in Iran follows that of the Irano-Turanian floristic region. Exceptionally, S. isophylla existed in transitional area between the Irano-Turanian and Euro-Siberian regions, and S. khuzistanica distributed on the rocky slopes of mesic areas between the Irano-Turanian and Saharo-Arabian regions. S. mutica is distributed in both the Irano-Turanian and Euro-Siberian regions (Floristic regions followed Zohary’s classification in 1973). PHYLOGENETIC RELATIONSHIPS . — Satureja is more closely related to the monotypic genus Gontscharovia ( Doroszenko 1986 ; Bräuchler et al. 2010 ). The two genera can be distinguished by the acute nutlet apex in Gontscharovia opposed to an obtuse (rounded) apex in Satureja s.s. ( Bräuchler et al. 2010 ). DESCRIPTION Suffruticose, rarely caespitose or fruticose, aromatic or non-aromatic, perennial or annual, sometimes with stout woody stock. stems erect, erect-procumbent or out-spreading, many-branched from base to above, covered with short retrorse or spreading, rarely antrorse hairs and sessile glands on young parts, rarely glabrous. Leaves entire, sessile, attenuate at base rarely with a short petiole, cauline leaves oblanceolate, narrow elliptic, linear, obovate, rarely obovate-spathulate and lanceolate, recurved or out-spreading, conduplicate, rarely flattened, covered with short and/or long hairs and subsessile or rarely capitate glands on both surfaces, floral leaves similar in shape to the cauline leaves and shorter. Verticillasters distant, approximate or in lax cymes, often with up to six flowers, rarely up to 20, flowers mostly sessile, or pedunculate. Calyx green to dark purple, straight, bilabiate, with three superior teeth clearly shorter than the inferior two, tubular, campanulate, or tubular-campanulate, covered with short hairs and sessile glands on the outside, throat usually with a sparse fringe of hairs, calyx teeth unequal, wide to narrowly triangular, recurved or rarely straight. Corolla two-lipped, white or violet, rarely with yellow lines, violet spots or smears, 4-20 mm long, covered with hirsute or puberulent hairs, tube straight, not annular inside, with few sparse hairs, throat ciliate, upper lip erect, shortly emarginated, lower lip spreading with three subequal lobes. Stamens 4, exserted from the corolla tube, rarely included, the two anterior longer, thecae divergent. Style shortly or longly exserted from the tube, two-branched, branches equal. Nutlets mostly elliptic, rarely obovate or widely obovate, rounded at the apex, ± smooth or sculptured, usually nervate, light to dark brown, 1-2 mm long, 0.5-1.5 mm wide. MORPHOLOGICAL CHARACTERS ( Figs 1 ; 2 ) Habit and stem Among the taxa studied, S. hortensis is annual, whereas the majority of species are herbaceous perennials. Perennial herbs sometimes have woody stems at the base or woody stocks and sometimes are recorded as low shrubs or subshrubs. Stems are erect (in S. hortensis , S. khuzistanica , Fig. 2B ), erect-procumbent (in S. bachtiarica , S. kermanica , S. intermedia C.A.Mey. , S. longiflora , S. mutica Fisch. & Mey. , S. sahendica Bornm. , Fig.1C, I ) and erect to out-spreading (in S. avromanica , S. macrantha C.A.Mey. , S. macrosiphonia , Fig. 1A ). Exceptionally, S. isophylla exhibits a caespitose habit with short internodes ( Fig. 1L ). The height of stem is variable throughout the range of distribution of S. bachtiarica ( Fig. 1C, E ). Leaves Most leaves are entire, usually sessile and attenuate at base. They are rarely with a short petiole up to 2 mm in S. khuzistanica and S. intermedia . Leaf shape is variable, from linear to lanceolate or oblanceolate, rarely obovate-oblong or obovate-spathulate ( S. intermedia , Fig. 1K ), and obovate, ovate or wide elliptic ( S. khuzistanica , Fig. 2B ). The lower leaves in S. longiflora are variable from elongated elliptic or obovate to oblong-oblanceolate ( Fig. 1 H-J). It seems that the leaves of Satureja species in more humidity are larger and wider. The middle and upper leaves are often conduplicate. Inflorescence Inflorescence is formed of terminal verticillaters composed of often 2-6 flowers with reduced or very short peduncles and pedicels, rarely with up to 20 flowers in S. hortensis or in lax cymes ( S. avromanica and S. macrosiphonia , Figs 1B ; 2G ). Verticillaters are distant or approximate. Sometimes both states are observed in one species (e.g. S. longiflora , S. macrantha , Fig. 1 G-H). Bract and bracteole Floral leaves in shape are similar to the cauline leaves but shorter. Calyx Calyx is straight and bilabiate with three superior teeth shorter than the inferior two. It is usually green, but sometimes dark purple in fruiting (e.g. S. avromanica , S. bachtiarica , S. macrantha ). Typically, the calyx is tubular-campanulate however S. avromanica , S. longiflora and S. macrantha are distinct by tubular ( Figs 1B , G-H; 2E, F) and S. bachtiarica by campanulate calyces ( Fig. 1D, F ). Calyx teeth vary in shape from wide triangular (e. g. S. bachtiarica ) to narrow linear (e.g. S. mutica and S. hortensis ). Moreover, several Satureja spp. have slightly recurved calyx teeth, while the erect state is observed in S. longiflora ( Fig. 1H ). Calyx length is ranged from 2.5 mm in S. bachtiarica up to 9 mm in S. khuzistanica . TABLE 1. — Overview of Satureja L. species belonging to flora of Iran in literature.

Flora Orientalis , Boissier (1879)	Flore de l’Iran , Parsa (1949)	Flora Iranica , Rechinger (1982)	Flora of Iran , Jamzad (2012)	Present study
S. atropatana Bunge	S. atropatana	S. atropatana	S. atropatana	S. avromanica Maroofi
S. bachtiarica Bunge	S. bachtiarica	S. bachtiarica	S. bachtiarica	S. bachtiarica Bunge
S. hortensis L.	S. boissieri Hausskn. ex Boiss.	S. boissieri	S. boissieri	S. edmondii (syn. of S. longiflora )
S. intermedia C.A.Mey. (syn. of S. subdentata Boiss.)		S. edmondii Briq. (syn. of S. longiflora )	S. edmondii (syn. of S. longiflora )
	S. hortensis		S. intermedia (syn. of S. subdentata )
	S. longiflora	S. intermedia (syn. of S. subdentata )	S. isophylla	S. intermedia (syn. of S. subdentata )
S. longiflora Boiss. & Hausskn.	S. macrantha		S. kallarica Jamzad
	S. macrosiphonia Bornm.	S. isophylla Rech.f.	S. kermanshahensis Jamzad
S. macrantha C.A.Mey.	S. mutica	S. laxiflora K.Koch	S. khuzistanica Jamzad	S. isophylla
S. mutica Fisch. & C.A.Mey.	S. sahendica Bornm.	S. macrantha	S. laxiflora	S. kermanica
S. spicigera (C.Koch) Boiss.	S. mutica	S. macrantha	S. macrantha
	S. sahendica	S. macrosiphonia	S. macrosiphonia
S. subdentata Boiss.	S. spicigera	S. mutica	S. mutica
S. rechingeri Jamzad	S. sahendica
S. spicigera
S. sahendica

Corolla The color ranges from white or violet, rarely lilac, yellowish, pink, and cream. The colors can vary within the same species in natural habitats (e.g. S. hortensis , S. khuzistanica ( Fig. 2C, D ), and S. bachtiarica , S. mutica ). The corolla is two-lipped (two lobes forming the upper lip, three lobes forming the lower lip) with the posterior lip straight and emarginated and the anterior lip spreading. The tube is straight, covered with hirsute or puberulent hairs, not annular inside, with few sparse hairs, and throat ciliate. Corolla in S. avromanica is conspicuous with the slender tube ( Fig. 1B ). The tube of the corolla is longer than the calyx in S. avromanica , S. longiflora , S. khuzistanica , S. macrantha , S. macrosiphonia and S. sahendica , while it is nearly of the same length as the calyx in S. bachtiarica , S. hortensis , S. intermedia , S. isophylla S. kermanica and S. mutica . The stamens, style and stigma have no taxonomic value for the Satureja as they have a very constant morphology. The four stamens are didynamous, curved, included in or exserted from corolla, and thecae divergent. The style is bifid with subequal branches. Nutlet The shape of nutlets are mostly elliptic, rarely obovate ( S. isophylla ) and widely obovate ( S. khuzistanica ). The nutlets are rounded at the apex, usually nervate, and light to dark brown in color. USES AND CHEMICAL PROPERTIES The species are widely used as flavor in foods, herbal tea, and traditional medicine to treat various ailments, such as cramps, muscle pains, nausea, indigestion, diarrhea, and infectious diseases. Chemical analyses have revealed that Satureja species are rich in terpenoids, such as carvacrol, γ- terpinene, thymol, p -cymene, β- caryophyllene, linalool, and known to have antimicrobial, antiprotozoal, antioxidant as well as anti-inflammatory and anti-nociceptive properties ( Amanlou et al. 2005 ; Sadeghi-Nejad et al. 2011 ; Alizadeh 2015 ; Mazandarani & Monfaredi 2017 ).