Labiatae) in Turkey

Biologia 65/2: 219—227, 2010 Section Botany DOI: 10.2478/s11756-010-0013-y

Morphology, anatomy, palynology and nutlet micromorphology of Salvia macrochlamys (Labiatae) in Turkey

Ahmet Kahraman,FerhatCelep &MusaDogan˘

Middle East Technical University, Department of Biological Sciences, 06531, Ankara/Turkey; e-mail: [email protected]

Abstract: Salvia macrochlamys Boiss. & Kotschy, which belongs to sect. Salvia Hedge, is a perennial herb usually growing on rocky limestone slopes in eastern Anatolia. In this study, the anatomical features of the root, the stem, the leaf and the petiole, as well as micromorphological characteristics of the pollen and the nutlet of this species have been investigated using Scanning Electron Microscopy (SEM). Moreover, its emended and expanded description, phenology, current distribution and ecology are also given. Anatomical characters such as the number of ray lines in the root, the distance between vascular bundles in the stem, the structure of mesophyll in the leaf and the structure of vascular bundles in the petiole have been found to have taxonomic value. The pollen grains are hexacolpate, radially symmetrical, isopolar and oblate-spheroidal. Their exine sculpturing is bireticulate and perforate. The nutlets are rounded-trigonous in transverse sections, broadly ovate to rotund in shape, light brown, glabrous and slightly rough to protuberances. This species often grows in habitats in which the main soil type is sandy-clayish with low organic content (1.32 ± 1.06%) and slightly alkaline (pH 7.75 ± 0.25). The amount of CaCO3 is 0.94% to 34.91%. The contents of P and K are 8.68 ± 0.96 and 110.12 ± 4.86 ppm, respectively. Key words: Salvia macrochlamys; Labiatae; anatomy; morphology; nutlet; palynology;

Introduction Benth., Horminum Benth., Drymosphace Benth. and Hemisphace Benth. Later on, sect. Eusphace was des- Salvia L., the largest genus of Labiatae, represents ignated as the type section of genus Salvia by Hedge an enormous and cosmopolitan assemblage of nearly (1972). 1000 species displaying a remarkable range of varia- Section Salvia Hedge (synonym: Eusphace Benth.) tion. It has undergone marked species radiations in is composed of shrubs and herbs with a relatively prim- three regions of the world: Central and South Amer- itive staminal structure. The characteristic features of ica (500 spp.), Central Asia/Mediterranean (250 spp.) this section are pinnatisect or simple leaves, herbaceous and Eastern Asia (90 spp.) (Walker et al. 2004). Salvia or suffruticose stems, calyx little enlarging after anthe- species are used in traditional medicines all around sis and lips not diverging, upper lip of corolla more the world, possessing antibacterial, antioxidant, antidi- or less straight and corolla tube annulate. Staminal abetic and antitumor properties (Ulubelen 2003). In ad- connectives are equal or slightly longer than filaments dition, many Salvia species are used as herbal tea and and lower theca fertile (Hedge 1972). The largest num- for food flavouring, as well as in cosmetics, perfumery ber of species in section Salvia grows in Turkey. Ac- and the pharmaceutical industry (Demirci et al. 2003). cording to Bentham’s (1833) sectional delimitation, S. The genus Salvia has been subject to a number of macrochlamys was placed in the section Salvia (= Eu- studies based mainly on morphological (Hedge 1982a, sphace) by Boissier (1879). 1982b), anatomical (Metcalfe & Chalk 1950, 1972; The first revision of Salvia in Turkey was made Kahraman et al. 2009), numerical (Reales et al. 2004), by Hedge (1982a), who recognized 86 species, one hy- karyological (Nakipo˘glu 1993), palynological (Hender- brid and one doubtful species. Between 1982 and 2005, son et al. 1968), phylogenetical (Walker et al. 2004) and four more species were published from Turkey (Huber- evolutionary data (Classen-Bockhoff et al. 2004). In ad- Morath. 1982; Vural & Adıgüzel 1996; Dönmez 2001; dition, the taxonomic importance of the nutlet shape Hamzao˘glu et al. 2005). Since 2005, as a part of a re- and surface micromorphology of Salvia have been stud- vision of the genus Salvia in Turkey, the authors have ied by Hedge (1970) and Marin et al. (1996). carried out extensive field studies and collected a large Boissier (1879) recognized 107 species of Salvia number of specimens. Furthermore, population size and in his Flora Orientalis, 75 of which were recorded phenological and ecological properties were noted in from Turkey. He placed these species under seven sec- the field. The studies have revealed two new species, tions using Bentham’s (1833) sectional delimitation. namely S. marashica A. Il¸˙ cim, F. Celep & Dogan (2009) These sections are as follows: Eusphace Benth., Hy- and S. ekimiana F. Celep & Dogan (2009) and two menosphace Benth., Aethiopis Benth., Plethiosphace new records, namely S. macrosiphon Boiss. (Kahra-

c 2010 Institute of Botany, Slovak Academy of Sciences 220 A. Kahraman et al.

Fig. 1. Distribution map of S. macrochlamys (). man et al. 2009) and S. viscosa Jacq. (Celep et al. prepared according to Wodehouse (1935) technique. For LM 2009). studies, measurements and observations were made using This study aims to give a detailed account of mor- the Leica DM1000 binocular light microscope with the Le- phological and anatomical characteristics and micro- ica DFC280 camera. The polar length (P), the equatorial morphological properties of pollen grains and nutlets of length (E), the colpus length (CLG), the exine and the intine thickness of 30 pollen grains were measured under the S. macrochlamys. The taxonomic significance of these light microscope (× 1000) and P/E ratios were calculated. findings is also discussed with the previous studies on For SEM, the pollen grains were observed and photographed the genus Salvia. with a JEOL-6060 scanning electron microscope to deter- After reading the paper written on anatomy and mine their exine ornamentation. Pollen terminology of Fae- morphology of S. napifolia by Baran & Ozdemir¨ (2006) gri & Iverson (1975) has been used. and examining relevant herbarium specimens, we de- The nutlets were first examined using a Leica S8AP0 cided that the plant was S. verbenaca.Itisaverycom- stereomicroscope to ensure that they were of normal size and mon species in Turkey and easily recognized by its small maturity. In order to determine the average seed sizes, 30 calyx and corolla, straight to subfalcate upper lip of mature nutlets were measured. For SEM, the mature nutlets were placed on stubs directly and covered with gold. After corolla and by its small number of flowers in its verti- that, they were observed and photographed with a JEOL cillasters. Therefore, this paper is based on a misidenti- JSM-6400 scanning electron microscope (Do˘gan 1988). fication. For this reason, the findings of the paper were For ecological studies, soil samples were taken from not compared to our results. suitable habitats of S. macrochlamys. All soil samples were analysed at the Soil, Fertilizer and Water Resources Cen- tral Research Institute, Ankara. Soil texture, pH, CaCO3, Material and methods P, K and organic matter analysis were made using standard techniques (Bayraklı 1987) and the results were evaluated Plant samples were collected from the following differ- according to Ka¸car (1972). ent localities in Turkey: B9 Bitlis: foot of Kambos Moun- tain, Direkta¸sı, 38◦18028 N41◦59015 E, 1351 m, 8 vii 2007, AKahraman 1426! Van: N.W. of Artos Mountain, Results 38◦18882 N42◦06737 E, 1662 m, 10 vii 2007, AKahra- man 1444! C9 Siirt: Eruh to S¸ırnak, Yanılmazlar village, ◦ ◦ Morphological characteristics 37 42 269 N42 15 804 E, 1195 m, 5 vi 2008, AKahraman S. macrochlamys is a perennial herb. Stems are 30– 1540! C10 Hakkari: junction of Yüksekova-Hakkari road, ◦ ◦ 50 cm, numerous, procumbent to almost erect and 37 42 768 N4403 357 E, 1625 m, 7 vi 2008, AKahra- man 1565! A distribution map of S. macrochlamys is given branched above. The stem indumentum is densely glan- (Fig. 1). The specimens have been stored in the Middle East dular pilose and sparsely eglandular villous. Leaves are Technical University (METU) Department of Biological Sci- 3.5–9 × 2–7.5 cm, simple, ovate to elliptic, cordate or ences and Ankara University Herbarium (ANK). rounded, and crenulate to serrulate. The leaf indumen- Anatomical studies were carried out on specimens kept tum is mostly glandular pilose with few sessile glands. in 70% alcohol. The paraffin method was used for the cross Petiole is 2–5 cm. Inflorescence is 10–35 cm, shortly sections of the leaves, stems, roots and petiole. The speci- paniculate or not. Verticillasters are 2-flowered, and mens were embedded in paraffin and then sectioned with approximating. Bracts are 25–45 × 15–25 mm, clearly a Leica RM2125RT rotary microtome. All sections were longer than calyces, almost enclosing flowers, ovate to stained with safranin-fast green and then mounted with Canada Balsam or Entellan (Johansen 1944). Measurements ovate-oblong, densely glandular pilose and sparsely eg- and photos were taken using a Leica DM1000 binocular light landular villous, membranous, prominently veined and microscope with a Leica DFC280 camera. greenish-white. Bracteoles were present. Pedicel is 1–3 For palynological investigations, pollen material was (–4) mm, glandular pilose, rigid and erecto-patent. Ca- obtained from herbarium samples. The pollen slides were lyx is tubular-ovate in shape, 14–20 mm, greenish-white Some morphological and anatomical characteristics of Salvia macrochlamys in Turkey 221

Fig. 2. General appearance of S. macrochlamys. and glandular pilose to villous. Its upper lip is triden- tate and acuminate. Calyx teeth are 2–5 mm. Fruiting calyx is thick-textured. Corolla is 23–35 mm. Its upper lip is white and its lower lip is pink. Corolla tube is 18–25 mm, straight below, widening above (not squa- mulate) and annulate. Its upper lip is almost straight and truncate. Stamens are A type, including the upper lip of corolla. Staminal connectives are shorter than ﬁla- ments. Filaments are 5–9 mm. Fertile and sterile anters are 3–5 mm and 1–2 mm, respectively. Upper thecae are 3–6 mm and lower thecae are 3–6 mm. Style is glabrous and 25–40 mm (Fig. 2).

Phenology, habitat and ecology S. macrochlamys ﬂowers in June to August. This species grows on rocky limestone slopes and on open slopes in Quercus sp. scrub between 900 and 2400 m. Other species in its habitat are S. trichoclada Ben- Figs 3–4. The transverse section of the root of S. macrochlamys. tham, S. verticillata L. ssp. verticillata and ssp. amasi- Pe – Periderm, C – Cortex, S – Sclerenchyma, Ph – Phloem, X aca (Freyn. & Bornm.) Bornm., S. poculata Náb., S. – Xylem, Pr – Pith ray, T – Trachea. indica L., Clypeola jonthlapsi L., Ptilostemon sp., Bro- mus sp., Echinops sp., Euphorbia sp., Onosma sp., Gal- ium sp., Convolvulus sp., Hordeum sp. and Achillea xylem occupies the pith, it is very narrow (Table 1). sp. According to the soil samples collected from the Stem suitable habitats of S. macrochlamys, this species usu- The transverse sections of the stems show that the epi- ally prefers sandy-clay soils which are slightly alkaline dermis is covered by a thin cuticle including unicellu- (pH 7.75 ± 0.25) with low organic content (1.32 ± lar or multicellular glandular and eglandular hairs. The 1.06%). CaCO3 in the soil is varying between 0.94% epidermis is single layered and consists of oval, squarish and 34.91%. The amounts of P and K present are 8.68 or rectangular cells (Fig. 5). The upper and lower walls ± 0.96 and 110.12 ± 4.86 ppm, respectively. of the epidermis cells are thicker than the lateral walls. The collenchyma tissue, which is placed immediately Anatomical characteristics under the epidermis, is composed of regular cells. The Root cortex under the collenchyma is composed of 2–6 layers The transverse sections of the roots reveal that the of squashed oval or almost rectangular parenchymatous periderm layer on the outermost surface is thick and cells with intercellular spaces (Fig. 6). Vascular bundles its cells are squashed or breaking up. Under the perid- are next to each other. Those located at the corners are erm, a multilayered parenchymatic cortex is present. slightly bigger in size than the others. Sclerenchymatous Schlerenchyma groups above several layered phloem cells above the phloem are (2–) 4–15 or more layered comprise 6–10 or more cells which are usually angular (Fig. 7). Cambium is not distinguishable. The xylem (Fig. 3). Cambium is not distinguishable. The xylem consists of trachea and tracheids. The cells underlying rays consist of 1–4 (–6) rowed cells (Fig. 4). As the the xylem are thick-walled. The trachea cells are hexag- 222 A. Kahraman et al.

Table 1. Anatomical measurement of various tissues of S. macrochlamys.

Width (µm) Length (µm)

Min.–Max. Mean ± S.D. Min.–Max. Mean ± S.D.

Root Peridermis Cell 11.2–69.9 36.7 ± 17.4 9.6–45.2 24.1 ± 12.7 Cortex Cell 15.7–49.0 33.0 ± 8.7 7.4–20.4 13.8 ± 3.8 Pith Ray 3.5–16.3 8.4 ± 4.1 9.2–21.9 14.3 ± 4.4 Trachea Cell 20.5–51.2 47.7 ± 19.5 23.2–63.3 70.3 ± 25.1 Stem Cuticle 3.1–6.2 4.3 ± 0.9 Epidermis Cell 12.9–41. 3 26.8 ± 9.0 11.5–25.4 15.9 ± 4.0 Trachea Cell 19.0–50.8 35.4 ± 11.9 19.0–60.0 37.8 ± 13.2 Pith Cell 39.6–160.1 102.4 ± 43.6 Leaf Cuticle 1.8–3.7 2.6 ± 0.7 Upper Epidermis Cell 16.4–38.9 25.4 ± 7.5 15.2–20.5 18.0 ± 1.8 Lower Epidermis Cell 16.4–40.5 23.9 ± 8.2 11.4–20.1 17.0 ± 3.7 Mesophyll Region 154.2–219.7 184.3 ± 19.1 Palisade Parenchyma 7.1–13.3 10.0 ± 2.2 18.9–33.8 25.7 ± 4.4 Spongy Parenchyma 9.9–13.5 12.0 ± 1.4 Petiole Adaxial Epidermis 17.5–35.1 24.4 ± 8.0 16.8–29.4 20.8 ± 4.7 Abaxial Epidermis 30.0–50.1 37.7 ± 6.1 13.4–26.6 21.7 ± 6.1 Cortex Cell 33.0–150.2 81.3 ± 47.1 44.8–177.0 96.3 ± 50.4 Trachea Cell 14.6–32.2 22.8 ± 6.3 10.0–34.0 25.3 ± 9.0

S.D.: Standard Deviation.

Figs 5–7. Transverse section of the stem of S. macrochlamys. Cu – Cuticle, Eg – Eglandular hair, E – Epidermis, Co – Collenchyma, C – Cortex, S – Sclerenchyma, Ph – Phloem, X – Xylem, T – Trachea, Pi – Pith region. Some morphological and anatomical characteristics of Salvia macrochlamys in Turkey 223

Figs 8–11. The transverse section of the leaf of S. macrochlamys. Cu – Cuticle, G – Glandular hair, Eg – Eglandular hair, Ue – Upper epidermis, Le – Lower epidermis, Pc – Parenchymatic cells, Vb – Vascular bundle, X – Xylem, K – Cambium, Ph – Phloem, Pp – Palisade parenchyma, Sp – Spongy parenchyma, St – Stomata. onal or orbicular. The pith is large and is composed of hexagonal or orbicular parenchymatic cells, often with abundant intercellular spaces (Table 1). Leaf The transverse sections of the lamina and midrib reveal that the upper and lower epidermises are covered with a thin cuticle layer. Both epidermises consist of uniseriate, oval, squarish or rectangular cells. The size of the upper epidermis cells is nearly equal to that of the lower epidermis cells. The stomata cells are present on the surfaces of both epidermal layers and are often located at the same level as the epidermal cells (Fig. 10). The leaf is of equifacial type. The palisade cells are elongated rectangular, 2–3 layered above and 1–2 layered below. The spongy parenchyma cells are 2– 3 layered with large intercellular spaces (Fig. 11). The Fig. 12. The transverse section of the petiole of S. macrochlamys. midrib region forms a projecting part (Fig. 8). It is Ad – Adaxial epidermis, Ab – Abaxial epidermis, C – Cortex, X abaxially convex and adaxially slightly convex. Below – Xylem, Ph – Phloem. the uniseriate epidermis, there are several layers of col- lenchymatous cells which provide support. Arc-shaped vascular bundles are surrounded by parenchymatic cells Petiole (Fig. 9). There are one or two major vascular bundles The transverse sections of the petiole show that adax- in the centre and one or two small lateral bundles on ial surface is ﬂat to slightly convex and abaxial surface sides (Table 1). is convex. The epidermis is composed of a single layer 224 A. Kahraman et al.

Figs 13–14. SEM photos of the pollen of S. macrochlamys. 13 – Equatorial view of the pollen with an aperture, 14 – Pollen surface in detail.

Figs 15–16. SEM photos of the nutlet of S. macrochlamys. 15 – General appearance of the nutlet, 16 – Nutlet surface in detail. of cells. The epidermal cells of both surfaces are oval Table 2. Morphological characters of S. macrochlamys based on and almost rectangular. Most hairs on both adaxial and our study and Flora of Turkey (Hedge 1982a). abaxial surfaces are glandular. The collenchyma tissue Our measurements Hedge 1982a is located under the epidermis. Parenchyma cells are hexagonal or nearly circular in shape and they include Stems 30–50 cm 30–50 cm abundant intercellular spaces. The vascular bundle ap- Leaves 3.5–9 × 2–7.5 4–7 × 2–4 cm Petiole 2–5 cm 2–4 cm pears as a shallow arc and is of collateral type. The Infloresence 10–35 cm – sclerenchyma tissue is surrounding the phloem, not the Verticillasters 2–flowered 2–flowered xylem. There is one large and lobed vascular bundle in Bracts 25–45 × 15–25 mm 30–40 × 15–20 mm the middle of the petiole. Also, there are two small sub- Calyx 14–20 mm 20 mm Pedicel 1–3 (–4) mm – sidiary vascular bundles in the petiolar wings (Fig. 12, Corolla 23–35 mm 35 mm Table 1). Filaments 5–9 mm – Pollen characteristics Fertile anther 3–5 mm – The pollen grains of S. macrochlamys are hexacol- Sterile anther 1–2 mm – Upper thecae 3–6 mm – pate, radially symmetrical and isopolar. Their shape Lower thecae 3–6 mm – is oblate-spheroidal (Fig. 13). The dimensions of the Style 25–40 mm – polar axis and equatorial axis are 47.77 ± 3.63 µmand Nutlets 4.7–5.2 × 3.8–4.1 mm 5 × 4mm 52.87 ± 5.23 µm, respectively. The ratio of P/E is 0.90. Colpus length is 40.07 ± 3.59 µm. The exine thickness is 1.80 ± 0.14 µm and intine thickness is 0.80 ± Nutlet micromorphology 0.13 µm. Exine sculpturing is bireticulate and perforate The nutlets of S. macrochlamys are rounded-trigonous (Fig. 14). in transverse section and broadly ovate to rotund in Some morphological and anatomical characteristics of Salvia macrochlamys in Turkey 225

Table 3. Comparison of anatomical, palynological and nutlet characters of S. macrochlamys with Salvia species previously examined.

S. macro- S. halophila S. blepharo- S. huberi S. sclarea S. forskalei chlamys (Kaya et al chlaena (Ozdemir¨ and (Ozdemir¨ and S¸enel (Ozdemir¨ and 2007) (Ozkan¨ and Altan. 2005) 1999, Hedge 1970, S¸enel 2001) Soy 2007) Marin et al. 1996)

Root The number of ray 1–4 (–6) – 1–2 1–3 1–8 2–40 lines Stem The distance of vas- Next to each Separate from – – Separate from each – cular bundles between other each other other the corners Leaf Mesophyll structure Equifacial Monofacial – Bifacial Bifacial – The number of vas- 1–2 in the cen- 2inthecentre 1inthecentre –– – cular bundles in the tre and 1–2 on and 2 on the midrib the sides sides Petiole Presence of sclerenchy- below phloem above xylem ––– – matic tissue and below phloem The number of vascu- 1inthecentre 6inthecentre 1inthecentre 1 in the centre 2 in the centre and – lar bundles and 2 in the and 6 in the and 2 in the and 5 in the 3 in the petiolar petiolar wings petiolar wings petiolar wings petiolar wings wings Pollen Shape Oblate- Prolate – – – – spheroidal The exine sculpturing Bireticulate- Bireticulate- ––– – perforate perforate Seed Size 4.7–5.2 × 3.8– –––3× 2mm – 4.1 mm Shape rounded- –––rounded-trigonous– trigonous and broadly ovate to rotund Ornementation glabrous –––Protuberances– and slightly rough to protuberances

their outline (Fig. 15). Its mature nutlets are 4.7– and the petiole was 2–4 mm in length. According to our 5.2 mm long and 3.8–4.1 mm wide. The attachment study, the leaf is 3.5–9 × 2–7.5 cm, the bract is 25–45 scar diameter ranges from 1.1 mm to 1.3 mm. The nut- × 15–25 mm and the nutlet is 4.7–5.2 × 3.8–4.1 mm let surface is glabrous and slightly rough with protuber- in size, while the calyx is 14–20 mm, the corolla is 23– ances, the epidermal cells of the pericarp appear regular 35 mm and the petiole is 2–5 mm in length. We also polygonal, and the mature nutlet colour is light brown measured other morphological characters of the species (Fig. 16). (Table 2). Research findings reveal that the morphological features of S. macrochlamys provide some additional Discussion information to data reported in Flora of Turkey (Hedge 1982a). Morphological characteristics such as the length of Metcalfe and Chalk (1972) gave some useful infor- corollas and bracts, the number of verticillasters and mation about the root anatomy of the family Labiatae. the stem indumentum are taxonomically significant to They pointed out that the pith rays of roots of this identify S. macrochlamys. The species morphologically family are composed of 2–12 or more rowed cells. The differs from the other members of Salvia in terms of pith rays of S. sclarea L. (sect. Aethiopis) consist of its large flowers and bracts, two flowered verticillasters 1–8 rowed cells (Ozdemir¨ & S¸enel 1999) and those of and thickly glandular villous stems. Even though our S. forskahlei L.(sect. Drymosphace) are composed of 2– results usually correspond with the description in Flora 40 rowed cells (Ozdemir¨ & S¸enel 2001). The rays of S. of Turkey (Hedge 1982a), several differences were found huberi Hedge (sect. Salvia) are 1–3 rowed (Ozdemir¨ & here. It was reported that the leaf was 4–7 × 2–4 cm, the Altan 2005) and those of S. blepharochlaena Hedge & bract was 30–40 × 15–20 mm, the nutlet was 5 × 4mm Hub.-Mor. (sect. Hymenosphace) are 1–2 rowed (Ozkan¨ in size, the calyx was 20 mm, the corolla was 35 mm & Soy 2007). Our studies on the transverse sections of 226 A. Kahraman et al. the root of S. macrochlamys revealed that the species the classification of all genera in Labiatae and placed comprises 1–4 (–6) rowed ray cells. The number of lines it within the subfamily Nepetoideae because the genus of pith rays is a taxonomically considerable significant Salvia has hexacolpate pollen grains. The pollen of S. feature to distinguish sections of the genus. macrochlamys is oblate-spheroidal and its exine sculp- Metcalfe and Chalk (1950) indicated that the turing is bireticulate and perforate. The pollen in S. stems of Labiatae species are quadrangular and have halophila is prolate and its exine sculpturing was ob- collenchyma cells occupying a broad area of the corners. served as bireticulate and perforate (Kaya et al. 2007). They also reported that scleranchymatic tissue encir- S. anatolica Hamzao˘glu & A. Duran and S. bracteata cles the vascular tissue. We observed the same anatom- Banks & Sol. (Hamzao˘glu et al. 2005) were observed to ical features in S. macrochlamys. We also found that have oblate-spheroidal shaped pollen grains. While the the distance of vascular bundles between the corners in sculpturing in S. anatolica is eurireticulate, the sculp- its stems has taxonomic value. All vascular bundles in turing in S. bracteata is suprareticulate. The shape of the stems of S. macrochlamys are next to each other. the pollen and the sculpturing of the exine in the genus However, those of S. blepharochlaena (Ozkan¨ & Soy Salvia may be significant in distinguishing the species. 2007), S. halophila (Kaya et al. 2007) and S. sclarea Seed surface micromorphology was found to have (Ozdemir¨ & S¸enel 1999) are separated from each other systematic significance at generic and specific levels by parenchymatous cells. (Hedge 1970). The nutlets of S. macrochlamys are The leaf mesophyll of Salvia species is entirely rounded-trigonous and broadly ovate to rotund shaped. parenchymatic and the midrib is surrounded by col- ThesizeofmaturenutletsofS. macrochlamys is 4.7– lenchymatous cells (Metcalfe & Chalk 1972). Ac- 5.2 × 3.8–4.1 mm. The nutlet surfaces are glabrous cording to the mesophyll structure, the leaves of S. and slightly rough with protuberances. The nutlets macrochlamys are equifacial, those of S. halophila of S. sclarea L. (sect. Aethiopis) were observed to be (Kaya et al. 2007) are monofacial, and those of S. hu- rounded-trigonous, 3 × 2 mm (Hedge 1982a) and their beri (Ozdemir¨ & Altan 2005) and S. sclarea (Ozdemir¨ surfaces are protuberant (Marin et al 1996). The nutlets &S¸enel 1999) are bifacial. The structure of vascular of S. verticillata L. (sect. Hemisphace)wereanalysedto bundles in the leaf anatomy of Salvia species can be be c. 2.2 × 1.3 mm (Hedge 1982a) and their surfaces are used as a very useful key for distinguishing the species. reticulo-papillosae (Marine et al. 1996). Seed size, shape In the midrib of S. macrochlamys, there are one or two and ornamentation of the genus Salvia have taxonomic large vascular bundles along with one or two small sub- importance in distinguishing the species. Results of the sidiary bundles on sides. However, S. blepharochlaena anatomical, palynological and nutlet characteristics of as indicated by Ozkan¨ & Soy (2007) has one vascular S. macrochlamys and previously examined species are bundle in the middle region and two small bundles on summarized (Table 3). the sides, while S. halophila has two large bundles in the middle of the midrib (Kaya et al. 2007). In their monumental study, Metcalfe and Chalk Acknowledgements (1972) pointed out that the structure of the vascu- We wish to thank Prof. Dr. Cihat Toker, Assoc. Prof. Dr. lar bundles in the petiole of the family Labiatae could H. Nurhan Büyükkartal from Ankara University and As- be used as a diagnostic feature. In the petiole of S. soc. Prof. Dr. Dr. Hatice Çölge¸cen from Zonguldak Karael- macrochlamys, there is a single and lobed large bun- mas University for useful discussions and guidance, Prof. dle and there are two small subsidiary bundles in peti- Dr. Sevil Pehlivan, Dr. Birol Ba¸ser and Dr. Hülya Ozler¨ for olar wings. S. halophila, as indicated by Kaya et al. contributing to the palynological studies and Technical Re- (2007), has six broad vascular bundles in the middle of search Council of Turkey (TUB¨ ITAK-TBAG-104˙ T 450) for the petiole and six small bundles in its wings, S. huberi their financial assistance. (Ozdemir¨ & Altan 2005) has a single large vascular bundle in the centre of the petiole and five small lateral bun- References dles, two of which are located in one petiolar wing and three in the other, S. sclarea (Ozdemir¨ & S¸enel 1999) Baran P. & Ozdemir¨ C. 2006. The Morphological and Anatomical has two large bundles in the centre and three small Characters of Salvia napifolia Jacq. in Turkey. Bangladesh J. 35(1): bundles in the wings, and S. blepharochlaena (Ozkan¨ & Bot. 77–84. Bayraklı F. 1987. Analysis of Soil and Plant. Ondokuz Mayıs Soy 2007) has one large bundle in the centre and two University Agricultural Faculty Publications 17, Samsun. small bundles in the wings. 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