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Department of Biology Education, Ahmet Keleso˘glu Education Faculty, Necmettin Erkaban University, .... permanent slides as described by Vardar (1987).
Biologia 67/4: 663—672, 2012 Section Botany DOI: 10.2478/s11756-012-0049-2

Anatomical and micromorphological studies on Teucrium sect. Isotriodon (Lamiaceae) in Turkey with a taxonomic note ˘ u2 Muhittin Dinc ¸ 1 & S¨ uleyman Dog 1

Department of Biology Education, Ahmet Kele¸so˘glu Education Faculty, Necmettin Erkaban University, 42090 Meram, Konya, Turkey; e-mail: [email protected] 2 Department of Science Education, Ahmet Kele¸so˘glu Education Faculty, Necmettin Erkaban University, 42090 Meram, Konya, Turkey

Abstract: In this study, the anatomical features of the leaf and stem, besides the nutlet characteristics of some Teucrium sect. Isotriodon (Lamiaceae) taxa in Turkey, T. montbretii Betham subsp. montbretii, T. montbretii subsp. pamphylicum P. H. Davis, T. odontites Boiss. & Bal., T. cavernarum P. H. Davis, T. antitauricum T. Ekim, along with an isolated population of T. montbretii (T. montbretii subsp.) were investigated. The anatomical studies revealed that the taxa share generally similar anatomical characters, such as thicker upper leaf cuticles and larger upper leaf epidermal cells compared to lower ones and diacytic to anomocytic stomata on the leaves. However, the portion of the mesophyll occupied by palisade parenchyma and the occurrence of mucilage cells in leaf epidermis shows difference among the taxa. Furthermore, the studied taxa have general stem characteristics of the Lamiaceae family, except for having poorly developed collenchyma at the corners. With the amphistomatic leaves and developed sclerenchymatic tissue in the leaf median vein, T. cavernarum is seperated from the other taxa. Trichome types on the vegetative organs and nutlet shape and sculpturing are generally the same or similar in the studied taxa, but trichomes on the nutlets are different among them. Based on nutlet characteristics and some morphological ones, it was revealed that the isolated population of T. montbretii represent a new subspecies, T. monbretii subsp. yildirimlii M.Din¸c & S.Do˘ gu subsp. nov. Key words: Teucrium; Isotriodon; anatomy; micromorphology; taxonomy; Lamiaceae; Turkey

Introduction Family Lamiaceae has a cosmopolitan distribution and consists of 236 genera and about 7000 species (Stevens 2001). Many species of Lamiaceae family have economic importance because of the essential oil production. Lamiaceae was divided into seven subfamilies: Symphorematoideae, Viticoideae, Ajugoideae, Prostantheroideae, Nepetoideae, Scutellarioideae, and Lamioideae (Harley et al. 2004). Teucrium L. is one of the largest genera of the subfamily Ajugoideae. It is polymorphic genus including about 300 species in the world (Cramer 1981). The genus has cosmopolitan distribution, mainly in Europe, North Africa and in the temperate parts of Asia (K¨ astner 1989; Abu-Assab & Cantino 1993). It has been divided into ten sections, identifiable through the calyx shape and the inflorescence structure (McClintock & Epling 1946; Tutin & Wood 1972). These sections are Teucropsis Benth., Teucrium Benth., Chamaedrys (Mill.) Schreb., Polium (Mill.) Schreb., Isotriodon Boiss., Pycnobotrys Benth., Scorodonia (Hill) Schreb., Stachybotrys Benth., Scordium (Mill.) Benth., and Spinularia Boiss. The genus Teucrium L. (Lamiaceae) is represented by 46 taxa included in 34 species and 8 sections in Turkey. Twelve species and two subspecies are endemic c 2012 Institute of Botany, Slovak Academy of Sciences 

¨ (Duman 2000; D¨ onmez 2006; Ozhatay & K¨ ult¨ ur 2006; Parolly & Eren 2007; D¨ onmez et al. 2010; Din¸c et al. 2011a). Sect. Isotriodon is characterized by dwarf, suffrutescent, fragile, saxatile, and herbaceous habitus; terete stems with dentate or entire margined leaves; simple and secund recemes; gibbous-based and bilabiate calyces; and small nutlets (Ekim 1982). It is represented by seven species in Turkey (Din¸c et al. 2011a). Studies on nutlet in Lamiaceae have shown it to be useful to varying degrees at different levels of the taxonomic hierarchy (Marin et al. 1994; Navarro & El Oualidi 2000; Moon & Hong 2006; Din¸c & Do˘gan 2006; Kaya & Kutluk 2007; Kaya & Dirmenci 2008; Din¸c et al. 2009a; Eshratifar et al. 2010). The absence or presence and types of trichomes on the nutlets and the vegetative parts have taxonomic value in clarification of sectional boundaries in Teucrium. Teucrium sect. Isotriodon is characterised by the elongated thick-walled trichomes on the vegetative parts and subsessile glandular trichomes on the nutlets (Navarro & El Oualidi 2000). In addition, in description of Teucrium pseudaroanium as a new species fom sect. Teucrium, trichome type and density were used as additional diagnostic character from its close relatives (Parolly & Eren 2007). Many anatomical studies were carried out in the family Lamiaceae. The anatomical characters have shown it to be important at different systematic levUnauthenticated Download Date | 4/9/18 8:12 PM

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664 Table 1. Studied materials of sect. Isotriodon taxa.

T. montbretii subsp.

T. montbreti subsp. montbretii

T. montbretii subsp. pamphylicum T. odontites T. cavernarum

T. antitauricum

Locality

Collection data

C6 ADANA: Feke, Feke-S¨ uphandere arası, S¨ uphandere’ye 5 km kala, Sencan Dere kenarı, kayalık yama¸clar, 750–850 m, 6 Jul 2010 C6 ADANA: Feke, Feke-S¨ uphandere arası, S¨ uphandere’ye 5 km kala, Sencan Dere kenarı, kayalık yama¸clar, 750–850 m, 28 Jul 2010 C6 HATAY: Antakya, St. Peter Kilisesi civarı, kayalık yama¸clar, 100 m, 07 Jul 2010 C6 HATAY: Samanda˘ g, sahile bakan kayalık yama¸clar, 50 m, 07 Jul 2010 C3 ANTALYA: Konyaaltı, T¨ uneller B¨ olgesi, kayalık yama¸clar, 100 m, 23 Jul 2010 ˙ A¸sa˘ C3 MERSIN: gıke¸slik K¨ oy¨ u, G¨ uzeldere, kayalık yama¸clar, 280 m, 25 Jul 2010 C4 KARAMAN: Ermenek, Teke¸catı mevkii, 1400 m, kayalık yama¸clar, 19 Aug 2009 C4 KARAMAN: Ermenek, Teke¸catı mevkii, 1400 m, kayalık yama¸clar, 11 Oct 2010 ˙ C6 ADANA: S ¸ aimbeyli, Obruk Ielalesi civarı, kayalık yama¸clar, 1100 m, 06. Jul 2010 C6 ADANA: Saimbeyli, Saimbeyli-Feke arası, Saimbeyli ¸cıkı¸sı, kayalık yama¸clar, 1000 m, 06 Jul 2010

M.Din¸c 3310 & S. Do˘ gua,d M.Din¸c 3344b,c,d M.Din¸c 3322 & S. Do˘ gua,b,c,d M.Din¸c 3315 & S. Do˘ gua,d M.Din¸c 3336 & S. Do˘ gua,b,c,d S. Do˘ gu 2456a,b,c S. Do˘ gu 2052a S. Do˘ gu 2476 & M. Din¸cb,c M.Din¸c 3308 & S. Do˘ gua M.Din¸c 3309 & S. Do˘ gub,c

a The

specimens used for anatomical studies specimens used for the nutlet studies using stereo-microscope c The specimens used for the nutlet studies using SEM d The specimens used for the general morphological studies b The

els. The systematic value of leaf epidermal micromorphological characteristics was useful only subtribal level in Mentheae. But, the combination of the anatomical characters has taxonomic value in segregation of some related species in Lamiaceae. These characters have diagnostic value in separation of the related Teucrium species from the same section (Din¸c et al. 2009b). Moreover, they were used as additional diagnastic characters in reinstatement of Teucrium andrusi Post from its sibling species T. paederotoides Boiss. & Hausskn. in Teucrium sect. Isotriodon. The objective of this study is to investigate the anatomical and nutlet macro-micromorphological characters of some species of sect. Isotriodon and to determine the systematic value of these characters in specific and infraspecific segregation of the taxa. Beside, it is to clarify the taxonomic position of T. montbretii samples collected from an isolated area and showing some morphological differences from Teucrium montbretii subspecies in Turkey.

taxon. The cross-sections were stained with basic fuchsine. All sections were covered by glycerin gelatin and made into permanent slides as described by Vardar (1987). Preparats were observed through an Olympus BX-50 microscope and photographed using “kameram” apparatus attached to the microscope. Anatomical characteristics and trichome morphology was defined by observations of the preparats under the light microcope. Trichome terminology follows Navarro & El Oualidi (2000). The stomatal index was calculated according to Meidner & Mansfield (1968). When the stomata and the epidermal cells number is given, the unit are refers the view area under the objective × 20. Each quantitive anatomical character was measured 20 times, and the data was given as minimum-maximum (mean ± standart deviation). For micromorphological studies on the nutlets, mature nutlets were mounted directly on stubs with double-side adhesive tape and coated with gold. The nutlets were examined by SEM, and the best representatives were photographed in general and in detail. For each taxon, 20 nutlets were measured, and the data was given as minimummaximum (mean ± SD). Nutlet sculpture terminology follows Punt et al. (1994). Features similar in the studied taxa were not emphasized when presenting the results.

Material and methods The samples belonging to Teucrium sect. Isotriodon were collected from different parts of Turkey in 2010. The collection data of the populations is given in Table 1. The samples were dried according to standard herbarium technics and conserved at the herbarium of the Necmettin Erkaban University Ahmet Kele¸so˘ glu Education Faculty. Some of the specimens were put in 70% alcohol for anatomical studies. Anatomical studies were carried out on 10 samples, on average, of each population. In these samples, leaves and stem cross-sections were studied with the lower and upper surface sections of the leaves. On average, twenty preparations were made of each type of sections for each

Results Anatomy Stem anatomy: The stem is more or less terete in transverse section. The epidermis is single-layered and consists of angular and ovoid cells whose walls are thickened, and is covered by a thick layer of cuticle. On the epidermis there are glandular and non-glandular trichomes. The type of glandular trichome is simple short and long clavate and subsessile glandular. Clavate glandular trichomes have a single secretory head cell, Unauthenticated Download Date | 4/9/18 8:12 PM

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Table 2. Important anatomical characters of the studied taxa (1. T. montbretii subsp., 2. T. montbreti subsp. montbretii, 3. T. montbretii subsp. pamphylicum, 4. T. odontites, 5. T. cavernarum, 6. T. antitauricum) Stem

Leaf midvein

1 more or less terete Sclerenchymatic cells with poorly devel- constitute a poorly oped collenchyma developed cluster or exists separately 2 more or less terete Sclerenchymatic cells with poorly devel- constitute a poorly oped collenchyma developed cluster or exists separately 3 more or less terete Sclerenchymatic cells with poorly devel- constitute a poorly oped collenchyma developed cluster or exists separately 4 more or less terete Sclerenchymatic cells with poorly devel- constitute a poorly oped collenchyma developed cluster or exists separately 5 more or less terete Sclerenchymatic cells with poorly devel- constitute a richly deoped collenchyma veloped cluster

6 more or less terete Sclerenchymatic cells with poorly devel- constitute a poorly oped collenchyma developed cluster or exists separately

Leaf mesophyll

Leaf surfaces

Stomatal distribution and type

Dorsiventral with 60–70 % palisade parenchyma

With a few mucilage cells, the stomatal index for the lower surface 21.4, upper epidermal cells are 2–4 times larger than the lower ones With abundant mucilage cells, the stomatal index for the lower surface 19.6, upper epidermal cells are 2–4 times larger than the lower ones With a few mucilage cells, the stomatal index for the lower surface 18.3, upper epidermal cells are 2–4 times larger than the lower ones With a few mucilage cells, the stomatal index for the lower surface 19.27, upper epidermal cells are 2–4 times larger than the lower ones With a few mucilage cells, the stomatal index for the upper surface is 5.26, for the lower surface 21.21, upper epidermal cells are 2–4 times larger than the lower ones With a few mucilage cells, the stomatal index for the lower surface is 35.5, upper epidermal cells are 2–4 times larger than the lower ones

Hypostomatic, xeromorphic, anomocytic to diacytic

Dorsiventral with 60–65 % palisade parenchyma Dorsiventral with 65–73% palisade parenchyma Dorsiventral with 60–65 % palisade parenchyma Dorsiventral with 50–55 % palisade parenchyma

Dorsiventral with 55–60 % palisade parenchyma

but subsessile glandular trichomes have 4-celled secretory head. The type of non-glandular trichome is simple thick-walled with 1-4-celled. There is a subepidermal poorly developed collenchyma. The collenchyma consists of 1-2-layered cells with thickened walls. The cortex between the collenchyma and the endodermis is composed of more or less circular and rectangular and 4-5-layered parenchymatic cells. The endodermis is regular, single layered and composed of quadrate, flattened and ovoid cells. A pericycle layer is present under the endodermis. It is sclerenchymatic, continuous or discontinuous and 1–4 sheets. Phloem and xylem elements can be distinguished in the vascular tissue. The cambium between xylem and phloem is inconspicous. In the whole centre of the stem, there is pith which is filled with circular, slightly pentagon and hexagon angular parenchymatical medullar cells (Table 2, Fig. 1). Leaf anatomy: Mesophyll anatomy: The epidermis consisting of ovoid, quadrate and rectangular cells is present at both surfaces of the leaves. It is one-layered, made of cells whose external walls are thickened. T. montbretii subsp. montbretii has more abundant mucilage cells in the epiderma as compared to the other studied taxa. There are sometimes hypodermal cells under the epidermis especially midvein region. Both surface is covered by a thick cuticula layer. The upper cuticula out of the midvein is quite thick compared to the lower cuticula. The upper cuticula layer is 10.0–25.0 (18.2 ± 2.0) µm thickness, and the lower one 5.0–11.0 (8.0 ± 1.8) µm. The size of epidermal cells in the transection are not significantly different among the taxa,

Hypostomatic, xeromorphic, anomocytic to diacytic Hypostomatic, xeromorphic, anomocytic to diacytic Hypostomatic, xeromorphic, anomocytic to diacytic Amphistomatic, xeromorphic, anomocytic to diacytic

Hypostomatic, xeromorphic, anomocytic to diacytic

but the upper ones are 2–4 times larger than the lower ones. Both leaf surfaces are covered by the indumentum built of glandular and non-glandular hairs of the same type as seen on the stem. The indumentum is densest in T. montbretii subsp., sparsest in T. cavernarum. Leaves are dorsiventral. Under the upper epidermis, there is a palisade parenchyma formed by 2–3 layers. The portion of the mesophyll occupied by palisade parenchyma is 60–70% in T. montbretii subsp., 60–65% in T. montbretii subsp. montbretii, 65–73% in T. montbretii subsp. pamphylicum, 60–65% in T. odontites, 50– 55% in T. cavernarum, 55–60% in T. antitauricum. There is a 2-4-layered spongy parenchyma under the palisade parenchyma. The mesophyll is thicker around the central vein. There are collenchyma under the epidermis in this region. While the collenchyma under the lower epidermis is always well-developed, the one under the upper epidermis is often poorly-developed or absent. Vascular bundle is present in the central part of the vein. It is encircled by vascular bundle sheet which is sometimes inconspicous. The phloem faced the upper epidermis and the xylem faced the lower epidermis are present in the vascular bundle. There is a sclerenchymatic tissue below the phloem. The sclerenchymatic cells constitute a cluster or exists separately. The sclerenchymatic tissue is most developed in T. cavernarum (Table 2, Fig. 2). Surface anatomy: The studied surface sections showed that the leaves are hypostomatic in T. montbretii, T. odontites, T. antitauricum, but amphistomatic in T. cavernarum. The stomata are anomocytic to diacytic. Unauthenticated Download Date | 4/9/18 8:12 PM

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Fig. 1. Cross-section of the stems. A – T. montbretii subsp.; B – T. montbretii subsp. montbretii; C – T. montbretii subsp. pamphylicum; D – T. odontites; E – T. cavernarum; F –T. antitauricum. t, trichome; gt, glandular trichome; e, epidermis; cl, collenchyma; co, cortex; en, endodermis; sc, sclerenchyma; ph, phloem; x, xylem; p, pith.

Fig. 2. Cross-section of the leaves including the midribs. A – T. montbretii subsp.; B – T. montbretii subsp. montbretii; C – T. montbretii subsp. pamphylicum; D – T. odontites; E – T. cavernarum; F –T. antitauricum. t, trichome; gt, glandular trichome; cu, cuticle; ue, upper epidermis; h, hypoderma; co, collenchyma; sc, sclerenchyma; ph, phloem; x, xylem; pp, palisade parenchyma; sp, spongy parenchyma; le, lower epidermis; mc, mucilage cell.

The stoma cells usually form a row slightly below the epidermis on the both surface. Namely, they are more or less xeromorphic type. The upper epidermal cells in surface sections are 67.0–170.0 (114.0 ± 15.0) µm long and 39.0–108.0 (71.0 ± 12.0) µm wide, the lower ones are 44.0–133.0 (81.0 ± 16.0) µm long and 11.0–76.0 (42.0 ± 14.0) µm wide. The anticlinal wall of the upper epidermal cell is thicker than of the lower epidermal cell. The lower epidermal cell walls are more undulated than the upper ones (Table 2, Figs 3, 4). In T. montbretii subsp., the upper epidermal cells are 88.0–170.0 (121.82 ± 13.46) µm long and 44.0–

108.0 (72.86 ± 8.56) µm wide, the lower ones are 64.0–133.0 (95.14 ± 17.68) µm long and 12.0–41.0 (26.46 ± 7.97) µm wide. On the lower surface, the number of stomata is 11 ± 3 per unit area, and that of the epidermal cells is 45 ± 5 per unit area. The stomatal index for the lower surface is 19.6 (Table 2, Figs 3, 4). In T. montbretii subsp. montbretii, the upper epidermal cells are 91.0–156.0 (110.91 ± 10.56) µm long and 51.0–71.0 (61.38 ± 3.80) µm wide, the lower ones are 61.0–103.0 (81.93 ± 15.61) µm long and 19.0–60.0 (34.14 ± 9.56) µm wide. On the lower surface, the number of stomata is 15 ± 4 per unit area, and that of the Unauthenticated Download Date | 4/9/18 8:12 PM

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Fig. 3. Upper surface-sections of the leaves. A – T. montbretii subsp.; B – T. montbretii subsp. montbretii; C – T. montbretii subsp. pamphylicum; D – T. odontites; E – T. cavernarum; F –T. antitauricum. st, stoma; ec, epidermal cell; gt, glandular trichome.

Fig. 4. Lower surface-sections of the leaves. A – T. montbretii subsp.; B – T. montbretii subsp. montbretii; C – T. montbretii subsp. pamphylicum; D – T. odontites; E – T. cavernarum; F –T. antitauricum. st, stoma; ec, epidermal cell.

epidermal cells is 55 ± 6 per unit area. The stomatal index for the lower surface is 21.4 (Table 2, Figs 3, 4). In T. montbretii subsp. pamphylicum, the upper epidermal cells are 89.0–160.0 (120.21 ± 14.34) µm long and 39.0–73.0 (52.31 ± 12.65) µm wide, the lower ones are 61.0–103.0 (80.56 ± 11.02) µm long and 11.0–31.0 (21.35 ± 5.87) µm wide. On the lower surface, the number of stomata is 18 ± 4 per unit area, and that of the epidermal cells is 80 ± 6 per unit area. The stomatal index for the lower surface is 18.3 (Table 2, Figs 3, 4). In T. odontites, the upper epidermal cells are 85.0–109.0 (97.87 ± 4.75) µm long and 55.0–96.0 (70.45 ± 9.21) µm wide, the lower ones are 74.0–101.0 (84.50 ± 6.37) µm long and 18.0–48.0 (31.32 ± 7.99) µm

wide. On the lower surface, the number of stomata is 15 ± 5 per unit area, and that of the epidermal cells is 68 ± 8 per unit area. The stomatal index for the lower surface is 19.27 (Table 2, Figs 3, 4). In T. cavernarum, the upper epidermal cells are 67.0–159.0 (109.61 ± 13.77) µm long and 41.0– 87.0 (61.38 ± 6.89) µm wide, the lower ones are 84.0–141.0 (116.73 ± 16.32) µm long and 13.0–76.0 (28.74 ± 3.02) µm wide. On the upper surface, the number of stomata is 2 ± 1 per unit area, and that of the epidermal cells is 36 ± 4 per unit area. The stomatal index for the upper surface is 5.26. On the lower surface, the number of stomata is 14 ± 4 per unit area, and that of the epidermal cells is 52 ± 8 per unit area. The Unauthenticated Download Date | 4/9/18 8:12 PM

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Fig. 5. SEM micrographs of nutlets. A,B – T. montbretii subsp.; C,D – T. montbretii subsp. montbretii; E,F – T. montbretii subsp. pamphylicum; G,H – T. odontites; I,J – T. cavernarum; K,L – T. antitauricum.

stomatal index for the lower surface is 21.21 (Table 2, Figs 3, 4). In T. antitauricum, the upper epidermal cells are 97.0–135.0 (111.48 ± 7.88) µm long and 44.0–84.0

(56.69 ± 9.21) µm wide, the lower ones are 44.0–121.0 (83.24 ± 21.23) µm long and 20.0–55.0 (31.49 ± 4.33) µm wide. On the lower surface, the number of stomata is 32 ± 5 per unit area, and that of the epidermal cells is Unauthenticated Download Date | 4/9/18 8:12 PM

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Table 3. Micromorphological nutlet characters of the studied taxa.

T. montbretii subsp. T. montbreti subsp. montbretii T. montbretii subsp. pamphylicum T. odontites T. cavernarum T. antitauricum

Nutlet length (mm)

Nutlet width (mm)

Nutlet colour and shape

Nutlet hairs

1.11–1.56 (1.39±0.12) 1.20–1.39 (1.29±0.08) 0.95–1.42 (1.20±0.13)

0.60–1.00 (0.77±0.11) 0.61–0.93 (0.75±0.10) 0.65–0.84 (0.73±0.07)

Densely non-glandular and subsessile glandular Subsessile glandular

1.10–1.36 (1.27±0.08) 0.81–1.45 (1.11±0.16) 1.09–1.43 (1.27±0.11)

0.60–0.85 (0.70±0.08) 0.51–0.75 (0.63±0.10) 0.72–0.84 (0.76±0.05)

Light-brown, narrowly ovate-oblong, with inconspicuous alveoles Blackish-brown, ovate-oblong, with regular, deep and thin-walled alveoles Blackish-brown, ovate-elliptic, with irregular, shallow and thick-walled alveoles Blackish-brown, ovate-oblong, with inconspicuous alveoles Light-brown, narrowly ovate-oblong, with inconspicuous alveoles Light-brown, ovate-oblong, with regular, deep and thin-walled alveoles

58 ± 8 per unit area. The stomatal index for the lower surface is 35.5 (Table 2, Figs 3, 4). Nutlet morphology: The size of nutlets ranges from 1.10 to 1.40 mm in length and 0.65 to 0.75 mm in width. The nutlet colour is light-brown in T. montbretii subsp., T. antitauricum and T. cavernarum, blackish-brown in T. montbreti subsp. montbretii, T. montbretii subsp. pamphylicum, and T. odontites. The nutlet shapes are narrowly ovate-oblong, ovate-oblong or ovate-elliptic. The nutlet surface is generally alveolate, but alveoles are inconspicuous in T. montbretii subsp., T. odontites and T. cavernarum. It is conspicuous in T. montbreti subsp. montbretii, T. montbretii subsp. pamphylicum and T. antitauricum, but has some specific differences in depth and shape of alveoles among these taxa. They are regular, deep and thin-walled in T. montbreti subsp. montbretii, irregular, shallow and thick-walled in T. montbretii subsp. pamphylicum and regular, deep and thin-walled in T. antitauricum. Subsessile glandular trichomes are present in the studied taxa, but eglandular trichomes are present only in T. montbretii subsp. and T. cavernarum (Table 3, Fig. 5). Taxonomic Treatment The present study showed that T. montbretii subsp. has enough differences from the other subspecies to be treated as separate subspecies. Teucrium montbretii subsp. yildirimlii M. Din¸c & S. Do˘gu subsp. nov. (Fig. 6A) Type: Turkey. C6 Adana: Feke, Feke-S¨ uphandere arası, S¨ uphandere’ye 5 km kala, Sencan Dere kenarı, kayalık yama¸clar, 750–850 m, 6 Jul 2010, M.Din¸c 3310 & S. Do˘ gu (holotype: KNYA, isoypes: GAZI, HUB) Diagnosis: subsp. montbretii similis sed plantis cinereo tomentosis-villosis (non viridis), caulis semper ramosus (non raro), nuculis cum pilis eglandulosis (non sine) differt. Description: Dwarf, fragile, suffruticose, densely greyish tomentose-villous and viscid, covered with glandular and non-glandular trichomes all over. Flowering stems

Subsessile glandular

Subsessile glandular Densely non-glandular and subsessile glandular Subsessile glandular

5–15 cm long, much-branched; Leaves petiolate with 2–8 mm petiole, densely tomentose-villous with simple short and long clavate glandular, subsessile glandular and simple thick-walled non-glandular trichomes; lamina 5–20 × 5–18 mm, ovate to ovate-lanceolate, broadest below the middle, slightly acute at apex, lower ones crenate with 4–9 teeth each side, subcordate to truncate at base, the upper ones indistinctly crenate to entire, cuneate at base; nerves fine, clearly distinct below; Inflorescence terminal and lateral raceme, dense, densely tomentose-villous with simple short and long clavate glandular, subsessile glandular and simple thick-walled non-glandular trichomes, nearly capitate, ovate or pyramidal, not more than 3 cm in length. Verticillasters biflowered, many, densely congested. Bracts linear to oblanceolate, acute at apex, attenuate at base, 5–7 mm, overtopping the calyx above the raceme, equalling below. Pedicels about 1–4 mm. Calyx 4.0–5.0 mm, clearly bilabiate, gibbous at base, tube 2.5–3.5 mm, teeth distinctly acute, equal or shorter than the tube, upper 3 teeth ovate, 2.4–2.6 mm long, the lower two teeth linear to linear-lanceolate, 2.6–3.0 mm, slightly longer than the upper ones. Corolla longer than calyx, pinkish to creamy-white, pale. Phenology: Flowering July, fruiting late July-August. Distribution: South Anatolia. Endemic, Medit. element. Ecology: Teucrium montbretii subsp. yildirimlii grows on the rocky slopes in conifer forest clearings faced to stream banks an altitude of more than 750 m. Conservation status: Teucrium montbretii subsp. yildirimlii is stenoendemic subspecies restricted to the presently known locality. Its estimated area of occupancy is less than 1 km2 (criterion B). The population includes less than 250 indivuduals (criterion C). The area is near to road (criterion D). Therefore, the new subspecies should be classified as “Critically endangered (CR)” based on the criteria of the IUCN Red List Categories (IUCN 2001). Identification Key General appearances of the subspecies of Teucrium Unauthenticated Download Date | 4/9/18 8:12 PM

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. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . subsp. yildirimlii – Plant greenish tomentose-villous; stem rarely branched; flowers lilac to purplish; nutlets without eglandular trichomes . . . . . . . . . . . . . . . . . . . subsp. montbretii Discussion

Fig. 6. General appearances of T. montbretii subspecies in Turkey. A – T. montbretii subsp. yildirimlii; B – T. montbretii subsp. montbretii; C – T. montbretii subsp. pamphylicum.

montbretii in Turkey show some differences (Fig. 6). In the light of the data from Ekim (1982) and the present study, the identification key to T. montbretii subspecies in Turkey can be constructed as follows; 1 Inflorescence lax, 5–20 cm . . . subsp. pamphylicum – Inflorescence dense, less than 5 cm . . . . . . . . . . . . . . . . 2 2 Plant greyish tomentose-villous; stem always branched; flowers generally white, rarely pinkish, nutlets with densely eglandular trichomes . . . . . . . . . . . . . . . . . .

The morphological polymorphism in the genus Teucrium is reflected to the anatomical characters. Therefore, the anatomical characters show differences between the closed species in the same section (Din¸c & ¨ urk 2008; Din¸c et al. 2008; Din¸c et al. 2009b). MoreOzt¨ over, they were used as additional diagnastic characters in reinstatement of Teucrium andrusi Post from its sibling species T. paederotoides Boiss. & Hausskn. in Teucrium sect. Isotriodon (Din¸c et al. 2011a). But, this study showed that the studied taxa generally exhibit anatomical homogeneity with regard to stem and leaf anatomy (Table 2). Only T. cavernarum, with the developed sclerenchymatic tissue in leaf midvein and amphistomatic distribution of the stomata, and T. montbretii subsp. montbretii with the abundant mucilage cells in the epidermis are distinguished from the other studied taxa. The portion of the mesophyll occupied by palisade parenchyma shows difference among the studied taxa. But, these differences can be based on the ecological plasticity among the populations of the same Teucrium species (Lakuši´c et al. 2007). Metcalfe & Chalk (1950) pointed out that the stems of the family Lamiaceae species are rectangular and the collenchymatic tissue covers broad area at the corners, and a developed sclerenchymatic tissue surrounds the vascular tissue. The anatomical studies on Lamiaceae showed that the species from the family had the same anatomical characteristics (Kaya et al. 2000; Kandemir 2003; Uysal 2002, 2003). The present anatomical studies revealed that the taxa share generally similar anatomical characters with the family, but the collenchymatic tissue do not cover broad area at the corners of the stem in the studied taxa. The stems with poorly developed collenchymatic tissue at the corners are present in the section Polium as well (Din¸c et al. 2011b). The sections Polium and Isotriodon are closely related in the genus Teucrium according to Turkish Flora (Ekim 1982) and characterised by their terete stems, as well as other morphological features. The anatomical features of the stems in both sections support their relationships in terms of stem morphology. The absence or presence and types of trichomes on the nutlets and the vegetative parts have taxonomic value in clarification of sectional boundaries in Teucrium (Navarro & El Oualidi 2000). In addition, in description of Teucrium pseudaroanium as a new species fom sect. Teucrium, trichome type and density were used as additional diagnostic character from its close relatives (Parolly & Eren 2007). With the non glandular trichomes on its nutlets, T. montbretii subsp. yildirimlii is readily distinguished from the other subspecies of T. montbretii. Unauthenticated Download Date | 4/9/18 8:12 PM

Anatomical and morphological studies on Teucrium sec. Isotriodon in Turkey Teucrium sect. Isotriodon is characterised by the elongated thick-walled trichomes on the vegetative parts and subsessile glandular trichomes on the nutlets (Navarro & El Oualidi 2000). This opinion is not supported by the present study. Because, the endemic taxa T. montbretii subsp. yildirimlii and T. cavernarum have both subsessile glandular and non glandular trichomes on their nutlets. The samples from the type locality of T. montbretii subsp. yildirimlii were previously identified as T. montbretii subsp. montbretii (Ekim 1982). But, both genuine subsp. montbretii and subsp. pamphylicum grow on the rocky slopes in maquis vegetation faced to the sea from the sea level to an altitude of about 250 m according to our observations. Whereas, the new subsp. yildirimlii grows on the rocky slopes in conifer forest clearings faced to Sencan Dere stream banks an altitude of more than 750 m. In conclusion, the description of the new subspecies is supported by the present ecological isolation as well. The results showed that the studied taxa from Teucrium sect. Isotriodon are generally the same or similar to each other anatomical and nutlet micromorphological point of view, but trichome types on their nutlets show some differences among them. In addition, it was concluded that the isolated population of T. montbretii from Adana-Feke show enough differences to be described as a new subspecies. References Abu-Assab M.S. & Cantino P.D. 1993. Phylogenetic implications of pollen morphology in tribe Ajugeae (Labiatae). Syst. Bot. 18: 100–122. Cramer L.H. 1981. Teucrium L. In: Dassanayake M.D. & Fosberg F.R. (eds), A revised handbook to the flora of Ceylon 3: 193194. Oxford and IBH Publishing Co. Pvt. Ltd., New Delhi. Din¸c M. & Do˘ gan H.H. 2006. Stachys yildirimlii M.Din¸c (Lamiaceae), a new species from South Anatolia, Turkey. Ann. Bot. Fenn. 43: 143–147. ¨ urk M. 2008. Comparative morphological, anatomDin¸c M. & Ozt¨ ical and palynological studies on the genus Stachys L. sect. Ambleia Bentham (Lamiaceae) species in Turkey. Turk. J. Bot. 32: 113–121. ¨ urk M. 2008. Anatomy, Din¸c M., Duran A., Pınar N.M. & Ozt¨ palynology and nutlet micromorphology of Turkish endemic Teucrium sandrasicum (Lamiaceae). Biologia 63: 637–641. Din¸c M., Pınar N.M., Do˘ gu S. & Yıldırımlı S ¸ . 2009a. Micromorphological studies of Lallemantia L. (Lamiaceae) species growing in Tuekey. Acta Bio. Crac. Ser. Bot. 51: 45–54. Din¸c M., Do˘ gu S., Bilgili B. & Duran A. 2009b. Comparative anatomical and micromorphological studies on Teucrium creticum and Teucrium orientale var. orientale (T. sect. Teucrium, Lamiaceae). Nord. J. Bot. 27: 251–256. Din¸c M., Do˘ gu S. & Ba˘ gcı Y. 2011a. Taxonomic reinstatement of Teucrium andrusi from T. paederotoides based on morphological and anatomical evidences. Nord. J. Bot. 29: 148–158. Din¸c M., Do˘ gu S., Do˘ gru Koca A. & Kaya B. 2011b. Anatomical and nutlet differentiation between Teucrium montanum and T. polium from Turkey. Biologia 66: 448–453. D¨ onmez A.A. 2006. Teucrium chasmophyticum Rech. f. (Lamiaceae): a new record for the flora of Turkey. Turk. J. Bot. 30: 317–320. ¨ celik A.D. 2010. Teucrium melisD¨ onmez A.A., Mutlu B. & Oz¸ soides Boiss. & Hausskn. ex Boiss. (Lamiaceae): A New

671

Record for Flora of Turkey. Hacettepe J. Biol. & Chem. 38: 291–294. Duman H. 2000. Teucrium L., pp. 197–198. In: G¨ uner A., ¨ Ozhatay N., Ekim T. & Ba¸ser K.H.C. (eds), Flora of Turkey and the East Aegean Islands, vol. 11 (Supplements), Edinburg Univ. Press, Edinburgh. Ekim T. 1982. Teucrium L., pp. 53–75. In: Davis P.H. (ed.), Flora of Turkey and the East Aegean Islands, vol. 7, Edinburgh Univ. Pres, Edinburgh, Royal Botanic Gardens, Kew. Eshratifar M., Attar F. & Mahdigholi K. 2010. Micromorphological studies on nutlet and leaf indumentum of genus Teucrium L. (Lamiaceae) in Iran. Turk J. Bot. 35: 25–35. Harley R.M., Atkins S., Budantsev A.L., Cantino P.D., Conn B.J., Grayer R., Harley M.M., De Kok R., Krestovskaja T., Morales R., Paton A.J., Ryding O. & Upson T. 2004. Labiatae, pp. 167–275. In: Kadereit J.W. (ed.), The families and genera of vascular plants, vol. 7, Springer Verlag, New York. IUCN 2001. IUCN Red List Categories and Criteria, Version 3.1. Prepared by the IUCN Species Survival Commission. Kandemir N. 2003. The morphological, anatomical and karyological properties of endemic Salvia hypargeia Fich. & Mey. (Lamiaceae) in Turkey. Pak. J. Bot. 35: 219–236. Kaya A., Ba¸ser K.H.C., Satıl F. & T¨ umen G. 2000. Morphological and anatomical studies on Cyclotrichium origanifolium (Labill.) Manden. & Scheng. (Labiatae). Turk. J. Bot. 24: 273–278. Kaya A. & Kutluk H. 2007. Pollen morphology of Acinos Miller species growing in Turkey. J. Integr. Plant Biol. 49: 1386– 1392. Kaya A. & Dirmenci T. 2008. Nutlet surface micromorphology and taxonomy of species of the genus Nepeta L. (Lamiaceae) in Turkey. Turk. J. Bot. 32: 103–112. K¨ astner A. 1989. Ubersicht zur systematischen Gliederung der Gattung Teucrium L. Biocosme Mesogeen (Nice) 6: 63–77. Lakuši´ c B., Lakuši´c D., Slavkovska V., Stevanovi´c V. & Stevanovi´ c B. 2007. Morpho-anatomical differentiation of the Balkan endemic species Teucrium arduini L. (Lamiaceae). Arch. Biol. Sci. 59: 369–381. Marín D.P., Petkovic B. & Duletic S. 1994. Nutlet sculpturing of selected Teucrium species (Lamiaceae): a character of taxonomic significance. Plant Syst. Evol. 192: 199–214. McClintock E. & Epling C. 1946. A revision of Teucrium in the New World, with observations on its variation, geographical distribution and history. Brittonia 5: 491–510. Metcalfe C.R. & Chalk L. 1950. Anatomy of the dicotyledons I. Oxford University Press, pp. 1041–1053. Meidner H. & Mansfield T.A. 1968. Physiology of stomata. McGraw-Hill, pp. 138–141. Moon H.K. & Hong S.P. 2006. Nutlet morphology and anatomy of the genus Lycopus (Lamiaceae: Mentheae). J. Plant Res. 119: 633–644. Navarro T. & El Oualidi J. 2000. Trichome morphology in Teucrium L. (Labiatae), a taxonomic review. Ann. J. Bot. Madrid 57: 277–297. ¨ Ozhatay N. & K¨ ult¨ ur S ¸ . 2006: Check-List of additional taxa to the Supplement Flora of Turkey III. Turk. J. Bot. 30: 281– 316. ¨ 2007. Teucrium (sect. Teucrium) pseuParolly G. & Eren O. daroanium Parolly, Erda˘ g & Nordt, sp. nov. In: Parolly G. & ¨ (eds), Contributions to the flora of Turkey, 2. WilldeEren O. nowia 37: 252–258. Punt W., Blackmore S., Nilson S. & Le Tomas A. 1994. Glossary of pollen and spore terminology. LPP Foundation, Utrecht. Stevens P.F. 2001. Angiosperm phylogeny website. Version 6. May 2005. Available at http://www.mobot.org/MOBOT/ research/APweb/ Tutin G. & Wood D. 1972. Teucrium L., pp. 129–135. In: Tutin T.G. Heywood V.H., Burges N.A., Moore D.M., Valentine D.H., Walters S.M. & Webb D.A. (eds), Flora Europaea, vol. 3, Cambridge University Press., Cambridge. ˙ 2002. Stachys cretica L. subsp. smyrnaea Rech Fil. Uysal I. Endemik Taksonunun Morfolojisi, Anatomisi ve Ekolojisi ¨ Uzerinde Ara¸stırmalar. Ekoloji 11: 16–20.

Unauthenticated Download Date | 4/9/18 8:12 PM

672 ˙ 2003. Stachys thirkei C.Koch (Kekikgiller) t¨ Uysal I. ur¨ un¨ un morfolojisi, anatomisi ve ekolojisi u ¨zerine ara¸stırmalar. Ot Sistematik Bot. Dergisi 10: 129–141. ¨ Vardar Y. 1987. Botanikte Preparasyon Tekni˘ gi. Ege Universitesi Fen Fak¨ ultesi Basımevi, pp. 25–26.

˘u M. Dinc ¸ & S. Dog Received June 1, 2011 Accepted September 29, 2011

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