Lamiaceae) in Turkey with a Taxonomic Note

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

Muhittin Dinc¸1 &S¨uleyman Dogu˘ 2

1Department of Biology Education, Ahmet Kele¸so˘glu Education Faculty, Necmettin Erkaban University, 42090 Meram, Konya, Turkey; e-mail: [email protected] 2Department 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 (Duman 2000; Dönmez 2006; Ozhatay¨ & Kültür 2006; Parolly & Eren 2007; Dönmez et al. 2010; Din¸cet Family Lamiaceae has a cosmopolitan distribution al. 2011a). Sect. Isotriodon is characterized by dwarf, and consists of 236 genera and about 7000 species suffrutescent, fragile, saxatile, and herbaceous habitus; (Stevens 2001). Many species of Lamiaceae family terete stems with dentate or entire margined leaves; have economic importance because of the essential oil simple and secund recemes; gibbous-based and bilabiate production. Lamiaceae was divided into seven sub- calyces; and small nutlets (Ekim 1982). It is represented families: Symphorematoideae, Viticoideae, Ajugoideae, by seven species in Turkey (Din¸c et al. 2011a). Prostantheroideae, Nepetoideae, Scutellarioideae, and Studies on nutlet in Lamiaceae have shown it to Lamioideae (Harley et al. 2004). be useful to varying degrees at different levels of the Teucrium L. is one of the largest genera of the sub- taxonomic hierarchy (Marin et al. 1994; Navarro & El family Ajugoideae. It is polymorphic genus including Oualidi 2000; Moon & Hong 2006; Din¸c&Do˘gan 2006; about 300 species in the world (Cramer 1981). The Kaya & Kutluk 2007; Kaya & Dirmenci 2008; Din¸cetal. genus has cosmopolitan distribution, mainly in Eu- 2009a; Eshratifar et al. 2010). The absence or presence rope, North Africa and in the temperate parts of Asia and types of trichomes on the nutlets and the vegetative (Kästner 1989; Abu-Assab & Cantino 1993). It has been parts have taxonomic value in clarification of sectional divided into ten sections, identifiable through the ca- boundaries in Teucrium. Teucrium sect. Isotriodon is lyx shape and the inflorescence structure (McClintock characterised by the elongated thick-walled trichomes & Epling 1946; Tutin & Wood 1972). These sections on the vegetative parts and subsessile glandular tri- are Teucropsis Benth., Teucrium Benth., Chamaedrys chomes on the nutlets (Navarro & El Oualidi 2000). (Mill.) Schreb., Polium (Mill.) Schreb., Isotriodon In addition, in description of Teucrium pseudaroanium Boiss., Pycnobotrys Benth., Scorodonia (Hill) Schreb., as a new species fom sect. Teucrium,trichometype Stachybotrys Benth., Scordium (Mill.) Benth., and and density were used as additional diagnostic char- Spinularia Boiss. acter from its close relatives (Parolly & Eren 2007). The genus Teucrium L. (Lamiaceae) is represented Many anatomical studies were carried out in the by 46 taxa included in 34 species and 8 sections in family Lamiaceae. The anatomical characters have Turkey. Twelve species and two subspecies are endemic shown it to be important at different systematic lev-

c 2012 Institute of Botany, Slovak Academy of Sciences 664 M. Dinc¸ &S.Dogu˘

Table 1. Studied materials of sect. Isotriodon taxa.

Locality Collection data

T. montbretii subsp. C6 ADANA: Feke, Feke-Süphandere arası, Süphandere’ye 5 km kala, M.Din¸c 3310 & S. Do˘gua,d Sencan Dere kenarı, kayalık yama¸clar, 750–850 m, 6 Jul 2010 C6 ADANA: Feke, Feke-Süphandere arası, Süphandere’ye 5 km kala, M.Din¸c 3344b,c,d Sencan Dere kenarı, kayalık yama¸clar, 750–850 m, 28 Jul 2010 T. montbreti subsp. C6 HATAY: Antakya, St. Peter Kilisesi civarı, kayalık yama¸clar, 100 M.Din¸c 3322 & S. Do˘gua,b,c,d montbretii m, 07 Jul 2010 C6 HATAY: Samanda˘g, sahile bakan kayalık yama¸clar,50m,07Jul M.Din¸c 3315 & S. Do˘gua,d 2010 T. montbretii subsp. C3 ANTALYA: Konyaaltı, Tüneller Bölgesi, kayalık yama¸clar, 100 m, M.Din¸c 3336 & S. Do˘gua,b,c,d pamphylicum 23 Jul 2010 T. odontites C3 MERSIN:˙ A¸sa˘gıke¸slik Köyü, Güzeldere, kayalık yama¸clar, 280 m, S. Do˘gu 2456a,b,c 25 Jul 2010 T. cavernarum C4 KARAMAN: Ermenek, Teke¸catı mevkii, 1400 m, kayalık yama¸clar, S. Do˘gu 2052a 19 Aug 2009 C4 KARAMAN: Ermenek, Teke¸catı mevkii, 1400 m, kayalık yama¸clar, S. Do˘gu 2476 & M. Din¸cb,c 11 Oct 2010 T. antitauricum C6 ADANA: S¸aimbeyli, Obruk Ielalesi˙ civarı, kayalık yama¸clar, 1100 M.Din¸c 3308 & S. Do˘gua m, 06. Jul 2010 C6 ADANA: Saimbeyli, Saimbeyli-Feke arası, Saimbeyli ¸cıkı¸sı, kayalık M.Din¸c 3309 & S. Do˘gub,c yama¸clar, 1000 m, 06 Jul 2010 aThe specimens used for anatomical studies bThe specimens used for the nutlet studies using stereo-microscope cThe specimens used for the nutlet studies using SEM dThe specimens used for the general morphological studies els. The systematic value of leaf epidermal micromor- taxon. The cross-sections were stained with basic fuchsine. phological characteristics was useful only subtribal level All sections were covered by glycerin gelatin and made into in Mentheae. But, the combination of the anatomical permanent slides as described by Vardar (1987). Preparats characters has taxonomic value in segregation of some were observed through an Olympus BX-50 microscope and related species in Lamiaceae. These characters have di- photographed using “kameram” apparatus attached to the microscope. Anatomical characteristics and trichome mor- agnostic value in separation of the related Teucrium phology was defined by observations of the preparats under species from the same section (Din¸c et al. 2009b). More- the light microcope. Trichome terminology follows Navarro over, they were used as additional diagnastic characters & El Oualidi (2000). The stomatal index was calculated ac- in reinstatement of Teucrium andrusi Post from its sib- cording to Meidner & Mansfield (1968). When the stomata ling species T. paederotoides Boiss. & Hausskn. in Teu- and the epidermal cells number is given, the unit are refers crium sect. Isotriodon. the view area under the objective × 20. Each quantitive The objective of this study is to investigate the anatomical character was measured 20 times, and the data anatomical and nutlet macro-micromorphological char- was given as minimum-maximum (mean ± standart devia- acters of some species of sect. Isotriodon and to deter- tion). For micromorphological studies on the nutlets, mature mine the systematic value of these characters in spe- nutlets were mounted directly on stubs with double-side cific and infraspecific segregation of the taxa. Beside, adhesive tape and coated with gold. The nutlets were ex- it is to clarify the taxonomic position of T. montbretii amined by SEM, and the best representatives were pho- samples collected from an isolated area and showing tographed in general and in detail. For each taxon, 20 nut- some morphological differences from Teucrium mont- lets were measured, and the data was given as minimum- bretii subspecies in Turkey. maximum (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 Results collected from diﬀerent parts of Turkey in 2010. The collection data of the populations is given in Table 1. The sam- Anatomy ples were dried according to standard herbarium technics Stem anatomy: The stem is more or less terete in trans- and conserved at the herbarium of the Necmettin Erkaban verse section. The epidermis is single-layered and con- University Ahmet Kele¸so˘gluEducation Faculty. sists of angular and ovoid cells whose walls are thick- Some of the specimens were put in 70% alcohol for ened, and is covered by a thick layer of cuticle. On anatomical studies. Anatomical studies were carried out on 10 samples, on average, of each population. In these samples, the epidermis there are glandular and non-glandular leaves and stem cross-sections were studied with the lower trichomes. The type of glandular trichome is simple and upper surface sections of the leaves. On average, twenty short and long clavate and subsessile glandular. Clavate preparations were made of each type of sections for each glandular trichomes have a single secretory head cell, Anatomical and morphological studies on Teucrium sec. Isotriodon in Turkey 665

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 Leaf mesophyll Leaf surfaces Stomatal distribution and type

1 more or less terete Sclerenchymatic cells Dorsiventral with With a few mucilage cells, the stom- Hypostomatic, xero- with poorly devel- constitute a poorly 60–70 % palisade atal index for the lower surface 21.4, morphic, anomocytic oped collenchyma developed cluster or parenchyma upper epidermal cells are 2–4 times to diacytic exists separately larger than the lower ones 2 more or less terete Sclerenchymatic cells Dorsiventral with With abundant mucilage cells, the Hypostomatic, xero- with poorly devel- constitute a poorly 60–65 % palisade stomatal index for the lower surface morphic, anomocytic oped collenchyma developed cluster or parenchyma 19.6, upper epidermal cells are 2–4 to diacytic exists separately times larger than the lower ones 3 more or less terete Sclerenchymatic cells Dorsiventral with With a few mucilage cells, the stom- Hypostomatic, xero- with poorly devel- constitute a poorly 65–73% palisade atal index for the lower surface 18.3, morphic, anomocytic oped collenchyma developed cluster or parenchyma upper epidermal cells are 2–4 times to diacytic exists separately larger than the lower ones 4 more or less terete Sclerenchymatic cells Dorsiventral with With a few mucilage cells, the stom- Hypostomatic, xero- with poorly devel- constitute a poorly 60–65 % palisade atal index for the lower surface 19.27, morphic, anomocytic oped collenchyma developed cluster or parenchyma upper epidermal cells are 2–4 times to diacytic exists separately larger than the lower ones 5 more or less terete Sclerenchymatic cells Dorsiventral with With a few mucilage cells, the stom- Amphistomatic, xero- with poorly devel- constitute a richly de- 50–55 % palisade atal index for the upper surface is morphic, anomocytic oped collenchyma veloped cluster parenchyma 5.26, for the lower surface 21.21, to diacytic upper epidermal cells are 2–4 times larger than the lower ones 6 more or less terete Sclerenchymatic cells Dorsiventral with With a few mucilage cells, the stom- Hypostomatic, xero- with poorly devel- constitute a poorly 55–60 % palisade atal index for the lower surface is morphic, anomocytic oped collenchyma developed cluster or parenchyma 35.5, upper epidermal cells are 2–4 to diacytic exists separately times larger than the lower ones but subsessile glandular trichomes have 4-celled secre- but the upper ones are 2–4 times larger than the lower tory head. The type of non-glandular trichome is sim- ones. Both leaf surfaces are covered by the indumentum ple thick-walled with 1-4-celled. There is a subepider- built of glandular and non-glandular hairs of the same mal poorly developed collenchyma. The collenchyma type as seen on the stem. The indumentum is densest in consists of 1-2-layered cells with thickened walls. The T. montbretii subsp., sparsest in T. cavernarum.Leaves cortex between the collenchyma and the endodermis are dorsiventral. Under the upper epidermis, there is a is composed of more or less circular and rectangular palisade parenchyma formed by 2–3 layers. and 4-5-layered parenchymatic cells. The endodermis is The portion of the mesophyll occupied by palisade regular, single layered and composed of quadrate, flat- parenchyma is 60–70% in T. montbretii subsp., 60–65% tened and ovoid cells. A pericycle layer is present un- in T. montbretii subsp. montbretii, 65–73% in T. mont- der the endodermis. It is sclerenchymatic, continuous bretii subsp. pamphylicum, 60–65% in T. odontites, 50– or discontinuous and 1–4 sheets. Phloem and xylem el- 55% in T. cavernarum, 55–60% in T. antitauricum. ements can be distinguished in the vascular tissue. The There is a 2-4-layered spongy parenchyma under the cambium between xylem and phloem is inconspicous. palisade parenchyma. The mesophyll is thicker around In the whole centre of the stem, there is pith which is the central vein. There are collenchyma under the epi- filled with circular, slightly pentagon and hexagon an- dermis in this region. While the collenchyma under the gular parenchymatical medullar cells (Table 2, Fig. 1). lower epidermis is always well-developed, the one under the upper epidermis is often poorly-developed or ab- Leaf anatomy: Mesophyll anatomy: The epidermis con- sent. Vascular bundle is present in the central part of sisting of ovoid, quadrate and rectangular cells is the vein. It is encircled by vascular bundle sheet which present at both surfaces of the leaves. It is one-layered, is sometimes inconspicous. The phloem faced the upper made of cells whose external walls are thickened. T. epidermis and the xylem faced the lower epidermis are montbretii subsp. montbretii has more abundant mu- present in the vascular bundle. There is a sclerenchy- cilage cells in the epiderma as compared to the other matic tissue below the phloem. The sclerenchymatic studied taxa. There are sometimes hypodermal cells un- cells constitute a cluster or exists separately. The scle- der the epidermis especially midvein region. Both sur- renchymatic tissue is most developed in T. cavernarum face is covered by a thick cuticula layer. The upper (Table 2, Fig. 2). cuticula out of the midvein is quite thick compared to the lower cuticula. The upper cuticula layer is 10.0–25.0 Surface anatomy: The studied surface sections showed (18.2 ± 2.0) µm thickness, and the lower one 5.0–11.0 that the leaves are hypostomatic in T. montbretii, T. (8.0 ± 1.8) µm. The size of epidermal cells in the tran- odontites, T. antitauricum, but amphistomatic in T. section are not significantly different among the taxa, cavernarum. The stomata are anomocytic to diacytic. 666 M. Dinç &S.Dogu˘

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 108.0 (72.86 ± 8.56) µm wide, the lower ones are epidermis on the both surface. Namely, they are more 64.0–133.0 (95.14 ± 17.68) µm long and 12.0–41.0 or less xeromorphic type. The upper epidermal cells in (26.46 ± 7.97) µm wide. On the lower surface, the num- surface sections are 67.0–170.0 (114.0 ± 15.0) µmlong ber of stomata is 11 ± 3 per unit area, and that of the and 39.0–108.0 (71.0 ± 12.0) µm wide, the lower ones epidermal cells is 45 ± 5 per unit area. The stomatal are 44.0–133.0 (81.0 ± 16.0) µm long and 11.0–76.0 index for the lower surface is 19.6 (Table 2, Figs 3, 4). (42.0 ± 14.0) µm wide. The anticlinal wall of the upper In T. montbretii subsp. montbretii, the upper epi- epidermal cell is thicker than of the lower epidermal dermal cells are 91.0–156.0 (110.91 ± 10.56) µmlong cell. The lower epidermal cell walls are more undulated and 51.0–71.0 (61.38 ± 3.80) µm wide, the lower ones than the upper ones (Table 2, Figs 3, 4). are 61.0–103.0 (81.93 ± 15.61) µm long and 19.0–60.0 In T. montbretii subsp., the upper epidermal cells (34.14 ± 9.56) µm wide. On the lower surface, the num- are 88.0–170.0 (121.82 ± 13.46) µm long and 44.0– ber of stomata is 15 ± 4 per unit area, and that of the Anatomical and morphological studies on Teucrium sec. Isotriodon in Turkey 667

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 wide. On the lower surface, the number of stomata is index for the lower surface is 21.4 (Table 2, Figs 3, 4). 15 ± 5 per unit area, and that of the epidermal cells is In T. montbretii subsp. pamphylicum, the upper 68 ± 8 per unit area. The stomatal index for the lower epidermal cells are 89.0–160.0 (120.21 ± 14.34) µmlong surface is 19.27 (Table 2, Figs 3, 4). and 39.0–73.0 (52.31 ± 12.65) µm wide, the lower ones In T. cavernarum, the upper epidermal cells are 61.0–103.0 (80.56 ± 11.02) µm long and 11.0–31.0 are 67.0–159.0 (109.61 ± 13.77) µm long and 41.0– (21.35 ± 5.87) µm wide. On the lower surface, the num- 87.0 (61.38 ± 6.89) µm wide, the lower ones are ber of stomata is 18 ± 4 per unit area, and that of the 84.0–141.0 (116.73 ± 16.32) µm long and 13.0–76.0 epidermal cells is 80 ± 6 per unit area. The stomatal (28.74 ± 3.02) µm wide. On the upper surface, the num- index for the lower surface is 18.3 (Table 2, Figs 3, 4). ber of stomata is 2 ± 1 per unit area, and that of the In T. odontites, the upper epidermal cells are epidermal cells is 36 ± 4 per unit area. The stomatal 85.0–109.0 (97.87 ± 4.75) µm long and 55.0–96.0 index for the upper surface is 5.26. On the lower sur- (70.45 ± 9.21) µm wide, the lower ones are 74.0–101.0 face, the number of stomata is 14 ± 4 per unit area, and (84.50 ± 6.37) µm long and 18.0–48.0 (31.32 ± 7.99) µm that of the epidermal cells is 52 ± 8 per unit area. The 668 M. Dinc¸ &S.Dogu˘

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, (56.69 ± 9.21) µm wide, the lower ones are 44.0–121.0 Figs 3, 4). (83.24 ± 21.23) µm long and 20.0–55.0 (31.49 ± 4.33) In T. antitauricum, the upper epidermal cells are µm wide. On the lower surface, the number of stomata 97.0–135.0 (111.48 ± 7.88) µm long and 44.0–84.0 is 32 ± 5 per unit area, and that of the epidermal cells is Anatomical and morphological studies on Teucrium sec. Isotriodon in Turkey 669

Table 3. Micromorphological nutlet characters of the studied taxa.

Nutlet length Nutlet width Nutlet colour and shape Nutlet hairs (mm) (mm)

T. montbretii subsp. 1.11–1.56 0.60–1.00 Light-brown, narrowly ovate-oblong, Densely non-glandular (1.39±0.12) (0.77±0.11) with inconspicuous alveoles and subsessile glandular T. montbreti subsp. 1.20–1.39 0.61–0.93 Blackish-brown, ovate-oblong, with Subsessile glandular montbretii (1.29±0.08) (0.75±0.10) regular, deep and thin-walled alveoles T. montbretii subsp. 0.95–1.42 0.65–0.84 Blackish-brown, ovate-elliptic, with Subsessile glandular pamphylicum (1.20±0.13) (0.73±0.07) irregular, shallow and thick-walled alveoles T. odontites 1.10–1.36 0.60–0.85 Blackish-brown, ovate-oblong, with Subsessile glandular (1.27±0.08) (0.70±0.08) inconspicuous alveoles T. cavernarum 0.81–1.45 0.51–0.75 Light-brown, narrowly ovate-oblong, Densely non-glandular (1.11±0.16) (0.63±0.10) with inconspicuous alveoles and subsessile glandular T. antitauricum 1.09–1.43 0.72–0.84 Light-brown, ovate-oblong, with regu- Subsessile glandular (1.27±0.11) (0.76±0.05) lar, deep and thin-walled alveoles

58 ± 8 per unit area. The stomatal index for the lower 5–15 cm long, much-branched; Leaves petiolate with surface is 35.5 (Table 2, Figs 3, 4). 2–8 mm petiole, densely tomentose-villous with simple short and long clavate glandular, subsessile glandular Nutlet morphology: The size of nutlets ranges from 1.10 and simple thick-walled non-glandular trichomes; lam- to 1.40 mm in length and 0.65 to 0.75 mm in width. ina 5–20 × 5–18 mm, ovate to ovate-lanceolate, broad- The nutlet colour is light-brown in T. montbretii subsp., est below the middle, slightly acute at apex, lower ones T. antitauricum and T. cavernarum, blackish-brown in crenate with 4–9 teeth each side, subcordate to truncate T. montbreti subsp. montbretii, T. montbretii subsp. at base, the upper ones indistinctly crenate to entire, pamphylicum,andT. odontites. The nutlet shapes are cuneate at base; nerves fine, clearly distinct below; In- narrowly ovate-oblong, ovate-oblong or ovate-elliptic. florescence terminal and lateral raceme, dense, densely The nutlet surface is generally alveolate, but alveoles tomentose-villous with simple short and long clavate are inconspicuous in T. montbretii subsp., T. odontites glandular, subsessile glandular and simple thick-walled and T. cavernarum. It is conspicuous in T. montbreti non-glandular trichomes, nearly capitate, ovate or pyra- subsp. montbretii, T. montbretii subsp. pamphylicum midal, not more than 3 cm in length. Verticillasters and T. antitauricum, but has some specific differences biflowered, many, densely congested. Bracts linear to in depth and shape of alveoles among these taxa. They oblanceolate, acute at apex, attenuate at base, 5–7 mm, are regular, deep and thin-walled in T. montbreti subsp. overtopping the calyx above the raceme, equalling be- montbretii, irregular, shallow and thick-walled in T. low. Pedicels about 1–4 mm. Calyx 4.0–5.0 mm, clearly montbretii subsp. pamphylicum and regular, deep and bilabiate, gibbous at base, tube 2.5–3.5 mm, teeth dis- thin-walled in T. antitauricum. Subsessile glandular tri- tinctly acute, equal or shorter than the tube, upper 3 chomes are present in the studied taxa, but eglandular teeth ovate, 2.4–2.6 mm long, the lower two teeth linear trichomes are present only in T. montbretii subsp. and to linear-lanceolate, 2.6–3.0 mm, slightly longer than T. cavernarum (Table 3, Fig. 5). the upper ones. Corolla longer than calyx, pinkish to creamy-white, pale. Taxonomic Treatment Phenology: Flowering July, fruiting late July-August. Distribution: South Anatolia. Endemic, Medit. element. The present study showed that T. montbretii subsp. Ecology: Teucrium montbretii subsp. yildirimlii grows has enough differences from the other subspecies to be on the rocky slopes in conifer forest clearings faced to treated as separate subspecies. stream banks an altitude of more than 750 m. Teucrium montbretii subsp. yildirimlii M. Din¸c&S. Conservation status: Teucrium montbretii subsp. yil- Do˘gu subsp. nov. (Fig. 6A) dirimlii is stenoendemic subspecies restricted to the Type: Turkey. C6 Adana: Feke, Feke-Süphandere arası, presently known locality. Its estimated area of occu- Süphandere’ye 5 km kala, Sencan Dere kenarı, kayalık pancy is less than 1 km2 (criterion B). The popula- yama¸clar, 750–850 m, 6 Jul 2010, M.Din¸c 3310 & S. tion includes less than 250 indivuduals (criterion C). Do˘gu (holotype: KNYA, isoypes: GAZI, HUB) The area is near to road (criterion D). Therefore, the Diagnosis: subsp. montbretii similis sed plantis cinereo new subspecies should be classified as “Critically en- tomentosis-villosis (non viridis), caulis semper ramosus dangered (CR)” based on the criteria of the IUCN Red (non raro), nuculis cum pilis eglandulosis (non sine) List Categories (IUCN 2001). differt. Description:Dwarf,fragile,suffruticose,denselygrey- Identification Key ish tomentose-villous and viscid, covered with glandular and non-glandular trichomes all over. Flowering stems General appearances of the subspecies of Teucrium 670 M. Dinç &S.Dogu˘

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

Discussion

The morphological polymorphism in the genus Teu- crium is reflected to the anatomical characters. There- fore, the anatomical characters show differences between the closed species in the same section (Din¸c& Ozt¨¨ urk 2008; Din¸cet al. 2008; Din¸c et al. 2009b). More- over, they were used as additional diagnastic characters in reinstatement of Teucrium andrusi Post from its sib- ling species T. paederotoides Boiss. & Hausskn. in Teu- crium 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 (Lakušić 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. Fig. 6. General appearances of T. montbretii subspecies in The anatomical features of the stems in both sections Turkey. A – T. montbretii subsp. yildirimlii;B–T. montbretii support their relationships in terms of stem morphol- subsp. montbretii;C–T. montbretii subsp. pamphylicum. ogy. The absence or presence and types of trichomes on the nutlets and the vegetative parts have taxonomic montbretii in Turkey show some differences (Fig. 6). In value in clarification of sectional boundaries in Teu- the light of the data from Ekim (1982) and the present crium (Navarro & El Oualidi 2000). In addition, in de- study, the identification key to T. montbretii subspecies scription of Teucrium pseudaroanium as a new species in Turkey can be constructed as follows; fom sect. Teucrium, trichome type and density were 1 Inflorescence lax, 5–20 cm ...subsp. pamphylicum used as additional diagnostic character from its close – Inflorescence dense, less than 5 cm ...... 2 relatives (Parolly & Eren 2007). With the non glandular 2 Plant greyish tomentose-villous; stem always branch- trichomes on its nutlets, T. montbretii subsp. yildirimlii ed; flowers generally white, rarely pinkish, nutlets is readily distinguished from the other subspecies of T. with densely eglandular trichomes ...... montbretii. Anatomical and morphological studies on Teucrium sec. Isotriodon in Turkey 671

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