Biologia, Bratislava, 62/2: 148—156, 2007 Section DOI: 10.2478/s11756-007-0023-6

Trichome micromorphology in () growing in Croatia

Renata JurišicGrubeši´ c´*, Sanda Vladimir-Kneževic´,DarioKremer, Zdenka Kalođera &JadrankaVukovic´

Faculty of Pharmacy and Biochemistry, University of Zagreb, Ante Kovači´ca 1,HR–10000 Zagreb, Croatia

Abstract: Micromorphological investigation of the types, dimensions and distribution of characteristic trichomes in and stems in Teucrium L. species (T. arduini L., T. chamaedrys L., T. flavum L., T. montanum L., T. polium L., and T. scordium L. subsp. scordioides Schreb.) distributed in Croatia was carried out as part of the taxonomical study of the genus Teucrium. Secretory types of , peltate and capitate hairs were observed on the of stems and leaves of all investigated species. Non-secretory, acicular hairs were almost completely lacking on stems of T. scordium subsp. scordioides. Flagelliform hairs were not found in T. flavum and T. polium. Cladose hairs were present only in T. polium.Thelargest micromorphological variability was established between wild and cultivated samples of T. arduini and T. scordium subsp. scordioides, while cultivated and wild specimens of T. polium were almost identical. Differences were primarily observed in trichome dimensions and much less in micromorphological features. Key words: Teucrium L.; micromorphology; hairs

Introduction with Pycnobotrys, Scorodonia with Stachyobotrys,and Scordium with Spinularia.K¨astner (1989) later im- Teucrium species have been used for centuries in folk proved his own proposal by creating subsections that medicine as a cholagoga, as well as antispasmodic, di- include groups formerly considered at the section level uretic, antidiabetic, antiphlogistic, antirheumatic, anti- (for instance, subsect. Polium and Chamaedrys within septic, anthelmintic, carminative and flavoring agents sect. Chamaedrys). Other important research on the (Gharabeih et al. 1988, 1989). From a taxonomical per- of the genus and species belonging to dif- spective, the genus Teucrium has been divided into sev- ferent sections were published by Puech (1978), Valdés eral sections, identifiable through calyx shape and in- Bermejo & Sanchez Crespo (1978), K¨astner (1979, florescence structure. The most important initial works 1981, 1985, 1986), Manzanares et al. (1983), Bini Maleci on this topic were by Schreber (1774), Bentham (1835) & Servettaz (1991) and Servettaz et al. (1992) in order and Boissier (1879, according to Bini Maleci & Servet- to obtain better insight into this complex genus. taz 1991). Schreber (1774) evaluated variation on the Plant hairs (trichomes) are of great interest to de- basis of width and dentate leaf tips and described scriptive and experimental botanists and data on these five taxa at the variety level. In further studies, these and indumenta are routinely included in many types characters were found to show great variation and are of of studies. As simple morphological tools, trichomes virtually no taxonomical significance (Mártonfi 1995). are useful due to the ease with which they are ex- These studies were resumed by McClintock & Epling amined and their almost universal occurrence, partic- (1946) for the flora of the New World and by Tutin & ularly among the and flowering . Beyond (1972) in Europaea. These authors usually their purely descriptive use, comparative data may be subdivided the genus Teucrium into ten sections: Teu- important for the study of evolution and relationships, cropsis Benth., Teucrium Benth., Chamaedrys (Mill.) and for the roles of hairs in various aspects of physio- Schreb., Polium (Mill.) Schreb., Isotriodon Boiss., Py- logical and ecological adaptation (Payne 1978). Many cnobotrys Benth., Scorodonia (Hill) Schreb., Stachy- authors, such as Behnke (1984), Güemes et al. (1992), obotrys Benth., Scordium (Mill.) Benth., and Spin- Servettaz et al. (1994), Bini Maleci et al. (1995) and ularia Boiss. Based on a comparative study of in- Mráz (1998) emphasize the great value of trichomes florescences and flowers, K¨astner (1978) made a re- in modern taxonomy. According to Mráz (1998), tri- markable modification by proposing only six sections chome micromorphology in the genus Teucrium has by uniting sect. Chamaedrys with Polium, Isotriodon been studied mainly in the sections Chamaedrys and

* Corresponding author e-mail: [email protected]

c 2007 Institute of Botany, Slovak Academy of Sciences Trichome micromorphology in Teucrium species 149

Fig. 1. Croatian localities of the collect Teucrium L. species: Krk island (• Omišalj: T. flavum; Garica: T. polium;  Baška Draga: T. scordium subsp. scordioides), pass Gornje Jelenje (T. chamaedrys and T. montanum), and Velebit mountain, near Starigrad Paklenica (T. arduini).

Scorodonia. Works on these topics are from Grzybek ies and for the roles of hairs in ecological adapta- (1965, 1967), Antunes & Sevinato-Pinto (1991), Bini tion. Maleci & Servettaz (1991), Bini Maleci et al. (1992), Servettaz et al. (1992), Servettaz et al. (1994) and Bini Material and methods Maleci et al. (1995). Puech (1984) and Navarro (1995) Material evaluated species in sect. Polium from the Mediter- Randomly selected samples of wild plants were collected at ranean area on the basis of characters of clothing hairs. several locations in Croatia (Fig. 1) in August 2001 (Ta- The structure and development of the glandular tri- ble 1). Among the studied species, T. arduini,whichisen- chomes of T. montanum (sect. Polium) were researched demic to Croatia, Montenegro and northern Albania (Kušan in detail by Mráz (1998). The same author did not find 1969) had the narrowest natural distribution area. The re- qualitative differentiation of glandular trichomes (pres- maining investigated species have larger natural distribu- ence or absence of distinct types on various plant parts) tion areas, as described by Tutin & Wood (1972). The cul- among three subspecies of T. montanum (T. montanum tivated samples were derived from of the same wild populations and cultivated under the same conditions in the subsp. montanum, T. montanum subsp. pannonicum ‘Fran Kušan’ Pharmaceutical Botanical Garden, Faculty of (A. Kern) Domin, and T. montanum subsp. jaileae Pharmacy and Biochemistry, University of Zagreb, Croatia. (Juz.) Soó.). Voucher specimens (No. 9801–9812) are deposited in the The current study was based on comparative re- of the Department of Pharmacognosy, Faculty search of different types, dimensions and distribution of Pharmacy and Biochemistry, Zagreb, Croatia. of characteristic leaf and stem trichomes of wild and Light microscopy cultivated specimens of Teucrium taxa growing in Croa- Ten wild plants of each species were taken and two speci- tia (T. arduini L., T. chamaedrys L., T. flavum L., mens of leaves as well as of stems, were prepared from each T. montanum L., T. polium L., and T. scordium L. wild plant (number of independent analyses, n = 20). The subsp. scordioides Schreb.). The objective of compar- same procedure was also performed for cultivated samples ing wild and cultivated samples was to study the de- of each plant species. Leaves and stems were soaked in wa- pendence of trichome development on ecological con- ter to soften the cuticle and facilitate further treatment and ditions. This is connected with the fact that cer- then subjected to micromorphological analysis. Cross sec- tain differences among taxa can be determined only tions were prepared for microscopy by placing them into a drop of glycerol and gently heating. A light microscope by cultivation in the same environment. The culti- with a built in camera was used in the work. Characteris- vation of different taxa in the same habitat is com- tic trichomes of Teucrium species were photographed and monly in use in many genetic fields, such as forest measured. Determination of trichomes was carried out ac- genetics (Wright 1976) and agriculture. Such com- cording to Uphof (1962), Metcalfe & Chalk (1972), Payne parative data may be important for taxonomic stud- (1978), and Bini Maleci & Servettaz (1991). 150 R. Grubešic´ et al.

Table 1. Studied material of Teucrium L. species.

Section Species Origin and collection data Origin and collection data (wild plants) (cultivated plants)

sect. Chamaedrys (Mill.) Schreb. T. chamaedrys L. pass Gornje Jelenje, 800 m a.s.l.1, coll. ‘F.K.’ P.B.G.3, August 2001 Juriši´cGrubeši´candBrki´c, August 2001 T. flavum L. Omišalj (Krk island), at s.l.2, coll. Juriši´c ‘F.K.’ P.B.G., August 2001 Grubeši´candBrki´c, August 2001

sect. Polium (Mill.) Schreb. T. montanum L. pass Gornje Jelenje, 800 m a.s.l., coll. ‘F.K.’ P.B.G., August 2001 Juriši´cGrubeši´candBrki´c, August 2001 T. polium L. Garica (Krk island), 30 m a.s.l., coll. Ju- ‘F.K.’ P.B.G., August 2001 riši´cGrubeši´candBrki´c, August 2001

sect. Stachyobotrys Benth. T. arduini L. Starigrad Paklenica (Velebit mountain), ‘F.K.’ P.B.G., August 2001 600 m a.s.l., coll. Juriši´cGrubeši´cand Brki´c, August 2001

sect. Scordium (Mill.) Benth. T. scordium L. Baška Draga (Krk island), at s.l., coll. ‘F.K.’ P.B.G., August 2001 subsp. scordioides Juriši´cGrubeši´candBrki´c, August 2001 Schreb.

1 a.s.l. – above sea level; 2s.l. – sea level; 3 ‘F.K.’ P.B.G. – ‘Fran Kušan’ Pharmaceutical Botanical Garden, Faculty of Pharmacy and Biochemistry, University of Zagreb, Croatia.

Statistics Differences between wild and cultivated samples were sta- tistically treated with the use of the Student’s t-test (Miller & Miller 2000).

Results

Types of trichomes Micromorphological investigation of Teucrium species growing in Croatia showed the presence of different types on leaves and stems of the examined wild and cultivated specimens (Table 2). Five different hair types were observed on leaves and stems: Atype:Peltate hairs with one basal epidermal , one neck cell and four to eight secretory cells (Fig. 2/A and Plate 1/1–2). Btype:Capitate hairs with one- to five-celled stalk and one to four-celled head (Fig. 2/B and Plate 1/3–5). Ctype:Acicular hairs, non-secretory multicellular clothing trichomes, unbranched, uniseriate with one to six cells arranged in a single row (Fig. 2/C and Plate 1/6–7). Dtype:Flagelliform hairs, multicellular, uniseri- ate trichomes with distal end of the terminal cells deli- cate and greatly elongated (Fig. 2/D and Plate 1/8). Etype:Cladose hairs, multicellular, branched (Fig. 2/E and Plate 1/9). Fig. 2. Schemes of hair types in the studied Teucrium L. species. Peltate hairs (A type); Capitate hairs (B type); Acicular hairs (C Distribution of trichomes type); Flagelliform hairs (D type); Cladose hairs (E type). Peltate (type A) and capitate (type B) hairs were ob- served on the epidermis of stems and leaves in all stud- ied species (Table 2). Acicular hairs (type C) were lack- flagelliform hairs were not observed on the leaves, while ing only on stems of wild plants of T. scordium subsp. in T. montanum they were present only on the leaves of scordioides (there were only a few acicular hairs on the cultivated plants. In T. arduini and T. scordium subsp. stems of the cultivated specimens). Flagelliform hairs scordioides flagelliform hairs were abundant. T. polium (type D) were absent from the leaves and stems of T. differed from the other species in the presence of cladose flavum and T. polium.InthecaseofT. chamaedrys, hairs (type E) on its stems and leaves. Trichome micromorphology in Teucrium species 151

Table 2. Trichome distribution on leaves and stems in Teucrium L. species.

AB C DE Species Plant Plant Peltate hairs Capitate hairs Acicular hairs Flagelliform hairs Cladose hairs material origin (stalk: 1–5 cells) (1–6 cells) (5–9 cells) 1234 5 123456

N + +– – – – ++++–– – – leaf C + +– – – – ++++–– + – T. arduini stem N + +– – – – ++++–– + – C + ++++ – ++++ – – + –

N + +– – – – ++++–– – – leaf C + +– – – – ++++–– – – T. chamaedrys stem N + + – – – – + – + + + + + – C++––+–+–+++++ –

N + ++ – – – ++++ – – – – leaf C + +– – – – ++++–– – – T. flavum stem N + + – – – – +++++ – – – C + + – – – – – ++++ – – –

N + ++ – – – – +++ – – – – leaf C + +++ – – – +++ – – + – T. montanum stem N + ++ – – – – +++ – – – – C + +++ – – – ++ – – – – –

N + ++ – – – +++ – – – – + leaf C + ++ – – – +++ – – – – + T. polium stem N + ++ – – – ++++ – – – + C + ++ – – – +++ – – – – +

N + +–++ – +–++–– + – leaf C + +++ – – – +++ – – + – T. scordium subsp. scordioides stem N + ++–++ –––––– + – C + +–++ – – – –+– – + –

N – native (wild) plant; C – cultivated plant; + indicates the presence of certain hair type; – indicates the absence of certain hair type.

Statistical analysis of the results based on measure- missing in the cultivated specimens (Table 4), while ments of different hair types showed significant variabil- the epidermises of leaves differed in the size of peltate ity between wild and cultivated specimens of the same trichomes (p<0.001), capitate hairs with one- or two- species (Tables 3–5). celled stalk (Table 3), as well as in one-, two- and four- The leaves of native and cultivated specimens of T. celled acicular hairs (Table 4). arduini were statistically different in the size of peltate The four-celled acicular hairs of the leaves of trichomes (p<0.02) and one- to four-celled acicular T. montanum wild plants (Table 4) were consider- hairs (Table 4). The leaves of T. arduini also differed ably shorter than those of the cultivated specimens from flagelliform hairs, which were observed only in the (p<0.001) while capitate hairs with three-celled stalks cultivated samples (Table 5). The stems of wild and cul- (Table 3) were missing on the leaf epidermis of the tivated plants of T. arduini were statistically different wild plants. Capitate hairs with three-celled stalk were in the length of peltate hairs (p<0.05), capitate hairs present only on stems of cultivated specimens of T. (Table 2), acicular hairs with one to four cells (Table 4), montanum (Table 3). The stems of cultivated samples and flagelliform hairs (Table 5). of T. montanum were not covered with four-celled aci- Wild and cultivated specimens of T. chamaedrys cular hairs, which were observed in wild samples (Ta- showed a great variability in stem epidermal features ble 4). possessing capitate hairs with one- and four-celled Wild and cultivated specimens of T. polium re- stalks, followed by one-, three-, four-, five- and six- vealed an almost negligible difference. If there was a dif- celled acicular hairs, and flagelliform hairs (Tables 3– ference, it concerned only the four-celled acicular hairs 5), while leaves were variable in terms of peltate hairs of the stem epidermis, which were lacking in the cul- (p<0.05) and one- to four-celled acicular hairs (Ta- tivated specimens (Table 4). Cladose hairs (size: 910– ble 4). 1220 µm) were found only in the case of T. polium.The Native and cultivated samples of T. flavum differed obvious gradation of these long, branched hairs was ob- in one-celled acicular hairs of the stem, which were served with significant variations in their size. 152 R. Grubešic´ et al. ; 65.16 11.12 14.66 ± ± ± scordioides – – – – – – – ∗ ∗ m) m) µ µ a 26.39 213.00 6.19 6.14 6.77 16.29 455.01 = 20); ** subsp. ± ± ± ± ± – 194.57 n – – – – 70.75 70.45 a c 2.06 6.65 98.72 6.10 270.37 2.78 9.49 8.39 ± ± ± ± ± ± ––– ––– – – 144.90 – – 05, respectively). 40.65 70.21 . 176.13 0 p< b a d d a a b 1.49 1.65 40.80 1.63 2.45 2.09 1.90 1.66 1.85 3.28 39.27 1.91 1.69 4.74 41.55 SC: Capitate hairs (stalk with 1–4 cells); length ( LC: Capitate hairs (stalk with 1–4 cells); length ( ± ± ± ± ± ± ± ± ± ± ± ± SD (number of independent analyses, 02, and . 0 ± p< ean 01, . 0 8.70 26.10 ± p< – 27.70 – 28.60 51234 51234 – 30.30 –– 30.98 34.78 – 31.69 – 30.75 – 20.23 – 24.40 – 32.63 001, . 0 ∗ ∗ p< m) m) µ µ 14.50 282.40 13.53 – 26.28 ± ± – – – – – – – – – – 205.95 a L. species. 7.76 ± – – – – – – – – Teucrium in SN – stem, native plant; SC – stem, cultivated plant; * m cultivated plant of the same species ( a c 16.99 – 211.04 14.192.96 – 10.77 – 9.42 2.86 ± ± ± ± ± ± – – 120.80 – – – – 43.74 37.65 122.57 SN: Capitate hairs (stalk with 1–5 cells); length ( LN: Capitate hairs (stalk with 1–5 cells); length ( b a d d a a b 1.23 1.68 1.62 1.62 1.65 2.10 4.23 41.06 3.26 39.87 1.18 1.61 1.26 1.12 38.70 1234 1234 ± ± ± ± ± ± ± ± ± ± ± ± 33.25 28.82 24.85 24.45 31.11 28.89 21.42 25.59 33.15 29.29 31.95 29.10 ∗∗ ∗∗ T. flavum T. montanum Table 3. Size of capitate hairs (stalk with 1–5 cells) Species T. arduini T. arduini LN – leaf, native plant; LC – leaf, cultivated plant; T. polium T. polium T. flavum a, b, c, d – significant difference between wild and T. chamaedrys T. chamaedrys T. scordium T. montanum T. scordium Trichome micromorphology in Teucrium species 153

Plate 1. Hair types in the studied Teucrium L. species (Figs 1 to 5 and 7: bar = 10 µm; Figs 6, 8 and 9: bar = 50 µm). Figs 1, 2: Peltate hairs (A type); 3–5: Capitate hairs (B type); 6, 7: Acicular hairs (C type); 8: Flagelliform hairs (D type); 9: Cladose hairs (E type).

T. scordium subsp. scordioides showed an excep- approaches applied by different authors in describing tional degree of variability. The stems of wild and cul- certain hair types. In this paper, non-secretory multicel- tivated specimens differed greatly in capitate type of lular, unbranched, uniseriate trichomes were specified trichomes with two-, three- and five-celled stalk (Ta- as acicular hairs, based on plant hair terminology by ble 3), as well as according to four-celled acicular hairs Payne (1978), while extraordinary long, whip-like hairs (Table 4). The leaves of wild and cultivated plants of were considered and described as flagelliform hairs. T. scordium subsp. scordioides were different in terms Long capitate hairs were absent from the leaves of of peltate hairs (p<0.05), capitate hairs with two-, the investigated T. flavum specimens. This fact con- three- and four-celled stalk, one- to four-celled acicular firmed the results by Bini Maleci et al. (1995). The hairs, as well as of flagelliform hairs (Tables 3–5). authors also did not find this type of trichomes in T. flavum subsp. flavum and T. flavum subsp. glaucum. Discussion Peltate hairs and non-secretory multicellular trichomes were found by Bini Maleci et al. (1995) on the leaves of The results obtained in this work, did not show visible both subspecies, though non-secretory trichomes were link between the appearance of certain hair types on the very rare in the subsp. glaucum. Based on these facts, stems and leaves of the investigated Teucrium species it could be concluded that specimens examined in this and their belonging to specific sections of the genus work actually presented T. flavum subsp. flavum. Teucrium. On the leaves of the studied T. chamaedrys,long The greatest difference between our data and the capitate hairs were not observed and this fact coinci- literature (Bini Maleci & Servettaz 1991; Bini Maleci et dences with the work of Bini Maleci & Servettaz (1991). al. 1995; Mráz 1998) for T. chamaedrys, T. flavum and Long capitate hairs were rarely found only on the stems T. montanum was the presence of flagelliform hairs in of the abovementioned species. the studied specimens. This is likely due to the varying Our results showed the greatest difference between 154 R. Grubešic´ et al.

Table 4. Size of acicular hairs (1–6 cells) in Teucrium L. species.

LN: Non-secretory hairs (1–6 cells); length (µm)* 1234 5 6

T. arduini 82.71 ± 13.05a 182.49 ± 22.16a 299.90 ± 28.14d 497.51 ± 31.61a –– T. chamaedrys 97.47 ± 14.76c 264.38 ± 28.05a 478.92 ± 35.61a 957.60 ± 44.05a –– T. flavum 73.78 ± 17.59a 195.40 ± 21.29a 328.09 ± 33.46 530.12 ± 31.68a –– T. montanum – 111.45 ± 15.65 235.00 ± 13.46 359.44 ± 31.50 – – T. polium 54.15 ± 14.18 126.14 ± 4.32 173.45 ± 12.92 – – – T. scordium** 76.16 ± 7.37 – 270.96 ± 11.32a 331.36 ± 22.66a ––

Species LC: Non-secretory hairs (1–6 cells); length (µm)* 1234 5 6

T. arduini 95.44 ± 8.03a 139.20 ± 39.83a 279.56 ± 31.90d 373.74 ± 44.66a –– T. chamaedrys 110.67 ± 18.85c 161.10 ± 27.41a 310.66 ± 42.63a 473.29 ± 65.90a –– T. flavum 117.22 ± 10.45a 136.75 ± 16.92a 321.39 ± 36.75 410.00 ± 30.82a –– T. montanum – 110.79 ± 15.59 244.62 ± 17.90 444.61 ± 21.87a –– T. polium 49.39 ± 5.26 124.36 ± 5.25 167.79 ± 7.84 – – – T. scordium** – 106.95 ± 32.15 160.05 ± 16.90a 222.57 ± 45.58a ––

SN: Non-secretory hairs (1–6 cells); length (µm)∗ 1234 5 6

T. arduini 67.24 ± 9.08a 101.15 ± 10.98a 269.35 ± 14.18a 308.81 ± 18.17a –– T. chamaedrys 138.84 ± 17.58a – 265.09 ± 15.86a 675.38 ± 54.56a 930.24 ± 71.91a 1100.55 ± 77.23a T. flavum 171.96 ± 13.44 231.46 ± 7.97 322.21 ± 24.50 480.24 ± 36.23c 679.44 ± 40.99 – T. montanum – 111.39 ± 16.26 217.60 ± 13.88c –– – T. polium 55.93 ± 4.87 107.70 ± 11.59d 121.85 ± 9.43d 282.63 ± 28.60 – – T. scordium**–––– – –

SC: Non-secretory hairs (1–6 cells); length (µm)* 1234 5 6

T. arduini 103.89 ± 8.85a 191.48 ± 10.74a 198.12 ± 28.32a 380.06 ± 14.17a –– T. chamaedrys 73.19 ± 8.37a – 210.31 ± 30.82a 372.63 ± 33.40a 592.80 ± 102.58a 803.61 ± 98.32a T. flavum – 226.58 ± 7.71 326.42 ± 28.10 453.59 ± 35.21c 667.52 ± 57.36 – T. montanum – 117.20 ± 20.24 229.31 ± 15.86c 415.95 ± 28.46 – – T. polium 53.55 ± 6.12 114.24 ± 5.71d 129.41 ± 12.92d –– – T. scordium** – – – 222.57 ± 45.58 – –

LN – leaf, native plant; LC – leaf, cultivated plant; SN – stem, native plant; SC – stem, cultivated plant; * mean ± SD (n = 20); ** subsp. scordioides; a, b, c, d – significant difference between wild and cultivated plant of the same species (p<0.001, p<0.01, p<0.02, and p<0.05, respectively). wild and cultivated specimens of T. arduini and T. hybrids, the type of calyx hair was the most valuable scordium subsp. scordioides, while the native and cul- character. They could distinguish between those speci- tivated specimens of T. polium differed inconsiderably. mens that only had short (0.05–0.6 mm) appressed eg- Since the examined cultivated plant material was ob- landular hairs (S. officinalis aggregate) and those that tained from the seeds of the same native population, had glandular and long (0.6–2 mm) subpatent eglandu- it could be presumed that the micromorphological dif- lar hairs (S. fruticosa/S. blancoana aggregate). Inter- ferences were ecologically conditioned as certain new mediate characters were found in hybrids between the characteristics appeared during cultivation. Numerous species from both groups. Aedo et al. (2005) investi- studies have been published addressing the effects of gated the Geranium L. section Dissecta (Geraniaceae) environmental influence on trichome morphology. Bini which consists of four species in Eurasia. A multivari- Maleci & Servettaz (1991) have also ascribed the pres- ate morphometric study clearly distinguished the an- ence of a large number of acicular hairs in T. marum nual G. dissectum L. The characters contributing most L. and T. subspinosum Pourr. ex Willd. to special en- to separation of the three remaining perennial species vironmental conditions, such as highly wind-exposed (G. asphodeloides Burm. f., G. sintenisii Freyn and G. places. Kofidis et al. (2003) found that the leaf stomata crenophilum Boiss.) were width and the length and non-glandular hairs increase in number in Orig- of glandular hairs on stems and pedicels. Syros et al. anum vulgare L. (Lamiaceae) from lowland to upland (2006) found that the density of non-glandular hairs habitats, whereas glandular hairs decrease in number. was higher for both leaflet surfaces in the ecotype A of Reales et al. (2004) elucidated the taxonomic relation- Ebenus creticus L. (Fabaceae) compared to ecotype C. ship between officinalis L., the type species of This fact was corroborated by physiological data that the genus, and S. fruticosa Mill. Both taxa were in- suggested that plants of ecotype A were better adapted cluded in sect. Salvia (Lamiaceae). They found that to the xerothermic environment of the island of Crete. for the differentiation of ten taxa including ssp. and Laboratory research gave similar results. For example, Trichome micromorphology in Teucrium species 155

Table 5. Size of flagelliform hairs (5–9 cells) in Teucrium L. Bentham G. 1835. Labiatarum genera et species. London, pp. species. 660–690. Bini Maleci L. & Servettaz O. 1991. Morphology and distribution Species LN: Flagelliform hairs LC: Flagelliform hairs of trichomes in Italian species of Teucrium sect. Chamaedrys length (µm)* length (µm)* (Labiatae) – a taxonomical evaluation. Pl. Syst. Evol. 174: 83–91. T. arduini – 374.01 ± 56.29 Bini Maleci L., Pinetti A. & Servettaz O. 1992. Micromorpholog- T. chamaedrys ––ical and phytochemical researches on Teucrium massiliense T. flavum ––L., pp. 349–355. In: Harley R.M. & Reynolds T. (eds), Ad- T. montanum – 760.00 ± 24.16 vances in Labiatae science, Royal Botanical Garden, Kew, T. polium ––568 pp. T. scordium** 918.38 ± 52.07a 508.60 ± 43.87a Bini Maleci L., Pinetti A. & Servettaz O. 1995. Micromorpho- logical and phytochemical characters of the two subspecies SN: Flagelliform hairs SC: Flagelliform hairs of Teucrium flavum (Labiatae) from the Italian flora. Flora length (µm)* length (µm)* 190: 237–242. Gharaibeh M.N., Elayan H.H. & Salhab A.S. 1988. Hypogly- T. arduini 734.67 ± 43.05a 552.78 ± 50.11a caemic effects of Teucrium polium. J. Ethnopharmacol. 24: T. chamaedrys 2075.46 ± 196.63a 1279.79 ± 145.58a 93–99. T. flavum ––Gharaibeh M.N., Elayan H.H. & Salhab A.S. 1989. Anorexic ef- T. montanum ––fect of Teucrium polium in rats. Int. J. Crude Drug Res. 27: T. polium ––201–210. T. scordium** 1205.39 ± 89.85 1153.33 ± 93.22 Grzybek J. 1965. Pharmaco-botanical investigation in Teucrium chamaedrys L. I. Morphology and anatomy. Acta Biol. Cra- LN – leaf, native plant; LC – leaf, cultivated plant; SN – stem, cov., Ser. Bot. 8: 147–154. native plant; SC – stem, cultivated plant; * mean ± SD (n = 20); Grzybek J. 1967. Morphological and anatomical comparison of ** subsp. scordioides; a, b, c, d – significant difference between species of the genus Teucrium L. indigenous in Poland. Acta wild and cultivated plant of the same species (p<0.001, p<0.01, Biol. Cracov., Ser. Bot. 10: 39–54. p<0.02, and p<0.05, respectively). Güemes J., Mateo G. & Crespo M.B. 1992. Importancia de los tri- comas en la taxonomía del grupo Teucrium buxifolium Schre- ber, pp. 155–159. In: Conesa J.A. & Recasens (eds), Actes del Simposi International de Botřnica Pius Font i Quer, 1988, 2, leaf stalks from tomato plants grown in the light showed Fanerogamia. larger trichome heads (Nihoul 1993). K¨astner A. 1978. Beitr¨age zur Wuchsformenanalyse und system- atischen Gliederung von Teucrium L. I. Die Infloreszenzen The differences between wild and cultivated speci- und Blüten. Flora 167: 485–514. mens of the investigated Teucrium species were appar- K¨astner A. 1979. Beitr¨age zur Wuchsformenanalyse und systema- ent mainly within the dimensions of certain hair types tischen Gliederung von Teucrium L. II. Anatomie der Sprosse and much less in their presence or absence on leaves und Bl¨ater. Flora 168: 431–467. K¨astner A. 1981. Beitr¨age zur Wuchsformenanalyse und system- or stems. This is due to the fact that the environment atischen Gliederung von Teucrium L. III. Wuchsformen und could have a much greater influence on trichome di- Verbreitung von Arten der Sektionen Teucropsis und Teu- mension than on their formation or shape. Moreover, crium.Flora171: 466–519. the characteristics associated with the shape better de- K¨astner A. 1985. Beitr¨age zur Wuchsformendifferenzierung und scribe phylogenetic and genetic relations between living systematischen Gliederung von Teucrium L. IV. Wuchsfor- men und Verbreitung von Arten der Sektionen Isotriodon. creatures (Reyment 1985). Flora 176: 73–93. K¨astner A. 1986. 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Received Aug. 25, 2005 Accepted Nov. 11, 2006