132 ACTA BOT. CROAT. 76 (2), 2017 Acta Bot. Croat. 76 (2), 132–137, 2017 CODEN: ABCRA 25 DOI: 10.1515/botcro-2017-0010 ISSN 0365-0588 eISSN 1847-8476 Morphological, anatomical and karyological investigations of the Turkish endemic species Lathyrus woronowii Bornm. (Fabaceae) Fatma Güneş1, Çiler Meriç2 1Department of Pharmaceutical Botany, Faculty of Pharmacy, Trakya University, Edirne 22100, Turkey. 2Department of Biology, Faculty of Sciences, Trakya University, Edirne 22100, Turkey Abstract – Lathyrus woronowii Bornm., an endemic species of Turkey, is threatened with extinction due to dam construction. It exists only in the Çoruh valley, Artvin. This annual species is in the critically endangered (CR) category according to the International Union of Conservation of Nature (IUCN) criteria. Its morphol- ogy, anatomy and karyology are studied here for the first time. A detailed description is given and the general appearance of the species has been drawn; cross sections from the stem and leaf have been taken and exam- ined; and the diploid chromosome number (2n = 14) has been reported and illustrated for the first time. Key words: anatomy, endemic, karyology, Lathyrus, morphology, Turkey. * Corresponding author, e-mail: drgunes@gmail.com Introduction The genus Lathyrus L. (Fabaceae) is represented by 13 sections and about 187 subspecies worldwide (Allkin et al. 1983), most of which are annual or perennial plants, pre- dominantly centred in the Mediterranean region (Kupicha 1983). In Turkey, Lathyrus is divided into 10 sections with 76 species, 26 of which are endemic (Davis 1970, Davis et al. 1988, Güneş and Özhatay 2000, Genç and Şahin 2011, Güneş and Çırpıcı 2012, Güneş 2014). Lathyrus woronowii Bornm. belongs to the section Oro- bastrum, which contains eight species, two of which (L. woronowii and L. tauricola P. H. Davis) are endemic to Turkey. The main characteristics of this section are that they are annual; are procumbent to climbing; have angled or sometimes very narrowly winged stems; have tendrilous or aristate leaves with single to several-paired, linear to el- liptic, parallel-veined leaflets; have peduncles that are sin- gle-flowered; and have legumes that are glabrous or pubes- cent, with the upper suture not winged. Only L. saxatilis (Vent.) Vis., L. vinealis Boiss. & Noё and L. woronowii have more than one pair of 2 or 3 leaflets, whereas all the other species in the section have only one pair. Leaves of L. saxatilis and L. woronowii are mucronate and shortly aris- tate, respectively, whereas all other species have leaves with simple tendrils, at least on the upper parts of the plant. The flower colour is creamy to yellow in L. saxatilis and L. woronowii; red in L. vinealis, L. sphaericus Retz. and L. setifolius L.; and lavender in L. inconspicuus L. and L. tau- ricola. Pods are hairy in L. inconspicuus and L. setifolius but glabrous in other Lathyrus species (Davis 1970). Pollen morphology and seed morphology of the Oro- bastrum section have been investigated by Chernoff et al. (1992), Pastor-Cavada et al. (2011), Güneş (2011a, 2012, 2013) and Güneş and Çırpıcı (2011). It was previously re- ported that the pollen type is 3-zonocolporate and pollen shape is prolate in L. woronowii and that the shape is subp- rolate or spheroidal in eight other studied taxa. The pollen structure of all studied taxa has a tectate infrastructure and reticulate ornamentation; the seed shape is spheroidal in L. woronowii and subprolate in L. vinealis and L. sphaericus; and the seed surface is mostly smooth. Some karyology studies were previously conducted within this genus, including the studies of Senn (1938), Ress and Hazarika (1969), Fouzdar and Tandon (1975), Yamamoto and Fujiwara (1984), Kuriyan and Narayan (1988), Kumar and Sinha (1989), Kakoli and Sumitra (1991), Şahin and Babaç (1995), Şahin et al. (2000), Klamt and Schifino-Wittmann (2000), Seijo and Fernandez (2003), Ayaz and Ertekin (2008), and Güneş (2011b). These studies indicated that the genus Lathyrus generally has a diploid chromosome number of 2n = 14, as reported in ap- proximately 95 Lathyrus species worldwide. Tetraploid counts of 2n = 28 have been reported in L. pratensis L. (Darlington and Wylie 1955), L. venosus Sweet. (Başaran et al. 2007) and L. brachypterus (Güneş 2011b). Hexaploid counts of 2n = 42 have been reported in L. palustris L. sub- sp. palustris (Ball 1968, Güneş and Çırpıcı 2008). Several InvesTIgATIOns Of Lathyrus woronowii In TuRkey ACTA BOT. CROAT. 76 (2), 2017 133 studies were conducted on the species of the Orobastrum section in the Elâzığ area by Şahin and Altan (1990), in the Black Sea region by Özcan et al. (2006) and in the Europe- an region of Turkey by Güneş and Çırpıcı (2008). Karyo- type analysis is an important tool for distinguishing contro- versial species, especially when there are very similar morphological characters, such as those in L. spatulatus Čel. and L. elongatus (Bornm.) Sirj. (Güneş 2011b). There are several anatomical studies on Lathyrus spe- cies. Lathyrus latifolius L. was investigated for its morpho- logical, anatomical and physiology characteristics by Krstic et al. (2002). The study by Çildir (2011) of the morphology, anatomy and systematics of the 25 taxa belonging to the genus Lathyrus in central Anatolia (Turkey) has been pre- sented. In addition, there are a few studies on the anatomy of Turkish Lathyrus (Mantar et al. 2002, Mantar et al. 2003, Celep et al. 2011, Kahraman et al. 2014). Since there have been no previous detailed morphological, anatomical or karyological studies on the Turkish endemic L. woronowii, the goal of this study is to determine the morphological, anatomical and karyological properties of L. woronowii, which is under threat of extinction due to several dam con- struction projects in the Çoruh valley, and to use this infor- mation to aid in identification of the species and raise awareness about its threatened status. Materials and methods In 2009, 72 specimens of Lathyrus woronowii were col- lected from four different localities in Artvin Province of north-eastern Turkey during their flowering time between April and May. The specimens were deposited in the Her- barium of the Faculty of Pharmacy. Morphological obser- vations and biometric measurements were made on fresh material and herbarium specimens. They were examined and drawn using an Olympus SZX7 microscope. For anatomical studies, the material was stored in 70% alcohol. Transverse sections of the stems and leaves and the surface sections of leaves from fixed samples were made with a razor blade. Some sections were stained with alcian blue (Sigma) for pectic substances and others with safranin (Sigma) for lignin, using 20 μL alcian blue + 20 μL safranin in 10 mL of 25% glycerine for 4 hours. Stained and un- stained sections were mounted in glycerine–gelatin to pre- serve the preparations, according to the methods of Jensen (1962). Slides were examined using an Olympus BH2 mi- croscope fitted with a digital camera. Images were taken with a ProgRes C12 Plus digital camera. For karyological studies, germinating root tips were col- lected and pre-treated in a saturated solution of α-bromo- naphthalene for 24 hours at 4 °C and then fixed in Carnoy’s solution. The root tips were hydrolysed in 1 N HCl for 10– 20 minutes at 60 °C and stained with Feulgen solution. Semi-permanent slides were examined with a light micro- scope. Mitotic chromosomes were photographed on a Nikon E600 trinocular photomicroscope using a 100 × magnifica- tion oil immersion objective. Chromosome measurements were made using 4000 × magnified photographs of five well-spread metaphase plates. Short arm lengths (S) and long arm lengths (L) were measured, and the total length of the chromosomes (C), arm ratio (R = L/S), centromeric in- dex (I) and relative length (R) were calculated. The classifi- cation of chromosomes by centromeric indices followed the methods of Levan et al. (1964). Results The habitat (Fig. 1) is given, while Tab. 1 shows the morphological features of L. woronowii. Previously, seed and pollen properties were studied by Güneş (2012, 2013). The general appearance and flower parts have been drawn (Fig. 2). Three-paired leaflets were observed in some speci- mens of L. woronowii in this study. Although peduncles were reported as being single-flowered in the Flora of Tur- key (Davis 1970), 2-flowered peduncles have also been found here. The standard colour of L. woronowii is yellow when it is fresh, turning violet when pressed in herbarium specimens. Stems are covered with a single-layered epidermis, with a 5–6-layered parenchymatic cortex underneath. The endo- dermis is not distinct. The central cylinder contains four units of collateral vascular bundles. The bundles have a scle- renchyma cap above the phloem. The vascular cambium is not distinct between the phloem and xylem. The pith is filled with parenchymatous cells (Fig. 3a). The leaves are covered with a single layer of epidermis. The mesophyll is bifacial and differentiated as palisade and spongy parenchyma. Un- Fig. 1. Habit and general view of Lathyrus woronowii, (a) flower, and (b) pod (Güneş 2190). Güneş F., Meriç ç 134 ACTA BOT. CROAT. 76 (2), 2017 Tab. 1. Morphological features of Lathyrus woronowii. P – polar axis, E – eguatorial diameter, M – median point, m – median region, sm – submedian region, st – subterminal region. Features P. H. Davis 1970 Our results Habit low, slender annual and glabrous low, slender annual and glabrous Stem stems are ascending or procumbent stems are ascending or procumbent Stem length (cm) 8–18 8–57; some stems drying blackish Stem width (mm) not winged 0.5–1.5; angled Leaves apex (mm) shortly aristate shortly aristate, 1–2 Leaflets, length and width (mm) ovate-elliptic, the lowest often with 1-paired leaflets the remainder 2-paired, c. 12 × 5 ovate-elliptic, the lowest with 1-paired leaflets, the remainder 2 or 3-paired, 3-12 × 1–6 Vein type unspecified 5–7 parallel-veined Petiol length (mm) unspecified 3–15 Stipules, length × width (mm) lanceolate-subulate, semi-sagittate lanceolate-subulate, semi-sagittate, 1–5 × 0.1 Peduncles (mm) 1-flowered, shorter than leaf 1 (–2) flowered, 4–15 Pedicel (mm) unspecified 3–4 Calyx (mm) c. 5 mm, glabrous 5–6 (–7) glabrous Calyx tube (mm) unspecified 2–2.5 (–3) Calyx teeth (mm) subequal, lanceolate, a little longer than the tube subequal, lanceolate, upper teeth 3–4, the lowest 4–5 Flowers length (mm) unspecified 8–14 Standard colour and length (mm) standard violet standard yellow with red veins, drying violet or purple, 9–14 × 8–12 Wings (mm) wings cream yellow, 11–13 × 3–5 Keel (mm) unspecified cream, 11–14 × 4–6 Style (mm) unspecified 5–6 × c.1 Stigma (mm) unspecified hairy, enlarged Legume, length × width (mm) 3.5 × 0.6 cm Altan (2001) glabrous, reticulate nerved, upper sture not winged, 18–28 × (3–)4–4.5 (–5), with 6–10 seeds Seeds, length × width [min. value (Mean) max. value (mm)], color and surface structure unspecified spheroidal (P/E = 1.04), smooth, pressed, 1.71 (2.21) 2.66 × 1.69 (2.13) 2.58, yellow-green tones and speckled (Güneş 2013) Hilum length × width [min. value (Mean) max. value (mm)] unspecified 0.32 (0.44) 0.55 × 0.18 (0.24) 0.33 (Güneş 2013) Pollen shape length × width (µm) unspecified prolate (P/E = 1.36), P × E = 35.78 x 26.31 (Güneş 2012) Chromosome number and type unspecified 2n = 14; m, m, sm-st, M, sm, m-sm, msat Flowering time May April – May Habitat schictone screes schictone screes Tab. 2. Chromosome characters of Lathyrus woronowii. MC – mean chromosome, MR – mean ratio, M – median point, m – median re- gion, sm – submedian region, st – subterminal region Chromosome pairs Total length (C, µm) Long arm (L, µm) Short arm (S, µm) Arm ratio R=L/S Centromeric index (I) Relative length (R) Chromosome type I 2.56 1.31 1.25 1.05 48.79 8.83 m II 2.25 1.38 0.88 1.57 38.88 7.76 m III 2.00 1.50 0.50 3.00 25.00 6.90 sm-st IV 2.00 1.00 1.00 1.00 50.00 6.90 M V 2.00 1.38 0.63 2.2 31.25 6.90 sm VI 2.00 1.25 0.75 1.67 37.50 6.90 m-sm VII 1.44 0.75 0.69 1.09 47.92 4.97 m Average MC=2.04 MR=1.65 InvesTIgATIOns Of Lathyrus woronowii In TuRkey ACTA BOT. CROAT. 76 (2), 2017 135 der the upper epidermis, the mesophyll contains two layers of palisade cells, which are elongated cells. Under the lower epidermis, spongy parenchyma is composed of two layers of oval-shaped cells with intercellular spaces. The vascular bundles of leaves are collateral (Fig. 3b). The leaves are am- phistomatic, and stomata are found along the upper and low- er epidermis. The stomata index is 23.38 for the upper epi- dermis and 28.02 for the lower epidermis (Figs. 3c, d). The chromosome number was determined to be 2n = 14, and the detailed characteristics of chromosomes are shown in Tab. 2 and Fig. 4. Discussion Lathyrus woronowii is a critically endangered endemic species [CR C2a(i)] (IUCN 2001), restricted to the prov- ince of Artvin (A8 Çoruh valley) in north-eastern Turkey (Ekim et al. 2013). It is faced with the threat of flooding due to 15 dams having been planned along the Çoruh river. Its distribution area is approximately 30 km in diameter. Speci- mens of living plants and seeds are preserved in the Neza- hat Gökyiğit Botanical Garden (NGBB) in İstanbul. Altan (2001) reported L. woronowii stem length (60 cm) and pod size (3.5 × 0.6 cm). Our results supported the stem length (8–57 cm; Tab. 1) but not the pod size (18–28 × (3–) 4–4.5 (–5) mm). The anatomical features of L. woronowii are reported for the first time in this study. Metcalfe and Chalk (1950) sug- gested that the presence or absence and structure of the Fig. 2. Lathyrus woronowii schematic view: A) general habit, B) flower, C) calyx, D) corolla (a – standard, b – wing, c – keel), E) androecium, F) gynoecium and G) fruit. Author: F. Güneş (Güneş 2190, Güneş 2244). Fig. 3. Anatomical sections of Lathyrus woronowii. a) cross-sec- tion of stem; b) cross-section of leaf; c) upper surface section of leaf; d) lower surface section of leaf. (co – cortex, ep – epidermis, le – lower epidermis, ph – phloem, pi – pith, pp – palisade paren- chyma, sc – sclerenchyma, sp – spongy parenchyma, st – stomata, ue – upper epidermis, xy – xylem. Table 3. Chromosome number and chromosome type of Orobastrum. M – median point, m – median region, sm – submedian region, st – subterminal region Taxa Chromosome number (2n) Chromosome types Total length of largest and smallest chromosome (µm) Reference L. saxatilis (Vent.) Vis. 14 sm,stsat,sm,sm,st,st,M 4.89 and 2.86 (Şahin and Altan 1990) L. vinealis Boiss. & Noe 14 msat,sm,sm,sm,sm,m,M 4.66 and 2.66 (Şahin and Altan 1990) L. sphaericus Retz. 14 m,m,m,sm,m,m,m m,msat,sm,m,m,m,sm M,m,sm,m,sm,m,M 6.92 and 4.68 3.96 and 2.72 4.50 and 2.83 (Genç and Şahin 2001) (Özcan et al. 2006) (Güneş and Çırpıcı 2008) L. inconspicuus L. 14 Msat,sm,sm,m,sm,sm,m msat,m,sm,m,sm,m,sm 4.44 and 2.64 4.66 and 3.28 (Şahin and Altan 1990) (Genç and Şahin 2001) L. tauricola P. H. Davis 14 msat, sm,m,sm,sm,m,m 6.06 and 3.98 (Genç and Şahin 2001) L. setifolius L. 14 msat,sm,st,sm,sm,m,M msat,sm,sm,sm,sm,m,m msat,sm,st,sm,sm,m,M 6.87 and 2.66 7.16 and 3.46 5.25 and 3.00 (Şahin and Altan 1990) (Genç and Şahin 2001) (Güneş and Çırpıcı 2008) Güneş F., Meriç ç 136 ACTA BOT. CROAT. 76 (2), 2017 wings in the stem are taxonomically important in the dis- tinction of Lathyrus species. The stem of L. woronowii has no wings like those of L. cilicicus Hayek & Siehe (Celep et al. 2011); L. inconspicuus L.; L. vinealis (Mantar et al. 2002); and L. nissolia L. (Kahraman et al. 2014). Stems of L. maritimus Bigel, L. pratensis, L. sylvestris L. (Metcalfe and Chalk 1950), L. latifolius L. (Krstic et al. 2002), L. hir- sutus L. and L. sativus L. (Mantar et al. 2003) contain wings. Çildir (2011) reported that L. incurvus (Roth.) Willd., L. cas- sius Boiss., L. cicera L. and L. chloranthus Boiss. have winged stems, while stems of L. aureus (Steven) Brandza, L. brachypterus, L. haussknechtii Širj., L. armenus (Boiss & Huet) Širj., L. digitatus (M. Bieb.) Fiori, L. tukhtensis Czec- zott, L. spathulatus Čelak., L. pratensis, L. laxiflorus (Desf.) O. Kuntze subsp. laxiflorus, L. czeczottianus Bässler, L. roseus Steven subsp. roseus, L. tuberosus L., L. saxatilis, L. sphaericus, L. aphaca L. var. biflorus Post and L. aphaca L. var. affinis (Guss.) Arc. contain no wings. The pith is composed of large parenchyma cells in L. woronowii., L. aureus, L. armenus, L. laxiflorus subsp. laxi- florus, L. roseus subsp. roseus, L. cilicicus, L. maritimus and L. pratensis, while L. latifolius stems are hollow at the centre. However, the stems of L. incurvus, L. brachypterus, L. haussknechtii, L. digitatus, L. tukhtensis, L. spathulatus, L. pratensis, L. czeczottianus, L. tuberosus, L. cassius, L. hirsutus, L. chloranthus, L. sativus, L. cicera, L. saxatilis, L. inconspicuus var. inconspicuus, L. sphaericus, L. nisso- lia, L. aphaca var. biflorus and L. aphaca var. affinis are hollow (Metcalfe and Chalk 1950, Krstic et al. 2002, Man- tar et al. 2002, 2003, Celep et al. 2011, Çildir 2011, Kahra- man et al. 2014). L. woronowii has amphistomatic leaflets, like those of L. incurvus, L. pratensis, L. czeczottianus, L. sphaericus, L. cassius, L. cicera, L. latifolius and L. cilici- cus, whereas L. roseus subsp. roseus and L. laxiflorus sub- sp. laxiflorus have hypostomatic leaves (Krstic et al. 2002, Celep et al. 2011, Çildir 2011). The chromosome measurements of Lathyrus species be- longing to section Orobastrum are shown in Tab. 3. The largest and smallest chromosomes of L. woronowii were measured as 2.56 and 1.44 µm, respectively. The total lengths of the largest and smallest chromosomes of other species belonging to the Orobastrum section were stated by the authors. In this section, L. woronowii has the smallest chromosome set. The largest set belongs to L. setifolius, and the chromosomal numbers of Lathyrus species in the present study (Tab. 3) agree with previous records (Şahin and Altan 1990, Genç and Şahin 2001, Özcan et al. 2006, Güneş and Çırpıcı 2008). 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