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Nutlet Micromorphology of Indian Leucas (Lamioideae: Lamiaceae) with Emphasis on Its Systematic Implication

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Nutlet Micromorphology of Indian Leucas (Lamioideae: Lamiaceae) with Emphasis on Its Systematic Implication Rheedea Vol. 28(1) 16–28 2018 ISSN: 0971-2313 https://dx.doi.org/10.22244/rheedea.2018.28.1.02 Nutlet micromorphology of Indian Leucas (Lamioideae: Lamiaceae) with emphasis on its systematic implication Mannethody Sajna and P. Sunojkumar* Department of Botany, University of Calicut, Calicut University P.O., Malappuram, Kerala – 673 635, India *E-mail: [email protected] Abstract Nutlet micromorphological characteristics in 40 taxa of Asian Leucas and two related species of African Leucas found in India were examined by light and scanning electron microscopy. Six types of basic ornamentation patterns can be distinguished: scalariform, verrucate, reticulate, ruminate, colliculate, and rugose. Of these, scalariform type was the most common and was found in all the sections of Asian Leucas, but the variation in alignment, size and shape of its composing cells provide further evidence in distinguishing the taxa. Verrucate, the second most frequent type was found in six species while other types were represented by only one to three species. However, the presence of colliculate sculpturing is more useful for separating African species from Asian Leucas. In Asian Leucas, nutlet size may be an important character rather than ornamentation for sectional division. Again, the type of glandular hairs on the nutlet apex can be used as a distinctive taxonomic character at specific level to separate L. martinicensis (Jacq.) R.Br. and L. urticifolia (Vahl) Sm. The results revealed high similarity of these taxa to the sect. Leucas. This study supports earlier observations on pericarp structure and subsequent phylogenetic analysis but is not in corroboration with the recent classification of Indian Leucas. Keywords: Asian Leucas, India, micromorphology, nutlet Introduction Leucas R.Br. comprises about 100 species important (Barthlott, 1981; Demissew & Harley, distributed mainly on dry or disturbed ground in 1992; Ryding, 1993, 1995; Oran, 1996; Guerin, 2005; tropical to southern Africa, Arabia, tropical and Moon & Hong, 2006; Kahraman et al., 2011). In subtropical parts of Asia and Australia (Harley et some genera of Lamiaceae, nutlet characters were al., 2004). Highest number of taxa is reported in potentially useful in infrageneric classification the northeast tropical Africa (Ryding, 1998), and (Demissew & Harley, 1992) as well as at specific in India (Singh, 2001). Species prefers low (±800 and subspecific levels (Marin et al., 1994; Navarro m) to high (800–2600 m) altitudes and grows in & El Oualidi, 2000; Sampath Kumar & Murthy, ecologically divergent localities such as sea coasts, 2012; Satıl et al., 2012; Krawczyk & Glowacka, paddy fields, near wet lands, road sides, open 2015). In Stachys L., one of the largest genera of habitats, farm land, plantations, grasslands, rocky Lamiaceae, it has been shown that the sculpturing hills, scrub jungles and shola grasslands. pattern of nutlet surface is more useful to separate Surface micromorphology of nutlets have become species within sections, rather than characterizing one of the strongest evidence reported to be useful at large natural groups as sections (Salmaki et al., various taxonomic levels in Lamiaceae and related 2008). Studies on Salvia L. showed that gross families, such as Verbenaceae and Scrophulariaceae morphology of nutlets and their sculpturing (Wojciechowska, 1966; Sivarajan, 1981; Husain et pattern are variable and taxonomically useful for al., 1990; Budantsev & Lobova, 1997; Jamzad et species delimitation (Oran, 1996; Kahraman et al., al., 2000; Attar et al., 2007; Krawczyk & Glowacka, 2011). The importance of nutlet morphological 2015). In addition to surface sculpturing, pericarp characteristics shown here has proven to be of structure, types of exocarp cells and indumentum great use in species discrimination and studying features were also considered to be taxonomically taxonomic relationships. Table 1. Details of the voucher specimens examined for the present study Taxa Section Collection details L. angularis Benth. Leucas Kerala, Mukkali, way to Silent Valley, Shinoj & Vimal CU135573 L. aspera (Willd.) Link Plagiostoma Kerala, Alappuzha, Punnapra, Shinoj CU135568 L. beddomei (Hook.f.) Sunojk. & P.Mathew* Astrodon Wayanad, Chembra Peak, Sunojkumar CU139543 L. biflora (Vahl) R.Br. ex Sm. Leucas Kerala, Munnar, Shinoj CU135590 L. cephalotes (Roth) Spreng. Plagiostoma Maharashtra, Sant Gadge Baba Amravati University, Prasad CU290102 L. chinensis (Retz.) R.Br. ex Sm. Leucas Kerala, Nelliyampathy, Seetharkundu, Smitha CU135538 L. ciliata Benth. Astrodon Kerala, Nelliyampathy, Manpara, Smitha CU135542 L. decemdentata (Willd.) R.Br. ex Sm. Leucas Andhra Pradesh, Vimal CU139692 L. deodikarii Billmore & Hemadri* Leucas Maharashtra, Junnar, Vimal CU139519 L. diffusa Benth.* Plagiostoma Tamil Nadu, Tirunelveli, Prasad CU139564 L. eriostoma Hook.f.* Astrodon Wayanad, Kurichiyarmala, Shinoj CU135577 L. eriostoma var. lanata Hook.f.* Astrodon Karnataka, Chikmagalur, Baba Budan Hills, Sunojkumar CU88101 L. helianthemifolia Desf.* Astrodon Kerala, Munnar, Devikulam, Shinoj & Vimal CU135939 L. helicterifolia Haines* Leucas Bihar, Valmiki Tiger reserve, Vimal CU139 652 L. hirta (B.Heyne ex Roth) Spreng.* Astrodon Palamala, Dhoni Hills, Shinoj & Vimal, CU135557 L. lamiifolia Desf.* Astrodon Tamil Nadu, Kodaikanal, Alfred Joe CU134225 L. lanata Benth. Leucas Uttarakhand, Dehradun, Mussoorie lake, Shinoj & Vimal CU139661 L. lanata var. candida Haines* Leucas Tamil Nadu, Ooty, Burliar road, Shinoj & Vimal CU135909 L. lanceaefolia Desf.* Astrodon Tamil Nadu, Ooty, Doddabetta, Vimal CU135924 Sajna &Sunojkumar L. lavandulifolia Sm. Plagiostoma Tamil Nadu, Ooty, Kodanad View point, Shinoj & Vimal CU135922 L. longifolia Benth. Plagiostoma Maharashtra, Junnar, Prasad & Sunojkumar CU88173 L. manipurensis N.P.Singh* Astrodon Meghalaya, Shillong, Mawphlang, Shinoj & Vimal CU139635 L. marrubioides Desf. Leucas Kerala, Wayanad, Muthanga forest, Ponkuzhi, Shinoj & Vimal CU135519 L. marrubioides var. pulneyensis Hook.f. Leucas Kerala, Nayikatty, on the way to Muthanga, Smitha CU290192 17 18 Nutlet micromorphologyofIndian Taxa Section Collection details L. martinicensis (Jacq.) R.Br. Hemistoma Kerala, Marayoor, Shinoj & Vimal CU135927 L. mathewiana Sunojk.* Plagiostoma Andhra Pradesh, Ananthapur, Kalasamudram, Sunojkumar CU49664 L. montana (Roth) Spreng.* Leucas Andhra Pradesh, Tirupati, Talakona waterfall, Vimal CU139682 L. mukerjiana Subba Rao & Kumari* Leucas Andhra Pradesh, Araku Valley, Vimal CU139694 L. nepetifolia Benth.* Leucas Tamil Nadu, Kanyakumari, Maruthwamalla, Thoiba CU138475 L. nutans (Roth) Spreng. Plagiostoma Andhra Pradesh, Ananthapur Dist., SK University campus, Sunojkumar CU49659 L. prostrata (Hook.f.) Gamble* Astrodon Tamil Nadu, Ooty, Kodanad View point, Shinoj & Vimal CU135921 L. pubescens Benth.* Leucas Kerala, Munnar, Vimal CU135751 Leucas L. rosmarinifolia Benth.* Astrodon Tamil Nadu, Ooty to Coonoor road, Ketti town, Shinoj CU135788 L. sebaldiana Sunojk.* Astrodon Kerala, Wayanad, Shinoj & Vimal CU135504 L. sivadasaniana Sunojk.* Astrodon Karnataka, Kudajadri Hills, Shinoj & Vimal CU135959 L. stelligera Wall. ex Benth. Astrodon Maharashtra, Vimal CU138401 L. stricta Benth. Plagiostoma Maharashtra, Prasad & Vimal CU290105 L. suffruticosa Benth.* Astrodon Kerala, Silent Valley, Prasad & Vimal CU135978 L. urticifolia (Vahl) Sm. Hemistoma Tamil Nadu, Coimbatore, TNAU, Soumya CU135321 L. vestita Benth.* Astrodon Karnataka, Kudachadri, Sunojkumar CU88122 L. wightiana Wall. ex Benth.* Plagiostoma Tamil Nadu, Tirunelveli, Kallidaikurichi, Shinoj CU138466 L. zeylanica (L.) R.Br. ex Sm. Plagiostoma Kerala, Malappuram, Sajna CU138483 *Species endemic to India Sajna & Sunojkumar 19 Based on Wallich’s collections from Asian Materials and Methods countries, Bentham (1832–1836, 1848) recognized six sections in Leucas. Recently, Scheen and Albert The details on specimens analyzed, collection data (2007) suggested segregation of this genus into two and the vouchers used are provided in Table 1. different groups, Leucas s.s. and ‘African Leucas’ Nutlets were collected from natural populations and based on their molecular phylogenetic studies the herbarium specimens deposited in the Calicut (Scheen & Albert, 2009). Leucas s.s. is monophyletic University Herbarium (CALI). Initially, nutlets whereas ‘African Leucas’ is an informal name given were observed using Leica M80 stereomicroscope to the paraphyletic group comprising African to check that they are of normal size and maturity. Leucas species and related genera such as Isoleucas, The dorsal surface of nutlet was considered for Leonotis, Otostegia and Acrotome. Morphologically describing surface sculpturing, to ensure certainty abrupt demarcation between Asian and African about the stability of characters among different species was earlier suggested by Sebald (1980) and accessions of one species. The measurements were Ryding (1998). Both of these groups are apparently made on 20 nutlets per species randomly obtained related, and resembling each other mainly in the from five to eight populations of each species. The absence of branched trichomes, and presence of ranges of nutlet measurements were presented in a beard along the margin of upper lip of corolla box plot graphics generated in R. (Hedge, 1992). For scanning electron microscopy (SEM), According to Bentham’s classification (1832– dried mature nutlets were mounted directly 1836, 1848), Asian Leucas consists of three natural on stubs using double-side adhesive tape, and sections:
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