Research Article Morphological and Anatomical Study on Plantaginaceae Juss. and Some Related Taxa of Scrophulariaceae Juss
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Egypt. J. Exp. Biol. (Bot.), 10(2): 135 – 146 (2014) © The Egyptian Society of Experimental Biology RESEARCH ARTICLE Karima A. Hamed Soad A. Hassan AL-Safa H. Mohamed Nareman. K. Hosney MORPHOLOGICAL AND ANATOMICAL STUDY ON PLANTAGINACEAE JUSS. AND SOME RELATED TAXA OF SCROPHULARIACEAE JUSS ABSTRACT: In recent taxonomic literature, Plantaginaceae Karima Abdel Khalek Hamed has been enlarged to include several genera AL-Safa Hassan Mohamed that were assigned to Scrophulariaceae. In the Nareman. Kmal Hosney present study, the morphology of the whole plant and the anatomy of stem, leaf and petiole Botany Department, Faculty of Science, Ain of the six wild Egyptian Plantago species as Shams University, Cairo, Egypt representatives of Plantaginaceae s.s., as well as 12 species belonging to ten Scrophulariaceae genera (Anarrhinum, Antirrhinum, Digitalis, Linaria, Kickxia, Nemesia, Russelia, Scrophularia, Verbascum, Veronica) were studied to determine whether such characters would be useful in transferring or retaining the Scrophulariaceous species in its original family. The data were analyzed by the NTSYS- PC program package using the UPGMA clustering method to clarify the interrelationships among the studied species. Three dendrogram were constructed. The first one was based on morphological criteria, the second on anatomical criteria while the third one represents a combination of both. The most obvious finding was the separation of the six Plantago species from the other 12 taxa. This finding substantiates the earlier taxonomic views in the isolation of the six studied Plantago species in a distinct monogeneric family (Plantaginaceae s.s.) away from the other Scrophulariaceae species based on the combination of both morphological and anatomical criteria. ARTICLE CODE: 15.02.14 INTRODUCTION: Plantaginaceae is a cosmopolitan family represented by three genera Plantago KEY WORDS: L., Littorella L., Bougueria L. and about 275 Anatomy, morphology, Plantaginaceae, species distributed in diverse habitats Scrophulariaceae throughout the world (Bentham and Hooker, 1876; von W ettstein, 1891; Bellini, 1907; Dahlgren, 1975; Cronquist, 1981; Heywood, 1993; Takhtajan, 1997). However, Rahn (1996) considered it as a monogeneric CORRESPONDENCE: family with only Plantago. Several authors stressed on the close relationship between Soad Abdullah Hassan Plantaginaceae and Scrophulariaceae Botany Department, Faculty of Science, Ain (Hallier, 1912; Warming, 1913; von Shams University, Cairo, Egypt Wettstein, 1935; Takhtajan, 1980; Heywood, E-mail: [email protected] ISSN: 1687-7497 On Line ISSN: 2090 - 0503 http://www.egyseb.org 136 Egypt. J. Exp. Biol. (Bot.), 10(2): 135 – 146 (2014) 1993; Mabberley, 1997). Moreover, in the Plantaginaceae (Judd, 1999; Albach et al., APG (2009) classification and in several 2005; APG 2009) and the traditional studies based on molecular criteria taxonomic systems (Bentham and Hooker, (Olmstead et al., 1992a&b and 1993; 1876). Olmstead and Reeves, 1995; Wagstaff and Olmstead, 1997), the concept of MATERIAL AND METHODS: Plantaginaceae was broadened to include In the present study, 18 taxa were many genera that were formerly assigned to collected from different localities (Table 1). Scrophulariaceae (e.g. Anarrhinum, The studied taxa include six Plantago Antirrhinum, Digitalis, Kickxia, Linaria, and species belonging to Plantaginaceae s.s. Veronica). However, the relationships either and 12 related taxa belonging to between the family and other families or Scrophulariaceae s.l.; Anarrhinum between their genera are still unclear and pubescens, Antirrhinum majus, Digitalis need to be clarified (Heywood, 1993; Albach purpurea, Kickxia acerbiana, K. aegyptiaca, et al., 2004 & 2005). Linaria maroccana, Nemesia strumosa, The main objective of the present Russelia equisetiformis, Scrophularia study is to determine the interrelationships deserti, Verbascum sinaiticum, V. sinuatum, among Plantago species, as well as the and Veronica anagallis-aquatica. The related Scrophulariaceous species to collected taxa represent 13 wild and five support the transferring or retaining these cultivated species; the wild species were taxa through studying the different identified according to Boulos (2002) and morphological and anatomical characters. the cultivated species according to Bailey The results are discussed in the light of (1949). both the current system of classification of Table 1. Locality information of the examined taxa Taxa Localities 1. Anarrhinum pubescens Fresen., Mus. Senckenb. 1: 89 (1833). Wadi El-Arbeen, Saint Katherine, Egypt Botanical Garden, Faculty of science, Ain Shams 2. *Antirrhinum majus L., Sp. Pl. 617 (1753). University, Abbassia, Egypt 3. *Digitalis purpurea L., Sp. Pl.2: 621 (1753). Orman Botanical Garden, Giza, Egypt 4. Kickxia acerbiana (Boiss.) Täckh. & Boulos, Publ.Cairo Univ. Wadi Hagol, Suez - Cairo Road, Egypt Herb. 5: 98 (1974). 5. Kickxia aegyptiaca (L.) Nábelek, Publ . Fac. Sci. Univ. Masaryk Wadi Hagol, Suez - Cairo Road, Egypt Brno 70 : 31 (1926). 6. * Linaria maroccana Hook. f., Bot. Mag. 2:18 (1872). El-Zohria Botanical Garden, Doki, Egypt 7. * Nemesia strumosa Benth., Bot. Mag. 2:18 (1836). Orman Botanical Garden, Giza, Egypt 8. Plantago albicans L., Sp., Pl., ed.1, 114 (1753). International Matrouh El-Alameen Road, Egypt 9. P. crypsoides Boiss., Fl. Orient. 4: 888 (1879). International Matrouh El-Alameen Road, Egypt 10. P. lagopus L., Sp. Pl., ed. 1, 114 (1753). International Matrouh El-Alameen Road, Egypt 11. P. lanceolata L., Sp., Pl., ed. 1, 113 (1753). International Matrouh El-Alameen Road, Egypt 12. P. major L., Sp. Pl., ed. 1, 112 (1753). Orman Botanical Garden, Giza, Egypt 13. P. ovata Forssk., Fl. Aegypt. -Arab. 31 (1775). International Matrouh El-Alameen Road, Egypt 14. *Russelia equisetiformis Schltdl. & Cham., Botanical Garden, Faculty of science, Ain Shams Linnaea 6: 377 (1831). University, Abbassia, Egypt 15. Scrophularia deserti Delile, Derscr. Egypte, Hist. Nat.240, t.33, f.1 Wadi El-Arbeen, Saint Katherine, Egypt (1814). 16. Verbascum sinaiticum Benth. in A.DC.,Prodr. 10: 236 (1846). Wadi El-Arbeen, Saint Katherine, Egypt 17. V. sinuatum L.,Sp., Pl., ed. 1, 178 (1753) . Wadi El-Arbeen, Saint Katherine, Egypt 18. Veronica anagallis-aquatica L., Sp. Pl., ed. 1, 12 (1753). Around water irrigation canal. Kafr El-Sheikh, Egypt * Cultivated species Morphological observations were hand microtome at 13-16 µm, then stained performed on the vegetative and floral organs and mounted according to the traditional of the fresh specimens by the aid of a methods (Johanson, 1940). The sections were stereomicroscope. Photos of the whole plant examined using BEL: B103 T-PL light were captured by the aid of A100 IS Digital microscope and the photos were taken using Camera (4 x). Canon Power Shot G12 digital camera. Anatomical observations were The recorded morphological and performed on transverse sections of stem, anatomical characters were coded as a binary petiole and lamina of the studied taxa using character either present (1) or absent (0) for all species to obtain a data matrix for ISSN: 1687-7497 On Line ISSN: 2090 - 0503 http://www.egyseb.org Hassan et al., Morphological and Anatomical Study on Plantaginaceae Juss. and Some Related Taxa of Scrophulariaceae Juss. 137 numerical analysis by NTSYS pc version 2.02 ii. Presence of radical leaves in rosette software using unweight pair group method form in all Plantago species while in the with arithmetic mean (UPGMA) (Rohlf, 2000). other taxa leaves are either cauline or basal The relationships between the studied taxa and cauline together. were demonstrated as a dendrogram. iii. Tetramerous flowers in the Plantago species and Veronica and pentamerous RESULTS AND DISCUSSION: flowers in the other taxa. Morphological characters: iv. Bracts and sepals in Plantago species are membranous brownish and The measurements and description of the morphological characters of the petals are scarious while otherwise in the other taxa. examined 18 taxa are presented in table 2 and figure 1 (A-F). The most obvious v. Stamens are exerted in the Plantago variations in such characters are as follows: species and Veronica while included in the corolla tube in the rest of taxa. i. Absence of stem (acaulescent) in all Plantago species except P. albicans where vi. Type of inflorescence spike in all a dwarf stem is present (caulescent) while studied Plantago species while either simple the stem is long in the other 12 taxa. raceme or cymose in the other taxa. Table 2. Data matrix of morphological characters for studied taxa (0 = absent, 1 = present): - Organ tagolagopus aquatica Characters Plantagoovata Plantago Plantago major Character state Plan Kickxiaacerbiana Plantagoalbicans Digitalis purpurea Linariamaroccana Antirrhinum majus Kickxiaaegyptiaca Nemesiastrumosa Plantagolanceolata Veronica anagallis Scrophulariadeserti Plantagocrypsoides Verbascumsinuatum Verbascumsinaiticum Anarrhinumpubescens Russeliaequisetiformis Annual 0 0 0 0 0 1 1 0 1 1 0 0 1 0 0 0 0 0 Life form Biennial 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 Perennial 1 1 0 1 1 0 0 1 0 0 1 1 0 1 1 0 0 1 Herb 1 1 1 0 0 1 1 1 1 1 1 1 1 0 0 0 0 1 Habit Woody herb 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 Subshrub 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 0 Glabrous 0 1 0 0 0 1 1 0 0 1 0 0 0 1 1 0 0 1 Whole plant Sparsely hairy 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 Texture Hairy 1 0 1 1 1 0 0 1 1 0 1 0 1 0 0 0 0 0 tomentose 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 Caulescent 1 1 1 1 1 1 1 1 0 0 0 0 0 1 1 1 1 1 Type Acaulescent 0