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Home , Anarrhinum, Kickxia, Linaria maroccana, Plantago, Russelia, Russelia equisetiformis

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 JUSS. AND SOME RELATED TAXA OF 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 and the anatomy of stem, and Botany Department, Faculty of Science, Ain of the six wild Egyptian as Shams University, Cairo, Egypt representatives of Plantaginaceae s.s., as well as 12 species belonging to ten Scrophulariaceae genera (Anarrhinum, Antirrhinum, Digitalis, , Kickxia, Nemesia, , 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., L., Bougueria L. and about 275 Anatomy, morphology, Plantaginaceae, species distributed in diverse Scrophulariaceae throughout the (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 , 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. * 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 . 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 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 in the Plantago species and Veronica and pentamerous RESULTS AND DISCUSSION: flowers in the other taxa. Morphological characters: iv. Bracts and 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. 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 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. or cymose in the other taxa. Table 2. Data matrix of morphological characters for studied taxa (0 = absent, 1 = present):

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Organ tagolagopus aquatica Characters Plantagoovata Plantago 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 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 0 0 0 0 0 0 0 1 1 1 1 1 0 0 0 0 0 Long 1 1 1 1 1 1 1 0 0 0 0 0 0 1 1 1 1 1 Length Dwarf 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0

Branched from 1 0 0 1 1 1 1 0 0 0 0 0 0 1 1 1 1 0 Branching base Unbranched 0 1 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 Stem Internal Solid 1 1 1 1 1 1 1 1 0 0 0 0 0 1 1 1 1 0 appearance Hollow 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 External Terete 1 1 0 1 1 1 0 1 0 0 0 0 0 0 0 1 1 1 appearance Angled 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 (cross section) Square 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 Homomorphic 0 1 0 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 Heterophylly Heteromorphic 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 Radical 0 0 0 0 0 0 0 1 1 1 1 1 1 0 0 0 0 0 Type Basal 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 Cauline 1 1 1 1 1 1 1 0 0 0 0 0 0 1 1 1 1 1 Absent 0 0 0 0 0 0 0 1 1 0 0 0 1 0 0 0 0 0 Petiole Present 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0

ves Linear 0 0 0 0 0 0 0 1 1 0 0 0 1 0 0 0 0 0

lea Oblanceolate 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 Leaf Shape Ovate 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 Lanceolate 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0

Radical Radical Entire 0 0 0 0 0 0 0 1 0 1 1 1 1 0 0 0 0 0 Margin Paired-teeth 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 Acute 0 0 0 0 0 0 0 1 0 1 1 1 1 0 0 0 0 0 Apex Acuminate 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 Absent 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 Petiole Present 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Shape Oblanceolate 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 ves Elliptic 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 lea

Leaf Dentate 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Margin

Basal Crenate 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 Acute 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 Apex Emarginated 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

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Table 2 : (Cont'd)

s

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ter state Organ Organ aquatica Characters Plantagoovata Plantago Plantago major Charac Kickxiaacerbiana Plantagolagopus Plantagoalbicans Digitalis purpurea Linariamaroccana Antirrhinum majus Kickxiaaegyptiaca Nemesiastrumosa Plantagolanceolata Veronica anagallis Plantagocrypsoides Scrophulariadeserti Verbascumsinuatum Verbascumsinaiticum Anarrhinumpubescens Russeliaequisetiformi Alternate 1 1 0 1 1 1 0 0 0 0 0 0 0 0 0 1 1 0 Arrangement Opposite 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 1 Whorled 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 Absent 1 1 0 0 0 1 1 0 0 0 0 0 0 1 1 1 1 1 Petiole

present 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 Linear 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 ves

Shape Lanceolate 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 Oblanceolate 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Leaf Ovate 0 0 0 1 1 0 0 0 0 0 0 0 0 1 0 1 1 1 Obovate 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 Cauline Cauline lea Entire 1 1 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 1 Toothed 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 Margin Serrate 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 Dentate 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 Crenate 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 Solitary 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 Aggregation Inflorescence 1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 Spike raceme 1 0 0 0 0 0 0 1 1 1 1 1 1 0 0 0 0 0 Inflorescence Simple raceme 0 1 1 0 0 1 1 0 0 0 0 0 0 0 0 1 1 1 type Cymose 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 Absent 1 0 0 0 0 0 0 1 1 1 1 1 1 0 0 0 0 0 Pedicel Present 0 1 1 1 1 1 1 0 0 0 0 0 0 1 1 1 1 1 Number of Tetramerous 0 0 0 0 0 0 0 1 1 1 1 1 1 0 0 0 0 1 floral parts Pentamerous 1 1 1 1 1 1 1 0 0 0 0 0 0 1 1 1 1 0

Filiform 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Cymbiform 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Lanceolate 0 0 1 0 0 1 1 0 0 0 0 0 0 0 0 0 0 1

Flower Shape Hastate 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 Ovate 0 0 0 0 0 0 0 1 1 1 1 1 1 1 0 1 1 0 Linear 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 Glabrous 0 0 0 0 0 0 0 0 0 0 1 1 0 1 1 0 0 1 Bract Texture Hairy 1 1 1 1 1 1 1 1 1 1 0 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 Green 1 1 1 1 1 1 1 0 0 0 0 0 0 1 1 0 0 1 Membrance- Colour 0 0 0 0 0 0 0 1 1 1 1 1 1 0 0 0 0 0 brownish Yellow-green 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 Free 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1 1 1 Union United 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 Glabrous 0 0 0 0 0 0 0 0 0 0 1 1 0 1 1 0 0 1 Texture Hairy 1 1 1 1 1 1 1 1 1 1 0 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

Lanceolate 1 0 0 1 1 0 1 0 0 0 0 0 0 0 0 1 1 0 Ovate 0 1 1 0 0 0 0 0 0 0 0 0 0 1 1 0 0 1 Shape Linear 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0

Sepals Cymbiform 0 0 0 0 0 0 0 1 1 1 1 1 1 0 0 0 0 0 Green 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 1 Membrance- 0 0 0 0 0 0 0 1 1 1 1 1 1 0 0 0 0 0 brownish Colour Yellow-green 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 Red-green 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 White-margined 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 Bilabiate 1 1 1 1 1 1 1 0 0 0 0 0 0 0 1 0 0 0 Shape

Rotate 0 0 0 0 0 0 0 1 1 1 1 1 1 1 0 1 1 1 Glabrous 1 1 1 0 0 0 1 1 0 1 1 1 1 1 1 0 0 1 Texture Hairy 0 0 0 1 1 1 0 0 1 0 0 0 0 0 0 1 1 0

Flower Absent 1 1 1 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 Spur Present 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 White 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Pale pink 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Purple with black 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 spots

Corolla Corolla yellow 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 1 1 0 Pink with yellow 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 Colour patches 4 with red varigation& 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 yellow orange Colorless 0 0 0 0 0 0 0 1 1 1 1 1 1 0 0 0 0 0 Red 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 Violet 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 Gynoecium Equal 1 0 1 0 0 1 1 1 1 1 1 1 1 0 1 1 1 1 locules Unequal 0 1 0 1 1 0 0 0 0 0 0 0 0 1 0 0 0 0

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Table 2 : (Cont'd)

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lbicans Organ Organ aquatica Characters Plantagoovata Plantago major Plantago Character state Character Plantagolagopus Kickxiaacerbiana Plantagoa Digitalis purpurea Digitalis Kickxiaaegyptiaca Linariamaroccana Antirrhinum majus Antirrhinum Nemesiastrumosa Plantagolanceolata anagallis Veronica Scrophulariadeserti Plantagocrypsoides Verbascumsinuatum Verbascumsinaiticum Russeliaequisetiformis Anarrhinumpubescens

2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 Number of 4 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 stamens

5 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0

Length of Didynamous 1 1 1 1 1 1 1 0 0 0 0 0 0 1 1 0 0 0 stamens Equal 0 0 0 0 0 0 0 1 1 1 1 1 1 0 0 1 1 1 Androecium Included in 1 1 1 1 1 1 1 0 0 0 0 0 0 1 1 1 1 0 Stamens corolla tube inclusion Exerted from 0 0 0 0 0 0 0 1 1 1 1 1 1 0 0 0 0 1 corolla tube Two-seeded 0 0 0 0 0 0 0 1 0 1 1 0 1 0 0 0 0 0 No. of seeds Three-seeded 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 Many-seeded 1 1 1 1 1 1 1 0 0 0 0 1 0 1 1 1 1 1 Ovoid 1 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 0

Rhomboid 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Claviform 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Fruit Oblong-ovoid 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 1 Seed Reniform 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 shape Cylindrical 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 Cymbiform 0 0 0 0 0 0 0 1 0 1 1 0 1 0 0 0 0 0 Ellipsoid 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 Rounded-angular 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0

Fig. 1 (A-F). General view of some investigated species: A-Digitalis purpurea, B- Nemesia strumosa, C- Plantago lagopus, D- P. ovata, E- Verbascum sinaiticum, F- Veronica anagallis-aquatica Numerical analysis of morphological that the taxa under investigation are divided characters: into two main series (SI, SII). Series I includes the six studied Plantago species only, while The 130 morphological character states series II includes all the remaining studied and their binary codes (0, 1) are given in table species. The same result obtained by Hassan 2. The cluster analysis obtained is represented et al. (2014) based on palynological criteria. as dendrogram in figure 2. The results clarified

I ISSN: 1687-7497 On Line ISSN: 2090 - 0503 http://www.egyseb.org 140 Egypt. J. Exp. Biol. (Bot.), 10(2): 135 – 146 (2014)

Fig. 2. Dendrogram indicating the relationships among the studied taxa based on morphological characters analysis. The isolation of the six Plantago species Anatomical characters: from the other Scrophulariaceous taxa in The anatomical characters of the accordance with many earlier taxonomic studied taxa are presented in tables 3, 4, & 5 systems in maintaining Plantago as a and figures 3 - 17. The results showed that, separate monogeneric family (Bentham and the most characteristic anatomical feature Hooker, 1876; von Wettstein, 1891; Bellini, was the presence of foliar endodermis in the 1907; Dahlgren, 1975; Cronquist, 1981; six Plantago species while absent in the rest Heywood, 1993; Takhtajan, 1997) while of the studied taxa. However, this feature is contradictory with other works based on very rare in the Angiosperms (Dickson and molecular evidence (Judd, 1999; APG, 2009). Weitzman, 1996). Table 3. Data matrix of stem anatomical characters for studied taxa (0 = absent, 1 = present) - escens hulariadeserti aquatica

Characters Plantagoovata

Plantago major Plantago Plantagolagopus Kickxiaacerbiana Plantagoalbicans Digitalis purpurea Digitalis

Kickxiaaegyptiaca Linariamaroccana Antirrhinum majus Antirrhinum Character state Nemesiastrumosa Plantagolanceolata anagallis Veronica Scrop

Plantagocrypsoides

Verbascumsinuatum

Verbascumsinaiticum

Russeliaequisetiformis Anarrhinumpub

Terete 1 1 0 1 1 1 0 1 1 0 1 1 1 0 0 1 1 1

Shape in cross section Angled 0 0 1 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 Square 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 Radial 1 1 1 1 1 1 0 0 0 1 1 1 0 1 0 0 1 1

Cells arrangement Tangential 0 0 0 0 0 0 0 1 1 0 0 0 1 0 1 1 0 0 Radial & tangential 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 Absent 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Bulliform cells idermis Present 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Ep Undulate 1 0 1 1 1 0 0 1 0 0 1 0 1 1 1 0 0 1 Cuticle shape Straight 0 1 0 0 0 1 1 0 1 1 0 1 0 0 0 1 1 0 Thin 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 Cuticle thickening Thick 1 0 1 1 1 1 1 1 1 1 0 1 1 1 1 1 1 0 Absent 1 1 1 1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 Hypodermis

Dermal systemDermal Present 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 Absent 1 1 0 0 0 1 1 0 0 1 0 1 0 1 1 0 0 1 Trichomes Present 0 0 1 1 1 0 0 1 1 0 1 0 1 0 0 1 1 0 Unbranched 0 0 1 1 1 0 0 1 1 0 1 0 1 0 0 0 0 0 Branching Stellate 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 Glandular 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 Type Eglandular 0 0 0 0 0 0 0 1 1 0 1 0 1 0 0 1 1 0 Glandular & eglandular 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Absent 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 Idioblasts Present 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0

m One 1 1 1 0 1 1 1 1 0 0 0 0 0 0 0 0 0 1 No. of tissues Two 0 0 0 1 0 0 0 0 1 1 1 1 1 0 1 1 1 0 Three 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 Absent 0 0 1 0 0 0 0 1 1 0 0 0 0 0 0 0 0 1 Cortex Chlorenchyma Present 1 1 0 1 1 1 1 0 0 1 1 1 1 1 1 1 1 0 Ground Ground syste

Polyhedral Absent 1 1 0 1 1 1 1 0 0 1 1 1 1 1 1 1 1 1 tissues Type of parenchyma Present 0 0 1 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 Table 3: (Cont'd)

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Characters Character state majus um aquatica

Plantagoovata bascumsinaiticum Plantago major Plantago Plantagolagopus Kickxiaacerbiana Plantagoalbicans Digitalis purpurea Digitalis

Kickxiaaegyptiaca Linariamaroccana Antirrhin Nemesiastrumosa Plantagolanceolata anagallis Veronica Scrophulariadeserti

Plantagocrypsoides

Verbascumsinuatum

Ver

Russeliaequisetiformis Anarrhinumpubescens

Elongated Absent 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 parenchyma Present 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 Absent 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 aerenchyma Present 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1

Lacunar Absent 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 collenchyma Present 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0

Type tissues of Angular Absent 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 0 0 1

collenchyma Present 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 1 0 Cortex

Absent 1 1 1 0 1 1 1 1 0 0 0 0 0 1 1 1 1 1 Sclerenchyma Present 0 0 0 1 0 0 0 0 1 1 1 1 1 0 0 0 0 0 Absent 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 Crystals Present 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 Ground Ground system

Ill-defined 0 0 1 1 0 0 0 1 0 0 0 0 0 1 0 1 1 0 Endodermis Well-defined 1 1 0 0 1 1 1 0 1 1 1 1 1 0 1 0 0 1 Hollow 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 Internal appearance Solid 1 1 1 1 1 1 1 1 0 1 1 1 1 1 1 1 1 0 Narrow 0 0 0 1 1 0 0 0 0 0 0 0 1 0 0 0 0 0 Width Pith Wide 1 1 1 0 0 1 1 1 1 1 1 1 0 1 1 1 1 1

Thin-walled 1 1 1 0 1 0 0 0 1 0 0 1 1 1 0 1 1 1 Type of parenchyma Thick-walled 0 0 0 1 0 1 1 1 0 1 1 0 0 0 1 0 0 0

Absent 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 Protophloem fibers Present 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0

Continuous 1 1 1 1 1 1 1 1 1 0 1 0 1 1 1 1 1 1 Type Dissected 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0

Absent 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 2ry xylem components in Fibers only 1 1 0 1 1 1 1 1 0 0 1 0 1 1 1 0 1 1 interfascicular region Vascular system

Vessels & fibers 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0

Cambium in interfascicular Absent 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 region present 1 1 1 1 1 1 1 1 1 0 1 0 1 1 1 1 1 1 Table 4. Data matrix of petiole anatomical characters for studied taxa (0=absent, 1 = present)

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ocrypsoides aquatica Characters Plantagoovata Plantago major Plantago Plantagolagopus Character state Character Kickxiaacerbiana Plantagoalbicans Digitalis purpurea Digitalis Kickxiaaegyptiaca Linariamaroccana Antirrhinum majus Antirrhinum Nemesiastrumosa Plantagolanceolata anagallis Veronica Scrophulariadeserti Plantag Verbascumsinuatum Verbascumsinaiticum Russeliaequisetiformis Anarrhinumpubescens

Half-circle 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Shape in cross section Winged triangular 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Crescent-shaped 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 Thin 0 0 1 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 Cuticle thickness

Thick 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0

Undulate 1 0 1 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 Cuticle shape Straight 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 Absent 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 Dermal Dermal system Trichomes Glandular 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Glandular & eglandular 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Table 4: (Cont'd)

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aquatica Characters kxiaaegyptiaca Plantagoovata Plantago major Plantago Plantagolagopus Character state Character Kickxiaacerbiana Plantagoalbicans Digitalis purpurea Digitalis Linariamaroccana Kic Antirrhinum majus Antirrhinum Nemesiastrumosa Plantagolanceolata anagallis Veronica Scrophulariadeserti Plantagocrypsoides Verbascumsinuatum Verbascumsinaiticum Russeliaequisetiformis Anarrhinumpubescens

Two 1 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 No. of tissues Three 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 Polyhedral Absent 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

parenchyma Present 1 0 1 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 Absent 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 tissues Type of of Type Ground system chlorenchyma Present 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Angular Absent 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 collenchyma Present 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0

Lacunar Absent 1 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 collenchyma Present 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 Arc-shaped 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Shape of bundle Rounded 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 One 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 No. of bundles Seven 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 Many 0 0 1 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 ar system Supporting angular Absent 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 collenchyma Present 1 0 1 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 Vascul Absent 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Starch sheath Present 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 Absent 0 0 1 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 Crystals present 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Table 5. Data matrix of lamina anatomical characters for studied taxa (0 = absent, 1 = present) - aquatica

Characters Plantagoovata antagolanceolata major Plantago Plantagolagopus Kickxiaacerbiana Character state Character Plantagoalbicans Digitalis purpurea Digitalis

Kickxiaaegyptiaca Linariamaroccana Antirrhinum majus Antirrhinum Nemesiastrumosa Pl anagallis Veronica Scrophulariadeserti

Plantagocrypsoides

Verbascumsinuatum

Verbascumsinaiticum

Russeliaequisetiformis Anarrhinumpubescens

Cells Tangential 1 0 0 0 0 0 1 0 1 1 1 0 1 0 0 1 0 0 arrangement Radial 0 1 1 1 1 1 0 1 0 0 0 1 0 1 1 0 1 1 Undulate 0 1 1 1 1 0 1 1 1 0 1 1 1 1 0 1 1 1 Cuticle shape Straight 1 0 0 0 0 1 0 0 0 1 0 0 0 0 1 0 0 0 Upper

epidermis Cuticle Thin 0 1 1 1 1 0 1 0 1 1 1 1 1 1 0 1 0 1 thickness Thick 1 0 0 0 0 1 0 1 0 0 0 0 0 0 1 0 1 0 Cells Tangential 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 arrangement Radial 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

Epidermis Undulate 0 1 1 1 1 0 1 1 1 1 1 0 0 1 0 1 1 1

Cuticle shape Straight 1 0 0 0 0 1 0 0 0 0 0 1 1 0 1 0 0 0 Lower Lower

epidermis Cuticle Thin 0 1 1 1 1 0 1 0 1 0 1 0 0 1 0 0 0 1 thickness Thick 1 0 0 0 0 1 0 1 0 1 0 1 1 0 1 1 1 0 Absent 1 1 1 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 Bulliform cells Present 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0

Dermal system Absent 0 1 0 0 0 1 1 0 0 1 1 0 0 1 1 0 0 1 Occurrence Present 1 0 1 1 1 0 0 1 1 0 0 1 1 0 0 1 1 0 Unbranched 1 0 1 1 1 0 0 1 1 0 0 1 1 0 0 0 0 0 Branching Stellate 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 Glandular 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 Type Eglandular 0 0 0 0 0 0 0 1 1 0 0 1 1 0 0 1 1 0

Trichomes Glandular & eglandular 1 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Unicellular 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 No. of cells Bicellular 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 Multicellular 1 0 1 1 1 0 0 1 0 0 0 1 0 0 0 1 1 0 Dorsiventral 1 1 0 0 0 0 1 0 0 0 1 0 0 1 0 0 0 1 Type Isobilateral 0 0 1 1 1 1 0 1 1 1 0 1 1 0 1 1 1 0 Absent 0 0 1 1 1 1 0 1 1 0 0 0 0 0 1 1 1 0 Two 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 Three 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 0 No. of rows Four 0 1 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 1 tissue Spongy Six 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Seven 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 Absent 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 Mesophyll Mesophyll

Shape of cells Cubic 0 1 1 0 1 0 0 0 0 1 1 0 0 1 0 0 0 1 Elongated 1 0 0 1 0 1 1 1 1 0 0 0 1 0 1 1 1 0 Two 1 1 0 0 0 0 1 0 0 0 1 0 0 1 0 0 0 1 Four 0 0 0 1 1 0 0 1 1 1 0 0 1 0 0 0 0 0 No. of rows Five 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 Palisade tissue

Six 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 Table 5 : (Cont'd)

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- jor aquatica

Characters Plantagoovata Plantago ma Plantago Plantagolagopus Kickxiaacerbiana Character state Character Plantagoalbicans Digitalis purpurea Digitalis

Kickxiaaegyptiaca Linariamaroccana Antirrhinum majus Antirrhinum Nemesiastrumosa Plantagolanceolata anagallis Veronica Scrophulariadeserti

Plantagocrypsoides

Verbascumsinuatum

Verbascumsinaiticum

Russeliaequisetiformis Anarrhinumpubescens

Eight 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0

Many 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Absent 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 Idioblasts Present 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 U-shaped 1 1 0 1 1 0 0 0 1 1 1 1 1 1 0 0 0 0 Half-circle 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 1 1 Shape Triangular 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Midrib

Curved ribbon-shaped 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 Flattened ribbon-shaped 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 One 0 0 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 No. of tissues Two 1 1 0 0 0 0 0 1 1 1 1 1 1 1 1 0 0 1 Three 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 Absent 1 1 1 0 0 0 1 0 1 1 1 1 0 1 0 1 1 1 Palisade Present 0 0 0 1 1 1 0 1 0 0 0 0 1 0 1 0 0 0 Absent 1 1 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 Spongy Present 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0

drib Absent 1 0 1 1 1 1 1 0 0 1 1 1 1 1 1 1 1 1 Chlorenchyma Mi

Present 0 1 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0

Polyhedral Absent 0 0 0 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 parenchyma Present 1 1 1 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 Type of tissuesType of

Lacunar Absent 0 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1 1 collenchyma Present 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0

Angular Absent 1 1 1 1 1 1 1 1 1 0 0 1 1 0 1 0 0 1 collenchyma Present 0 0 0 0 0 0 0 0 0 1 1 0 0 1 0 1 1 0 Absent 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 Sclerenchyma Present 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 Arc-shaped 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1

Shape of vascular Rounded 0 0 0 1 1 1 0 0 1 0 0 1 0 0 1 0 0 0 bundles Triangular 0 0 0 0 0 0 1 1 0 1 1 0 1 0 0 0 0 0 Half-circle 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 One 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 No. of vascular Five 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 bundles Many 1 1 1 1 1 1 1 0 1 1 1 1 1 0 1 1 1 1

Vascular system Absent 0 0 0 1 1 1 0 0 0 0 0 0 0 1 1 1 1 1 Supporting angular collenchyma Present 1 1 1 0 0 0 1 1 1 1 1 1 1 0 0 0 0 0 Starch sheath around Absent 1 1 1 1 1 1 1 0 0 0 0 0 0 1 1 1 1 1 vascular bundles (Endodermis) present 0 0 0 0 0 0 0 1 1 1 1 1 1 0 0 0 0 0 Absent 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 Crystals Present 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0

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Figs 3-5: Stem micrographs: 3- Scrophularia deserti, 4- Russelia equisetiformis, & 5-Kickxia acerbiana Figs 6-8: Petiole micrographs: 6- Anarrhinum pubescens, 7-Digitalis purpurea, & 8-Plantago major Figs 9-13: Lamina micrographs:9-Verbascum sinaiticum,10- Digitalis purpure,11- Plantago albicans,12- Linaria maroccana, & 13- Anarrhinum pubescens Figs 14-17: Different types of trichomes, 14- Plantago crypsoides, 15- Plantago albicans, 16- Anarrhinum pubescens, & 17- Verbascum sinaiticum Numerical analysis of the anatomical based on molecular phylogenetic relationships. characters: On the other hand, the result disagrees with the traditional taxonomic systems. The 168 anatomical character states and their binary codes (0, 1) are given in tables 3, 4, Numerical analysis of the combined and 5. The cluster analysis obtained is morphological and anatomical data: represented in the dendrogram (Fig. 18). The The dendrogram illustrating the relationships among the studied taxa are relationship among the examined taxa is discussed. shown in figure 19. The species are divided into two main series (SI and SII). Series I The results obtained from the dendrogram includes the six studied Plantago species, in figure 18 show that the six studied Plantago while series II includes the rest of the species are clustered along with remaining species. Therefore, this dendrogram Scrophulariaceous species in two different clearly separated the six Plantago species clusters (C , C ) but in three different groups. 2 4 from the 12 Scrophulariaceous species. So, it This finding may be support to some extent the is concluded that the most accurate taxa previous aspects of Olmstead et al. (1993); delimitation will be attained when the Olmstead and Reeves (1995); Judd (1999), and morphological and anatomical characters are APG (2009) who added many genera previously combined. placed in Scrophulariaceae to Plantaginaceae

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Fig. 18. Dendrogram indicating the relationships among the studied taxa based on anatomical characters analysis

Fig. 19. Dendrogram indicating the relationships among the studied taxa based on combined morphological and anatomicaldata analysis This result is congruent with the most included in Plantaginaceae s.1. according to traditional taxonomic treatments (Bentham molecular data. and Hooker, 1846; von Wettstein, 1891; Thus, from the present study the Bellini, 1907; Dahlgren, 1975; Cronquist, authors suggest the retention of 12 1981; Heywood, 1993; Rahn, 1996; Scrophulariaceous species in family Takhtajan, 1997) in remaining Scrophulariaceae s.1. and maintaining of Plantaginaceae as a separate family than the Plantago in a separate monogeneric family Scrophulariaceae. However, this result is Plantaginaceae. For more accurate contradictory with APG (2009) which stated assignment of the Scrophulariaceae to that Anarrhinum, Antirrhinum, Digitalis, Plantaginaceae or not, more studies on a Kickxia, Linaria, Russelia and Veronica large number of taxa are needed. which belonging to Scrophulariaceae are

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دراﺳﺔ ﻣﻮرﻓﻮﻟﻮﺟﯿﺔ وﺗﺸﺮﻳﺤﯿﺔ ﻋﻠﻰ اﻟﻔﺼﯿﻠﺔ اﻟﺤﻤﻠﯿﺔ وﺑﻌﺾ اﻟﻮﺣﺪات ذات اﻟﺼﻠﺔ ﻣﻦ ﻓﺼﯿﻠﺔ ﺣﻨﻚ اﻟﺴﺒﻊ ﻛﺮﻳﻤﺔ ﻋﺒﺪ اﻟﺨﺎﻟﻖ ﺣﺎﻣﺪ، ﺳﻌﺎد ﻋﺒﺪ ﷲ ﺣﺴﻦ، اﻟﺼﻔﺎ ﺣﺴﻦ ﻣﺤﻤﺪ، ﻧﺎرﻳﻤﺎن ﻛﻤﺎل ﺣﺴﻨﻰ ﻗﺴﻢ اﻟﻨﺒﺎت، ﻛﻠﯿﺔ اﻟﻌﻠﻮم، ﺟﺎﻣﻌﺔ ﻋﯿﻦ ﺷﻤﺲ، اﻟﻘﺎھﺮة، ﻣﺼﺮ ﺷﻤﻠﺖ ھﺬه اﻟﺪراﺳﺔ اﻟﺼﻔﺎت اﻟﺸﻜﻠﯿﺔ اﻟﻈﺎھﺮﻳﺔ (included). ادت ﻧﺘﺎﺋﺞ اﻟﺘﺤﻠﯿﻞ اﻟﻌﺪدي واﻟﺸﻜﻞ اﻟﺸﺠﺮي واﻟﺘﺸﺮﻳﺤﯿﺔ ﻟﺜﻤﺎﻧﯿﺔ ﻋﺸﺮ ﻧﻮﻋﺎ ً ﺗﻤﺜﻞ إﺣﺪى ﻋﺸﺮة ﺟﻨﺴﺎ ً ﻣﻦ اﻟﺬي اﻋﺘﻤﺪ ﻋﻠﻰ اﻟﺼﻔﺎت اﻟﺸﻜﻠﯿﺔ اﻟﻤﻈﮫﺮﻳﺔ (130 ﺣﺎﻟﺔ اﻟﻔﺼﯿﻠﺔ اﻟﺤﻤﻠﯿﺔ وﻓﺼﯿﻠﺔ ﺣﻨﻚ اﻟﺴﺒﻊ ﺑﮫﺪف إﺳﺘﻨﺒﺎط دﻻﺋﻞ ﺻﻔﺔ) اﻟﻰ ﻓﺼﻞ اﻧﻮاع اﻟﺒﻼﻧﺘﺎﺟﻮ اﻟﺴﺘﺔ ﻋﻦ ﺑﺎﻗﻲ اﻻﻧﻮاع ﻗﯿﺪ ﺗﺼﻨﯿﻔﯿﺔ ﻳﻤﻜﻦ ﻣﻦ ﺧﻼﻟﮫﺎ اﻟﻔﺼﻞ ﺑﯿﻦ اﻷﻧﻮاع ﻗﯿﺪ اﻟﺪراﺳﺔ. اﻟﺪراﺳﺔ. وﻗﺪ ﺗﻮاﻓﻘﺖ ھﺬه اﻟﻨﺘﯿﺠﺔ ﻣﻊ ﺑﻌﺾ اﻟﻨﻈﻢ ﻓﻘﺪ ﺗﻢ ﺗﺠﻤﯿﻊ اﻟﻌﯿﻨﺎت اﻟﻨﺒﺎﺗﯿﺔ اﻟﻤﺰھﺮة اﻟﺒﺮﻳﺔ واﻟﻤﻨﺰرﻋﺔ ﻣﻦ اﻟﺘﺼﻨﯿﻔﯿﺔ اﻟﺘﻘﻠﯿﺪﻳﺔ ﻓﻲ اﻻﺑﻘﺎء ﻋﻠﻰ ﺟﻨﺲ اﻟﺒﻼﻧﺘﺎﺟﻮ ﻓﻲ اﻣﺎﻛﻦ ﻣﺘﻔﺮﻗﺔ ﺑﻤﺼﺮ. وﺗﻢ ﺗﻌﺮﻳﻔﮫﺎ طﺒﻘﺎ ً ﻟﻠﻤﺮاﺟﻊ اﻟﺨﺎﺻﺔ ﺑﺬﻟﻚ. ﻓﺼﯿﻠﺔ ﻣﺴﺘﻘﻠﺔ وﺣﯿﺪة اﻟﺠﻨﺲ (monogeneric) وﺧﺎﻟﻒ ذﻟﻚ وﻟﻘﺪ ﺗﻢ ﻓﺤﺺ وﺗﺴﺠﯿﻞ اﻟﺼﻔﺎت اﻟﺸﻜﻠﯿﺔ اﻟﻈﺎھﺮﻳﺔ ﻟﻠﻨﺒﺎﺗﺎت ﻧﺘﺎﺋﺞ (APG (2009 اﻟﺘﻰ اﻋﺘﻤﺪت ﻋﻠﻰ دﻻﺋﻞ ﺟﺰﻳﺌﯿﺔ ﻓﻘﻂ. ﻗﯿﺪ اﻟﺪراﺳﺔ ﺑﺼﻮرﺗﮫﺎ اﻟﺤﯿﺔ ﺑﺎﻟﻌﯿﻦ اﻟﻤﺠﺮدة وﺑﺈﺳﺘﺨﺪام 2- اﻟﺨﺼﺎﺋﺺ اﻟﺘﺸﺮﻳﺤﯿﺔ: اﻟﻤﯿﻜﺮوﺳﻜﻮب اﻟﺒﺴﯿﻂ (binocular). واﻳﻀﺎ ً ﺗﻢ ﻓﺤﺺ اﻟﺼﻔﺎت اوﺿﺤﺖ اﻟﻨﺘﺎﺋﺞ ﺗﻤﯿﺰ اﻧﻮاع اﻟﺒﻼﻧﺘﺎﺟﻮ اﻟﺴﺘﺔ ﻋﻦ ﺑﺎﻗﻲ اﻻﻧﻮاع ﻗﯿﺪ اﻟﺘﺸﺮﻳﺤﯿﺔ ﻟﻠﺴﯿﻘﺎن واﻷوراق واﻻﻋﻨﺎق ﺑﺈﺳﺘﺨﺪام اﻟﻤﺠﮫﺮ اﻟﺪراﺳﺔ ﺑﻮﺟﻮد اﻧﺪوﺳﺒﯿﺮم ﻧﺸﻮى ﻣﺤﯿﻂ ﺑﺎﻟﺤﺰﻣﺔ اﻟﻮﻋﺎﺋﯿﺔ ﻟﻠﻮرﻗﺔ اﻟﻀﻮﺋﻲ، وﺗﻢ إﻟﺘﻘﺎط ﺻﻮرا ً ﻟﮫﺎ ﺑﺈﺳﺘﺨﺪام اﻟﻜﺎﻣﯿﺮا اﻟﺮﻗﻤﯿﺔ (foliar endodermis) وھﻲ ﺧﺎﺻﯿﺔ ﻧﺎدرة ﻓﻲ ﻛﺎﺳﯿﺎت اﻟﺒﺬور Gl2) canon power shot) ﺛﻢ ﺗﻢ ﺗﺠﻤﯿﻊ ﺣﺎﻻت اﻟﺼﻔﺎت (Dickson & Weitzman, 1996). اﻟﺸﻜﻞ اﻟﺸﺠﺮى اﻟﻤﺴﺘﺨﻠﺺ ﻟﻠﺸﻜﻞ اﻟﻈﺎھﺮي واﻟﺘﺸﺮﻳﺤﻲ ﻓﻲ ﺟﺪاول وﻟﻮﺣﺎت ﻣﺠﻤﻌﺔ. ﻣﻦ 168 ﺣﺎﻟﺔ ﺻﻔﺔ ﺗﺸﺮﻳﺤﯿﺔ اظﮫﺮ ﺑﻮﺿﻮح ﺗﺠﻤﻊ ﺛﻼﺛﺔ اﻧﻮاع ﻣﻦ ﺗﻢ إﺧﻀﺎع اﻟﻨﺘﺎﺋﺞ ﻟﻠﺘﺤﻠﯿﻞ اﻟﻌﺪدي ﺑﺈﺳﺘﺨﺪام ﺑﺮﻧﺎﻣﺞ -NTSYS ﺟﻨﺲ اﻟﺒﻼﻧﺘﺎﺟﻮ (P. albicans, P. crypsoides, P. ovata) ﻣﻊ PC version 2.02 software وأﻣﻜﻦ اﻟﺤﺼﻮل ﻋﻠﻰ ﺛﻼﺛﺔ اﻟﻮﺣﺪات اﻻﺧﺮى ﻣﻦ ﻓﺼﯿﻠﺔ ﺣﻨﻚ اﻟﺴﺒﻊ ( ,Antirrhinum, Kickxia أﺷﻜﺎل ﺷﺠﺮﻳﺔ (dendrograms) ; اﻻول ﻟﻠﺼﻔﺎت اﻟﺸﻜﻠﯿﺔ Linaria, Nemesia, Russelia, Scrophularia, Veronica) ﻓﻲ اﻟﻈﺎھﺮﻳﺔ واﻟﺜﺎﻧﻲ ﻟﻠﺼﻔﺎت اﻟﺘﺸﺮﻳﺤﯿﺔ واﻟﺜﺎﻟﺚ ﻟﻠﺼﻔﺎت ﻋﻨﻘﻮد واﺣﺪ cluster. وﻗﺪ ﺗﻮاﻓﻘﺖ ھﺬه اﻟﻨﺘﯿﺠﺔ ﻣﻊ (APG (2009 اﻟﻈﺎھﺮﻳﺔ واﻟﺘﺸﺮﻳﺤﯿﺔ ﻣﺠﺘﻤﻌﺔ. وﻟﻘﺪ ﻧﻮﻗﺸﺖ اﻻﺷﻜﺎل واﻟﺘﻰ ﺗﺪﻋﻢ ﻧﻘﻞ ﺑﻌﺾ اﻻﻧﻮاع ﻣﻦ ﻓﺼﯿﻠﺔ ﺣﻨﻚ اﻟﺴﺒﻊ اﻟﻰ اﻟﺸﺠﺮﻳﺔ وﻗﻮرﻧﺖ ﻧﺘﺎﺋﺠﮫﺎ ﺑﺎﻷﻧﻈﻤﺔ اﻟﺘﺼﻨﯿﻔﯿﺔ اﻟﻤﺨﺘﻠﻔﺔ اﻟﻔﺼﯿﻠﺔ اﻟﺤﻤﻠﯿﺔ ﺑﻤﻔﮫﻮﻣﮫﺎ اﻟﻮاﺳﻊ. وﺧﻠﺼﺖ اﻟﺪراﺳﺔ ﻟﻠﻨﺘﺎﺋﺞ اﻟﺘﺎﻟﯿﺔ: 3- ﻧﺘﺎﺋﺞ اﻟﺘﺤﻠﯿﻞ اﻟﻌﺪدى اﻟﻤﺠﻤﻊ ﻟﻜﻞ اﻟﺼﻔﺎت: 1- اﻟﺨﺼﺎﺋﺺ اﻟﻤﻈﮫﺮﻳﺔ: اﻳﺪت اﻟﻨﺘﺎﺋﺞ اﻟﺸﺠﺮﻳﺔ ﻟﺠﻤﯿﻊ اﻟﺼﻔﺎت اﻟﺸﻜﻠﯿﺔ اﻟﻈﺎھﺮﻳﺔ ﻛﺎن ﻣﻦ أﺑﺮز ھﺎ ﻣﺎ ﻳﻠﻲ ﻏﯿﺎب اﻟﺴﯿﻘﺎن (acaulescent) ﻓﻲ واﻟﺘﺸﺮﻳﺤﯿﺔ ﻣﻌﺎ ً ﻋﻠﻰ اﻻﺑﻘﺎء ﻋﻠﻰ ﺟﻨﺲ اﻟﺒﻼﻧﺘﺎﺟﻮ ﻓﻲ ﺟﻤﯿﻊ أﻧﻮاع ﺑﻼﻧﺘﺎﺟﻮ ﻣﺎ ﻋﺪا ﻧﻮع (P. albicans) ﻛﺎﻧﺖ اﻟﺴﺎق ﻓﺼﯿﻠﺔ ﻟﺴﺎن اﻟﺤﻤﻞ ﺑﻤﻔﮫﻮﻣﮫﺎ اﻟﻀﯿﻖ .Plantaginaceae s.s ﻣﻮﺟﻮدة وﻟﻜﻦ ﻣﺘﻘﺰﻣﺔ أﻣﺎ ﺑﺎﻗﻲ اﻻﻧﻮاع ﻗﯿﺪ اﻟﺪراﺳﺔ ﻓﻘﺪ وﺑﺎﻗﻲ اﻷﺟﻨﺎس ﻗﯿﺪ اﻟﺪراﺳﺔ ﻓﻲ ﻓﺼﯿﻠﺔ ﺣﻨﻚ اﻟﺴﺒﻊ ﺗﻤﯿﺰت ﺑﻮﺟﻮد ﺳﯿﻘﺎن طﻮﻳﻠﺔ. اﻻوراق ﻛﻠﮫﺎ أوراق ﺟﺬرﻳﺔ ﺑﻤﻔﮫﻮﻣﮫﺎ اﻟﻮاﺳﻊ .Scrophulariaceae s.l ﺗﻮاﻓﻘﺎ ﻣﻊ ﺑﻌﺾ (radical) ﺗﺸﺒﻪ ﺷﻜﻞ اﻟﻮردة ﻓﻲ ﺟﻤﯿﻊ أﻧﻮاع اﻟﺒﻼﻧﺘﺎﺟﻮ اﻷراء اﻟﺘﺼﻨﯿﻔﯿﺔ اﻟﺘﻘﻠﯿﺪﻳﺔ ﻋﻠﻰ ﺧﻼف APG 2009 وﺑﻌﺾ اﻟﻤﺪروﺳﺔ اﻣﺎ ﺑﺎﻗﻲ اﻻﻧﻮاع ﻣﻦ ﻓﺼﯿﻠﺔ ﺣﻨﻚ اﻟﺴﺒﻊ ﻓﻘﺪ اﻻﺑﺤﺎث اﻟﻤﻌﺘﻤﺪة ﻋﻠﻰ اﻟﺪراﺳﺎت اﻟﺠﺰﻳﺌﯿﺔ ﻓﻘﻂ. وﻓﯿﻤﺎ ﻳﺨﺘﺺ ﺗﺮاوﺣﺖ ﺑﯿﻦ أوراق ﺳﺎﻗﯿﺔ أو ﺳﺎﻗﯿﺔ وﻗﺎﻋﺪﻳﺔ ﻣﻌﺎ. ﻛﺎﻧﺖ ﺑﺎﻟﻮﺣﺪات اﻟﺘﺼﻨﯿﻔﯿﺔ ﻓﻲ ھﺬه اﻟﺪراﺳﺔ وﻟﻤﺰﻳﺪ ﻣﻦ اﻟﺪﻗﺔ اﻻزھﺎر رﺑﺎﻋﯿﺔ اﻻﺟﺰاء ﻓﻲ ﺟﻤﯿﻊ اﻧﻮاع اﻟﺒﻼﻧﺘﺎﺟﻮ ﺑﺎﻻﺿﺎﻓﺔ وﺗﺎﻛﯿﺪ ﻧﻘﻞ اﻻﻧﻮاع ﺑﯿﻦ اﻟﻔﺼﯿﻠﺔ اﻟﺤﻤﻠﯿﺔ وﻓﺼﯿﻠﺔ ﺣﻨﻚ اﻟﺴﺒﻊ اﻟﻰ ﺟﻨﺲ Veronica اﻣﺎ ﺑﺎﻗﻲ اﻻﻧﻮاع ﻓﻜﺎﻧﺖ ﺧﻤﺎﺳﯿﺔ. وﻛﺎن ﻓﺎﻻﻣﺮ ﻳﺘﻄﻠﺐ اﻻﻋﺘﻤﺎد ﻋﻠﻰ ﺷﻮاھﺪ ﺗﺼﻨﯿﻔﯿﺔ ﻋﺪﻳﺪة واﻳﻀﺎ ﻟﻮن اﻟﻘﻨﺎﺑﺔ واﻟﺴﺒﻠﺔ ﺑﻨﻰ ﻏﺸﺎﺋﻰ وﻛﺬا اﻟﺒﺘﻠﺔ ﻏﺸﺎﺋﯿﺔ ﻓﻲ ﻋﺪد اﻛﺒﺮ ﻣﻦ اﻻﻧﻮاع اﻟﻨﺒﺎﺗﯿﺔ. ﺟﻤﯿﻊ اﻧﻮاع اﻟﺒﻼﻧﺘﺎﺟﻮ وﻣﺘﻌﺪدة اﻻﻟﻮان ﻓﻲ ﺑﺎﻗﻲ اﻻﻧﻮاع اﻟﻤﺤﻜﻤﻮن: اﻟﻤﺪروﺳﺔ. ﻛﺎﻧﺖ اﻻﺳﺪﻳﺔ ﻣﻤﺘﺪة ﺧﺎرج اﻻﻧﺒﻮﺑﺔ اﻟﺒﺘﻠﯿﺔ (exerted) ﻓﻲ ﺟﻤﯿﻊ اﻧﻮاع اﻟﺒﻼﻧﺘﺎﺟﻮ ﺑﺎﻻﺿﺎﻓﺔ اﻟﻰ ﺟﻨﺲ أ.د. ﻋﺒﺪ اﻟﻔﺘﺎح ﺑﺪر ﻗﺴﻢ اﻟﻨﺒﺎت، ﻋﻠﻮم ﺣﻠﻮان Veronica ﺑﯿﻨﻤﺎ ﻓﻲ ﺑﺎﻗﻲ اﻻﻧﻮاع ﻛﺎﻧﺖ داﺧﻞ اﻻﻧﺒﻮﺑﺔ اﻟﺒﺘﻠﯿﺔ أ.د. ﻣﺠﺪي ﻣﺤﻤﺪ ﻣﺮاد ﻗﺴﻢ اﻟﻨﺒﺎت، ﻋﻠﻮم ﻋﯿﻦ ﺷﻤﺲ

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