BIOSYSTEMATIC STUDIES ON GENUS CLEOME L. FROM PAKISTAN
SANA
D/O RIAZ AHMED
Department of Botany
University of Karachi
Pakistan
2018
BIOSYSTEMATIC STUDIES ON GENUS CLEOME L. FROM PAKISTAN
By
SANA D/O RIAZ AHMED
THESIS
SUBMITTED TO THE FACULTY OF SCIENCE, UNIVERSITY OF KARACHI IN PARTIAL FULFILLMENT OF THE REQUIREMENT FOR THE DEGREE OF DOCTOR OF PHILOSOPHY
Department of Botany
University of Karachi
Karachi - 75270
2018
i
BIOSYSTEMATIC STUDIES ON GENUS CLEOME L. FROM PAKISTAN
THESIS APPROVED
SUPERVISOR ______
EXTERNAL EXAMINER ______
ii
Certificate of Approval
This is to certify that the research work presented in this thesis, entitled “ Biosystematic studies on genus Cleome L. from Pakistan ” was conducted by Ms. Sana D/O Riaz Ahmed under the supervision of Prof. Dr. Rubina Abid . No part of thesis has been submitted anywhere else for any other degree. The thesis is submitted to the Department of Botany in partial fulfillment of the requirement for the degree of Doctor of Philosophy in the field of Plant Taxonomy, Department of Botany, University of Karachi.
Student Name______Signature______
Supervisor Name______Signature______
Name of Dean/HOD______Signature______
iii
CONTENTS Pages
Abstract
i. English vii
ii. Urdu viii
General Introduction 1
Chapter 1- General Morphology 3
Chapter 2- Seed Micromorphology 54
Chapter 3- Palynology 63
Chapter 4- Leaf Epidermal Characters 72
Chapter 5- Phytochemistry 86
Chapter 6- Numerical Taxonomy 99
General Discussion 115
Acknowledgements 118
Appendices 119
References 123
Index 139
Addendum 141
iv
List of Figures Figure Page Details No. No. 1.1 Grid map for various localities of Pakistan 6 1.2 Distribution Pattern of C. ariana and C. oxypetala 11 1.3 a, b C ariana 13 -14 1.4 a, b C. brachycarpa 16 -17 1.5 a, b C. dolichostyla 21 -22 Distribution Pattern of C. fimbriata , C. rupicola and C. 1.6 23 dolichostyla 1.7a, b C. fimbriata 25 -26 1.8a, b C. karachiensis 28 -29 1.9 Distribution Pattern of C. pakistanica and C. karachiensis 31 1.10 a, b C. oxypetala 32 -33 1.11 a, b C. pakistanica 35 -36 1.12 Distribution Pattern of C. scaposa and C. brachycarpa 38 1.13 a, b C. rupicola 39 -40 1.14 a, b C. scaposa 42 -43 1.1 5a, b C. spinosa 46 -47 1.16 a, b C. viscosa 49 -50 1.17 Distribution Pattern of C. viscosa and C. spinosa 52 2.1 ‒2.3 Scanning electron micrographs of seeds 57 -59 3.1 – 3.2 Scanning electron micrographs of pollen 70 -71 4.1 – 4.4 Scanning electron micrographs of stomata 80 -83 4.5 – 4.6 Scanning electron micrographs of trichomes 84 -85 5.1 – 5.2 Elemental compos ition of seed coats of Cleome 88 -89 Dendrogram showing the relationship of the species of the 6.1 103 genus Cleome .
v
List of Tables Table Page Details No. No.
1 Seed micro morphological characteristics of the genus Cleome L. 56 2-3 Pollen charac teristics of the genus Cleome 65-66 4 Stomatal characteristics of the genus Cleome 75 5 Trichome characteristics of the genus Cleome 76 6 Elemental composition in seed coats of Cleome 92 7 Phenolic acids and flavonols in Cleome 93 Flavones, glycosyl flavones, chalcones, aurones, flavonones and 8 94 isoflavonones in Cleome Frequency of occurance of different phenolic compounds in 9 95 Cleome 10 Unknown compounds in Cleome 96 List of characters scored for cluster analysis for the species of 11 104-109 genus Cleome Data matrix of the genus Cleome for c haracters presented in table 12 110-114 11
vi
ABSTRACT
The genus Cleome L. is systematically revised from Pakistan and the genus is represented by 11 species including one new species. The genus Cleome is distributed in all four provinces of Pakistan, grown wild as well as cultivated. Globally it is found in tropical sub-tropical and warm regions of the world. The data from various aspects i.e., general morphology, seed and leaf micromorphology, palynology and chemistry have been obtained and analyzed numerically and distribution pattern of the genus is also studied.
The genus Cleome is generally characterized by the presence of complete and tertramerous flowers with parietal placentation, two valved capsule with persistent replum, grooved seeds, tricolporate and more or less trilobed pollen grains, presence of carbon and oxygen in seed coats and quercetin and its derivatives in leaves. While, the genus Cleome can be mainly divided into two groups usually based on simple and compound leaves and each of the species shows great diversity by having distinct combination of pollen tectum and colpal membrane surface. Similarly, a variety of trichomes like pubescent, hirsute or scabrous is also observed.
Artificial keys are constructed based on studied aspects and the data is also analyzed numerically by clustering to find out the relationship among the species of the genus Cleome from Pakistan.
vii viii
GENERAL INTRODUCTION
Cleome L. is the largest genus of the family Cleomaceae, commonly known as spider flowers. About 250 species of the genus Cleome are distributed among the tropical, sub- tropical and warm areas of the world but some species are also found in temperate regions. In Pakistan the genus Cleome is distributed from plains to hills and roadsides to stream banks, found in wild as well as in cultivated forms. Many species of the genus Cleome are reported to have medicinal values (Puri, 1971; Hameed, Ashraf, Al-Qurianty, Nawaz, Ahmed, Younis and Naz, 2011; Saroop and Kaul, 2015). While, some of the species like, C. spinosa and C. serrulata are cultivated in gardens for ornamental purposes. Similarly, C. viscosa and C. serrulata are also used as vegetable (Castetter, 1935; Ahouansinkpo, Atanasso, Dansi, Adjatin, Azize and Sanni, 2016).
Genus Cleome was first described by Linnaeus in 1753 since then many workers gave the attention to this genus and various species have been explored from different regions of the world (DE Candolle, 1824; Boissier, 1867; Oliver, 1868; Hooker, f., 1875; Blatter, 1919; Bobrov, 1939; Hutchinson and Dalziel, 1954; Gleason and Cronquist, 1963; Hedge and Lamond, 1970; Blakelock and Townsend, 1980; Thulin, 1993). Mostly, along with the gross morphology, leaf micromorphology, pollen, seed characters and chemical compounds have long been significantly used as an aid for taxonomic delimitation at various levels (Harborne and Green, 1980; Grosso, Saint-Martin and Vassal, 1994; Blatt, Santos and Salatino, 1998; Perveen and Qaiser, 2001; Khalik and Maesen, 2002; Abid and Qaiser, 2003; Abid, Ather and Qaiser, 2011, 2013, 2015, 2016). While, few workers exclusively paid the attention to micromorphology of seeds, pollen and leaves to strengthen the specific delimitation of the genus Cleome (Aleykutty and Inamdar, 1978; Sanchez-Acebo, 2005; Inda, Torrecilla, Catalan and Ruiz-Zapata, 2008; Edeoga, Omosun, Osuagwu, Mbaebie and Madu, 2009; Kamel, El-Ghani and El-Bous, 2010; Iltis, Hall, Cochrane and Sytsma, 2011; Riaz and Abid, 2018). However, no previous reports were found on seed coat elements and phenolic compounds on the genus Cleome .
Regarding to the genus Cleome from Pakistan, Stewart (1972) reported 10 species from Pakistan. While, Jafri (1973) included 10 species of Cleome in his Flora of Pakistan. Thus,
1 the most recent record of the genus from Pakistan is about 45 years old. During this period lot of nomenclatural changes have taken place and various new records, new combinations and the new species have also been introduced (Khatoon and Perveen, 2003).
Thus, in view of the current situation there is an urgent need to revise the genus from Pakistan. Detailed macromorphology, micromorphology of seeds, pollen, leaf epidermal studies like stomata, trichomes and chemistry have been carried out. Data obtained from these studies have been numerically analyzed and synthetic approach is carried out to find the specific relationship of the genus Cleome from Pakistan.
2
Chapter 1 General Morphology
Introduction
The genus Cleome L. was first described by Linnaeus (1753) with 8 species. He placed Cleome in class “Tetradynamia” under the family Capparidaceae with the following protologue:
“Perianthium tertraphyllum, minimum, patens: foliolo inferiori magis dehiscente, dedciduum. tetrapetala. Petala Omnia sursum inclinata, patentia: quorum intermedia proxima reliquis minora. Necƈteriferæ Glandulæ tres, subrotundæ, ad singulam divisuram calycis singulæ, excepta unica. Filamenta sex (rarius 12. ſ. 24.) subulata, incurva. Antheræ laterales, adscendentes. Stylus simplex. German oblongum, situ & longitudine staminum. Stigma crassiusculum, assurgens. Siliqua longa, cylindracea, stylo insidens, unilocularis, bivalvis. Sem. Plurima, subrotunda .”
Following the Linnaeus several workers studied this genus such as DE Candolle (1824) recognized Cleome on the basis of 6 (rarely 4) stamens. He (1824) divided Cleome into 2 sections i.e. Pedicellaria and Siliquaria and 53 species were described under these 2 sections. Bentham and Hooker (1862) placed the genus Cleome under Capparidaceae with 70 species. Oliver (1868) in his Flora of Tropical Africa treated Cleome under the family Capparidaceae with 20 species . Hooker f. (1875) in Flora of British India treated this genus in Capparideae and described 12 species out of which 5 species were also reported from Sindh, Punjab and Peshawar. Similarly, Boissier (1867) in Flora Orientalis also treated this genus under Capparideae with 7 species. While, Pax (1891) divided the family Capparidaceae into two subfamilies viz., Capparoideae and Cleomoideae. He placed the genus under the subfamily Cleomoideae.
Blatter (1919) placed Cleome under the family Capparidaceae and described 21 species in Flora Arabica out of which 3 were also reported from Punjab and Sindh. Post (1932) also treated Cleome under the family Capparidaceae. He described 8 species in Flora of Syria,
3
Palestine and Sinai. Bobrov (1939) in Flora of U.S.S.R described 11 species of the genus Cleome under the family Capparidaceae. Alafemar (1951) in Flore de I’iran reported 11 species of Cleome under the family Capparidaceae out of which 4 species were also reported from Baluchistan. Hutchinson and Dalziel (1954) placed the genus Cleome under the family Capparidaceae and reported 11 species of Cleome in Flora of West Tropical Africa. Wild (1960) also treated Cleome in Capparidaceae and described 13 species of Cleome in Flora Zambesiaca. Jacobs (1960) also treated this genus in family Capparidaceae and he recorded 8 species of Cleome in Flora Malesiana. Gleason and Cronquist (1963) treated Cleome under the family Capparidaceae and described 2 species in Manual of vascular plants of northeastern United States and adjacent Canada. Chater (1964) described 2 species of Cleome in Flora Europaea under the family Capparidaceae. Later on, Airy Shaw (1965) upgraded the rank of subfamily Cleomoideae as a family Cleomaceae. Zoharay (1966) included Cleome in Capparaceae and reported 3 species in Flora Palaestina. Hedge and Lamond (1970) also treated Cleome as a genus of Capparidaceae and reported 22 species in their Flora Iranica out of which 8 species were reported from area under consideration. Meikle (1977) reported 2 species of Cleome in Flora of Cyprus under Capparaceae. Jafri (1977) described 6 species of Cleome in Capparidaceae in Flora of Libya out of which 2 species were also reported from Pakistan. Whitmore (1979) described Cleome under Capparaceae with 3 species. Blakelock and Townsend (1980) treated Cleome under the family Cleomaceae and recognized 5 species in Flora of Iraq out of which 3 species were also reported from Pakistan.
Hewson (1982) recognized 10 species in the genus Cleome under the family Cleomaceae in his Flora of Australia. Grierson (1984) described 3 species of Cleome (Capparaceae) in Flora of Bhutan. Raghavan (1993) recognized 16 species of the genus Cleome under the family Capparaceae in Flora of India, out of these 6 species were also reported from Pakistan. Thulin (1993) recognized 17 species in Flora of Somalia amongst them 2 species were also reported from our area of study. Chaudhary (1998) reported 16 species in the genus Cleome under the family Capparaceae in Flora of the Kingdom of Saudi Arabia out of which 4 species were reported from Pakistan. Mathur (2002) in Flora of Jammu and Kashmir described 5 species under the genus Cleome. Mia, Ara and Khan (2007) described 5 species of Cleome under the family Capparaceae in Flora of Bangladesh.
4
According to angiosperm phylogeny group (APG) I- IV (1998, 2003, 2009, 2016) based on DNA studies, Cleome was treated under the family Cleomaceae. While, some taxonomists transferred the species of Cleome under the separate genus such as, Rafinesque (1838) placed C. viscosa under the genus Arivela mainly based on unequal petals and 8‒15 stamens. Similarly, Zhang and Tucker (2008) in Flora of China followed the treatment of Rafinesque by introducing two distinct genera Cleome and Arivela on the basis of absence or presence of gynophore and number of stamens (6 and 14‒25 respectively). While, Tucker and Vanderpool (2010) in Flora of North America treated the genus Cleome with [4] 6 stamens, pubescent capsule and 1‒25mm long gynophore and the genus Arivela was represented by the presence of 14‒35 stamens, hirsute capsule and obsolete gynophore.
Presently, the genus Cleome cannot be recognized on the basis of 4 or 6 stamens as described earlier. Since Cleome has great diversity in terms of number of stamens. e.g. 4 stamens are present in C. dolichostyla while, number of stamens ranges from 10‒20 in C. viscosa . Similarly gynophore is also absent in C. brachycarpa .
Currently the genus Cleome comprises ca. 250 species all over the world (Mabberley, 2008), out of which 10 species viz., C. ariana , C. brachycarpa , C. scaposa , C. dolichostyla , C. rupicola , C. fimbriata , C. heratensis subsp. pakistanica , C. oxypetala C. viscosa and C. spinosa are reported from Pakistan by Jafri (1973) under the family Capparidaceae. From the available reports it is evident that most recent record for the genus is outdated i.e. about 45 years old during this period various nomenclatural changes have taken place. So there seems an urgent need for the revision of the genus Cleome from Pakistan.
Materials and Method:
Genus Cleome (ca. 600 specimens) was studied from various herbaria viz., BM, E, ISL, K, KUH, PMNH and RAW (abbreviated as in Holmgren, Holmgren and Barnett, 1990). Various morphological characters have been studied (1) Habit (2) Stem (size, branching pattern, surface) (3) Leaves (type, size, shape, surface) (4) Flower (size, colour) (5) Sepals (colour, size, shape, surface) (6) Petals (colour, size, shape) (7) Androecium (number of stamens, length) (8) Gynoecium (size, surface, structure) (9) Capsule (shape,
5
Fig. 1.1. Grid map for various localities of Pakistan (after Ali and Qaiser, 1986).
6 size, surface) (10) seeds (surface, colour). Terminology used is in accordance with Lawrence (1970) and Stearn (1983).
Original descriptions of all the species were studied and types were traced out, in case of non-availability of herbarium specimens online photographs of type specimens were seen. An artificial key to the species of the genus Cleome has been constructed. Distribution pattern of all taxa has been traced out with the help of herbarium specimens and literature. Localities in Pakistan are mentioned according to the grid map (Fig. 1.1).
Results and Discussion:
The genus Cleome is represented by 11 species in Pakistan.
CLEOME L.
L., Sp. Pl. 671. 1753; Gen. Pl. ed. 5: 302. 1754; De Candolle, Prods. 238. 1824; Benth. and Hook., Gen. Pl. 1: 105. 1862; Boiss., Fl. Or. 1: 410. 1867; Oliver, Fl. Trop. Afr. 1: 74. 1868; Hook. f., Fl. Brit. Ind. 1: 168. 1875; Blatter, Fl. Arab. 8(1): 34.1919; Post, Fl. Syria, Palestine and Sinai. 1: 132. 1932; Bobrov in Komarov, Fl. U.S.S.R. 8: 5. 1939; Alafemar in Parsa, Flore de I’Iran. 1(1): 908. 1951; Hutchinson and Dalziel, Fl. W. Trop. Afr. 1(1): 86. 1954; Jafri, Kew Bull. 173-175. 1957; Kitamura., Fl. Afg. 137. 1960; Jacobs, Fl. Malesiana. 1(6): 99. 1960; Wild in Exell and Wild, Fl. Zambesiaca. 1(1): 195. 1960; Gleason and Cronquist, Manual of Vas. Pl. North. U. S. & adj. Canada. 349. 1963; Chater in Tutin, Heywood, Burges, Valentine, Walters and Webb, Fl. Europ. 1: 259. 1964; Zohary, Fl. Palaestina. 1: 245. 1966; Hedge and Lamond in Rech. f., Fl. Iran. 68: 13. 1970; Jafri in Nasir and Ali, Fl. W. Pak. 34: 20. 1973; Meikle, Fl. Cyprus. 1: 173. 1977; Jafri, Fl. Libya. 12: 11. 1977; Whitmore in Hara and Williams, An enum. Fl. Pl. Nepal. 2: 46. 1979; Blakelock and Townsend in Townsend and Guest, Fl. Iraq. 4(1): 145. 1980; Grierson in Grierson and Long, Fl. Bhutan. 1(2):
7
415. 1984; Thulin, Fl. Somalia. 1: 53. 1993; Raghavan in Sharma and Balakrishnan, Fl. Ind. 2: 299. 1993; Chaudhary, Fl. K.S.A. I: 439. 1998; Mathur in Singh, Singh and Uniyal, Fl. Jam. and Kash. 1: 600. 2002; Mia, Ara and Khan in Khanum and Ara, Fl. Bangladesh. 57: 7. 2007.
Herbs or undershrubs, glandular or eglandular hairy. Stem erect or semi-erect, branched or unbranched, 4–100cm tall. Leaves alternate, petiolate or subsessile, simple or compound, 3– 7 foliate, Inflorescence racemose, Flowers, complete, bisexual, small, pedicellate, yellow, yellow-brown, white, off white or pinkish. Sepals 4, free, green, small, glandular or eglandular. Petals 4, free, equal or unequal, monomorphic or dimorphic, imbricate. Stamens 4–20, free, equal or subequal, sessile or on androphore. Gynoecium bicarpellary, ovary sessile or on gynophore, unilocular with many ovules, parietal placentation, style 0.5‒4.5mm or absent, Capsule 2 valved with persistent replum, linear or oblong, Seeds many, usually brown, glabrous or hairy.
Key to the species of Cleome L. from Pakistan
1+ Leaves simple………………...………………………….……………..…………2
- Leaves compound …...………………………………….…………...... …………7
2+ Undershrub, leaves scanty and very small...………….….…….7. C. pakistanica
- Herbs, leaves many and comparatively large………..……….………….………3
3+ Capsule linear……………………………………...……………………………...4
- Capsule oblong….…………………………………………………………..……..5
4+ Stem and leaves glabrous, flowers yellow, 1cm across, seeds hairy…..………..
…………………………………………………...………….……..6. C. oxypetala
- Stem and leaves hairy glandular, flowers pale white or pinkish, 3–4mm across, seeds glabrous.…………………………………………..…..………9. C. scaposa
8
5+ Stamens 6, fruits 50mm long, seeds hairy…………………..….….8. C. rupicola
- Stamens 4, fruits 12–30mm long, seeds glabrous………….……………..…….6
6+ Leaf base truncate or cordate……………….………….….……3. C. dolichostyla
- Leaf base obtuse or cuneate……………………………….…….…4. C. fimbriata
7+ Petals linear-oblong, oblong, obovate or obovate-elliptic, stamens 6……..……8
- Petals oblanceolate, oblong-lanceolate or oblong-elliptic, stamens 6–20…..…10
8+ Stem semi-erect, branched from base, flowers yellow………...2. C. brachycarpa
- Stem erect, branched but not from base, flowers white or pinkish………..……9
9+ Leaves 3 foliate, flowers 3–5mm across, filaments 1–6mm long..…1. C. ariana
- Leaves 5–7 foliate, flowers 25mm across, filaments 25mm long……………… …………………………………………………………….…..……10. C. spinosa
10+ Stamens 6–8, capsule 10–29mm long………………….…….…5. C. karachiensis
- Stamens 10–20, capsule 37–80mm long………………….…….....11. C. viscosa
1. Cleome ariana Hedge and Lamond in Rech. f., Fl. Iran. 68: 17. 1970; Jafri in Nasir and Ali, Fl. W. Pak. 34: 24. 1973; Raghavan in Sharma and Balakrishnan, Fl. Ind. 2: 302. 1993; Mathur in Singh, Singh and Uniyal, Fl. Jam. & Kash. 1: 600. 2002 (Figs. 1.3a & b).
C. ornithopodioides auct. non L., Sp. Pl. 672. 1753; Jafri, Kew Bull. 175. 1957; C. iberica Aitch., J. Linn. Soc. 18: 35. 1880. (not of DC., Prodr., 1: 240. 1824).
Erect herb, 20–97cm tall, branched, hairy, sparsely glandular. Leaves compound, alternate, trifoliate, uppermost becoming unifoliolate, sessile to sub-sessile, lower petiolate, petiole 4– 12mm. Leaflets linear, lanceolate, linear–elliptic, oblong, obovate, apex obtuse, rarely acute, base obtuse-cuneate, glandular, 9–20 × 1–6mm. Flowers complete, 3–5mm across, white or pinkish, pedicel 4–10mm. Sepals 4, green, eglandular, ovate, oblong-elliptic or elliptic,
9 acute, 1–2mm long, 0.2–0.5mm broad, connate at base. Petals 4, oblong, linear-oblong, apex acute, obtuse, base attenuate, 1.5–3mm × 0.2–0.5mm. Stamens 6, free, filaments 1–6mm long. Ovary eglandular, 1–2.5mm long, style almost absent, gynophore upto 1mm. Capsule linear, eglandular, 7–22mm × 1mm, stalked, apex acute, base cuneate or acute. Seeds many, glabrous, brown.
Type: Solang pass, Afghanistan, 26.8.1965, Stainton 5072 (E!,W).
A-6 Chitral Dist.: Pasti, Chitral, steep rock slope, 2.7.2007, Haider Ali 6439 (KUH); A-7 Chitral Dist.: 5km from Booni on way to Mastuj about 2000m, 25.6.1987, A. Ghafoor & S. Omer 3019 (KUH); 13km from Chitral to Garam Chashma, Chitral, 18.7.1995, G.R. Sarwar & S. Omer 246 (KUH); About 13km from Chitral on way to Shoghor, 11.8.1992, Tahir Ali, Z.S. Hussain & Gohar Khan 2215 (KUH); Barum gol-mujain ghari, Chitral-Chitral, steep stony slope, 29.6.2007, Haider Ali 6358 (KUH); Sarigulbek, Chitral-Torikhoo, undulating gentle slope, 16.8.2006, Haider Ali 5041 (KUH); Sarigulbek, Torikhoo, Chitral, 16.8.2006, Haider Ali 5067 (KUH); Shehreshan towards shoghore, Chitral-Chitral, stony steep slope, 18.6.2007, Haider Ali 5776 (KUH); Agarak, Chitral Dist., ± 8000m, 24.6.1977, Muqqarab Shah & Dilawar 1717 (ISL); Chitral, 16.7.1979, Wali ur Rehman & Subhan 299 (ISL); Chitral, 20.7.1971, Sultan-ul-Abedin 8003 (KUH); Nishkoo Tirich, Molikhoo, Chitral, 6.8.2005, Haider Ali 2440 (KUH); Morkoh, 55 miles from Chitral city on way to Trichmir, 16.8.1981, Kamal Akhter Malik & S. Nazimuddin 1473 (KUH); Chumarkan, Mastooj, Chitral, 14.7.2005, Haider Ali 2063 (KUH); Chowinch, Mastooj, Chitral, 14.7.2005, Haider Ali 2007 (KUH); about 13km from Chitral on way to Shoghore ±1400m, 11.8.1992, Tahir Ali, S.Z. Hussain & Gohar khan 2215 (KUH); Isphedar CGNP, Chitral-Chitral, grows on moist slope, 21.6.2005, Haider Ali 951 (KUH); Mera, Chitral Gol, 30.6.1977, Hakim Khan 209 (RAW); B-6 Chitral Dist: Birmoghlasht, 2.8.1954, M.A. Siddiqui & A. Rehman s.n. (RAW); Birir gol, Chitral-Chitral, steep stony slope, 19.7.2007, Haider Ali 6633 (KUH); Birir Gol valley, Chitral Dist., 24.7.1976, M. Qaiser & A. Ghafoor 6592 (KUH); Kafiristan valley, 22.6.1970, M. Qaiser & A. Ghafoor 1948 (KUH); Shishi valley, Kashindel, Chitral- Drosh, Beside stream bank, 30.6.2005, Haider Ali 1370 (KUH); Bombrait, Chitral, 3.7.2005, Haider Ali 1464 (KUH); Rabat Arkari, Chitral-Lutkhoo, undulating rocky slope, 29.8.2005, Haider Ali 3060 (KUH); D-4 Kalmangalzai, Ziarat road, Baluchistan, 10.7.1980, Farrukh
10
Fig. 1.2. Distribution pattern of C. ariana (●) and C. oxypetala (▲).
11
Hussain 115 (RAW); Shingar, about 50km N. Fort Sandeman shale slopes below steep slopes, Quetta, 20.5.1965, Jennifer Lamond 1470 (E); Shingar, Kili Ibrahim Khel, about 54km from Zhob, 31.5.1995, Tahir Ali & G.R. Sarwar 2747 (KUH); D-5 Loralai Dist.: About 34 miles from Loralai towards Qila Saifullah, 13.6.1970, M. Qaiser & A. Ghafoor 1557 (KUH).
Distribtion: Iran, Afghanistan, Pakistan, Kashmir & Pamir, Alai, India (Fig. 1.2).
2. Cleome brachycarpa Vahl. ex DC., Prodr. 1: 240. 1824; Boiss., Fl. Or. 1: 412. 1867; Oliver, Fl. Trop. Afr. 1: 77. 1868; Hook. f., Fl. Brit. Ind. 1: 169. 1875; Blatter, Fl. Arab. 8(1): 35. 1919; Post, Fl. Syria, Palestine and Sinai 1: 133. 1932; Alafemar in Parsa, Flore de I’Iran 1(1): 912. 1951; Hutchinson and Dalziel, Fl. W. Trop. Afr. 1(1): 87. 1954; Hedge and Lamond in Rech. f., Fl. Iran. 68: 20. 1970; Jafri in Nasir and Ali, Fl. W. Pak. 34: 23. 1973; Jafri, Fl. Libya 12: 17. 1977; Western, Fl. U.A.E. 57. 1989; Thulin, Fl. Somalia 1: 56. 1993; Chaudhary, Fl. K.S.A. 1: 441. 1998 (Figs. 1.4a & b).
C.ruta Camb. In Jacquem., Voy. Bot. 19, 1844; C. moschata Stocks ex. T. Anders. in J. Linn. Soc. Suppl. 5, 1: 5. 1860.
Semi-erect herb, 10–35cm tall, branched from base, glandular. Leaves alternate, compound, 3–5 foliate, glandular, petiole 2–30mm. Leaflets elliptic, obovate, oblanceolate, oblong or orbicular, acute or obtuse, base cuneate, 6–20 × 2–8mm. Flower complete, 0.8–1.2cm across, yellow, pedicellate. Sepals 4, green, glandular, oblong, lanceolate, elliptic or oblong- elliptic, acute, 1.5–2.5 rarely upto 4 × 0.5–2mm. Petals 4, obovate or obovate-elliptic, acute, base cuneate, 3–7 × 1–3mm. Stamens 6, free, 2–6mm long, basifixed. Ovary glandular, 1– 5mm, ovules many, style persistent, 1–4.5mm, gynophore absent. Capsule 5–10 × 1.5–2mm, oblong, glandular. Seeds many, glabrous, rust brown or maroon.
Type: Ex. Orient, Asia, Forsskal s.n. (BM!).
B-6 Khyber Agency Dist.: Landi Kotal, 20.4.1964, Siddiqui & Hassan ud din 2671 (RAW); S.E. of Cherat & west of Nizampur on road to Mir Kalan, Kohat, 12.8.1958, B.L. Burtt 1046 (E); C-6 Bannu Dist.: Bannu, NWFP, 23.4.1954, A. Rahman s.n. (RAW); Dogra, Bara, 6.5.1979, M. Tanvir & Dilawar 1007 (ISL); Tanda Dam, Kohat Dist., N.W.F.P., 30.4.1977,
12
Fig. 1.3a. C ariana : Photograph showing the habit.
13
Fig. 1.3b. C. ariana : A, Habit; B, Fruit.
14
Hafizullah & Dilawar 1052 (ISL); C-7 Jhelum Dist.: Choa Saidan Shah, ± 763,24.9.1976, M.A. Siddiqui , Akram & Lal Khan 53 (ISL); About 25km from Khushab on way to Naushehra, Tahir Ali & Tufail Ahmed 1798 (KUH); Dhamyal, Rawalpindi, 17.4.1943, Baldev Raj Khosla s.n. (KUH); C-8 Lahore Dist.: Khagah Dogran, Gujranwala Dist., 15.3.1917, R.R. Stewart 1475 (RAW); D-4 Old Much about 63km from Quetta on way to Sibi, across the river, 29.5.1995, Tahir Ali & G.R. Sarwar 2637 (KUH); E-4 Kalat Dist.: 50km from Kalat on way to Swabi, RCD highway near gasoline station,15.5.1990, A. Ghafoor & Steve M. Goodman 5258 (E, KUH); About 85km from Khuzdar city on way to Kalat, 27.5.1995, Tahir Ali & G.R. Sarwar 2548 (KUH); E-5 D.G. Khan Dist.: 25 miles from Kashmor on way to Rojhan, 20.5.1972, Sultan-ul-Abedin & Abrar Hussain 9542 (KUH); About 6 miles before Sakhi Sarwar on way to D.G. Khan, D.G. Khan-Loralai road, 17.9.1970, M. Qaiser 2562 (KUH); E-6 Bahawalpur Dist.: Cholistan Desert, Bahawalpur, S. Punjab, 4.11.1996, Dr. Rubina Akhter & Dr. M. Reidl 25 (RAW); E-7 Bahawalpur Dist.: Near WAPDA grid station, 25.9.1988, A. Ghafoor & Tahir Ali 3556 (KUH); F-4 Dadu Dist.: 4 miles from Sehwan on Hyderabad road, Dry lime stone hill, 6.9.1970, S.A. Farooqi & M. Qaiser 2203 (KUH); Near Bagolare railway station, Dadu Dist., 30.4.1976, Nazim, S. Abedin & M. Qaiser 36 (KUH); about 2 miles from Bela on way to Khuzdar, 11.10.1978, Kamal Akhter Malik & S. Nazimuddin 934 (KUH); Lak Pass about 5km from Bela on way to Awaran, near Asant, 20.9.1986, A. Ghafoor & S. Omer 1681 (KUH); 15km W. of Bela, E. side of Jan paas, Baluchistan, 9.4.1965, Jennifer Lamond 292, 301 (E); 30 miles from Bela on way to Awaran, 25.6.1971, Sultan ul Abedin & M. Qaiser 7463 (KUH); F-5 Nawabshah College Campus, 25.5.1965, S.M.H. Jafri 4026 (KUH); Seeta Goth near Kutte ji kabar, Kirthar Range, Larkana, 18.3.1983, Kamal A. Malik , Saood Omer & Abdul Wahid 2322 (KUH); G-2 Agor, Makran Coast, S. Baluchistan, 10.5.1997, Dr, Rubina Akhter 65 (RAW); Ras Nuh, South of Gawadar, Makran, Baluchistan, 15.4.1965, Jennifer Lamond 479 (E); G- 3 about 40km before Aghore on Liyari-Ormara road Makran, Baluchistan, 2.3.1990, M. Qaiser , K.H. Rechinger , Jennifer Lamond & Tahir Ali 8196 (KUH); 6 miles from Lasbela on way to Awaran, 12.10.1978, Kamal Akhter & S. Nazimuddin 986 (KUH); Hab to Diwana, about 10km from Hab, Bela, Baluchistan, 30.4.1965, Jennifer Lamond 779 (E); About 10 miles from Bela on way to Khuzdar, 4.6.1970, M. Qaiser & A. Ghafoor 1150 (KUH); 32 miles from Awaran on way to Hoshab, 21.4.1970, M. Qaiser 793 (KUH); San
15
Fig. 1.4a. C. brachycarpa : Photograph showing the habit.
16
Fig. 1.4b. C. brachycarpa : A, Habit; B, Rhizome; C, flower (Courtesy Flora of Pakistan, 1973).
17
Pass 8 miles from Bela on way to Awaran, 2.4.1972, Abdul Ghafoor & M. Qaiser 3 (KUH); Between Awaran & Hoshab, 26.11.1988, M. Qaiser, S. Omer & T. Ahmed 8133 (KUH); G-4 Paradise point, Jennifer Lamond 5320 (E); Hawks bay, Karachi, 8.3.1952, S.M.H. Jafri s.n. (E); University of Karachi, Sindh, 13.3.1972, M.A. Siddiqui 6778 (PMNH); University Campus KU, K.S.A. Shah s.n. (KUH); Cape Monze, Karachi, 23.8.1984, Rizwan yousuf 9 (KUH); New Microbiology Block, University Campus, Karachi, 26.8.1971, Al Zohad 51 (KUH); University of Karachi Campus, Karachi, 10.9.1960, Mr. Shamim Ahmed s.n. (KUH); Kambhu, Kirthar, 20.10.2001, Dr. Rubina Akhter 616 (RAW); Kambhu, Kirthar, 17.11.2001, Dr. Rubina Akhter 607 (RAW); Kambhu, Kirthar, 25.9.2001, Dr. Rubina Akhter 595 (RAW); Kirthar National Park, 5.1.2002, Dr. Rubina Akhter & Tariq Khan 1185 (RAW); Dureji, Kirthar National Park, Karachi, 5.1.2002, Dr. Rubina Akhter & Tariq Khan 089 (RAW); Bundak hill, Kirthar National Park, Karachi, 5.1.2002, Dr. Rubina Akhter & Tariq Khan 088 (RAW); Benir, Kirthar National Park, 17.11.2001, Rubina Akhter 606 (RAW); Dumbar, Kirthar National Park, 28.2.2002, Dr. Rubina Akhter & Tariq Khan 1182 (RAW); Hills near Sandspit Coast, 24.9.2002, Rubina Akhter & Tariq Khan 050 (RAW); Tomato field 15km from Gadap Town, 2.10.2005, Sadaf Rahimi , Sahar Zaidi , M. Imran & Surraya Khatoon SR 743 (KUH); Karachi to Dadu, about 10–15km S. Sehwan, 5.5.1965, Jennifer Lamond 867 (E); Kirthar, Anjum Perveen & Ishtiaq Hussain s.n. (KUH); Mangopir, 3.8.1956, I.I. Chaudhary s.n. (KUH); Along Liyari, Behind Bihar Colony, 15.3.1955, S.M.H. Jafri 813 (KUH); Karachi, 3.9.1950, Imtiaz 68 (KUH); Rizvia colony, Karachi, 21.7.1954, S.R.B. Rizvi s.n. (KUH); Lalukhet, Karachi, 20.8.1956, S.A. Alvi s.n. (KUH); Kambu, Kirthar, 15.3.2005, Anjum Perveen & Ishtiaq Hussain 29 (KUH); About 15 miles from T.B. Khan on way to Karachi, 27.3.1970, M. Qaiser 451 (KUH); University campus, Karachi, 21.9.1960, M. Saeed Haider s.n. (KUH); Near primary school, Universuty campus, Karachi, 19.7.1969, Sultan-ul-Abedin 3570 (KUH); Near Baldia colony, Karachi, 11.1.1959, Mr. Shamshad, M. Naqvi & party s.n. (KUH); North Nazimabad, Karachi, 9.8.1958, T.R. Naqvi s.n. (KUH); Malir, Karachi, 9.8.1958, Shamim Akhter s.n. (KUH); Nazimabad, Karachi, 6.3.1958, Asif Zaidi s.n. (KUH); Rizvia colony, Karachi, 19.9.1955, S.R. Baqar Rizvi s.n. (KUH); University campus, Karachi, Tashmina Razzaki s.n. (KUH); University of Karachi, 16.2.1964, M. Aftab Ahmed Yousufi s.n. (KUH); University campus, 1961, A.S. Bhatti s.n. (KUH); University campus, Karachi, 10.8.1971, Tariq Haroon 228 (KUH);
18
University campus, 12.12.1960, S.A. Kheiri s.n. (KUH); University campus, A.S. Bhatti 461 (KUH); Karachi University campus, 18.6.1969, Sultan-ul-Abedin s.n. (KUH); Rahu jo attu, about 24km from Ghaibi dewar on way to Kutte jo Qabar, Kirthar range, 18.3.1983, Kamal A. Malik, Saood Omer & Abdul Wahid 2785 (KUH); Opposite National stadium, Karachi, 6.5.1956, S.M.H. Jafri 1412 (KUH); University campus, 1962, Kishwar Rehman s.n. (KUH); Darsano Chino, Malir, Karachi, 2.4.1987, Ismail s.n. (KUH); Botany Department, University campus, Karachi, 7.4.1986, Anjum Perveen 75 (KUH); Near Microbiology Department, Karachi University campus, 12.10.1986, Abrar Husain s.n. (KUH); Boys Hostel, University campus, Karachi, 26.8.1971, Ali Zohhad 59 (KUH); Hawks bay on way to Mubarak village, 6.3.2011, Shazia Mansoori & Anjum Perveen s.n. (KUH); Mubarak village, 24.10.2011, G.R. Sarwar, Anjum Perveen & students 2086 (KUH); Tikobaran, Kirthar National Park, 24.9.2009, Jan Alam, G.R. Sarwar 4293 (KUH); Malir, Karachi, 21.11.1958, Shamim Akhter s.n. (KUH); North Nazimabad, Karachi, 9.8.1958, Muhammad Tausif s.n. (KUH); Malir, Karachi, 3.12.1958, Muhammad Amanullah s.n. (KUH); Clifton, Karachi, 22.12.1957, Anwar Iqbal s.n. (KUH); Clifton, Karachi, 26.9.1956, S.A. Qadir s.n. (KUH); Near Baldia colony, Karachi, 11.1.1959, S.I. Ali & students 291 (KUH); University campus, 09.1960, Shamim Ahmed s.n. (KUH); University campus, 5.10.1960, Siraj A. Rizvi s.n. (KUH); Malir, Karachi, 28.11.1960, Zubair Ahmed s.n. (KUH); Roadside, Karachi city, 8.10.2003, Dr. Rubina Akhter 98 (RAW); G-5 Dadu Dist.: Laki Shah Sedan, 6.4.1959, S.M.H. Jafri 2749 (KUH).
Distribution: Mauritania, Chad, Egypt, Sudan, Saudi Arabia, Yemen, Ethiopia, Somaliland, Iran, Afghanistan, Pakistan, India (Fig. 1.12).
3. C. dolichostyla Jafri in Kew Bull. 174. 1957; Hedge and Lamond in Rech. f., Fl. Iran. 68: 23. 1970; Jafri in Nasir and Ali, Fl. W. Pak.34: 28. 1973; Western, Fl. U.A.E. 58. 1989 (Figs. 1.5a & b).
Erect herb, 30–80cm tall, hairy, branched, glandular. Leaves alternate, petiole 13–25mm, simple, cordate, broadly cordate, suborbicular, five veined, glandular, cordate, acute- acuminate, obtuse or obtuse-apiculate, base truncate, 14–23 × 12–20mm. Flower complete, 5-8mm across, yellow on 5–6mm pedicel. Sepals 4, green, hairy, oblong-elliptic, acute 3–5
19
× 1–1.5mm. Petals 4, lanceolate or lanceolate-oblong, apex acute, base truncate, 6–8 × 1– 2mm. Stamens 4, free, basifixed, 7–13mm long. Ovary glandular, gynophore absent, 5– 15mm, style persistent, 1–10mm. Capsule oblong, glandular, 20–24 × 2–5mm. Seeds many, glabrous, brown.
Type: Masjid-I-Sulaiman, Iran, 23.9.1949, A. J. Lee 66 (K!).
G-2 Makran Dist.: 30 miles from Mand on way to Teerandazuk, 11.4.1969, S.I. Ali, S.A. Farooqi & Sultan-ul-Abedin 1840 (KUH); Turbat on way to Pasni, 5.9.1986, A. Ghafoor & S. Omer 1804A (RAW); 2 miles from Turbat on way to Pasni, 25.4.1970, M. Qaiser, Asad Raza & Abrar Hussain 922 (KUH); Makran, 80km S. of Turbat, road to Pasni, Baluchistan, 13.4.1965, Jennifer Lamond 432 (E); About 30km from sandsar on way to Mand, Baluchistan, 3.10.1986, A. Ghafoor & S. Omer 1863 (KUH); about 10 miles from Mand on way to Sandsar, Makran Dist., 27.4.1970, M. Qaiser, Asad Raza & Abrar Hussain 1043 (KUH); G-4 Karachi Dist.: Baran valley, Kirthar, 20.10.2001, Rubina Akhter 094 (RAW).
Distribution: United Arab Emirates, Oman, Iran, Afghanistan, Pakistan (Fig. 1.6).
4. Cleome fimbriata Vicary in J. Asait. Soc. Beng. 16: 1158. 1847; Jafri in Nasir and Ali, Fl. W. Pak. 34: 29. 1973; Raghavan in Sharma and Balakrishnan, Fl. Ind., 2: 308.1993; Mathur in Singh, Singh and Uinyal, Fl. Jam. & Kash. 1: 600. 2002 (Figs. 1.7a & b).
C. noeana Boiss., Diagn. Pl. Or. Nov. Ser. 2. 1: 48. 1853; Boiss., Fl. Or. 1: 415. 1867; Hedge and Lamond in Rechinger f., Fl. Iran. 68: 22. 1970; C. quinquinervia var. noeana (Boiss.) Alafemar in Parsa, Flore de I’Iran. 1: 914. 1951; C.pentanervia Aitch., Catl. Punj. & Sind Plt. 9. 1869; C. droserifolia J. L. Stewart in J. As. Soc. Beng. 32: 219- 266. 1863 (not of Del., Fl. Egypt. 250. 1813).
Erect herb, 15–46cm tall, branched, glandular. Leaves simple, alternate, suborbicular to broadly ovate, obtuse-acute, base obtuse-cuneate, glandular, 3–5 nerved, 7–18 × 6–11mm, petiole 4–11mm. Flower complete, 5–7cm across, yellow, pedicel 4–6mm long. Sepals 4, free, glandular, 3–4 × 1.5–2mm, linear or lanceolate, acute. Petals 4, free, 6–8 × 1.5–2.5mm, dimorphic, 2 lanceolate and 2 broadly ovate, acute or obtuse, base attenuate. Stamens 4,
20
Fig. 1.5a. C. dolichostyla : Photograph showing the habit.
21
Fig. 1.5b. C. dolichostyla : Habit (Courtesy Flora of Pakistan, 1973).
22
Fig. 1.6. Distribution pattern of C. fimbriata (●), C. rupicola (▲) and C. dolichostyla (■) .
23 free, 10–12mm. Ovary glandular, gynophore absent, style 1.5–4mm. Capsule oblong, glandular, 12–30 × 2–3mm. Seeds many, glabrous, brown.
Type: W. Pakistan; Sindh, Lower hills, Vicary (K!).
C-6 Peshawar Dist.: Jamrud near Peshawar, 9.11.1963, Hassan ud din 3019 (RAW); G-4 Karachi Dist.: 8 miles from Karachi, Kirthar National Park, 15.1.1982, Kamal Akhter & Malik M. Sharif 1911 (KUH); F-4 Dadu Dist.: On rock near sulphur bed, Laki Shah, 5.5.1959, S.I. Ali 540 (KUH); Laki Shah Sedan, near Sehwan, 6.4.1959, S.M.H. Jafri 2754 (E).
Distribution: Turkmenistan, Iraq, Iran, Afghanistan, Pakistan, Kashmir (Fig. 1.6).
5. Cleome karachiensis (sp. nov.) S. Riaz, R. Abid and M. Qaiser (Figs. 1.8a & b).
Annual herb, stem erect, soft, unbranched, 4–17cm tall, glandular hairy. Leaves alternate, petioled, palmately compound, 3–5 foliate, leaflets entire, obovate-oblanceolate, acute, base cuneate, 6–16 × 3–9mm. Flower yellow, complete, 5–8mm across, pedicel 0.5–1.5cm; Sepals, 4, green, dorsally glandular, oblong–obovate, acute, base truncate, 2–4 × 0.5–1mm; petals 4, yellow, oblanceolate, obtuse, cuneate, attenuate base, 2.5–6 × 1–2.5mm. Stamens 6–8, 2-5mm long. Ovary glandular, upto 5mm long, style persistent, 0.5–1mm long; Capsule linear, glandular, many seeded, 10–29 × 1–2mm. Seeds many, glabrous, light brown, myrtle green.
Type: G-4 Karachi Dist.: Near Department of visual studies, Karachi University campus, 28.8.2014, Sana 8 (KUH!).
G-4 Karachi Dist.: Near Department of visual studies, Karachi University campus, 28.8.2014, Sana 9 (KUH); Near Department of visual studies, Karachi University campus, 4.9.2014, Sana & Rubina Abid 61, 64, 65 (KUH); Near Department of visual studies, Karachi University campus, 16.8.2015, Sana 101, 102, 103 (KUH); Near Department of visual studies, Karachi University campus, 8.9.2015, Sana 108 (KUH); Hub River road, 28.9.2017, Sana & Rubina Abid 149 (KUH).
24
Fig. 1.7a. C. fimbriata : Photograph showing the habit.
25
1.7b. C. fimbriata : Habit (Courtesy Flora of Pakistan, 1973).
26
Distribution: Known from type locality only (Fig. 1.9).
6. Cleome oxypetala Boiss., Diagn. Pl. Or. Nov., Ser. I, 6: 20. 1846; Boiss., Fl. Or. 1: 414. 1867; Blatter, Fl. Arab. 8(1): 37. 1919; Alafemar in Parsa, Flore de’ Iran. 1(1): 913. 1951; Hedge and Lamond in Rechinger f., Fl. Iran. 68: 25. 1970; Jafri in Nasir and Ali, Fl. W. Pak. 34: 30. 1973 (Figs. 1.10a & b).
C. hotsonii Blatt. and Hallb. in J. Ind. Bot., 1: 58. 1919.
Erect herb, 40–60cm tall, branched, eglandular. Leaves simple, alternate, 1–3 nerved, 16–25 × 4–10mm, glabrous, elliptic or elliptic-obovate, acute, obtuse or base cuneate, petiole 5mm. Flower complete, 1cm across, yellow. Sepals 4, free, green, linear, acute, glandular, 1.5 × 0.5mm. Petals 4, free, 6–8 × 1–2mm, lanceolate or ovate, acute or acuminate, base attenuate. Stamens 6, free, 4–8mm long. Ovary 3–5mm, glabrous, gynophore inconspicuous, style 1– 2mm long. Capsule pendulous, glabrous, linear, 45–80 × 3–5mm. Seeds many, hairy, golden brown, dull brown.
Type: Pr. Dalechi, Iran, 3.1842, Kotschy 142 (K!, W, E!, BM!).
G-2 Makran Dist.: 5.5 miles from Mand on way to Turbat, 13.2.1969, S.I. Ali , S.A. Farooqi & S. Abedin 1033 (KUH).
Distribution: Saudi Arabia, Iran, Afghanistan, Pakistan (Fig. 1.2).
7. Cleome pakistanica (Jafri) S. Khatoon and A. Perveen status novum, Pak. J. Bot., 35(2): 145- 146, 2003 (Figs. 1.11a & b).
Cleome heratensis Bunge et Bien. ex Boiss., subsp. pakistanica Jafri in Nasir and Ali, Fl. W. Pak. 34:25, 1973.
Undershrub, 90cm tall, branched, hairy. Leaves simple, very few in number, 8 × 2mm, elliptic, acute or obtuse, base cuneate, hairy, petiole upto 1mm long. Flower, complete, 5mm across, pedicel 4–6mm long. Sepals 4, free, 1–2 × 0.5mm, linear, acute, eglandular. Petals 4, free, linear-oblong, obtuse, base truncate, 2–3 × 0.5mm. Stamens 6, free, filament 5mm
27
Fig. 1.8a. C. karachiensis : Photograph showing the type specimen.
28
Fig. 1.8b. Cleome karachiensis : Type specimen. A, Habit; B, Flower.
29 long, basifixed; ovary glandular, 5mm long, gynophore upto 1mm, style < 1mm, ovules many. Capsule linear, 20–40 × 1–2mm, glandular. Seeds many, hairy, light brown.
Type: V.P. Dutta 5 (RAW!, KUH!).
E-4 Chaghi Dist.: Ahamedwal near Nushki, Baluchistan 11.10. 1934, Ved Parkash Datta 5 (RAW, KUH).
Distribution: Endemic to Pakistan (Fig. 1.9).
8. Cleome rupicola Vicary in J. As. Soc. Bengal 16: 1158. 1847; Jafri in Kew Bull. 174. 1957; Hedge and Lamond in Rechinger f., Fl. Iran. 68: 26. 1970; Jafri in Nasir and Ali, Fl. W. Pak. 34: 33. 1973; Western, Fl. U.A.E. 58.1989; Chaudhary, Fl. K.S.A. 1: 443. 1998 (Figs. 1.13a & b).
C. stocksiana Boiss., Diag. Pl. Or. Nov., Ser.2, 1: 47.1853; Boiss., Fl. Or., 1: 414. 1867; Hook. f. Fl. Brit. Ind., 1: 169.1875; Burtt and Lewis in Kew Bull. 300. 1949.
Erect herb, upto 40cm tall, branched from base, glandular. Leaves simple, 1–3nerved, alternate, ovate-elliptic, elliptic-lanceolate, acute, base cuneate, glandular, 13–19 × 3– 11mm, petiole 4–7mm. Flower complete, 6–7mm across, yellow-brown, pedicel 1–2cm. Sepals 4, green, glandular, linear-oblong, acute, 1.5 × 0.5mm. Petals 4, spathulate, obtuse, base attenuate, 4–5 × 1.5–3mm. Stamens 6, free, 5mm long. Ovary 4mm, glabrous, style persistent, 1mm, gynophore about 1mm. Capsule oblong, sparsely glandular, 50 × 5mm. Seeds many, hairy, dust brown.
Type: Boogta Hills, Scinde, Vicary s.n. (K!).
D-4 Sibi Dist.; Zahree, Baluchistan, 1851, Stocks, J.E. 1110 (K); Kohar, Baluchistan, 1.5.1987, Rubina Akhter s.n. (RAW); D-5 Loralai Dist.; Loralai, 28.3.1943, Mohindarnath 6044 (RAW); E-4 Makran Dist.: Surab, 16.5.1978, S. Nazimuddin , S. Abedin & Hameedullah 557 (KUH); F-3 Makran Dist.: 22 miles from Panjgur on way to Nag, 4.4.1971, Sultan-ul-Abedin & Abrar Hussain 6801, 6802, 6803, 6804, 6805 (KUH); Basima road, 6.4.1971, Sultan ul Abedin &Abrar Hussain 6943 (KUH); 23 miles from Basima on
30
Fig. 1.9. Distribution pattern of C. pakistanica (▲) and C. karachiensis (●) .
31
Fig. 1.10a. C. oxypetala : Photograph showing the habit.
32
Fig. 1.10b. C. oxypetala : A, habit; B, flower; C, fruit (Courtesy Flora of Pakistan, 1973).
33 way to Surab, 6.4.1971, Sultan ul Abedin & Abrar Hussain 6972 (KUH); In way to Basima, 5.4.1971, Sultan ul Abedin & Abrar Hussain 6889 (KUH); F-4 Dadu Dist.: Between Sehwan and Laki Shah Sedan, 6.4.1959, S.M.H. Jafri 2689 (E, KUH) G-2 Makran Dist.: 55 miles from Mand on way to Turbat, 13.2.1969, S.I. Ali, S.A. Farooqi, S. Abedin 1033 (KUH).
Distribution : United Arab Emirates, Oman, Iran, Pakistan, Saudi Arabia (Fig. 1.6).
9. Cleome scaposa DC., Prodr. 1: 239. 1824; Hutchinson and Dalziel, Fl. W. Trop. Africa. 1(1): 87. 1954; Hedge and Lamond in Rechinger f., Fl. Iran. 68: 15. 1970; Jafri in Nasir, E. and Ali, S.I., Fl. W. Pak. 34: 30. 1973; Jafri, Fl. Libya. 12: 12. 1977; Thulin, Fl. Somalia 1: 56. 1993; Raghavan in Sharma and Balakrishnan, Fl. Ind. 2: 314. 1993; Chaudhary, Fl. K.S.A. 1: 441. 1998 (Figs. 1.14a & b).
C.papillosa Stued., Nom. Bot. ed. 2, 1: 382. 1840; Boiss., Fl. Or. 1: 413. 1867; T. Anders. in J. Linn. Soc. 5, Suppl. 1: 3. 1860; Hook. f., Fl. Brit. Ind. 1: 168. 1875; C. gracilis Edgew. in J. As. Soc. Bengal. 16: 1212. 1847; C. linearis Stocks ex T. Anders. in J. Linn. Soc. 5 suppl. 1: 3. 1860.
Erect herb, 15–30cm tall, branched, hairy. Leaves simple, 10–21 × 8–18mm, glandular, alternate, elliptic, ovate or orbicular, acute or obtuse, base obtuse or cuneate, petiole 2– 20mm. Flower complete, 3–4mm across, white-yellow or sometimes pinkish. Sepals 4, free, green, glandular, rarely glabrous, lanceolate, acute, base truncate, 1.5–2.0 × 0.5mm. Petals 4, free, 3–5 × 1–2mm, oblanceolate-spathulate, obtuse or rarely acute-obtuse, base attenuate. Stamens 6, free, 1–5mm long. Ovary glabrous, 1–5mm long, gynophore absent, style 0.5– 1mm long. Capsule linear, glabrous, 7–35 × 1mm. Seeds many, glabrous, metal green or black.
Type: Egypt, Collector not Known (G).
B-6 Nowshera Dist.: 4.10.1988, A. Ghafoor & Tahir Ali 3910 (KUH); Jhagra forest nursery near Peshawar, Jan Muhammad s.n. (RAW); Thal, N.W. Frontier Desserts, 6.9.1956, Sadiq Masih 10 (KUH); B-7 Hazara Dist.: Khanpur, 14.9.1957, R. R. Stewart 28896 (E); C- 6 Kohat Dist.: N. W. Frontier deserts, 7.9.1956, Sadiq Masih 11 (KUH); Near the bank of Man, Indus, Kalabagh, 27.8.1971, Sultan-ul-Abedin & M. Qaiser 9328 (KUH);
34
Fig. 1.11a. C. pakistanica : Photograph showing the habit.
35
Fig. 1.11b. C. pakistanica : A, habit; B, flower; C, fruit (Courtesy Flora of Pakistan, 1973).
36
C-7 Rawalpindi Dist., Jafri Park, Rawalpindi, Punjab, 12.9.1927, R. R. Stewart 9491 (RAW); Rawalpindi-Kahuta, 15.8.1958, B.L. Burtt B IIII (E); Ayub Park, Rawalpindi, Punjab, 16.8.1962, M.A. Siddiqui 909 (RAW); About 10km from Chakwal on way to Chao Saidan Shah, 10.9.1989, Tahir Ali & Tufail Ahmed 1867 (KUH); Kala Chitta Hills, about 10km from Fateh Jang towards Attock, Dist. Attock, 12.10.1988, A. Ghafoor & Tahir Ali 4222 (KUH); 5 miles from T.B. Khan on way to Darband, 14.10.1977, Kamal Akhter & S. Nazimuddin 702 (KUH); Dry river bed near village Kotha between Khushab & Pail on Khushab-Chakwal road, 1.10.1988, A. Ghafoor & Tahir Ali 3769 (KUH); Between Hassan abdal & Attock, 4.10.1988, A. Ghafoor & Tahir Ali 3902 (KUH); About 45km from Mirpur on way to Kotli, 14.9.1987, Tahir Ali, M. Qaiser & M. Ajmal Khan 27 (KUH); 8km from Mirpur on way to Bhimber, 4.9.1987, Tahir Ali, M. Qaiser & M. Ajmal Khan 67 (KUH); Salt range ca. 12km from Choa Saidan Shah on road to Khewra & Pind Dadan Khan, A. Ghafoor & Tahir Ali 4266 (KUH); D-5 Zhob Dist.: between D.G. Khan & Zhob, 27.9.1988, M. Qaiser & A. Ghafoor 4867 (KUH); D-6 D.G. Khan Dist.: About 6 miles before Sakhi Sarwar on way to D.G. Khan, 17.9.1970, M. Qaiser 2563 (KUH); D-7 Jhelum Dist.: 1 mile from Khewra on way to Choa Sedan Shah, 25.9.1970, S.A. Farooqi & M. Qaiser 2843 (KUH); E-4 Kalat Dist.: about 20km W. of Bela, W. side of Jan pass, Baluchistan, 9.4.1965, Jennifer Lamond 309 (E); F-2 Makran Dist.: Between Turbat & Khoshab, 29.2.1988, S. Omer , M. Qaiser &Y. Nasir 2120 (RAW); F-3 Makran Dist.: Awaran to Hoshab, Baluchistan,10.4.1965, Jennifer Lamond 348 (E); Lak Pass ca. 5km from Bela on way to Awaran, near Asant, 20.9.1986, A. Ghafoor & S. Omer 1681 (KUH);6 miles from Bela on way to Awaran, 20.4.1970, M. Qaiser, A. Raza & Abrar Hussain 729 (KUH); Between Turbat & Hoshab, 29.2.1988, S. Omer, M. Qaiser & Y. Nasir 2120 (KUH); Shangla region, 1936, Sultan Ahmed 948 (KUH); Dry stream bed, about 6 miles from Awaran on way to Jhal Jhoa, Bella, 11.4.1972, A. Ghafoor & M. Qaiser 386 (KUH); F-4 Lasbella Dist.: Hub Duregi road, Kirthar, 23.6.2005, Anjum Perveen & Ishtiaq Hussain 139 (KUH); Gorakh, Kirthar, 30.5.2005, Anjum Perveen & Ishtiaq Hussain 107 (KUH); about 96 miles from Karachi on way to Dist. Bela, 10.10.1978, Kamal Akhter Malik & S. Nazimuddin 912 (KUH); About 10 miles from Sehwan on way to Jam shoro, 6.9.1970, S.A. Farooqi & M. Qaiser 2197 (KUH); 16 miles from Hub on way to Shah Noorani, M. Qaiser, Saeeda Qureshi & Asad Raza 508
37
Fig. 1.12. Distribution pattern of C. scaposa (●) and C. brachycarpa (▲) .
38
Fig.1.13a. C. rupicola : Photograph showing the habit.
39
Fig. 1.13b. C. rupicola : Habit.
40
(KUH); Between Bela and Uthal, Baluchistan, 13.9.1986, A. Ghafoor & S. Omer 1629 (KUH); About 5 miles from Laki on way to sehwan, 6.11.1982, Kamal A. Malik, M. Qaiser, Saood Omer & Gohar Khan 2123 (KUH); 4 miles from Shah Hassan on way to Barari, Dadu Dist., 1.5.2976, Nazim, S.Abedin & M. Qaiser 63 (KUH); About 48km from Lasbella on way to Wadh and Khuzdar, 1.5.1990, A. Ghafoor & Steve M. Goodman 4938 (KUH); Karachi to Dadu, ca. 10–15km, S. Sehwan, 5.5.1965, Jennifer Lamond 863a (E); Manachura, 10 miles from Naigaj to Karachi mountain, 8.11.1982, Kamal A. Malik, M. Qaiser, Saood Omer & Gohar Khan 2139 (KUH); Naigaj, Dist. Dadu, 4.5.1976, Nazim, S. Abedin & M. Qaiser 113 (KUH); About 2km from Laki Shah Saddar on way to Sehwan, 16.3.1983, Kamal A. Malik, Saood Omer & Abdul Wahid 2246 (KUH); Naigaj, Talika Johi, Dist. Dadu, 3.5.1976, Nazim, S. Abedin & M. Qaiser 99 (KUH); About 4 miles from Khanote on way to Sehwan, 6.11.1982, Kamal A. Malik, M. Qaiser, Saood Omer & Gohar Khan 2109 (KUH); F-5 Khairpur Dist.: Kot diji, Khairpur Division, 27.10.1955, S.M.H. Jafri 1083 (E, KUH); Dokri, Rice research station, 17.10.1957, Mushtaq Hussain s.n.(KUH); G-2 Makran Dist.: About 3km from village Kikin in Kikin Kor Nullah, between village Shahrak & Turbat on road to Hoshab, 13.3.1990, A. Ghafoor & Steve M. Goodman 4581 (KUH); G-3 Bela Dist.: About 60km from Wadh on way to Bela, 25.10.1986, A. Ghafoor & S. Omer 2247 (KUH); Between Jhal Jhao and Bela, 29.11.1988, M. Qaiser, S. Omer & T. Ahmed 8187 (KUH); 2 miles from Bela on way to Khuzdar, 11.10.1978, Kamal Akhter & S. Nazimuddin 938 (KUH); G-4 Karachi Dist.:Guest house, K.U. Campus, Karachi, 12.9.1967, Mr. Abrar Hussain s.n. (KUH); Darsano Chano, 28.8.1986, S. Khatoon, A. Perveen & Abrar Hussain 232 (KUH); About 28 miles from Karachi on way to Thatta, 23.8.1920, Sultan ul Abedin 5210 (KUH); Manghopir, Karachi, 18.7.1956, S.M.H. Jafri 1512 (KUH); Grand Hotel Malir, Karachi, 23.9.1967, Mr. Abrar Hussain s.n. (KUH); Kirthar, Kirthar National Park, S.W. Sindh, 22.11.2001, Dr. Rubina Akhter 096 (RAW); Kambhu, Kirthar National Park, Dadu Dist., South Sindh, 17.11.2011, Dr. Rubina Akhter 889 (RAW); Road side, Karachi city, 8.10.2003, Dr. Rubina Akhter 101 (RAW); Drig road, Karachi, 16.10.1952, S. M. H. Jafri s.n. (E); Near Karchat, Kirthar, Anjum perveen & Ishtiaq Hussain 94 (KUH); Kirthar, Anjum Perveen & Ishtiaq Ahmed 222 (KUH); Nursery, University campus, Karachi, 10.9.1969, Abrar Hussain s.n.(KUH); Malir, Karachi,
41
Fig. 1.14a. C. scaposa : Photograph showing the habit.
42
Fig. 1.14b. C. scaposa : Habit.
43
10.10.1958, S.I. Ali s.n. (KUH); Super Highway near tool plaza, Karachi, 2.9.2011, Arshad Gabol s.n. (KUH); University campus, Karachi, 14.11.1962, M. Faheem uddin s.n. (KUH); Malir fields, Karachi, 21.8.1967, Abrar Hussain s.n. (KUH); Gujar nala, Karachi, 5.10.1959, Tausif s.n. (KUH); Nazimabad, Karachi, 23.11.1958, Shamim Akhter s.n. (KUH); Drig road, Karachi, 8.1951, S.M.H. Jafri s.n. (KUH); G-5 Hyderabad Dist.: 5 miles from Sehwan on way to Hyderabad, 30.4.1976, Nazim, S. Abedin & M. Qaiser 14 (KUH); 10 miles from Sehwan on way to Hyderabad, 30.4.1976, Nazim, S. Abedin & M. Qaiser 33 (KUH); About 16 miles from Bulla on way to Khyber, 4.6.1970, M. Qaiser & A. Ghafoor 1153 (KUH); Thana Bulla Khan, Hyderabad, 20.2.1959, Ali 454 (KUH); G-6 Tharparker Dist.: Karanjuhill, Nagarparkar, 26.10.2002, G.R. Sarwar, Qurban & Zamarrud 1288 (KUH); Karanjuhill, Nagarparkar, 20.4.2003, Zammarrud, Adil & G.R. Sarwar 1413 (KUH).
Distribution: West to Mauritania, Sudan, Ethiopia, Kenya, N. Tanzania, Trop. Africa, N. Africa, Saudi Arabia, Pakistan, India (Fig. 1.12).
10. Cleome spinosa Jacq., Pl. Carib. 26. 1760; Oliver, Fl. Trop. Afr. 1: 78. 1868; Hutchinson and Dalziel, Fl. W. Trop. Afr. 1(1): 87.1954; Jacobs, Fl. Malesiana 6(1): 102. 1960; Gleason and Cronquist, Man. of Vas. Plts. of North. U. S. and adj. Canada 349. 1963; De Candolle, Prodr. 239. 1824; Jafri in Nasir and Ali, Fl. W. Pak. 34: 22. 1973; Iltis in Nicolson , DeFillips, Nicolson and others, Fl. Dominica 2: 62.1991 (Figs. 1.15a & b).
Erect herb, 60‒120cm tall, branched, hairy. Leaves alternate, compound, 5‒7 foliate, petiolate, lower petioles lagre upto 5.7cm, upper small ca. 2mm, usually with 2 small spiny stipules at the base of petioles upto 4mm long. Leaflets hairy, oblong-lanceolate, acute, base cuneate, 11‒80 × 3‒20mm. Flower complete, about 2.5cm across, white or pinkish, pedicel 1.5‒2cm. Sepals 4, green, 4‒8 × 3mm, linear-lanceolate, acute, glandular. Petals 4, obovate, clawed, base cuneate, 1.5 × 1cm. Stamens 6, free, 3.5cm long; Ovary glabrous, 7 × 0.5mm gynophore ca. 4cm long, style <1mm. Capsule linear, 10‒45 × 1‒3.5mm, glabrous, gynophore 3.3‒8cm. Seeds many, glabrous, reddish brown.
Type: Jamaica, North America, 1730, Person String and W. Houston , s.n. (BM!).
G- 4 Karachi Dist.: Cultivated in Garden as an ornamental, 20.6.1965, Jafri 4010 (KUH).
44
Distribution: Native to tropical South America, Cultivated in S.E. Asia, India (West Bengal), Pakistan, Malaysia, Java, sometimes wild in Java (Fig. 1.17).
11. Cleome viscosa L., Sp. Pl. 672. 1753; Oliver, Fl. Trop. Afr. 1: 80. 1868; Hook. f., Fl. Brit. Ind. I: 170. 1875; Blatter, Fl. Arab.8(1): 39. 1919; Hutchinson and Dalziel, Fl. W. Trop. Afr. 1(1): 87. 1954; Hedge and Lamond in Rechinger f., Fl. Iran. 68: 29. 1970; Jafri in Nasir and Ali, Fl. W. Pak. 34: 23. 1973; Whitmore in Hara and Williams, An Enum.Fl. Pl. Nepal, 2: 46. 1979; Blakelock and Townsend, Fl. Iraq 4(1): 149. 1980; Grierson in Grierson and Long, Fl. Bhutan 1(2): 416. 1984; Iltis in Nicolson, DeFillips, Nicolson and others, Fl. Dominica 2: 62. 1991; Raghavan in Sharma and Balakrishnan, Fl. Ind. 2: 317. 1993; Chaudhary, Fl. K.S.A. I: 444. 1998; Mathur in Singh, Singh and Uniyal, Fl. Jam. and Kash. I: 604. 2002; Mia, Ara and Khan in Khanum and Ara, Fl. Bangladesh. 57: 11. 2007 (Figs. 1.16a & b).
C. icosandra L., Sp. Pl. 938. 1753; Polinisia viscosa (L.) DC., Prodr. I: 242. 1824; Polinisia icosandra (L.) Wight and Arn., Prodr. Fl. Pen. Ind. Orient. 22. 1834. Arivela viscosa Refq., Sylva Tellurina. 110. 1838; Zhang and Tucker, Fl. China. 7: 429. 2008; Tucker and Vanderpool, Fl. N. America 7: 199-222. 2010.
Erect herb, upto 100cm tall, branched or unbranched with glandular hairs. Leaves compound, alternate, 3‒5 foliate, petiole 5–43mm, leaflets elliptic, broad elliptic, oblanceolate, lanceolate or obovate, acute or obtuse, base cuneate, attenuate, glandular, 14–32 × 3–17mm. Flower complete, 7–15mm across, yellow, pedicel 0.8–1.5cm. Sepals 4, green, glandular, linear-oblong, oblong or oblong-lanceolate, acute, 4–6 × 1–2mm. Petals 4, oblanceolate, oblong-lanceolate or oblong-elliptic, obtuse, base attenuate,6–10 × 1– 3mm. Stamens 10–20, free, 3–10mm long, unequal in length. Ovary glandular, 2.5–8.5mm long, style persistant, 0.5–1mm long, in fruiting stage 2–6mm long, gynophore absent. Capsule linear rarely slightly curved, acute, base cuneate, glandular, striate, 37–80 × 3mm, petiole 17–28mm. Seeds many, glabrous, rust brown to dark brown.
Type: Srilanka, Hermann s.n. (BM!, LINN).
45
Fig. 1.15a. C. spinosa : Photograph showing the habit.
46
Fig. 1.15b. C. spinosa : Habit (Courtesy Flora of Pakistan, 1973).
47
A-7 Chitral Dist.: Tukjoshu, Chitral, 22.7.1956, A. R. Beg 1451 (RAW); Chitral, 20.7.1971, Sultan-ul-Abedin 7989 (KUH); Chitral city, 15.9.2004, Jan Alam & Fazal-e-Khaliq 2920b (KUH); B-6 Chitral Dist.: Drosh, Chitral, 14.7.1963, M.A. Siddiqui 2252 (RAW); Gahreet, Drosh area, Chitral, 20.9.2000, M. Rashid Awan & Jamshed Saqib 1076 (PMNH); Shishi Gol, N.E. of Drosh, Chitral, 16.8.1958, J. D. A. stainton 3154 (E); Peshawar-Nowshera, 11.7.1958, B.L.Burtt 984 (E); Near Airport, Chitral, 21.7.1971, Sultan-ul-Abedin 8104 (KUH); B-7 Swat Dist.: 25 miles from Dir on way to Tamir Garh, 25.9.1971, Sultan-ul- Abedin 8174 (KUH); Tamir garh, 17.7.1971, Sultan-ul-Abedin 7790 (KUH); Near Dir, 20.7.1976, M. Qaiser & A. Ghafoor 6455 (KUH); 6 miles from Kawas on way to Naran, 30.8.1972, M. Qaiser & Azmat Ali 5177 (KUH); Barawal bandi, Dir, 23.7,1972, Farrukh Hussain 7085 (RAW); Kishan ganga Valley on the road to Nanga parbat, Titwal, Kashmir, 10.7.1931, R. R. & I. D. Stewart s.n. (RAW); Muzzafarabad, 28.8.1981, S. Malik & Ashfaq 1813 (PMNH); Patikka, Muzaffarabad, A.J.K, 1.8.1988, Saleem Ahmed & M. Ashfaq 104 (PMNH); Palas, Kohistan, NWFP, 23.9.2001, M. Rashid , Naeem & Jamshed 40367 (PMNH); Chakdara, 16.7,1971, Sultan-ul-Abedin 7737 (KUH); Sanghag Hills, Mardan, NWFP, 20.9.1985, Zabta Khan & Jawed Iqbal s.n. (PMNH); 45 miles from Farm on way to Gilgit, 14.8.1971, Sultan-ul-Abedin & M. Qaiser 9079 (KUH); Between Dasu & Pattan, 16.7.1990, M. Qaiser , S. Omer & S.Z. Hussain 8265 (KUH); Dasu, between Besham & Chilas, 1.10.1989, S. I. Ali, W. Sugong & Tahir Ali 3176 (KUH); C-7 Rawalpindi Dist.: Topi Park, Rawalpindi, Punjab, 28.7.1930, R.R. Stewart s.n. (RAW); Tarbela±1200’, 31.8.1975, E & Y. Nasir s.n. (RAW); Noorpur Shahan, Rawalpindi, Punjab, soil stony, 24.8.1977, Ayaz & Dilawar 23 (ISL); University to Malpur, Rawalpindi, Punjab, 22.10.1978, Nisar, Dilawar & Akram 431 (ISL); Dursha Khail ±2000’ Haripur-Rehana road Hazara, 5.9.1986, Y. Nasir , Rubina Akhter & Hanif 12055 (RAW); In compound of Mirpur rest house, Azad Kashmir, 14.9.1987, Tahir Ali, M. Qaiser & M. Ajmal Khan 17 (KUH); 8km from Mirpur on way to Bhimlar, 14.9.1987, Tahir Ali, M. Qaiser & M. Ajmal Khan 65 (KUH); Dist. Sargodha, Sakesar Range, 14.8.1972, M. Qaiser & Azhar Ali 4472 (KUH); About 30 miles from Sakesar on way to Musakhel, Punjab, 9.9.1989, Tahir Ali & Tufail Ahmed 1838 (KUH); 16 miles from Rawalpindi on way to Kahota, 7.8.1971, Sultan-ul-Abedin & M. Qaiser 8586 (KUH); Dist. Jhelum, Ismailwal Cement Factory, 9.10.1965, E. Nasir & M.A. Siddiqui 3639 (RAW); Islamabad, Punjab, 12.6.1982, Muqarrab Shah 3186 (PMNH); 8km from Mirpur
48
Fig. 1.16a. C. viscosa : Photograph showing the habit.
49
Fig. 1.16b. C. viscosa : Illustration showing A, habit; B, flower.
50 on way to Bhimlar, 14.9.1987, M. Qaiser & M. Ajmal 65 (KUH); D-5 Loralai Dist.: Gadebar Forest, Baluchistan, 19.8.1985, Qutabuddin & M. Ashfaq 1996, 1973 (PMNH); D-6 Multan Dist.: About 1km from Layyah on way to Chawk Azam, Chaubara road, Punjab, 30.9.1988, A. Ghafoor & Tahir Ali 3739 (KUH); D-7 Jhelum Dist.: 1 mile from Khewra on way to Choa Sedan Shah, 25.9.1970, S. A. Farooqi & M. Qaiser 2842 (KUH); D-8 Sialkot Dist.: About 250m near village Daurental between Zafarwal and Narowal, 15.10.1988, A. Ghafoor & Tahir Ali 4341 (KUH); Lahore Dist.: 5 miles from Changa Manga on way to Chunian, 9.7.1971, Sultan-ul-Abedin 7556 (KUH); E-6 Rahim Yar Khan Dist: Near primary health centre, Mian wali Qureshian, 25.5.1972, Sultan-ul-Abedin & Abrar Hussain 9745 (KUH); F-4 Ghotki Dist.: About 5km from Ghotki on way to Ubaro, in cultivated fields, 24.9.1988, A. Ghafoor & Tahir Ali 3511 (KUH); F-5 Khairpur Dist.: Khairpur, 20.10.1955, S.M.H. Jafri 80 (KUH); Khairpur, 20.9.1955, S.M.H. Jafri 1003 (KUH); Near village Ratta Tibba, between Luddan and Vehari in cultivated fields, 26.9.1988, A. Ghafoor & Tahir Ali 3593 (KUH); G-4 Karachi Dist.: Nursery, Karachi University campus, Karachi, 18.8.1967, Abrar Hussain s.n. (KUH); Behind Chemistry department, Karachi University campus, 8.8.1970, Sultan-ul-Abedin 5059 (KUH); Nearnursery, University campus, Karachi, 33.8.1967, Abrar Hussain s.n. (KUH); University campus, 1964, Rais Fatima s.n. (KUH); University campus, 19.2.1965, Asif Mohd. Khan s.n. (KUH); Near nursery, University campus, Karachi, 23.8.1967, Abrar Hussain s.n. (KUH); Karachi University campus, 2.8.1986, Anjum Perveen 136 (KUH); Near Chemistry department, University campus, Karachi, 5.10.1967, Khush ejul s.n. (KUH); Nazimabad, Karachi, 25.9.1958, Azhar A. Hussain s.n. (KUH); Kambu, Kirthar, 15.3.2005, Anjum Perveen & Ishtiaq Hussain 4 (KUH); Dumbhar, Kirthar National Park, Karachi Dist., S.W. Sindh, 8.1.2002, Rubina Akhter & Tariq Khan 086 (RAW); Drig road, Karachi, 9.1951, S.M.H. Jafri s.n. (KUH); Karachi, 3.4.1958, Asif Zaidi s.n. (KUH); 21 miles from Karachi on way to Thatta, 23.8.1970, Sultan-ul-Abedin 5231 (KUH); Hawks bay on way to Mubarak village, 10.2016, Anjum Perveen & Shazia Mansoori s.n. (KUH); Dist. Jamshoro, Ranikot, 9.10.2011, G.R. Sarwar, Anjum Perveen & Abrar Ali 2055 (KUH); G-5 Hyderabad Dist.: Campus Agricultural College, Tandojam, 22.8.1959, Asghar Jalis s.n. (RAW); 3 miles from Hyderabad on way to Lahore, 20.9.1969, Sultan-ul-Abedin 3902 (KUH); 3 miles from Hyderabad on way to Sanghar, 20.9.1969, Sultan-ul-Abedin 4007 (KUH); 21 miles from Nawabshah on way to Sanghar, 21.9.1969,
51
Fig. 1.17. Distribution Pattern of C. viscosa (●) and C. spinosa (▲) .
52
Sultan-ul-Abedin 4008, 4009 (KUH); 3 miles from Hyderabad on way to Sanghar, 20.9.1969, Sultan-ul-Abedin 3903 (KUH); 11 miles from Sakrand on way to Sukkur, 17.5.1972, Sultan-ul-Abedin & Abrar Hussain 9461 (KUH).
Distribution: America, West Indies, Tropical Africa, Egypt, Saudi Arabia, Iraq, Afghanistan, Pakistan, India, Nepal, Bangladesh, Bhutan, Burma, Malaysia, Mascarene Isles, Australia, Polynesia, also widespread in tropics of the world (Fig. 1.17).
53
Chapter 2
Seed Micromorphology
Introduction
Seeds are produced by all spermatophytes in the process of reproduction. They have important role in plant dispersal besides that seed morphological data could also be significantly used for plant taxonomy (McClure 1957; Berggren, 1962, 1969; Corner 1976; Hufford, 1995; Otto, 2002; Black, 2007) and to trace out the phylogenetic relationship of taxa (Otto, 2002).
Seed morphological characters have long been used as an important taxonomic tool by various workers and probably Gaertner (1788-1791) was the first to use the seed characters to classify the plants. Afterward, Berggren (1969, 1981) extensively used seed morphological data to delimit the taxa in various families of north European flora. Chuang and Heckard (1972) correlated seed coat morphology with taxonomy in Cordylanthus (Scrophulariaceae). Lersten and Gunn (1981) studied seed morphology and testa topography in Cicer and concluded that these seed characters also support the transfer of genus Cicer (Fabaceae) from Vicieae to Cicereae. El Naggar (2001) studied seed coat morphology of the family Malvaceae and found that these characters are quite enough to correlate the systematics. Khalik and Maesen (2002) studied seed morphology of 45 taxa and data was used in identification of species of the family Brassicaceae from Egypt. Kanwal, Abid and Qaiser (2009) studied the seed morphological characters for the family Aizoaceae and these characters were used to classify the taxa within the family. Akcin and Binzet (2011) studied micromorphology of nutlets of Onsoma L. (Boraginaceae) and concluded that nutlets characters could be useful to classify the species within the genus. Abid, Ather and Qaiser. (2013) studied the seed morphology of Fumariaceae and correlated it with gross morphology to provide the strength for taxonomic delimitation. Besides this Abid, Ather and Qaiser. (2011, 2015, 2016) also studied the seed morphological characters of many other families like Balsaminaceae, Papaveraceae, Urticaceae, and Abid, Kanwal and Qaiser (2014, 2015)
54 studied the seed morphological characters of cucurbitaceae and some monocot families from Pakistan for strengthening the taxonomic decisions.
Concerning to the seed morphology of the family Capparidaceae there are various reports such as; Iltis, Hall, Cochrane and Sytsma (2011) gave a brief account on seeds of Brassicaceae, Capparaceae and Cleomaceae and the data was used for separate recognition of these families. Sanchez-Acebo (2005) studied 35 species of Cleome L. for seed characteristics and data was significantly used for taxonomic delimitation of taxa within genus. The seeds of 35 species of Cleome were also studied by Inda, Torrecilla, Catalan and Ruiz-Zapata (2008). They gave the importance to seed coat, aril and seed cleft and data was correlated with molecular data to study the phylogeny of genus Cleome . Similarly, Kamel, Abd El-Ghani and El-Bous (2010) described the 10 species of Cleome . from Egypt, where seed characters were also studied for taxonomic delimitation.
Although from the above reports, it is evident that there is considerable work on seed morphology of the genus Cleome but neither the detailed account of seed characters was provided or nor covered the entire species of the genus Cleome found in Pakistan. Presently detailed seed morphological characters are studied to provide the strength to the taxonomic delimitation of the genus Cleome from Pakistan.
Materials and Method:
Mature and healthy seeds of 11 species of the genus Cleome were collected from fresh or herbarium specimens. About 5–10 plants per species and 10–15 seeds per plant (depending on availability of material; Appendix-1) were studied under stereomicroscope (Nikon XN model) and scanning electron microscope (JSM-6380 A). For scanning electron microscopy the mature dried seeds were mounted on metallic stub using double adhesive tape, then gold plated in sputtering chamber for a period of 6 minutes and observed under scanning electron microscope. Seed characters like size, shape, colour, surface, and hilum were studied. The terminology used is in accordance with Lawrence (1970), Radford, Dickison, Massey and Ritchie Bell (1974) and Stearn (1983) with slight modifications.
55
Table 1. Seed micromorphological characteristics of the genus Cleome L.
S. Name of Weight Size Angular/ Colour Shape Surface Hilum no species (g) (mm) Not angular . L × b 1 Cleome ariana 0.00029 1.5 X Not -angular Light brown or Ob -elliptic pyriform with Papillate, p apilae Lateral 1.0 chocolate brown small groove narrow towards lineate the edges 2 C. brachycarpa 0.00014 0.85 X Not -angular Rust brown or El liptic pyriform or o void with Reticulate, f avulariate Lateral 0.85 maroon small groove 3 C. dolichostyla 0.00009 0.95 X Angular Light b rown or Reniform or ob -elliptic Appressedly Lateral 0.9 maroon pyriformwith extremely reticulate narrow groove 4 C. fimbriata 0.000 1 0.8 X Not -angular Musturd brown Reniform with somewhat Reticulate, Lateral 0.5 circular groove pusticulate 5 C. karachiensis 0.00058 1.2 X Not -angular Light brown or Retort iform with long groove Concentrically Lateral 1.0 myrtle green narrow towards the edges ridged, foveate with appressedly colliculate 6 C. oxypetala 0.00139 2.5 X Not -angular Golden brown Obovate with minute Tomentose hairy Lateral 2.2 or dull brown uniformely wide groove 7 C. pakistanica 0.00013 1.5 X 1 Angular Orange -brown Elliptic pyriform with a small Densely p ubescent, Lateral groove hair lineate 8 C. rupicola 0.00151 2.5 X Not -angular Dust b rown Reniform with wide uniform Pillose hairy Lateral 2.25 long groove 9 C. scaposa 0.00018 0.75 X Not -angular Metal green or Re niform with small narrow Alveolate Lateral 0.7 black groove 10 C. spinosa 0.00017 1.0 X Angular Blackish b rown Retortiform with a wide Faintly r eticulate Lateral 1.0 rectangular groove 11 C. viscosa 0.0006 2.0 X Not -angular Rust brown or Retortiform , groove is widen, Concentrically Lateral 1.5 dark brown beaked ridged, foveate
56
Fig. 2.1. Scanning electron micrographs. Cleome ariana : A, seed; B, surface. C. brachycarpa : C, seed; D, surface. C. dolichostyla : E, seed; F, surface. C. fimbriara : G, seed; H, surface. (scale bars: A, E = 200µm; C, G = 100µm; B, F = 50µm; D = 20µm; H = 10µm)
57
Fig. 2.2. Scanning electron micrographs. Cleome karachiensis : A, seed; B, surface. C. oxypetala : C, seed; D, surface. C. pakistanica : E, seed; F, surface. C. rupicola : G, seed; H, surface. (scale bars: A, E = 200µm; B, D, F = 20µm; C, G= 500µm; H = 100µm)
58
Fig. 2.3. Scanning electron micrographs. Cleome scaposa : A, seed; B, surface. C. spinosa : C, seed; D, surface. C. viscosa : E, seed; F, surface. (scale bars: A = 100µm; B = 50µm; C, E = 200; D = 10µm; F = 20µ)
59
Observations and Results:
General seed characteristics of the genus Cleome L.
Seeds 0.75–2.5 × 0.5–2.25mm, angular or not angular, grooved, reniform, retortiform, obovate, elliptic-pyriform, ob-elliptic pyriform or ovoid; light brown, dark brown, chocolate brown, rust brown, dull brown, dust brown, mustard brown, orange- brown, golden brown, myrtle green, metal green, maroon, blackish brown or black; reticulate, appressedly reticulate, faintly reticulate, alveolate, foveate, favulariate, pusticulate, appressedly colliculate, concentrically ridged, glabrous or pubescent, pilose, tomentose or papillate, papilla lineate, hilum lateral, 0.00009–0.00151 g in weight (Table 1) (Figs. 2.1, 2.2, 2.3).
Key to the species
1+ Seeds angular…………………………………………………………………………2
- Seeds not angular..……………………………………………….…………………...4
2+ Seeds retortiform…………………………………………………………… C. spinosa
- Seeds reniform, elliptic pyriform or ob-elliptic pyriform…………………………….3
3+ Seeds surface appressedly reticulate, glabrous………………………. C. dolichostyla
- Seeds surface densely pubescent, hair lineate………………………… C. pakistanica
4+ Seeds pubescent, pilose, tomentose or papillate…………….……………………….5
- Seeds glabrous………………………………………………………………………..7
5+ Seeds ob-elliptic pyriform, groove narrow towards the edge……..……….. C. ariana
- Seeds obovate, reniform, groove uniform throughout the entire length……………..6
6+ Seeds reniform, pillose…..……………………………………………….. C. rupicola
- Seeds obovate, tomentose…..…………………………………………… C. oxypetala
60
7+ Seeds retortiform……………………………………………………………………8
- Seeds elliptic-pyriform, ovoid or reniform…………………………..…………….9
8+ Seeds concentrically ridged, foveate without appressedly colliculate…….C. viscosa
- Seeds concentrically ridged, foveate with appressedly colliculate….. C. karachiensis
9+ Seeds elliptic- pyriform or ovoid, surface reticulate, fovulariate…… C. brachycarpa
- Seeds reniform, surface alveolate, reticulate, pusticulate…………………………..10
10+ Seeds metal green or black, alveolate…………………………………….. C. scaposa
- Seeds mustard brown, reticulate and pusticulate...... C. fimbriata
Discussion:
The genus Cleome has great diversity in their seeds and the seed morphological data can be significantly used in delimiting the taxa within the genus Cleome . The genus Cleome is characterized by the presence of cleft in their seeds (Sanchez-Acebo, 2005; Aparadh and Karadge, 2010). Regarding to the diversity of seeds, there is a variety of shapes like, reniform, retortiform, elliptic pyriform, ovoid, obovate, ob-elliptic pyriform. Amongst these shapes reniform is most dominant, found in C. dolichostyla , C. fimbriata , C. rupicola and C.scaposa . The genus Cleome can be bifurcated on the basis of angular or not angular seeds. Angular seeds are present in C. dolichostyla , C. spinosa and C. pakistanica . Amongst this group C. spinosa remains distinct from other two species by having retortiform seeds and faintly reticulate seed surface pattern. While, C. dolichostyla and C. pakistanica are characterized by the presence of ob-elliptic pyriform and reniform seeds with appressedly reticulate surface or elliptic pyriform seeds with densely pubescent surface respectively. The taxa of the second group having non angular seeds i.e., C. ariana , C. rupicola , C. oxypetala , C. viscosa , C. brachycarpa , C. karachiensis (sp. nov.) ,C. scaposa and C. fimbriata could be further divided in to two subgroups on the basis of hairy or glabrous seeds. C.ariana , C. rupicola and C. oxypetala have hairy seeds. While, the remaining species having glabrous seeds. The species with hairy seeds could be
61
further differentiated from each other due to the presence of different seed shapes and surface patterns such as, C. ariana has ob-elliptic pyriform seeds and papillate surface pattern. While, C. oxypetala having obovate seeds with tomentose hairs and the remaining species, C. rupicola has reniform seeds with pillose hair. Other group with glabrous seeds including C. viscosa , C. karachiensis (sp. nov.), C. brachycarpa , C. scaposa and C. fimbriata could be further distinguished on the basis of different seed shapes. Amongst them C. viscosa and C. karachiensis sharing retortiform seeds while, concentrically ridged, foveate surface is characteristic for C. viscosa and concentrically ridged, foveate with appressedly colliculate surface is present in C. karachiensis (sp. nov.) (Riaz and Abid, 2018). However, C. brachycarpa having elliptic-pyriform and ovoid seeds with reticulate, favulariate surface pattern. The remaining two species C. scaposa and C. fimbriata having reniform seeds but both the species could be further splitted by having alveolate surface in C. scaposa and C. fimbriata is characterized by having reticulate pusticulate seed surface.
62
Chapter 3
Palynology
Introduction
Pollen grains are the most important part of plant not only because of its role in fertilization but also for many other uses in day to day life of humans. Many people around the globe used pollen as food (Linskens and Jorde, 1997) since they are rich in nutrients. Despite this fact, pollen grains are also the cause of many allergies and alleviate the symptoms of asthma. Study of pollen is always a piece of interest for scientists in different fields like pollen are studied in archeology for tracing the vegetation types of past. Palaeobotanists use pollen studies to know about the environment of past eras. Pollen studies are also useful in geochronology and industries like oil and gas industry. Pollen are also helpful in solving criminal cases i.e., in forensic sciences (Coyle, Ladd, Palmbach and Lee, 2001).
After the discovery of microscope and microscopic techniques, many scientists used pollen as a tool to solve taxonomic problems. Grew was the first to observe pollen under microscope in 1640s (Bradbury, 1967). While, Brown (1811) first used the pollen in taxonomic delimitation of taxa of family Proteaceae. After him several workers studied pollen of different families such as pollen of genus Cordia (Boraginaceae) were studied by Nowicke and Ridgway (1973). They significantly correlated pollen morphology with gross morphology. Zavada and De Villiers (2000) studied pollen of two species of Tubuliforidites from sediments of Palaeocene-Ecocene of south Africa. They analyzed that the surface structure of their pollen is similar to the pollen of Asteraceae. Similarly, Verhoeven and Venter (2001) studied the pollen of the family Apocyanaceae and on the basis of pollen morphology three tribes of this family i.e. Peroplocoideae, Secamonoideae and Asclepiadoideae were differentiated. Dawar, Qaiser and Perveen (2002) studied the pollen of Inula and its allied genera. They classified pollen grains of Inula and its allied genera into three types and used the pollen characters to delimit the taxa. Similarly, Furness and Rudall (2004) extensively studied the pollen grains of gymnosperms and angiosperms and correlate
63 pollen structure with the evolutionary success of eudicots. Pollen of Asteraceae were studied by Zafar, Ahmed and Khan (2007). They found that pollen morphological characters are important in taxonomy of this family. Perveen and Qaiser (2001, 2007) Studied the pollen for flora of Pakistan and published a series of papers on various families like Capparidaceae, Caprifoliaceae, Rubiaceae, Valerianaceae, Verbenaceae and data was extensively used to support the taxonomic delimitation.
Regarding to the pollen morphology of the family Capparidaceae, Pope (1925) studied the pollen morphology of 85 families from Colorado including Capparidaceae and the data was used to found out the relationships among various taxa. Erdtman (1952) studied the pollen morphology of angiosperms including Capparidaceae and data was correlated with gross morphology for taxonomic purposes. Mitra (1978) studied the pollen morphology of the family Capparidaceae including some species of the genus Cleome and found that pollen morphology was useful for delimiting taxa in relation to seed characters. Chaturvedi and Gupta (1983) studied pollen morphology of the genus Capparis for specific delimitation. Perveen and Qaiser (2001) also studied pollen morphological characters of the family Capparidaceae from Pakistan including 8 species of genus Cleome and concluded that pollen characters were useful in specific delimitation. Sanchez-Acebo (2005) studied pollen of 35 species in Cleome and concluded that pollen morphological characters were found useful in phylogenetic studies of the genus Cleome . Inda, Torrecilla, Catalan and Ruiz-Zapata (2008) studied pollen grains ornamentation of 35 species of Cleome . Similarly, Edeoga, Omosun, Osuagwu, Mbaebie and Madu (2009) studied 3 species of Cleome and importance was given to only shape and aperture type of pollen and it was suggested that pollen characters can be used in systematics of this genus.
In the present study 8 species of the genus Cleome viz., C. ariana , C. brachycarpa , C. dolichostyla , C. karachiensis , C. pakistanica , C. rupicola , C. scaposa and C. viscosa are investigated for their detail pollen characters like, shape, size, tectum and colpal membrane surface.
64
Table 2. Pollen characteristics of the genus Cleome L.
S. Length (µm) Breadth (µm) Colpus length Name of P/E Shape Apertures No. (P) (E) (µm) species 13.8(14.5)15.1 14.3(14.05)13.8 12.5(12.9)13.4 1 Cleome ariana 1.03 Prolate-spheroidal Tricolporate ±0.4737 ±0.1950 ±0.3018 24.8(25.7)26.1 17(17.15)17.8 21.6(21.9)22.2 2 C. brachycarpa 1.502 Prolate Tricolporate ±0.5249 ± 0.2408 ±0.1943 26.7(28.1)29.4 18.8(20.15)21.5 24.1(25.5)26.7 3 C. dolichostyla 1.39 Prolate Tricolporate ±0.9556 ±0.8945 ±0.8595 14.5(15.57)16.1 10.4(12.07)13.1 11.5(12.5)13.8 4 C. karachiensis 1.28 Subprolate Tricolporate ±0.72 ±1.16 ±0.86 19(19.5)20.1 16.5(17)18 16.01(16.6)17.1 5 C. pakistanica 1.56 Prolate Tricolporate ±0.3351 ±0.4515 ±0.3391 14(15.32)16.5 8.71(9.57)10.7 11.2(12.9)14.7 6 C. rupicola 1.60 Prolate Tricolporate ±0.9807 ±0.8857 ±1.3428 12.8(13.4)14.1 8.97(10.39)11.7 11.1(11.42)12.5 7 C. scaposa 1.28 Subprolate Tricolporate ±0.4909 ±1.1010 ±0.5423 20(22.5)24.3 19(19.1)19.2 19(19.6)20.1 8 C. viscosa 1.17 Subprolate Tricolporate ±1.3965 ±0.0942 ±0.2981
65
Table 3. Pollen characteristics of the genus Cleome L.
Polar S. Colpal Mesocolpium Apocolpium Name of area Exine thickness Tectum No. membrane (µm) (µm) species index Rugulate- 10(10.4)10.9 2.51(2.64)2.78 1.51(2.05)2.51 1 Cleome ariana 0.74 Rugulate-reticulate reticulate ±0.4055 ±0.0945 ±0.2485
8.2(8.55)9.1 5.0(5.71)6.2 1.15(2.21)2.56 Spinulose 2 C. brachycarpa Spinulose 0.49 ±0.3593 ±0.4507 ±0.5170
Densely 12.6(13.2)13.6 5.47(6.82)7.57 Reticulate, 3 C. dolichostyla 0.65 1.75(2.81)3.15 granulated ±0.3887 ±0.6153 ±0.5420 scrobiculous
6.02(6.76)7.51 4(4.13)4.3 1.43(1.95)2.51 4 C. karachiensis Granulated 0.56 Verrucate ±0.47 ±0.09 ±0.3817
12.8(13.3)13.9 3.98(4.13)4.5 2.1(2.80)3.5 5 C. pakistanica Sub-psilate 0.78 Rugulate-reticulate ±0.4269 ±0.2025 ±0.4822
3.86(5.28)6.04 2.10(2.45)2.68 1.31(1.59)2.05 6 C. rupicola Sub-psilate 0.55 Rugulate-striate ±0.8177 ±0.2519 ±0.2677
Verrucate, 6.39(7.11)7.84 3.06(4.25)5.19 1.81(1.94)2.08 Spinulose- 7 C. scaposa 0.68 scabrate ±1.0253 ±0.9442 ±0.1042 verrucate
15(15.33)15.8 5.94(6.37)6.77 2.24(2.66)3.48 8 C. viscosa Verrucate 0.80 Reticulate-rugulate ±0.2843 ±0.4162 ±0.4150
66
Materials and Method:
Polleniferous material was taken from fresh and herbarium specimens (Appendix II). Slides and stubs were prepared by acetolysis technique following Erdtman (1952). Pollen grains were mounted in unstained glycerin jelly for light microscopy. For Electron microscopy pollen material was directly mounted with capillary tubes on metallic stubs using double adhesive tape, dried at room temperature then gold plated in sputtering chamber and observed under scanning electron microscope. Following measurements were recorded: Polar length, equatorial diameter, colpi length, apocolpium, mesocolpium, thickness of exine, tectum surface and colpi surfaces were also observed. The terminologies used are in accordance with Erdtman (1952) and Punt, Hoen, Blackmore, Nilsson and Le Thomas (2007).
Observations and Results:
Pollen characteristics of the genus Cleome L.
Pollen grains prolate, sub prolate or prolate-spheroidal; tricolporate, trilobed or weakly trilobed; size (P) 12.8‒29.4µm, Equatorial diameter (E) 8.71‒21.5µm, P/E ratio 1.03–1.60; Colpi 11.1–26.7µm long, sunken with acute ends, colpal membrane regulate-reticulate, densely granulated, granulated, sub-psilate, spinulose, scabrate or verrucate; mesocolpium 3.86–15.8µm, apocolpium 2.10–7.57µm, Exine thickness 1.15–3.5µm; tectum rugulate– reticulate, spinulose, reticulate, scrobiculous, verrucate, regulate-reticulate, regulate-striate or spinulose-verrucate (Tables 2-3)(Figs. 3.1, 3.2).
Key to the species based on pollen characters of the genus Cleome L.
1 + pollen prolate-spheroidal ………………………………………...………… C. ariana
- Pollen prolate or subprolate.……………………………………….……………..….2
2 + pollen prolate……………………………………………...………….………………3
67
- Pollen subprolate…………………………………………….………….……………6
3 + Colpal membrane densely granulated or spinulose………….……...………………4
- Colpal membrane sub-psilate……………..……………………………….…………5
4 + Pollen length 24.8‒26.1µm, P.A.I. 0.49, tectum spinulose………….. C. brachycarpa
- Pollen length 26.7‒29.4µm, P.A.I. 0.65, tectum reticulate, suborbiculous………… ….………………………………………………..……….…………….., C. dolichostyla
5 + Pollen length 19‒20.1µm, P.A.I. 0.78, tectum rugulate-reticulate…….C. pakistanica
- Pollen length 14‒16.5µm, P.A.I. 0.55, tectum rugulate-striate……….….C. rupicola
6 + Tectum reticulate-rugulate………………………………………………… C. viscosa
- Tectum verrucate or spinulose-verrucate..……………………………………………7
7 + Pollen length 14.5‒16.1µm, P.A.I. 0.56, colpal membrane granulated………………
……………………………………………………………………..…C. karachiensis
- Pollen length 12.8‒14.1µm, P.A.I. 0.68, colpal membrane verrucate, scabrate……… ……………………………………………………………………………... C. scaposa
Discussion:
The genus Cleome is characterized by tricolporate pollen grains (Sanchez-Acebo, 2005), sunken colpi with acute apex and pollen are more or less trilobed. On the other hand, great diversity was observed in terms of tectum and colpal membrane surface. The genus Cleome can be classified into three groups on the basis of pollen shape classes. Group I having prolate-spheroidal pollen consists of a single species i.e., C. ariana . Group II is characterized by prolate pollen comprises four species such as, C. brachycarpa , C. dolichostyla , C. pakistanica and C. rupicola . Further more, these species can be distinguished from each other on the basis of tectum types i.e., C. brachycarpa is characterized by spinulose tectum. C. dolichostyla has reticulate and scrobiculous tectum,
68
C. pakistanica has regulate-reticulate and C. rupicola has regulate-striate tectum. However, regarding to the previous findings there are some contradictions such as, tetracolpate pollen grains (Perveen and Qaiser, 2001) and psilate tectum (Ahmed, Khan and Shaheen, 2010) were reported in C. brachycarpa . While, in the present findings tricolporate pollen with spinulose tectum are reported in C. brachycarpa . Group III of the genus Cleome has subprolate pollen. This group comprises C. karachiensis , C. scaposa and C. viscosa . Amongst them C. karachiensis has verrucate tectum, C. scaposa has spinulose-verrucate tectum and C. viscosa is characterized by reticulate-rugulate tectum and these findings are not in accordance with that of the previous findings where reticulate tectum was reported in C. viscosa (Joshi,Kumar and Kothiyal, 2015). Pollen of the genus Cleome also show differences in colpal membrane and quantitative measurements such as, apocolpium, mesocolpium, and colpi length. Thus, from the ongoing discussion it is evident that pollen characters are highly significant and they can be used in strengthening the taxonomic decision.
69
A B
Fig. 3.1. Scanning electron micrographs. C. ariana : A, pollen grain; B, surface. C. brachycarpa : C, pollen grain; D, surface. C. dolichostyla : E, pollen grain; F, surface. C. karachiensis : G, pollen grain; H, surface. (Scale bars: A, G = 2 µm; B, D, F, H = 1 µm; C, E = 5 µm)
70
A B
Fig. 3.2. Scanning electron micrographs. C. pakistanica : A, pollen grain; B, surface. C. rupicola : C, pollen grain; D, surface. C. scaposa : E, pollen grain; F, surface. C. viscosa : G, pollen grain; H, surface (Scale bars: A, C, E = 2 µm; B, D, F, H = 1 µm; G = 5 µm)
71
Chapter 4
Leaf Epidermal Characters
Introduction
Stomata are the small apertures mainly present in leaves either on one side or on both sides. Leaves can easily be classified in to three categories on the basis of presence or absence of stomata i.e., if stomata are present on both surfaces of a leaf then it will be an amphistomatic leaf, if stomata are present on upper surface only, the leaf is said to be epistomatic and if stomata are present only on lower surface, the leaf will be hypostomatic (Perveen, Abid and Fatima, 2007). Similarly, stomata are classified into various types on the basis of arrangement of subsidiary cells around the guard cells such as, Metcalfe and Chalk (1950) classified the stomata for the first time, he proposed four types viz., ranunculaceous, rubiaceous, cruciferous and caryophyllaceous. Later on, Metcalfe and Chalk (1979) recognized 25 types of stomata in dicot plants. Prabhakar (2004) observed 11 types with various subtypes of stomata.
Stomata plays vital role in plants life as they are the structures for gaseous exchange and transpiration. Besides this, plants can be classified on the basis of stomatal types and the data could be used to delimit the various taxa such as, Ahmed (1964) studied stomata of 26 species belonging to 17 genera of Solanaceae from India and South America. He suggested the regrouping of taxa on the basis of stomata and other epidermal characteristics. Grosso, Saint-Martin and Vassal (1994) analyzed the stomatal types in 102 species of the genus Acacia (Fabaceae) from Australia and the data was found useful for specific delimitation. Similarly, Kothresha and Seetharam (1995) studied stomatal types along with other epidermal characters in 12 species and 3 varieties of Bauhinia L. (Fabaceae) from south India and used the data to delimit the taxa. Shah and Kothari (1975) studied stomata and hairs on vegetative and floral parts of 12 species of tribe Trifolieae of the family Papilionaceae. They reported that type of stomata varies in different organs of same species but they did not use the data for taxonomic delimitation of the family. Gill, Olabanji and Husaini (1982) studied type, size and distribution of stomata in 74 species of legumes from
72
Nigeria. They studied the stomatal diversity and concluded that stomata type was not specific for any genus. Abid, Sharmeen and Perveen (2007) described stomatal types in 54 monocot species belonging to 10 families from Karachi, Pakistan and it was observed that each family has particular type of stomata.
Regarding to the stomatal studies for the family Capparidaceae few workers gave attention such as, Aleykutty and Inamdar (1978) described the taxonomic significance of stomata for 16 taxa within the family. Edeoga, Omosun, Osuagwu, Mbaebie and Madu, (2009) studied types of stomata and stomatal frequency in 3 species of Cleome L. along with other micromorphological characters from eastern Nigeria, but they did not use the data for taxonomic purposes. Similarly, Ahmed, Khan, Ahmad, Zafar, Arshad and Ahmad, (2009) studied taxonomic diversity of stomata in 36 dicot species of district tank (KPK, Pakistan) and concluded that the data was useful for taxonomic delimitation. While, Joshi, Kumar and Kothiyal (2015) studied the type and frequency of stomata in C. viscosa only.
Besides stomata, leaves may be covered by trichomes. Trichomes may vary in their structure, size and frequency. They can be glandular or eglandular, unicellular or multicellular. Trichomes may be present either on ventral or dorsal side of leaf or on both sides or sometimes may be present around the stomata. Trichomes are beneficial for plants in many ways. They protect plants from over drying in hot and arid conditions by covering stomata and lowering transpiration rate. While, glandular hairs provide chemical defense to the plant from over grazing by producing repellents and toxins (Theobald, Krahulik and Rollins, 1979). Aleykutty and Inamdar (1980) described the structure and ontogeny of trichomes in 14 families of order Ranales including 52 genera and 119 species where 14 types of trichomes were described based on their structure and contents.
Trichomes also play a significant role in plant classification. Plants can be classified on the basis of presence or absence, type and distribution pattern of trichomes. Prabhakar, Kumar, Ramayya and Leelavathi (1985) studied the morphology and distribution of trichomes on petals of 8 species of Indigofera L. (Fabaceae) and used the data for the specific delimitation within the genus. Khalik (2005) studied trichome morphology of 82 species of the family Brassicaceae from Egypt and data was found useful for taxonomic delimitation within this family. Adedeji, Ajuwon and Babawale (2007) studied trichomes of
73
9 species belonging to 4 genera of the family Solanaceae from Nigeria and found that trichome characteristics are important in differentiating species within this genus. Shaheen, Ajab, Yasmeen and Hayat (2009) studied morphology and distribution of trichomes in 7 species of genus Hibiscus (Malvaceae) from Pakistan and found that the data was important for classifying species within the genus. Similarly, Osman (2012) studied the trichome micromorphology of 5 taxa of Egyptian Ballota (Lamiaceae) and the data was used for specific delimitation. Khokhar, Rajput and Tahir (2012) studied the trichomes in 5 species of the genus Convolvulus (Convolvulaceae) and trichome characteristics were used to classify the species within the genus. Regarding to the trichomes many species of Cleome have received attention but neither the detailed study was undertaken or nor the trichome data was used for taxonomic purposes except that of the work of Aleykutty and Inamdar (1978) who described the structure, ontogeny and taxonomic significance of trichomes in 16 species of 5 genera of the family Capparidaceae where they concluded that the data do not support the separation of Cleomaceae and Capparidaceae. While, Puri (1971) studied the trichomes in C. viscosa only. Similarly, Edeoga, Omosun, Osuagwu, Mbaebie and Madu (2009) described trichomes of three species of Cleome from Nigeria but they did not use this data for taxonomic purpose.
Presently, 11 species of the genus Cleome L. i.e., C. ariana , C. brachycarpa , C. dolichostyla , C. fimbriata , C. karachiensis (sp. nov.), C. oxypetala , C. pakistanica , C. rupicola , C. scaposa , C. spinosa and C. viscosa are studied for their stomata and trichomes.
Materials and Method:
Leaves of 11 taxa were collected from fresh and herbarium specimens (Appendix- III) and studied under stereomicroscope (Nikon XN model) and scanning electron microscope (JSM-6380 A). For scanning electron microscopy, a small part of leaf was mounted on metal stub using double adhesive tape, then gold plated in sputtering chamber for the period of 6 minutes and observed under scanning electron microscope for stomata and trichomes. Types of stomata, size and density and for trichomes their types, structure, base and head cell were studied. Terminologies used with accordance to Metcalfe and Chalk (1950), Lawrence (1970) and Stearn (1983).
74
Table 4. Stomatal characteristics of the genus Cleome L.
Stomatal Aperture S. Type of Aperture frequency Aperture length Name of Species Position width no. stomata shape (No. of µm µm Stomata/mm 2) 5.58(7.64)8.21 2.87(3.01)4.01 1 Cleome ariana Anisocytic Sunken Ovate 231 ±0.8340 ±0.4608 18.6(20)21.4 7.54(8.43)9.21 2 C. brachycarpa Paractyic Sunken Ovate 462 ±1.0672 ±0.6355 At the level of epidermal 16.9(18.35)19.8 8.38(10.14)11.9 3 C. dolichostyla Actinocytic Elliptic 462 cells ±1.2812 ±1.3761 At the level of epidermal 6.58(8.27)9.89 3.1(4.15)5.3 4 C. fimbriara Actinocytic Oblong 539 cells ±1.1537 ±0.8843 At the level of epidermal 5.97(6.86)7.18 1.25(1.42)1.59 5 C. karachiensis Actinocytic Oblong 154 cells ±0.5858 ±0.1237 11.8(13.2)15.5 4.61(5.87)6.85 6 C. oxypetala Actinocytic Sunken Elliptic 308 ±1.3046 ±0.7730 5.56(6.73)7.84 3(3.88)4.96 7 C. pakistanica Actinocytic Sunken Elliptic 385 ±0.9233 ±0.6862 At the level of epidermal 12.3(14.95)17.6 7.76(8.05)8.34 8 C. rupicola Anomocytic Elliptic 385 cells 2.1637 ±0.2367 At the level of epidermal 13(13.6)14.6 3.81(4.47)5.14 9 C. scaposa Anomocytic Elliptic 692 cells ±0.5211 ±0.6223 At the level of epidermal 9.9(10.6)11.3 2.94(4.4)5.87 10 C. spinosa Anomocytic Elliptic 769 cells ±0.6955 ±1.1453 At the level of epidermal 7.5(8.1)9 1.95(2.34)3.48 11 C. viscosa Anomocytic Oblong 769 cells ±0.5429 ±0.6205
75
Table 5. Trichome characteristics of the genus Cleome L.
S. Type of Gladular/ Name of species Indumentum Location Head Trichome type Trichome base no. indumentum eglandular 1 + Mostly towards Multicellular, Cleome ariana Pubescent the midrib and Glandular Peltate Broad smooth margin 2 + Multicellular, C. brachycarpa Pubescent All over leaf Glandular Peltate Broad smooth 3 + Multicellular, C. dolichostyla Hirsute All over leaf Glandular Globular Broad smooth 4 + Irregularly Multicellular, C. fimbriata Scabrous All over leaf Glandular Slightly broad circular smooth 5 + Multicellular, C. karachiensis Pubescent All over leaf Glandular Capitate Broad smooth 6 C. oxypetala ------7 + Scanty, few Unicellular, C. pakistanica Pubescent hairs present all Eglandular Pointed Broad smooth over leaf 8 + Mul ticellular, C. rupicola Pubescent All over leaf Glandular Peltate Broad smooth 9 + Spiny C. scaposa Scabrous All over leaf Eglandular pointed projections on Swollen, broad surface 10 + Multicellular, C. spinosa Dense pubescent All over leaf Eglandular Irregular Broad smooth 11 + Multicellular, C. viscosa Dense pubescent All over leaf Glandular Capitate Broad smooth += present; -= absent
76
Observations and Results:
General stomatal characteristics of the genus Cleome L.
Stomata anisocytic, paracytic, actinocytic or anomocytic, sunken or at the level of epidermal cells, 154–769/mm 2, aperture ovate, elliptic or oblong, aperture 5.56–21.4µm long and 1.25– 11.9µm broad (Table 4) (Figs. 4.1–4.4).
Key to the species based on stomatal characteristics could not be constructed due to the overlapping of characters.
General Trichome Characteristics of the genus Cleome L.
Leaves usually hairy or sometimes glabrous, pubescent, scabrous or hirsute; unicellular or multicellular; glandular or eglandular, unbranched; hairs with or without spiny projections; base broad; head pointed, capitate, peltate, globular, circular or irregular (Table 5) (Figs. 4.5 & 4.6).
Key to species based on trichomes characteristics
1 + Hairs present on leaf….…………..………………………………………………….2
- Hairs absent on leaf…………………………………………………… …C. oxypetala
2 + Hairs glandular.………………………………………………………………………3
- Hairs eglandular.………………………………………………………………….….7
3 + Pubescent hairy………………………………………………………………………4
- Hairs not as above……..…..…………………………………………………………6
4 + Hairs concentrated towards the margin and midrib………………………… C. ariana
- Hairs distributed all over the lamina……………….…..…………………………….5
5 + Hairs head peltate………….…………………………… C. rupicola , C. brachycarpa
77
- Hairs head capitate……..………………………………… C. viscosa , C. karachiensis
6 + Hairs scabrous, head cell irregular in shape………..…………………… C. fimbriata
- Hairs hirsute, head cell globular……..………….…………………… C. dolichostyla
7 + Hairs scabrous with spiny projections……………………………………. C. scaposa
- Hairs pubescent smooth……………..…………..…………………………………...8
8 + Hair multicellular, head cell irregular in shape…….…………………..… C. spinosa
- Hair unicellular and apically pointed….……………………………… C. pakistanica
Discussion:
The genus Cleome is characterized by the presence of anisocytic, anomocytic, actioncytic and paracytic stomatal type. While, Metcalf and Chalk (1950) reported only anomocytic stomata throughout the Capparidaceae. Similar to the findings of Gill, Olabanji and Husaini (1982) leaves are amphistomatic i.e., stomata on both sides of leaf but in the genus Cleome stomata were mainly concentrated towards the dorsal side. Stomata are sunken in C. ariana , C. brachycarpa , C. oxypetala and C. pakistanica. In this group of species there seems to be a good correlation with sunken stomata and small sized leaves to reduce the rate of transpiration. While, the remaining species which have relatively larger leaves having stomata at the level of epidermal cells.
Regarding to the type of stomata there are few reports available on C. viscosa and C. brachycarpa . In these two species similar to the previous findings of Puri (1971) and Perveen, Abid and Fatima (2007) anomocytic and paracytic stomata are reported respectively. However, our findings on C. viscosa are in contrast to the findings of Joshi, Kumar and Kothiyal (2015) where they observed diacytic type of stomata.
Data obtained by the studies of trichomes could be significantly used as an additional tool to provide the strength for taxonomic delimitation of the genus Cleome. Amongst all the studied species C. oxypetala is the only glabrous species as also reported by Jafri (1973), while all the remaining species show diversity in trichomes. Such as, species may be divided
78 into two groups on the basis of glandular and eglandular hairs. The species viz., C. scaposa , C. spinosa and C. pakistanica having eglandular hairs and may be further distinguished i.e., C. scaposa is characterized due to the presence of scabrous hairs with spiny projections. While, C. spinosa and C. pakistanica having pubescent and smooth hairs but both the species remain distinct by the presence of multicellular and unicellular hairs respectively. The another group of taxa with glandular hairs viz., C. ariana , C. brachycarpa , C. dolichostyla , C. fimbriata , C. rupicola , C. viscosa and C. karachiensis may be further delimited on the basis of type, density and structure of trichome head such as, among these 7 species C. fimbriata has scabrous indumentum type with irregularly circular headed trichomes, C. dolichostyla having hirsute indumentum with globular headed hairs and rest of the 5 species have pubescent indumentum. Among these 5 species C. ariana could remain separate from others due to the hairs concentrated only toward the margins and midrib of leaves. While, C. rupicola and C. brachycarpa have peltate trichomes head and C. viscosa and C. karachiensis have capitate trichomes head.
From the preceding discussion it can be concluded that from leaf epidermal studies trichomes characters are more significant than stomatal data to strengthen the taxonomic delimitation of the genus Cleome from Pakistan.
79
A
B
Fig. 4.1. Scanning electron micrographs showing leaf stomata. A, C. ariana ; B, C. brachycarpa ; C, C. dolichostyla (Scale bars: A, B, C = 10 µm).
80
Fig. 4.2. Scanning electron micrographs showing leaf stomata. A, C. fimbriata ; B, C. karachiensis ; C, C. oxypetala (Scale bar: A, B, C = 10 µm).
81
Fig. 4.3. Scanning electron micrographsshowing leaf stomata. A, C. pakistanica ; B, C. rupicola ; C, C. scaposa (Scale bar: A, B, C = 10 µm).
82
Fig. 4.4. Scanning electron micrographs showing leaf stomata. A, C.spinosa ; B, C. viscosa (Scale bar: A, B = 10 µm).
83
A B
Fig. 4.5. Scanning electron micrographs showing trichomes. A, C. ariana ; B, C. brachycarpa ; C, C. dolichostyla ; D, C. fimbriata ; E, C. karachiensis ; F, C. pakistanica (Scale bar: A = 10 µm; B, E = 20 µm; C, D = 100 µm; F= 2 µm ).
84
A B
Fig. 4.6. Scanning Electron micrographs showing trichomes. A, C. rupicola ; B, C. scaposa ; C, C. spinosa ; D, C. viscosa (Scale bar: A, B, C, D = 50 µm).
85
Chapter 5
Phytochemistry
Introduction
Utilization of chemical data in systematics can be traced back by the work of Petiver (1699) and amongst all of the chemicals, phenolic compounds have received more attention for systematic delimitation (Harborne, 1973; Smith, 1976; Crawford, 1978) and to trace out the hybridization (Harborne, 1973). While, flavonoidal data as compared to phenols have considerable potentialities as a taxonomic marker (Fang, Bahorum and Khittoo, 2002).
Regarding to the use of phenolic compounds there are number of reports available such as, Harborne and Williams (1973) studied phenolic compounds for 344 species of Ericaceae and 37 species of related families where they concluded that the data supports the taxonomic delimitation. Similarly, Harborne and Green (1980) analyzed the flavonoids for 97 taxa of the family Oleaceae and data was found significant for classificatory purpose as well as for tracing evolutionary advancement.
Similarly, Haron (1992) studied the distribution and taxonomic significance of leaf flavonoids in 17 species of Eugenia L. (Myrtaceae). Blatt, Santos and Salatino (1998) detected the flavonoids of Bignoniaceae and data was used to delimit the various tribes within this family. While, Fang, Bahorum and Khittoo (2002) studied Eugenia species and a distinct flavonoid pattern in this genus was observed where they concluded that chemotaxonomy provide major source of characters for classification. Abid and Qaiser (2003) studied 21 species of Inula L. and its allied genera for their phenolic compounds and data was used for generic and specific delimitation. Similarly, Marzouk, Al-Nowaihi, Kawashty and Saleh (2010) isolated 38 compouds (flavonols and their derivatives) from 9 species of different tribes of the family Brassicaceae and the data was correlated with gross morphology and anatomy for taxonomic purposes.
Regarding to the chemical constituents for the genus Cleome L. various scientists have made attempts but no one gave the attention to utilize this data for taxonomic purposes.
86
Such as, Wollenweber, Valant-vetschera and Roitman (2007) studied 7 species of Cleome and isolated flavonoids including two novel compounds i.e., 5, 3´,4´-triOH-3, 6, 7, 5´-tetra OMe-flavone from C. felina L. f. and 5, 3´-diOH-3, 7, 8, 4´, 5´-penta OMe-flavone from C. viscosa L. Similarly, Sharaf, El-Ansari and Saleh (1997) also studied flavonoids of four Cleome species and 13 different flavonoid glycosides were identified. Aboushoer, Fathy, Abdel-Kader, Goetz and Omar (2010) isolated four sesquiterpene derivatives from C. droserifolia (Forssk.) Delile. Moreover, Kapoor and Mishra (2013) isolated quercetin and kaempferol from C. viscosa . While, Kasem and Fathy (2016) detected quercetin, kaempferol and their derivatives in 5 species of the genus Cleome .
From the above mentioned reports it is evident that the genus Cleome has been studied for their phenolic compounds but the data was not used for taxonomic delimitation. Similarly, no reports were found on the seed coat elements. The present study is first of its kind to correlate the data of phenolic compounds and seed coat elements for strengthening the specific delimitation of the genus Cleome from Pakistan.
Materials and Methods:
Energy-dispersive X-ray spectroscopy (EDS):
Mature and healthy seeds of 11 species of the genus Cleome were collected from fresh and herbarium specimens. About 5–10 plants per species (Appendix-I) and 10–15 seeds per plant (depending on availability of material) were studied for the seed coat elements. The mature dried seeds were directly mounted on metallic stub, using double adhesive tape, then gold plated in sputtering chamber for a period of 6 minutes and observed under scanning electron microscope for the detection of elements in seed coat (Table 6). Seed coat elements were detected by energy-dispersive X-ray spectroscopy (EDS) using EDS detector (EX-54175jMU) attached to scanning electron microscope (JSM-6380 A). Standardless quantitative analysis was performed by ZAF method.
87
F
Fig. 5.1. Energy-dispersive X-ray spectroscopy (EDS) graphs showing seed coat elemental composition. A, Cleome ariana ; B, C. brachycarpa ; C, C. dolichostyla ; D, C. fimbriata ; E, C. karachiensis (sp. nov.); F, C. oxypetala .
88
Fig. 5.2. Energy dispersive X-ray spectroscopy (EDS) graphs showing seed coat elemental composition. A, Cleome pakistanica ; B, C. rupicola ; C, C. scaposa ; D, C. spinosa ; E, C. viscosa .
89
Phenolic compounds:
Leaves of 7 species of the genus Cleome were analyzed for their phenolic compounds (Appendix-IV). For extraction, 1 gm dried leaves from fresh material or herbarium specimens were extracted with 70% ethanol for 24 hours at room temperature. Extracts were concentrated and chromatographed two dimensionally on thin layer chromatographic (TLC) plates using BAW (n-butanol : acetic acid : water, 4 : 1: 5) versus 15% acetic acid, following standard procedure of Harborne (1973). A list of specimens investigated is given in appendix IV.
Phenolic compounds were identified by comparing with the Rf values and their colour in ultra-violet light before and after ammonia fumigation (Tables 7-10).
Results and Discussion:
Key to the species of the genus Cleome based on seed coat elements
1+ Chlorine present.…..…………………………………………………………………2
- Chlorine absent……………………………………………………………………….5
2+ Silicon present.………………………………………………………… C. pakistanica
- Silicon absent.………………………………………………………………………..3
3+ Sodium present…………………………………………………………… C. spinosa
- Sodium absent….…………………………………………..………………………..4
4+ Calcium present…………………………………………………………… C. scaposa
- Calcium absent………………………………………….. C. rupicola , C. dolichostyla
5+ Copper present………………………………………………………………………..6
- Copper absent………..………………………………………………. C. brachycarpa
6+ Magnesium present……………………….………………………………………….7
90
- Magnesium absent…………………………………………..……………………….9
7+ Aluminum present…………………….…………………………………… C. ariana
- Aluminum absent……………………………………..…………………………….8
8+ Sulphur present..…………………………………………… C. karachiensis (sp.nov.)
- Sulphur absent…………………………………………………………… C. oxypetala
9+ Potassium, Aluminium, Sulphur present…………………………………… C. viscosa
- Potassium, Aluminium, Sulphur absent…………..……………………… C. fimbriata
Detailed analysis of seed coat elements (Table 6; Figs. 5.1 & 5.2) of the genus Cleome revealed that carbon and oxygen are present throughout the species within the genus. While, the species of the genus Cleome can be bifurcated on the basis of presence or absence of chlorine such as, chlorine is present in C. dolichostyla , C. pakistanica , C. rupicola , C. scaposa and C. spinosa . While, chlorine is absent from C. ariana , C. brachycarpa , C. fimbriata , C. karachiensis , C. oxypetala and C. viscosa . From the first group, C. pakistanica can be differentiated from all the other species by the presence of silicon. While, seed coat elements of C. rupicola and C. dolichostyla are similar but they can be differentiated on the basis of their quantitative values. The remaining species C. scaposa could not be separated from other species due to shared seed coat elements but their quantitative values differ from others. Similarly, C. spinosa having sodium also found in C. pakistanica but both the species could be separated due to the distinct quantitative value.
Among the species of another group C. brachycarpa remains distinct from rest of the species due to the absence of copper. The remaining species in this group can be bifurcated on the basis of presence or absence of magnesium such as, C. ariana , C. karachiensis (sp. nov.) and C. oxypetala having magnesium while, magnesium was absent in C. viscosa and C. fimbriata . Furthermore, the species having magnesium may remain distinct due to the presence of aluminum and sulphur in C. ariana and C. karachiensis (sp. nov.) respectively. While in C. oxypetala these two elements were missing. Similarly, the C. viscosa and C. fimbriata may further be separated due to the presence or absence of potassium, aluminum and silicon in respective manner
91
Table 6. Composition of elements in seed coat determined in the genus Cleome L. by using energy-dispersive X-ray spectroscopy (EDS) and the associated analytical program EDS analysis station .
Percentage of Elements in Seed Coat
S. Name of No C O Mg K Cu Zn Cl Ca Na Si Al S species . 1 Cleome ariana 56.58 37.68 0.60 0.96 1.31 - - 0.49 - - 2.39 -
2 C. brachycarpa 52.05 42.03 0.63 3.07 - - - 1.83 - - - 0.39
3 C. dolichostyla 56.70 26.22 0.67 2.09 8.73 4.76 0.82 - - - - -
4 C. fimbriata 66.38 18.09 - - 9.89 5.09 - 0.55 - - - -
5 C. karachiensis 55.58 32.08 1.98 2.43 3.96 - - 3.19 - - - 0.78
6 C. oxypetala 51.94 42.12 1.01 2.13 2.12 - - 0.69 - - - -
7 C. pakistanica 71.13 18.14 0.56 2.05 3.58 - 1.75 0.95 1.40 0.43 - -
8 C. rupicola 51.74 31.65 2.88 4.75 5.38 2.75 0.86 - - - - -
9 C. scaposa 38.65 24.29 1.55 4.64 12.07 6.07 0.52 12.2 - - - -
10 C. spinosa 55.01 32.62 1.24 0.48 3.39 1.57 0.51 1.28 3.89 - - -
11 C. viscosa 50.2 41.8 - 2.54 2.29 1.41 - 0.65 - - 0.62 0.49
Key: C = Carbon, O = Oxygen, Mg = Magnessium, K = Potassium, Cu = Copper, Zn = Zinc, Cl = Chlorine, Ca = Calcium, Na = Sodium, Si = Silicon, Al = Aluminum, S = Sulphur.
92
Table 7. Phenolic acids and flavonols in Cleome L.
S. No. Name of species 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 1 Cleome ariana ------+ + - - - + - + - 2 C. brachycarpa + + - - + - + - + - + - + + + - - 3 C. dolichostyla - - - - + + + + + + - - + + - - - 4 C. karachiensis - - - - - + - - - - + + + + + + + 5 C. rupicola - - - + - + - + - + - - - + - - - 6 C. scaposa ------+ - - + - - + - - + - 7 C. viscosa + + + - + - - + ------
Key: + = Present, - = Absent, 1 = p-coumaric acid, 2 = sinapic acid, 3 = 3, 4, 5’trimethoxy cinnamic acid, 4 = aesculin, 5 = Kaempferol, 6 = Quercetin, 7 = Quercetin 3- Arabinoside, 8 = Quercetin 3- xyloside, 9 = Quercetin 3- glucoside, 10 = Quercetin 3- galactoside, 11 = Quercetin 3- Rhamnoside, 12 = Quercetin 3- Glucuronide, 13 = Quercetin 3- Rutinoside, 14 = Quercetin 7- Glucoside, 15 = Quercetin 4’ Glucoside, 16 = Azaleatin, 17 = Gossypetin.
93
Table 8. Flavones, glycosyl flavones, chalcones, aurones, flavonones and isoflavonones in Cleome L.
Flavones Glycosylflavones Biflavonyl Chalcones Aurones Flavones Isoflavone
S. Name of 1 2 3 4 5 6 7 8 9 10 11 12 13 14 No. species
1 Cleome ariana ------+ - - - - 2 C. brachycarpa - - - + - + + + - + + - + -
3 C. dolichostyla ------+
4 C. karachiensis + - - - + ------5 C. rupicola - - + ------+ - - 6 C. scaposa ------7 C. viscosa - + + - - - + - + - + - - -
Key: + = Present, - = Absent, 1 = Apigenin, 2 = Luteolin, 3 = Chrysoeriol, 4 = Isovitexin, 5 = kayaflavone, 6 = Isoliquiritigenin, 7 = Isoliquiritigenin 4’ glucoside, 8 = Aureosidin, 9 = Aureousidin 4- glucoside, 10 = Aureousidin 6- glucoside, 11 = Hesperitin, 12 = Hesperidin, 13 = Naringin, 14 = Daidzein.
94
Table 9. Frequency of occurance of different phenolic compounds in Cleome L.
S. No. Compounds No. of species Percentage of species Phenolic acids 1 p- coumaric acid 2 28.57 % 2 Sinapic acid 2 28.57 % 3 3,4,5 Trimethoxy cinnamic acid 1 14.28 % 4 Aesculin 1 14.28 %
Flavonols 5 Kaempferol 3 42.85 % 6 Quercetin 7 100.00 % (Including glycosides) 7 Azaleatin 3 42.85 % 8 Gossypetin 1 14.28 %
Flavones 9 Apigenin 1 14.28 % 10 Luteolin 1 14.28 % 11 Chrysoeriol 2 28.57 %
Glycosylflavones 12 Isovitexin 1 14.2 8 %
Biflavonyl 13 Kayaflavone 1 14.28 %
Chalcones 14 Isoliquiritigenin 2 28.57 % (including glucoside)
Aurones 15 Aureusidin 3 42.85 % (including glucosides)
Flavanones 16 Hesperitin 2 28.57 % 17 Hesperidin 1 14.28 % 18 Naringin 1 14.28 %
Isoflavones 19 Daidzein 1 14.28 %
95
Table 10. Unknown compounds in Cleome L.
Rf Values Colour in UV S. No. Name of species BAW 15 % HoAc With Amonia Without Amonia 1 Cleome ariana 34.50 45.50 Fl. Blu. Blu. 21.17 85 .51 Br. Br .
2 C. brachycarpa 29.34 55.19 Br . Br . 46.16 66.47 Fl. Blu. Fl. Blu. 3 C. dolichostyla 35.22 55.19 Fl. Gr. Fl. Gr. 25.78 42.40 Gr. Gr. 4 C. karachiensis 44.38 91.97 Fl. Blu. Blu. 55.05 91.97 Fl. Blu. Blu. 69.10 91.97 Fl. Blu. Blu. 5 C. rupicola 25.1 3.33 Fl. Gr. Gr. 23.89 16.11 Fl. Blu. Blu. 6 C. scaposa 36.93 58.04 Blu. Blu. 7 C. viscosa 37.27 61.62 Blu. Blu.
Key: Fl. = Florescent; Gr. Green; Blu. = Blue; Br. = Brown.
96
Key to the species of the genus Cleome based on
phenolic compounds
1 + Kaempferol present…………………………………………………………………..2
- Kaempferol absent……………………………………………………………………4
2 + p- coumaric acid and sinapic acid present……………………………………………3
- p- coumaric acid and sinapic acid absent……………………………... C. dolichostyla
3 + Isovitexin, isoliquiritigenin, aureusidin and naringin present……….. C. brachycarpa
- Isovitexin, isoliquiritigenin, aureusidin and naringin absent………………. C. viscosa
4 + Aesculin and hesperidin present………………………………………….. C. rupicola
- Aesculin and hesperidin absent………………………………………………………5
5+ Quercetin, gossypetin, apigenin and kayaflavone present…C. karachiensis (sp.nov.)
- Quercetin, gossypetin, apigenin and kayaflavone absent…………………………….6
6 + Quercetin 3-glucoside, quercetin 7-glucoside and aureusidin 6-glucoside present…….
………………………………………………………………………………. C. ariana
- Quercetin 3-glucoside, quercetin 7-glucoside and aureusidin 6-glucoside absent…….. ……………………………………………………………………………… C. scaposa
Thin layer chromatography of aqueous ethanolic extracts of 7 species of genus Cleome resulted in the isolation of 44 phenolic compounds including 13 unknown compounds (Tables 7-10).
The result revealed that quercetin and its derivatives are the most widespread compounds within the genus Cleome . While, genus Cleome can be divided into two groups on the basis of presence or absence of kaempferol. The group of species with kaempferol comprises C. brachycarpa , C. dolichostyla and C. viscosa and the grouping of these three
97 species by sharing similar compounds may also be supported by comparatively larger pollen grains, mesocolpium and apocolpium. The other group of species, which lacks kaempferol viz., C. rupicola , C. karachiensis (sp.nov.), C. ariana and C. scaposa could be well correlated by having smaller pollen grains, mesocolpium and apocolpium. Amongst the species of group I, C. dolichostyla can be distinguished from other two species by the presence of quercetin and diadzein along with simple, suborbicular and cordate leaves. While, the remaining two species C. viscosa and C. brachycarpa are closely related with each other by having palmately compound leaves, elliptic petal shape and rust brown seeds. Furthermore, these two species remain distinct by the exclusive chemical data such as, isovitexin, isoliquiritigenin, aureusidin and naringin in C. brachycarpa and C. viscosa is characterized by luteolin, chrysoeriol, 3, 4, 5’-trimethoxycinamic acid and aureusidin 4- glucoside.
In group II, C. rupicola remains distinct from other three species by the presence of aesculin and hesperidin as well as linear sepals and prolate pollen with rugulate-striate tectum. Similarly, C. karachiensis (sp.nov.) can be differentiated from C. ariana and C. scaposa by the presence of unbranched stem and by having apigenin, gossypetin and kayaflavone. On the other hand, C. ariana and C. scaposa can be differentiated from each other by having different morphologies i.e., compound and simple leaves respectively. Moreover, C. ariana is characterized by having quercetin 3-glucoside, quercetin 7-glucoside and aureusidin 6-glucoside and C. scaposa having quercetin 3- arabinoside and rutin.
By the above discussion, it is evident that the data obtained from the analysis of seed coat elements and phenolic contents from leaves of genus Cleome can be utilized to strengthen taxonomic delimitation.
98
Chapter 6 Numerical Taxonomy
Introduction
Statistical procedures have long been used to analyze the data in various fields of studies as a decision indicating tool. While, Stearn (1968) was opined that this data should not interfere with decision making. Similarly, Cronquist (1964) suggested that the subjective data should not become objective when analyzed. Sokal and Sneath (1963) first developed the concepts of numerical taxonomy. Following them, many scientisits utilized the cladistics to study the relationship among plants such as Walsh and Hoot (2001) studied the phylogenetic relationships of Capsicum (Solanaceae), using DNA sequences from chloroplast and nuclear introns numerically. They proposed informal classification within the genus. Abid and Qaiser (2006) studied phenetic relationships among various taxa of Inula L. (s. str.) and its allied genera from Pakistan and Kashmir. They concluded that it is a heterogenus and paraphyletic genus. Similarly, Sarwar and Qaiser (2012) studied the numerical taxonomy of the genus Rosularia (DC.) Stapf. from Pakistan and Kashmir and concluded that there are two distinct groups present within the genus.
While, Fay and Christenhusz (2010) studied the phylogeny of the order Brassicales by using numerical data based on mustard oil precursor and concluded that Cleomaceae and Brassicaceae were closely related families. Similarly, Sanchez-Acebao (2005) numerically analyzed the phylogeny of the new world Cleome by utilizing DNA profile, seed and pollen data and it was observed that seeds and pollen characters were significant for specific delimitation of the genus. While, Inda, Torrecilla, Catalan and Ruiz-Zapata (2008) worked on the phylogeny of Cleome, Podandrogyne and polinisia and concluded that they are paraphyletic genera. Similarly, Kasem and Fathy (2013) also used the numerical method to study the correlation among Cleome species based on flavonoids and isoenzymes. Later on, Kasem (2016) numerically analyzed the Cleome species by using the data from anatomy, pollen and seed morphological studies to find out the phenetic relationship and it was
99 concluded that the data obtained from pollen and seed exomorphic studies were helpful in making taxonomic decisions at specific level.
Thus, from the above reports, it is evident that numerical data and cladistics have been used by various workers to study taxonomic correlation among the taxa of the family Cleomaceae and the genus Cleome but no comprehensive reports are available on species of Cleome found in Pakistan. Presently, detailed morphological and micromorphological characters are used to determine the phenetic relationship among the species of the genus Cleome present in Pakistan.
Materials and Method:
Numerical analysis of 11 species of the genus Cleome was carried out to study the phenetic relationship. Hierarchical clustering was performed using the Euclidean distance index with the computer package (IBM Corp., 2011). Each taxon was considered an operational taxonomic unit (OTU). Qualitative characters were recorded in binary state as 1, 2 and characters which were either absent or present were coded as 0 or 1 respectively. While, for quantitative characters average values were directly used (Table 11).
Results and Discussion:
Dendrogram obtained from numerical analysis of genus Cleome indicates the presence of two distinct clusters i.e., group-I and group-II (Fig. 6.1).
Group-I is represented by 7 species i.e., C. karachiensis , C. scaposa , C. brachycarpa , C. fimbriata , C. rupicola , C.viscosa and C. oxypetala . Their affinities may be due to average height of plants (up to 50cm). while, group-II attaining > 50cm plant height. Furthermore, group-II also having silicon and sodium in their seed coats, spiny stipules, cordate leaves, anisocytic stomata, irregular trichome head, hirsute hairs and truncate leaf base in one or more species. While, these features are totally missing from group-I.
Group-I is further splitted into two distinct clads viz., I A and I B. In clad I AC. karachiensis , C. scaposa , C. brachycarpa and C. fimbriata are grouped together due to comparatively smaller size of capsules and seeds. Amongst these taxa, C. fimbriata remains
100 separate from other species by having 4 stamens (Jafri, 1973), mustard brown and pusticulate seeds, and absence of magnesium and potassium in seed coat. While, C. karachiensis , C. brachycarpa and C. scaposa form common cluster due to 6 or more stamens and presence of magnesium and potassium in seed coat. Furthermore, C. karachiensis and C. scaposa are coupled together by having subprolate pollen and more or less similar size of pollen grains, colpus, apocolpium, exine thickness and petals and presence of azaleatin in leaves. Although C. karachiensis has morphological affinities with C. viscosa and C. brachycarpa. While, C. karachiensis remains distinct from C. scaposa due to compound leaves, actinocytic stomata, retortiform, concentrically ridged, foveate with appressedly colliculate seeds. C. scaposa having simple leaves, anomocytic stomata, reniform and alveolate seeds. On the other hand, C. brachycarpa falls separately from C. karachiensis and C. scaposa probably due to the presence of prolate pollen, with larger colpus, apocolpium and absence of azaleatin in the leaves.
Group-IB comprised of three species viz., C. rupicola , C. viscosa and C. oxypetala . These three species are linked together due to comparatively larger size of capsules and seeds. While, the close affinities of C. rupicola with C. oxypetala were earlier reported by Jafri (1973) on the basis of similar flower and capsule. The remaining species i.e., C. viscosa stands distinct from both of the species by having capitate trichome head, distinct petal and seed shapes.
Group-II comprised of C. ariana , C. dolichostyla , C. pakistanica and C. spinosa . Amongst them C. spinosa falls separately and characterized by the presence of spiny stipules at the base of leaves, larger leaf and flower. While the remaining taxa viz., C. ariana , C. dolichostyla and C. pakistanica form common cluster due to the presence of smaller leaves and flowers and absence of spiny stipules. Among these three species C. pakistanica could be separated from rest of the two species due to larger plant height, undershrub habit and presence of silicon in seed coats (Riaz and Abid, 2018). While, C. ariana and C. dolichostyla are coupled together by having more or less similar plant height, herbaceous habit and absence of silicon in seed coats. While, both the species could be separated by having compound leaves and 6 stamens in C. ariana and C. dolichostyla is characterized due to the presence of simple leaves and 4 stamens.
101
It is noteworthy that the conclusions obtained by numerical data did not support the close affiliation of species based on gross morphological data and the present findings could be supported with the opinion of Stearn (1968) and Cronquist (1964), where they suggested that numerical analysis represent the synthetic approach which could not always be correlated with subjective findings.
102
Fig. 6.1. Dendrogram showing the relationship of the species of the genus Cleome L.
103
Table 11. List of characters scored for cluster analysis for the species of genus Cleome .
Character s description S. No. Habit 1 Herb (1) undershru b (2) Stem 2 Length ( Cm ) 3 Suberect (1), erect (2) 4 Unbranched (0), branched (1) 5 Hairs: Absent (0), present (1) Leaves 6 Simple (1), Compound (2) Size 7 Length (mm) 8 Breadth (mm) 9 Spiny stipules at base: Absent (0), present (1) Shape: 10 Linear: Absent (0), present (1) 11 Lanceolate: Absent (0), present (1) 12 Oblong: Absent (0), present (1) 13 Ovate: Absent (0), present (1) 14 Broadly ovate: Absent (0), present (1) 15 Orbicular: Absent (0), present (1) 16 Suborbicular: Absent (0 ), present (1) 17 Elliptic: Absent (0), present (1) 18 Broad elliptic: Absent (0), present (1) 19 Cordate: Absent (0), present (1) 20 Oblanceolate: Absent (0), present (1) 21 Obovate: Absent (0), present (1) 22 Linear -elliptic: Absent (0), present (1 ) 23 Broadly cordate: Absent (0), present (1) 24 Suborbicular -broadly ovate: Absent (0), present (1) 25 Obovate -oblanceolate: Absent (0), present (1) 26 Elliptic -obovate: Absent (0), present (1) 27 Ovate -elliptic: Absent (0), present (1) 28 Elliptic -lanceolate: Absent (0), present (1) 29 Oblong -lanceolate: Absent (0), present (1) Apex 30 Acuminate: Absent (0), present (1) 31 Apiculate: Absent (0), present (1) 32 Obtuse: Absent (0), present (1) Base 33 Cuneate: Absent (0), present (1) 34 Obtu se: Absent (0), present (1)
104
35 Attenuate: Absent (0), present (1) 36 Truncate: Absent (0), present (1) Leaf Indumentum 37 Absent (0), present (1) 38 Eglandular (1), glandular (2) Vesture type: 39 Pubescent : Absent (0), present (1) 40 Dense pubesc ent: Absent (0), present (1) 41 Hirsute: Absent (0), present (1) 42 Scabrous: Absent (0), present (1) Trichome head 43 Peltate: Absent (0), present (1) 44 Capitate: Absent (0), present (1) 45 Globular: Absent (0), present (1) 46 Irregularly c ircular : Absent (0), present (1) 47 Pointed: Absent (0), present (1) 48 Irregular: Absent (0), present (1) Leaf stomata Aperture 49 Length (µm) 50 Breadth (µm) Type 51 Anisocytic: Absent (0), present (1) 52 Paracytic: Absent (0), present (1) 53 Actinocytic: Absent (0), present (1) 54 Anomocytic: Absent (0), present (1) Position 55 Sunken: Absent (0), present (1) 56 Slightly sunken: Absent (0), present (1) 57 At the level of epidermis: Absent (0), present (1) Apperture shape 58 Oblon g: Absent (0), present (1) 59 Elliptic: Absent (0), present (1) 60 Ovate: Absent (0), present (1) Flower 61 Size (mm) Sepals Size 62 Length (mm) 63 Breadth (mm) 64 Indumentum: Absent (0), present (1) Shape 65 Linear: Absent (0), present (1) 66 Lanceolate: Absent (0), present (1) 67 Oblong: Absent (0), present (1)
105
68 Ovate: Absent (0), present (1) 69 Elliptic: Absent (0), present (1) 70 Obovate: Absent (0), present (1) 71 Oblong -elliptic: Absent (0), present (1) 72 Oblong -obovate: Absen t (0), present (1) 73 Linear -oblong: Absent (0), present (1) 74 Linear -lanceolate: Absent (0), present (1) 75 Oblong -lanceolate: Absent (0), present (1) Petals Size 76 Length (mm) 77 Breadth (mm) Colour 78 White: Absent (0), present (1) 79 Yel low: Absent (0), present (1) 80 Pinkish: Absent (0), present (1) Shape 81 Linear: Absent (0), present (1) 82 Oblong: Absent (0), present (1) 83 Lanceolate: Absent (0), present (1) 84 Ovate: Absent (0), present (1) 85 Broadly ovate: Absent (0), pres ent (1) 86 Elliptic: Absent (0), present (1) 87 Obovate: Absent (0), present (1) 88 Oblanceolate: Absent (0), present (1) 89 Spathulate: Absent (0), present (1) 90 Linear -oblong: Absent (0), present (1) 91 Obovate -elliptic: Absent (0), present (1) 92 Lanceolate -oblong: Absent (0), present (1) 93 Oblanceolate -spathulate: Absent (0), present (1) 94 Oblong -elliptic: Absent (0), present (1) Apex 95 Acute: Absent (0), present (1) 96 Acuminate: Absent (0), present (1) 97 Obtuse: Absent (0), present (1) 98 Clawed: Absent (0), present (1) 99 Acute -obtuse: Absent (0), present (1) Base 100 Cuneate: Absent (0), present (1) 101 Attenuate: Absent (0), present (1) 102 Truncate: Absent (0), present (1) Gynoecium 103 Gynophore: Absent (0), present (1 ) 104 Hairs: Absent (0), present (1) 105 Carpel length (mm)
106
Androecium 106 Number of stamens Capsule Size 107 Length (mm) 108 Breadth (mm) Shape 109 Linear: Absent (0), present (1) 110 Oblong: Absent (0), present (1) 111 Slightly curved: Ab sent (0), present (1) Seeds Size 112 Length (mm) 113 Breadth (mm) 114 Nonangular (0), Angular (1) Colour 115 Light brown: Absent (0), present (1) 116 Chocolate brown: Absent (0), present (1) 117 Rust brown: Absent (0), present (1) 118 Dark br own: Absent (0), present (1) 119 Dust brown: Absent (0), present (1) 120 Dull brown: Absent (0), present (1) 121 Golden brown: Absent (0), present (1) 122 Mustard brown: Absent (0), present (1) 123 Orange brown: Absent (0), present (1) 124 Metal gree n: Absent (0), present (1) 125 Myrtle green: Absent (0), present (1) 126 Maroon: Absent (0), present (1) 127 Black: Absent (0), present (1) 128 Blackish brown: Absent (0), present (1) Shape 129 Ovoid: Absent (0), present (1) 130 Ob -elliptic pyrifor m: Absent (0), present (1) 131 Retortiform: Absent (0), present (1) 132 Elliptic pyriform: Absent (0), present (1) 133 Reniform: Absent (0), present (1) 134 Obovate: Absent (0), present (1) Surfaces 135 Concentrically ridged: Absent (0), present (1 ) 136 Appressedly reticulate: Absent (0), present (1) 137 Reticulate: Absent (0), present (1) 138 Faintly reticulate: Absent (0), present (1) 139 Favulariate: Absent (0), present (1) 140 Alveolate: Absent (0), present (1) 141 Foveate: Absent (0), pre sent (1)
107
142 Pusticulate: Absent (0), present (1) 143 Appressesdly colliculate: Absent (0), present (1) 144 Densely pubescent: Absent (0), present (1) 145 Papillae: Absent (0), present (1) 146 Hair pillose: Absent (0), present (1) 147 Hair tomentose: Absent (0), present (1) 148 Hair lineate: Absent (0), present (1) Pollen grains Size 149 Length (µm) 150 Breadth (µm) 151 Colpus length (µm) 152 Mesocolpium (µm) 153 Apocolpium (µm) 154 Exine thickness (µm) Shape 155 Prolate: Absent (0), pre sent (1) 156 Sub -prolate: Absent (0), present (1) 157 Prolate -spheroidal: Absent (0), present (1) Colpal membrane surface 158 Sub -psilate: Absent (0), present (1) 159 Scabrate: Absent (0), present (1) 160 Granulated: Absent (0), present (1) 161 Den sely granulated: Absent (0), present (1) 162 Rugulate -reticulate: Absent (0), present (1) 163 Spinulose: Absent (0), present (1) 164 Verrucate: Absent (0), present (1) Te ctum surface 165 Rugulate -reticulate: Absent (0), present (1) 166 Reticulate: A bsent (0), present (1) 167 Spinulose -verrucate: Absent (0), present (1) 168 Spinulose: Absent (0), present (1) 169 Rugulate -striate: Absent (0), present (1) 170 Verrucate: Absent (0), present (1) 171 Scrobiculous: Absent (0), present (1) Seed coat e lements 172 Carbon: Absent (0), present (1) 173 Oxygen: Absent (0), present (1) 174 Magnessium: Absent (0), present (1) 175 Potassium: Absent (0), present (1) 176 Copper: Absent (0), present (1) 177 Zinc: Absent (0), present (1) 178 Chlorine: Absent (0), present (1) 179 Calcium: Absent (0), present (1) 180 Sodium: Absent (0), present (1)
108
181 Silicon: Absent (0), present (1) 182 Aluminium: Absent (0), present (1) 183 Sulphur: Absent (0), present (1) Phenolic acids 184 Sinapic acid: Absent (0), present (1) 185 p-coumaric acid: Absent (0), present (1) 186 3, 4, 5, Trimethoxy cinnamic acid: Absent (0), present (1) 187 Aesculin: Absent (0), present (1) Flavonols 188 Kaempferol: Absent (0), present (1) 189 Quercetin: Absent (0), present (1) 190 Azaleatin: Absent (0), present (1) 191 Gossypetin: Absent (0), present (1) Flavones 192 Apigenin: Absent (0), present (1) 193 Luteolin: Absent (0), present (1) 194 Chrysoeriol: Absent (0), present (1) Glycosylflavones 195 Isovitexin: Absent (0 ), present (1) Biflavonyl 196 Kayaflavone: Absent (0), present (1) Quercetin glycosides 197 3- Arabinoside: Absent (0), present (1) 198 3- Xyloside: Absent (0), present (1) 199 3- Glucoside: Absent (0), present (1) 200 3- galactoside: Absent (0), present (1) 201 3- Rhamnoside: Absent (0), present (1) 202 3- Glucuronide: Absent (0), present (1) 203 3- Rutinoside: Absent (0), present (1) 204 7- Glucoside: Absent (0), present (1) 205 4’ - Glucoside: Absent (0), present (1) Chalcones 206 Isoliqu iritigenin: Absent (0), present (1) 207 Isoliquiritigenin 4’ - glucoside: Absent (0), present (1) Aurones 208 Aureusidin: Absent (0), present (1) 209 Aureusidin 4 - glucoside: Absent (0), present (1) 210 Aureusidin 6 - glucoside: Absent (0), present (1) Flavonones 211 Hesperitin: Absent (0), present (1) 212 Hesperidin: Absent (0), present (1) 213 Naringin: Absent (0), present (1) Isoflavones 214 Daidzein: Absent (0), present (1)
109
Table 12. Data matrix of the genus Cleome L. for characters presented in table 11.
Name of species 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 Cleome ariana 1 58.5 2 1 1 2 14.5 3 0 1 1 1 0 0 0 0 0 0 0 0 1 1 0 0 0 0 C. brachycarpa 1 22.5 1 1 1 2 13 5 0 0 0 1 0 0 1 0 1 0 0 1 1 0 0 0 0 0 C. dolichostyla 1 55 2 1 1 1 18.5 16 0 0 0 0 0 0 0 1 0 0 1 0 0 0 1 0 0 0 C. fimbriata 1 30.5 2 1 1 1 12.5 9 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 C. karachiensis 1 10.5 2 0 1 2 11 6 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 C. oxypetala 1 50 2 1 0 1 20.5 7 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 C. pakistanica 2 90 2 1 1 1 4 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 C. rupicola 1 20 1 1 1 1 16 7 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 C. scaposa 1 22.5 2 1 1 1 10.5 13 0 0 0 0 1 0 1 0 1 0 0 0 0 0 0 0 0 0 C. spinosa 1 90 2 1 1 2 55.5 12 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 C. viscosa 1 50 2 1 1 2 23 10 0 0 1 0 0 0 0 0 1 1 0 1 1 0 0 0 0 0
Table 12. (Contd.)
Name of species 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 Cleome ariana 0 0 0 0 0 1 1 1 0 0 1 2 1 0 0 0 1 0 0 0 0 0 7.6 3.01 1 C. brachycarpa 0 0 0 0 0 1 1 0 0 0 1 2 1 0 0 0 1 0 0 0 0 0 20 8.43 0 C. dolichostyla 0 0 0 1 1 0 0 0 0 1 1 2 0 0 1 0 0 0 1 0 0 0 18 10.1 0 C. fimbriata 0 0 0 0 0 1 1 1 0 0 1 2 0 0 0 1 0 0 0 1 0 0 8.3 4.15 0 C. karachiensis 0 0 0 0 0 0 1 0 0 0 1 2 1 0 0 0 0 1 0 0 0 0 6.9 1.42 0 C. oxypetala 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 13 5.87 0 C. pakistanica 0 0 0 0 0 1 1 0 0 0 1 1 1 0 0 0 0 0 0 0 1 0 6.7 3.88 0 C. rupicola 1 1 0 0 0 0 1 0 0 0 1 2 1 0 0 0 1 0 0 0 0 0 15 8.05 0 C. scaposa 0 0 0 0 0 1 1 1 0 0 1 1 0 0 0 1 0 0 0 0 1 0 14 4.47 0 C. spinosa 0 0 1 0 0 0 1 0 0 0 1 1 0 1 0 0 0 0 0 0 0 1 11 4.4 0 C. viscosa 0 0 0 0 0 1 1 0 1 0 1 2 0 1 0 0 0 1 0 0 0 0 8.1 2.34 0 110
Table 12. (Contd.)
Name of species 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 Cleome ariana 0 0 0 0 1 0 0 0 1 4 1.5 0.4 0 0 0 0 1 1 0 1 0 0 0 0 C. brachycarpa 1 0 0 0 1 0 0 0 1 10 2.8 1.3 1 1 0 1 0 1 0 1 0 0 0 0 C. dolichostyla 0 1 0 0 0 1 0 1 0 6.5 4 1.3 1 0 0 0 0 0 0 1 0 0 0 0 C. fimbriata 0 1 0 0 0 1 1 0 0 6 3.5 1.8 1 1 1 0 0 0 0 0 0 0 0 0 C. karachiensis 0 1 0 0 0 1 1 0 0 6.5 3 0.8 1 0 0 0 0 0 0 0 1 0 0 0 C. oxypetala 0 1 0 0 1 0 0 1 0 10 1.5 0.5 1 1 0 0 0 0 0 0 0 0 0 0 C. pakistanica 0 1 0 1 0 0 0 1 0 5 1.5 0.5 0 1 0 0 0 0 0 0 0 0 0 0 C. rupicola 0 0 1 0 0 1 0 1 0 6.5 1.5 0.5 1 0 0 0 0 0 0 0 0 1 0 0 C. scaposa 0 0 1 0 0 1 0 1 0 3.5 1.8 0.5 1 0 1 0 0 0 0 0 0 0 0 0 C. spinosa 0 0 1 0 0 1 0 1 0 25 6 3 1 0 0 0 0 0 0 0 0 0 1 0 C. viscosa 0 0 1 0 0 1 1 1 0 11 5 1.5 1 0 0 1 0 0 0 0 0 1 0 1
Table 12. (Contd.)
Name of species 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 Cleome ariana 2.25 0.35 1 0 1 0 1 0 0 0 0 0 0 0 1 0 0 0 0 1 0 1 0 0 C. brachycarpa 5 2 0 1 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 1 0 1 0 0 C. dolichostyla 7 1.5 0 1 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 C. fimbriata 7 2 0 1 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 C. karachiensis 4.25 1.75 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 C. oxypetala 7 1.5 0 1 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 C. pakistanica 2.5 0.5 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 C. rupicola 4.5 2.25 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 C. scaposa 4 1.5 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 1 C. spinosa 15 10 1 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 C. viscosa 8 2 0 1 0 0 0 0 0 0 0 0 1 0 0 0 1 0 1 1 0 0 0 0 111
Table 12. (Contd.)
Name of species 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 Cleome ariana 0 1 0 1 0 1.25 6 14.5 1 1 0 0 1.2 1 0 1 1 0 0 0 0 C. brachycarpa 1 0 0 0 1 2.5 6 7.5 1.75 0 1 0 0.82 0.82 0 0 0 1 0 0 0 C. dolichostyla 0 0 1 0 1 10 4 22 3.5 1 1 0 0.93 0.88 1 1 0 0 0 0 0 C. fimbriata 0 1 0 0 1 - 4 21 2.5 0 1 0 0.8 0.5 0 0 0 0 0 0 0 C. karachiensis 1 1 0 0 1 4.0 7 19.5 1.5 1 0 0 1.16 1.05 0 1 0 0 0 0 0 C. oxypetala 0 1 0 0 0 4.0 6 62.5 4 1 0 0 2.33 2.17 0 0 0 0 0 0 1 C. pakistanica 0 0 1 0 1 5.0 6 30 1.5 1 0 0 1.5 1 1 0 0 0 0 0 0 C. rupicola 0 1 0 0 0 4.0 6 50 5 0 1 0 2.36 2.22 0 0 0 0 0 1 0 C. scaposa 0 1 0 0 0 2.5 6 21 1 1 0 0 0.73 0.71 0 0 0 0 0 0 0 C. spinosa 1 0 0 1 0 7.0 6 22.5 2.25 1 0 0 0.99 0.95 1 0 0 0 0 0 0 C. viscosa 0 1 0 0 1 5.5 16 58.5 3 1 0 1 1.6 1.2 0 0 0 1 1 0 0
Table 12. (Contd.)
Name of species 121 122 123 124 125 126 127 128 129 130 131 132 133 134 135 136 137 138 139 140 141 142 Cleome ariana 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 C. brachycarpa 0 0 0 0 0 1 0 0 1 0 0 1 0 0 0 0 1 0 1 0 0 0 C. dolichostyla 0 0 0 0 0 1 0 0 0 1 0 0 1 0 0 1 0 0 0 0 0 0 C. fimbriata 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 1 C. karachiensis 0 0 0 0 1 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 1 0 C. oxypetala 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 C. pakistanica 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 C. rupicola 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 C. scaposa 0 0 0 1 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 C. spinosa 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 C. viscosa 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 1 0 112
Table 12. (Contd.)
Name of species 143 144 145 146 147 148 149 150 151 152 153 154 155 156 157 158 159 160 161 162 163 Cleome ariana 0 0 1 0 0 1 14.5 14.05 12.9 10.4 2.64 2.05 0 0 1 0 0 0 0 1 0 C. brachycarpa 0 0 0 0 0 0 25.7 17.15 21.9 8.55 5.71 2.21 1 0 0 0 0 0 0 0 1 C. dolichostyla 0 0 0 0 0 0 28.1 20.15 25.5 13.2 6.82 2.81 1 0 0 0 0 0 1 0 0 C. fimbriata 0 0 0 0 0 0 ------C. karachiensis 1 0 0 0 0 0 15.6 12.07 12.5 6.76 4.13 1.95 0 1 0 0 0 1 0 0 0 C. oxypetala 0 0 0 0 1 0 ------C. pakistanica 0 1 0 0 0 1 19.5 17 16.6 13.3 4.13 2.8 1 0 0 1 0 0 0 0 0 C. rupicola 0 0 0 1 0 0 15.3 9.57 12.9 5.28 2.45 1.59 1 0 0 1 0 0 0 0 0 C. scaposa 0 0 0 0 0 0 13.4 10.39 11.42 7.11 4.25 1.94 0 1 0 0 1 0 0 0 0 C. spinosa 0 0 0 0 0 0 ------C. viscosa 0 0 0 0 0 0 22.5 19.1 19.6 15.33 6.37 2.66 0 1 0 0 0 0 0 0 0
Table 12. (Contd.)
Name of species 164 165 166 167 168 169 170 171 172 173 174 175 176 177 178 179 180 181 182 183 184 Cleome ariana 0 1 0 0 0 0 0 0 1 1 1 1 1 0 0 1 0 0 1 0 0 C. brachycarpa 0 0 0 0 1 0 0 0 1 1 1 1 0 0 0 1 0 0 0 1 1 C. dolichostyla 0 0 1 0 0 0 0 1 1 1 1 1 1 1 1 0 0 0 0 0 0 C. fimbriata ------1 1 0 0 1 1 0 1 0 0 0 0 0 C. karachiensis 0 0 0 0 0 0 1 0 1 1 1 1 1 0 0 1 0 0 0 1 0 C. oxypetala ------1 1 1 1 1 0 0 1 0 0 0 0 0 C. pakistanica 0 1 0 0 0 0 0 0 1 1 1 1 1 0 1 1 1 1 0 0 0 C. rupicola 0 0 0 0 0 1 0 0 1 1 1 1 1 1 1 0 0 0 0 0 0 C. scaposa 1 0 0 1 0 0 0 0 1 1 1 1 1 1 1 1 0 0 0 0 0 C. spinosa ------1 1 1 1 1 1 1 1 1 0 0 0 0 C. viscosa 1 1 0 0 0 0 0 0 1 1 0 1 1 1 0 1 0 0 1 1 1 113
Table 12. (Contd.)
Name of species 185 186 187 188 189 190 191 192 193 194 195 196 197 198 199 200 201 202 203 204 Cleome ariana 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 1 0 0 0 1 C. brachycarpa 1 0 0 1 0 0 0 0 0 0 1 0 1 0 1 0 1 0 1 1 C. dolichostyla 0 0 0 1 1 0 0 0 0 0 0 0 1 1 1 1 0 0 1 1 C. fimbriata 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 C. karachiensis 0 0 0 0 1 1 1 1 0 0 0 1 0 0 0 0 1 1 1 1 C. oxypetala 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 C. pakistanica 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 C. rupicola 0 0 1 0 1 0 0 0 0 1 0 0 0 1 0 1 0 0 0 1 C. scaposa 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 1 0 0 1 0 C. spinosa 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 C. viscosa 1 1 0 1 0 0 0 0 1 1 0 0 0 1 0 0 0 0 0 0
Table 12. (Contd.)
Name of species 205 206 207 208 209 210 211 212 213 214 Cleome ariana 0 0 0 0 0 1 0 0 0 0 C. brachycarpa 1 1 1 1 0 1 1 0 1 0 C. dolichostyla 0 0 0 0 0 0 0 0 0 1 C. fimbriata 0 0 0 0 0 0 0 0 0 0 C. karachiensis 1 0 0 0 0 0 0 0 0 0 C. oxypetala 0 0 0 0 0 0 0 0 0 0 C. pakistanica 0 0 0 0 0 0 0 0 0 0 C. rupicola 0 0 0 0 0 0 0 1 0 0 C. scaposa 0 0 0 0 0 0 0 0 0 0 C. spinosa 0 0 0 0 0 0 0 0 0 0 C. viscosa 0 0 1 0 1 0 1 0 0 0 114
GENERAL DISCUSSION
Linnaeus (1753) first described the genus Cleome under the family Capparidaceae. While, Airy Shaw (1965) was the first person who placed the genus under the family Cleomaceae. Later on, many workers described the genus either by treating under the family Capparidaceae or Cleomaceae (already discussed in chapter I). According to the recent APG systems (I-IV) of classification, the genus Cleome is placed under Cleomaceae with about 250 species throughout the world. Thus, the genus Cleome is the largest and the most prevalent genus of the family. In Pakistan it is represented by 11 species including one new species. Presently, morphological data of the genus Cleome is correlated with various micromorphological features viz., pollen morphology, seed morphology, leaf epidermal features like, stomata and trichomes and chemistry of seeds and leaves is also carried out. While, for getting a synthetic approach, the data was analyzed numericaly.
Great diversity has been observed in the genus Cleome in terms of morphology and micromorphology. The most prevalent and type species of the genus Cleome is C. viscosa L. Although some of the workers (Rafinesque, 1838; Zhang and Tucker, 2008; Tucker and Vanderpool, 2010) treated C. viscosa under a separate genus Arivela, mainly due to the absence of gynophore and number of stamens. In the present studies it is found that in the genus Cleome number of stamens ranges from 4 to 20, similarly, presence and absence of gynophore and its length varies from species to species with in the genus. The species within the genus Cleome could be mainly bifurcated by having simple or compound leaves (Bobrov, 1939; Jafri, 1973; Blakelock and Townsend, 1980; Hedge and Lamond, 1970) such as, C. pakistanica , C. dolichostyla , C. fimbriata , C. scaposa , C. oxypetala and C. rupicola have simple leaves. While, C. ariana , C. brachycarpa , C. karachiensis sp. nov. C. spinosa and C. viscosa are characterized by having compound leaves.
On the other hand, when the genus was analyzed numerically, the usual alliance of species was quite different where quantitative characters were found to have more effect on specific delimitation as compared to qualitative characters. Mainly two groups were recognized on the basis of numerical analysis i.e., group-I comprises C. karachiensis sp.
115 nov., C. scaposa , C. brachycarpa , C. fimbriata , C. rupicola , C. viscosa and C. oxypetala . While, in group-II C. ariana , C. dolichostyla , C. pakistanica and C. spinosa were coupled.
Furthermore, every species of the genus Cleome could be easily distinguished from other species. Such as, C. spinosa can be distinguished from all other species of the genus by having spiny stipules and larger floral parts with distinct pollen and seed characteristics (Riaz and Abid, 2018). Similarly, C. dolichostyla is characterized by having cordate, broadly cordate or suborbicular leaves (Hedge and Lamond, 1970), hirsute hairs, appressedly reticulate seeds surface and reticulate or scrobiculous pollen tectum. While, C. oxypetala can be differentiated by its distinct seed characters i.e., obovate and tomentose hairy seeds (Riaz and Abid, 2018). Similarly, C. rupicola is characterized by having distinct tectum surface and presence of aesculin in leaves. The newly described species C. karachiensis is morphologically allied to C. brachycarpa and C. viscosa by having 3‒5 foliate, glandular leaves with acute apex and cuneate base, dorsally glandular sepals, yellow petals, tricolporate pollen and non angular seeds. While, it has more affinities to C. viscosa due to the presence of capitate glandular hairs on leaves, rounded petals, short style, subprolate pollen grains, linear capsule and retortiform seeds. It is also noteworthy that by applying numerical data, all the three species were gathered in same major clad of dendrogram but none of them showed close affinities may be due to differences in micromorphological characters, chemical constituents and quantitative characters. Present findings could be well supported by the suggestions of Cronquist (1964) and Stearn (1968) where they opined, that numerical analysis could not always be correlated with subjective findings. In dendrogram C. karachiensis made its affiliation with C. scaposa may be due to rather similar quantitative values and similarity in pollen structure. While, C. scaposa is apparently different from C. karachiensis due to its simple leaves, anomocytic stomata, reniform and alveolate seeds. Similarly, C. ariana and C.pakistanica were seem to be closely allied species in Flora of Pakistan (Jafri, 1973) but they remain apart in present studies on the basis of numerical analysis.
Thus, it is concluded that the data obtained from macromorphology could be significantly correlated with that of the data obtained from micromorphology of pollen
116 grains, seeds, trichomes and chemical constituents for specific delimitation of the genus Cleome from Pakistan.
117
Acknowledgements
First and foremost I am thankful to almighty Allah for providing me an opportunity to complete my Ph.D. thesis. I am also indebted to my supervisor Prof. Dr. Rubina Abid for her valueable and constructive suggestions and encouragement during the entire period of research.
I would like to express my gratitude to the Chairperson, Department of Botany, University of Karachi for providing departmental facilities. I am also thankful to the Director, Karachi University herbarium for providing herbarium and facilities for scanning electron microscopy. My gratitudes also extend to the directors and curators of BM, E, K, PMNH, RAW, ISL for providing herbarium facilities or herbarium specimens on loan.
I owe great debts of gratitude to Mr. Abrar Ali for illustrations and Ms. Sadaf Riaz for photographic enhancements. Special thanks are also for Dr. Afsheen Ather, Dr. Durdana Kanwal and Mrs. Syeda Darakhshan for their support and motivation in every possible way. I would like to express my great appreciation to Mr. Gohar and Mr. Arshad Gabol for providing laboratory facilities.
I am also thankful to my parents, sister and friends for being my strength throughout my life and specially in my research period.
118
Appendix‒I List of voucher specimens for seed micromorphology and EDS.
S. Species Collector, number and herbarium no. 1 Cleome ariana S. Abedin 8003 (KUH); M. Qaiser & A. Ghafoor 6592 (KUH); Kamal A. Malik & S. Abedin 1473 (KUH); Haider Ali 2063 , 6633, 6439, 1379 (KUH); A. Ghafoor & S. Omer 3019 (KUH) 2 C. brachycarpa M. Qaiser & A. Ghafoor 3 (KUH); S.M.H Jafri 813 (KUH); Shamim Akhter s.n. (KUH) 3 C. dolichostyla A.Ghafoor & S. Omer 1863 (KUH); A.G. Miller & J.A. Nyberg M9569 (KUH); M. Qaiser & Asad Raza Abbas 1043 (KUH) 4 C. fimbriata S.I.Ali 540 (KUH) 5 C. karachiensis Sana 9 (KUH); Sana & Rubina Abid 61 (KUH) 6 C. oxypetala Herbier J. Leonard 5856 (KUH); S.I. Ali, S.A. Farooqi & S. Abedin 1033 (KUH) 7 C. pakistanica V. P. Dutta 5 (KUH) 8 C. rupicola Sultan -ul -abedin & Abrar Hussain 6943, 6801 (KUH); S. Nazim Uddin, S. Abedin & Hameedullah 557 (KUH); S.M.H. Jafri 2689 (KUH) 9 C.scaposa Nazim, S. Abedin & M. Qaiser 63 (KUH); Kamal Akhter & Nazim Uddin 702 (KUH); Kamal A. Malik, M. Qaiser, Saood Omer & Gohar Khan 2123 (KUH); Sultan-ul-abedin & M. Qaiser 9328 (KUH); A. Ghafoor & M. Qaiser 386 (KUH); S.M.H. Jafri s.n. (KUH); Sadiq Masih 10 (KUH); A. Ghafoor & Tahir Ali 3620 (KUH); A. Ghafoor & Steve A. Goodman 4938 (KUH); M. Qaiser 2563 (KUH) 10 C. spinosa Jafri 4010 (KUH) 11 C. viscosa M. Qaiser & A. Ghafoor 4867 (KUH); Sultan -ul -abedin 5059, 4007, 4008, 4009 (KUH); A. Ghafoor & Tahir Ali 3739 (KUH)
119
Appendix‒II List of voucher specimens for pollen morphology.
S. Species Collector, number and herbarium no. 1 Cleome ariana Tahir Ali, Z.S. Huss ain & Gohar Khan 2215 (KUH); G. R. Sarwar & S. Omer 246 (KUH); M. Qaiser & A. Ghafoor 6597 (KUH); Haider Ali 6439 (KUH); Hakim khan 209 (RAW) 2 C. brachycarpa Tariq Hussain 228 (KUH); M.A. Siddiqui 6778 (PMNH); Sultan ul Abedin & M. Qaiser 7463 (KUH) 3 C. dolichostyla A. Ghafoor & S. Omer 1804,1863 (KUH); M. Qaiser, Asad Raza & Abrar Hussain 1043 (KUH); Rubina Akhter 094 (RAW); 4 C. karachiensis Sana 9 (KUH); Sana 101 (KUH) 5 C. pakistanica V. P. Dutta 5 (KUH) 6 C. rupicola Sultan -ul -abedin & Abrar Hussain 6803,6889 (KUH); Rubina Akhter s.n. (RAW) 7 C. scaposa A. Ghafoor & Steve M. Goodman 4938 (KUH); A. Ghaf oor & Tahir Ali 3769 (KUH); R. R. Stewart 9491 (RAW); Dr. Rubina Akhter 101 (RAW) 8 C. viscosa Sultan -ul -abedin 3969 (KUH); Rubina Akhter & Tariq Khan 086 (RAW); Ayaz & Dilawar 23 (ISL); , Muqarrab shah 3186 (PMNH); A. Ghafoor & Tahir Ali 4341 (KUH)
120
Appendix‒III List of voucher specimens for stomata and trichomes studies.
S. Species Collector, number and herbarium no. 1 Cleome ariana S. Abedin 8003 (KUH); M. Qaiser & A. Ghafoor 6592 (KUH); Kamal A. Malik & S. Abedin 1473 (KUH); A. Ghafoor & S. Omer 3019 (KUH) 2 C. brachycarpa M. Qaiser & A. Ghafoor 3 (KUH); S.M.H Jafri 813 (KUH); Shamim Akhter s.n. (KUH) 3 C.dolichostyla A.Ghafoor & S. Omer 1863 (KUH); A.G. Miller & J.A. Nyberg M9569 (KUH); M.Qaiser & Asad Raza 1043 (KUH) 4 C. fimbriata S.I.Ali 540 (KUH) 5 C. karachiensis Sana 9 (KUH); Sana & Rubina Abid 65 (KUH) 6 C. oxypetala Herbier J. Leonard 5856 (KUH); S.I.Ali 1033 (KUH) 7 C. pakistanica V. P. Dutta 5 (KUH) 8 C. rupicola Sultan -ul -abedin & Abrar Hussain 6801, 6943 (KUH); S. Nazimuddin, S. Abedin & Hameedullah 557 (KUH); S. M. H Jafri 2689 (KUH) 9 C.scaposa Kamal A. Malik, M. Qaiser, Saood Omer & Gohar Khan 2123 (KUH); Sultan-ul-abedin & M. Qaiser 9328 (KUH); A. Ghafoor & M. Qaiser 386 (KUH);S.M.H. Jafri (KUH); A. Ghafoor & Steve A. Goodman 4938 (KUH); M. Qaiser 2563 (KUH) 10 C. spinosa Jafri 4010 (KUH) 11 C. viscosa M. Qaiser & A. Ghafoor 4867 (KUH); Sultan -ul -abedin 5059, 4007, 4008, 4009 (KUH); A. Ghafoor & Tahir Ali 3739 (KUH)
121
Appendix‒IV List of voucher specimens for phenolic compounds.
S. Species Collector, number and herbarium no. 1 Cleome ariana G.R. Sarwas & S. Omer 3019 (KUH); Haider Ali 6358 (KUH); Muqqarab Shah & Dilawar 1717 (ISL); Wali ur Rehman & Subhan 299 (ISL); Sultan-ul-Abedin 8003 (KUH); Kamal Akhter Malik & S. Nazimuddin 1473 (KUH); Hakim Khan 209 (RAW); M. Qaiser & A. Ghafoor 6592 (KUH); Haider Ali 1370, 1464 (KUH). 2 C. brachycarpa Siddiqu i & Hassan ud din 2671 (RAW); M. Tanvir & Dilawar 1007 (ISL); M.A. Siddiqui, Akram & Lal Khan 53 (ISL); Tahir Ali & G.R. Sarwar 2548 (KUH); Dr. Rubina Akhter & Dr. M. Reidl 25 (RAW); A. Ghafoor & S. Omer 1681 (KUH); M. Qaiser, K.H. Rechinger, Jennifer Lamond & Tahir Ali 8196 (KUH); M. Qaiser, S. Omer & T. Ahmed 8133 (KUH); Rizwan yousuf 9 (KUH). 3 C.dolichostyla S.I. Ali, S.A. Farooqi & Sultan -ul -Abedin 1840 (KUH); A. Ghafoor & S. Omer 1804A (RAW); M. Qaiser, Asad Raza & Abrar Hussain 922 (KUH); A. Ghafoor & S. Omer 1863 (KUH); M. Qaiser, Asad Raza & Abrar Hussain 1043 (KUH). 4 C. karachiensis Sana & R ubina Abid 65 5 C. rupicola S. Nazimuddin, S. Abedin & Hameedullah 557 (KUH); Sultan - ul-Abedin & Abrar Hussain 6801, 6802, 6804, 6805, 6943 (KUH); S.M.H. Jafri 2689 (KUH); S.I. Ali, S.A. Farooqi, S. Abedin 1033 (KUH). 6 C.scaposa A. Ghafoor & Tahir Ali 3910 (KUH); Sultan -ul -Abedin & M. Qaiser 9328 (KUH); S.A. Farooqi & M. Qaiser 2843 (KUH); Anjum Perveen & Ishtiaq Hussain 107 (KUH); Kamal A. Malik, M. Qaiser, Saood Omer & Gohar Khan 2123 (KUH); Kamal Akhter & S. Nazimuddin 938 (KUH); Dr. Rubina Akhter 101 (RAW); Ali 454 (KUH); G.R. Sarwar, Qurban & Zamarrud 1288 (KUH). 7 C. viscosa A.R. Beg 1451 (RAW); Sultan -ul -Abedin 3902, 7989 (KUH); M. Rashid Awan & Jamshed Saqib 1076 (PMNH); Farrukh Hussain 7085 (RAW); Sultan-ul-Abedin & M. Qaiser 9079 (KUH); Ayaz & Dilawar 23 (ISL); Anjum Perveen 136 (KUH); Rubina Akhter & Tariq Khan 086 (RAW); Sultan-ul-Abedin & Abrar Hussain 9461 (KUH).
122
References:
Abid, R., and Qaiser, M. (2003). Chemotaxonomic study of Inula L. (S. Str.) and its allied genera (Inuleae-Compositae) from Pakistan and Kashmir. Pakistan Journal of Botany, 35(2), 127-140.
Abid, R., and Qaiser, M. (2006). Numerical analysis of Inula L. (s. str.) and its allied genera from Pakistan and Kashmir. Pakistan Journal of Botany , 38(3), 521-526.
Abid, R., Ather, A., and Qaiser, M. (2011). The seed atlas of Pakistan-V. Balsaminaceae. Pakistan Journal of Botany , 43(5), 2451-2456.
Abid, R., Ather, A., and Qaiser, M. (2013). The seed atlas of Pakistan-VIII. Fumariaceae. Pakistan Journal of Botany, 45(I), 87-92.
Abid, R., Ather, A., and Qaiser, M. (2015). The seed atlas of Pakistan-XI Urticaceae. Pakistan Journal of Botany, 47(3), 987-994.
Abid, R., Ather, A., and Qaiser, M. (2016). The seed atlas of Pakistan- Papavaraceae. Pakistan Journal of Botany, 48(3), 1035- 1044.
Abid, R., Kanwal, D., and Qaiser, M. (2014). Seed morphological studies on some monocot families (excluding gramineae) and their phylogenetic implications. Pakistan Journal of Botany, 46(4), 1309-1324.
Abid, R., Kanwal, D., and Qaiser, M. (2015). The seed atlas of Pakistan-X. Cucurbitaceae. Pakistan Journal of Botany, 47(2), 429-436.
Abid, R., Sharmeen, S., and Perveen, A. (2007). Stomatal types of monocots within Flora of Karachi, Pakistan. Pakistan Journal of Botany, 39(1), 15-21.
Aboushoer, M. I., Fathy, H. M., Abdel-Kader, M. S., Goetz, G., and Omar, A. A. Terpenes and flavonoids from an Egyptian collection of Cleome droserifolia . 2010. Natural Product Research, 24(7), 687-696.
123
Adedeji, O., Ajuwon, O. Y., and Babawale, O. O. (2007). Foliar epidermal studies, organographic distribution and taxonomic importance of trichomes in the family Solanaceae. International Journal of Botany , 3(3), 276-282.
Ahmad, K., Khan, M. A., Ahmad, M., Zafar, M., Arshad, M., and Ahmad, F. (2009). Taxonomic diversity of stomata in dicot flora of a district tank (N.W.F.P.) in Pakistan. African Journal of Biotechnology , 8(6), 1052-1055.
Ahmed, K. J. (1964). Cuticular studies in Solanaceae. Canadian Journal of Bot any, 42(7), 793-803.
Ahmed, K., Khan, M. A., and Shaheen, N. (2010). Palynological studies of the semi- desert plant species from Pakistan. African Journal of Biotechnology , 9(24), 3527-3535.
Ahouansinkpo, E., Atanasso, J., Dansi, A., Adjatin, A., Azize, O., and Sanni, A. (2016). Ethnobotany, phytochemical screening and toxicity risk of Cleome gynandra and Cleome viscosa , two traditional leafy vegetables consumed in Benin. International Journal of Current Microbiology and Applied Sciences, 5 (2), 813- 829.
Airy Shaw H. K. (1965). Diagnosis of new families, new names etc., for the seventh edition of Willis’s “Dictionary”. Kew Bulletin, 18(2), 256.
Aitchison, J. E. T. (1869). A Catalogue of the Plants of the Punjab and Sindh (pp. 9). London, Taylor and Francis.
Aitchison, J. E. T. (1880). On the Flora of the Kuram valley, & c., Afghanistan. The Journal of the Linnean Society , 18, 35.
Akcin, O. E., and Binzet, R. (2011). Micro morphological studies on nutlets of some Onsoma L. (Boraginaceae) species from Turkey. Pakistan Journal of Botany, 43(2), 743- 752.
Alafemar, T. (1951). Capparidaceae. In A. Parsa (Ed.), Flore de I’iran (Vol. 1, pp.908- 914). Tehran, Imprimerie Mazaheri.
124
Aleykutty, K. M., and Inamdar, J. A. (1978). Structure, ontogeny and taxonomic significance of trichomes and stomata in some Capparidaceae. Feddes Repertorium , 89, 19–30.
Aleykutty, K. M., and Inamdar, J. A. (1980). Structure, ontogeny and classification of trichomes in Ranales. Feddes Repertorium , 91(1-2), 95–108.
Ali, S.I., and Qaiser, M. (1986). A phytogeographical analysis of the phenerogems of Pakistan and Kashmir. Proceeding of The Royal Society of Edinburgh , 98B, 89- 101.
Anderson, T. (1860). Florula Andenensis. A systematic account with descriptions of the flowering plants hitherto found at Aden. The Journal of the Linnean Society , 5(s1), 3-43.
Aparadh, V.T., and Karadge, B. A. (2010). Seed ornamentation studies in some Cleome species. Bioinfolet , 7(1), 73-76.
APG (Angiosperm Phylogeny Group) I. (1998). An ordinal classification for the families of flowering plants. Annals of the Missouri Botanical Garden , 85, 531-553.
APG (Angiosperm Phylogeny Group) II. (2003). An update of the angiosperm phylogeny group for the orders and families of flowering plants. Botanical Journal of the Linnean Society , 141, 399-436.
APG (Angiosperm Phylogeny Group) III. (2009). An update of the angiosperm phylogeny group for the orders and families of flowering plants. Botanical Journal of the Linnean Society , 161, 105-121.
APG (Angiosperm Phylogeny Group) IV. (2016). An update of the angiosperm phylogeny group for the orders and families of flowering plants. Botanical Journal of the Linnean Society , 181, 1-20.
Bentham, G., and Hooker, J. D. (1862). Genera Plantarum ad exemplaria imprimis in herbariis kewensibus servata definite (Vol. 1). London, L Reeve & Co.
125
Berggren, G. (1962). Reviews on the taxonomy of some species of the genus Brassica based on their seeds. Svensk Botanisk Tidskrift , 56, 65-134.
Berggren, G. (1969). Atlas of Seeds and Small Fruits of Northwest European Plant Species, Cyperaceae (Part 2). Stockholm, Sewdish Museum of Natural History.
Berggren, G. (1981). Atlas of Seeds and Small Fruits of Northwest European Plant Species, Salicaceae-Cruciferae (Part 3). Stockholm, Sewdish Museum of Natural History.
Black, M. (2007). Seeds. In K. Roberts (ed.). Hand Book of Plant Science (Vol. 1, pp. 128-133). England, John Wiley & Sons. Ltd.
Blakelock, R. A., and Townsend, T. C. (1980). Capparidaceae. In: Townsend, C.C. and Evanguest. (Eds.), Flora of Iraq (Vol. 4, pp.145-153). Republic of Iraq, Ministry of Agriculture and Agrarian reform.
Blatt, C. T. T., Santos, M. D. D., and Salatino, A. (1998). Flavonoids of Bignoniaceae from the “cerrado” and their possible taxonomic significance. Plant Systematics and Evolution, 210(3), 289-292.
Blatter, E. (1919). Flora Arabica . (Vol. 8(1), pp. 34-39). Calcutta, India, Superintendent, Government Printing.
Blatter, E., Hallberg, P. F. and Mc Cann, C. (1919). Contributions towards a Flora of Baluchistan. The Journal of Indian Botany , 1, 54-59.
Bobrov, E. G. (1939). Capparidaceae. In V. L. Komarov (Ed.), Flora of U.S.S.R. , (Vol. 8, pp. 5-13). Israel, Israel program for scientific translations.
Boissier, E. (1846). Diagnoses Plantarum Orientalium Novarum . (Ser. 1, No. 6, pp. 20). Lipsiae, Apud B. Hermann.
Boissier, E. (1853). Diagnoses Plantarum Orientalium Novarum . (Ser. 2, No. 1, pp. 47- 48). Lipsiae, Apud B. Hermann.
126
Boissier, E. (1867). Flora Orientalis (Vol. 1, pp. 410-416) Reimpression facsimile. Geneva, A. Asher & Co.
Bradbury, S. (1967). The Evolution of the Microscope (pp. 375). New York, Pergamon Press.
Brown, R. (1811). On the Proteaceae of Jussieu. Transactions of the Linnean Society of London , 10, 15-226.
Burtt, B. L., and Lewis, P. (1949). On the Flora of Kuweit: I. Kew Bulletin , 4(3), 273- 308.
Cambessedes, J. (1844). In V. Jacquemount., Botanique: Voyage dans I’Inde (pp. 19). Paris, Firmin Didot Freres.
Castetter, E. F. (1935). Uncultivated native plants used as source of food. UNM Bulletins . http://digitalrepository.unm.edu/unm_bulletin/23 .
Chater, A. O. (1964). Cleome L. In T. G. Tutin, V. H. Heywood, N. A. Burges, D. H. Valentine, S. M. Walters, and D. A. Webb(Eds.), Flora Europaea - Lycopodiaceae to Platanaceae (Vol. 1, pp. 259). New York, Cambridge University Press.
Chaturvedi, M., and Gupta, S. (1983). Studies on the pollen morphology of some Capparis L. (Capparaceae) species. Proceedings: Plant Sciences , 92(1), 29-34.
Chaudhary, S. A. (1998). Flora of the Kingdom of Saudi Arabia illustrated. (Vol. 1, pp. 439-445). Riyadh, Ministry of agriculture and water, National herbarium, National Agriculture and water research center.
Chuang, T., and Heckard, L. R. (1972). Seeds coat morphology in Cordylanthus (Scrophulariaceae) and its taxonomic significance. American Journal of Botany , 59(3), 258-265.
Corner, E.J.H. (1976). The seeds of Dicotyledons (Vol. 1).Britain, Cambridge University Press.
127
Coyle, H. M., Ladd, C., Palmbach, T., and Lee H. C. (2001). The green revolution: Botanical contributions to forensics and drug enforcement. Croatian Medical Journal , 42(3), 340-345.
Crawford, D.J. (1978). Flavonoid chemistry and angiosperm evolution. The Botanical Review , 44, 431-455.
Cronquist, A. (1964). The old systematics. In C. A. Leone (ed.), Taxonomic Biochemistry and Serology . New York, Ronald Press.
Dawar, R., Qaiser, M., and Perveen, A. (2002). Pollen morphology of Inula L. (s.str.) and its allied genera (Inuleae- Compositae) from Kashmir and Pakistan. Pakistan Journal of Botany , 34(1), 9-22.
DE Candolle, A. P. (1824). Prodromus Systematics Naturalis Regni Vegetabilis , (Vol. 1, pp. 237-241). Paris, Sumptibus Sociorum Treuttel et Wurtz.
Delile, P. M. (1813). Flore D’Egypte. Explication des planches (pp. 106). Paris, Imprimerie Imperiale.
Edeoga, H. O., Omosun, G., Osuagwu, G.G.E., Mbaebie, B.O., and Madu, B.A. (2009). Micromorphological characters of the vegetative and floral organs of some Cleome species from Nigeria. American-Eurasian Journal of Scientific Research , 4(3), 124-127.
Edgeworth, M. P. (1847). A couple of hours’ Herborization at Aden. The Journal of the Asiatic Society of Bengal , 16(2), 1211-1220.
El Naggar, S. M. I. (2001). Systematic implications of seed coat morphology in Malvaceae. Pakistan Journal of Biological Sciences, 4(7), 822-828.
Erdtman, G. (1952). Pollen Morphology and Plant Taxonomy. Angiosperms. Waltham, Chronica Botanica Co.
Fang, N. R. L., Bahorum, T., and Khittoo, G. (2002). Chemosystematics: A new source of evidence for the classification of the endemic flora of Mauritius. Proceeding
128
of the Fifth Annual Meeting of Agricultural Science (73-79). Reduit, Mauritius, Food & Agri. Res. Council.
Fay, M. F., and Christenhusz, M. J. M. (2010). An Order of plants Characterised by shared chemistry. Curtis’s Botanical Magazine , 27(3), 165-196.
Furness, C.A., and Rudall, P. J.(2004). Pollen aperture evolution – a crucial factor for eudicot success?. TRENDS in Plant Science , 9(3), 154- 158.
Gaertner, J. (1788-1791). De Fructus et Seminibus Plantarum . Carolinae, Sumtibus Auctoris, Typis Academiae.
Gill, L. S., Olabanji, G. O., and Husaini, W. H. (1982). Studies on the structural variation and distribution of stomata in some Nigerian legumes. Willdenowia, 12, 87-94.
Gleason, H. A. and Cronquist, A. (1963). Manual of Vascular Plants of Northeastern United States and adjacent Canada (pp. 349). New York, D. Van Nostrand Co.
Grierson, A. J. C. (1984). Capparaceae. In A. J. C. Grierson and D. G. Long (Eds.), Flora of Bhutan (Vol. 1(2), pp. 415-416). Edinburgh, Royal Botanic Garden.
Grosso, B., Saint-Martin, M., and Vassal, J. (1994). Stomatal types of the genus Acacia (Fabaceae, Mimosoideae): an appraisal of diversity and taxonomic interest. Botanical Journal of the Linnean Society , 116(4), 325-341.
Hameed, M., Ashraf, M., Al-Qurianty, F., Nawaz, T., Ahmed, M. S. A., Younis, A., and Naz, N.(2011). Medicinal flora of the cholistan desert: A Review. Pakistan Journal of Botany , 43, 39-50.
Harborne, J.B. (1973). Phytochemical methods: A guide to modern techniques to plant analysis . London, Chapman and Hall.
Harborne, J.B., and Green, P. S. (1980). A chemotaxonomic survey of flavonoids in leaves of the Oleaceae. Botanical Journal of the Linnean Society , 81(2), 155-167.
129
Harborne, J.B., and Williams, C. A. (1973). A chemotaxonomic survey of flavonoids and simple phenols in leaves of the Ericaceae. Botanical Journal of the Linnean Society , 66(1), 37-54.
Haron, N. W., Moore, D. M., and Harborne, J. B. (1992). Distribution and taxonomic significance of flavonoids in the genus Eugenia (Myrtaceae). Biochemical Systematics and Ecology , 20(3), 266-268.
Hedge, I., and Lamond, J. (1970). Capparidaceae. In K. H. Rechinger (Ed.), Flora Iranica . (Vol. 68, pp.13-30). Austria, Academische Druck-U-verlagsanstalt Graz.
Hewson, H.J. (1982). Cleome L. Flora of Australia . (Vol. 8, pp 223-231). Canberra, Australian Government Publishing Service.
Holmgren, P. K., Holmgren, N. H., and Barnett, L. C. (1990). Index Herbariorum . Part I: The Herbaria of the World. 8 th ed. Regnum. Veg. New York.
Hooker, J. D. (1875). Flora of British India (Vol. 1, pp. 168-170). England, L.Reeve & Co. Ltd.
Hufford, L. (1995). Seed morphology of Hydraganaceae and its phylogenetic implicatiuons. International Journal of Plant Science , 156(4), 555-580.
Hutchinson, J., and Dalzeil, J. M. (1954). Flora of West Tropical Africa (Vol. 1, Part 1, pp. 86-87). London, Crown agents for oversea governments and administrations. Millbank.
IBM Corp. (2011). IBM SPSS Statistics for Windows, Version 20.0. Armonk, NY: IBM Corp.
Iltis, H. H. (1991). Capparaceae. In D. H. Nicolson, R. A. DeFillips, A. C. Nicolson and others (Eds.), Flora of Dominica, Part 2: Dicotyledoneae . (pp. 61-62). Washington, D.C., Smithsonian Institution press.
Iltis, H.H., Hall, J. C., Cochrane, T.S.,and Sytsma, K.J. (2011). Studies in the Cleomaceae I. on the separate recognition of Capparaceae, Cleomaceae and Brassicaceae. Annals of Missouri Botanical Garden , 98(1), 28- 36.
130
Inda, L. A., Torrecilla, P., Catalan, P., and Ruiz-Zapata, T. (2008). Phylogeny of Cleome L. and its close relatives Podandrogyne Ducke and Polanisia Raf. (Cleomoideae, Cleomaceae) based on analysis of nuclear ITS sequences and morphology. Plant Systematics and Evolution , 274, 111-126.
Jacobs, M. (1960). Capparidaceae. In C. G. C. J. Van Steenis (Ed.) Flora Malesiana (Vol. 6 1, Ser. 1, pp. 99-105). Djakarta, Noordhoff-Kolff N. V.
Jacquin, N. J., and Von, F. (1760). Enumeratio Systematica Plantarum: quas in insulis Caribaeis vicinaque Americes continente detexit novas, aut iam cognitas emendavit (pp. 26). London, Christians Lavnoy.
Jafri, S. M. H. (1957). Some species of Cleome . Kew Bulletin , 12(1), 173-175.
Jafri, S. M. H. (1977). Capparaceae. In Flora of Libya (Vol. 12, pp 11-18). Tripoli, Al faateh University, Faculty of Science, Department of Botany.
Jafri, S.M.H. (1973). Capparidaceae. In E. Nasir, and S. I. Ali (Eds.), Flora of Pakistan . (Vol. 34, pp. 20-32). Pakistan, The Herbarium, Dept. of Botany, University of Karachi.
Joshi, T., Kumar, N., and Kothiyal, P. (2015). A review on Cleome viscosa: An endogenous herb of Uttarakhand. International Journal of Pharma Research and Review , 4(7), 25-31.
Kamel, W. M., Abd El-Ghani, M.M., and El-Bous, M. M. (2010). Cleomaceae as a distinct family in the flora of Egypt. The African Journal of Plant Science and Biotechnology , 4(1), 11-16.
Kanwal, D., Abid, R., and Qaiser, M. (2009). The seed atlas of Pakistan- I. Aizoaceae. Pakistan Journal of Botany , 41(4), 1557-1564.
Kapoor, B.B.S., and Mishra, R. (2013). Flavonoid contents of some capparidaceous medicinal plants of North-West Rajasthan. Indian Journal of Pharmaceutical and Biological Research , 1(1), 9-11.
131
Kasem, W. T. (2016). Anatomical, pollen grains and seed exomorphic studies on five species of Cleome L. (Cleomaceae Bercht. & Presl)) collected from South west of Saudi Arabia. Journal of Plant Sciences , 4(2), 29-36.
Kasem, W. T., and Fathy, S. (2013). Flavonoids and isoenzymes as chemotaxonomic markers in Cleome L. (Cleomaceae Bercht and J. Presl). Current Botany , 4(3), 67-72.
Kasem, W.T., and Fathy, S. (2016). Flavonoids and isoenzymes as chemotaxonomic markers in Cleome L. (Cleomaceae Bercht and J. Presl). Current Botany , 7, 11- 16.
Khalik, K. A., and Maesen, L.J.G.V.D. (2002). Seed morphology of some tribes of Brassicaceae (Implications for taxonomy and species identification for the flora of Egypt). Blumea , 47, 363-383.
Khalik, K.A. (2005). Morphological studies of trichomes of Brassicaeae in Egypt and taxonomic significance. Acta Botanica Croatica , 64(1), 57-73.
Khatoon, S., and Perveen, A. (2003). Cleome pakistanica (Capparidaceae) – A new endemic species from Pakistan. Pakistan Journal of Botany , 35(2), 145-146.
Khokhar, A.L., Rajput, M.T., and Tahir, S. S. (2012). Taxonomic study of the trichomes in the some members of the genus Convolvulus (Convolvulaceae). Pakistan Journal of Botany , 44(4), 1219-1224.
Kitamura, S. (1960). Flora of Afghanistan (pp137). Japan, Kyaoto University, Nissha Printing Co., Ltd.
Kotresha, K., and Seetharam, Y. N. (1995). Epidermal studies in some species of Bauhinia L. (Caesalpinioideae). Phytomorphology, 45(1&2), 127-137.
Lawrence, G.H.M. (1970). Taxonomy of Vascular Plants , Toronto, Ontario, New York, The Macmillan Company, Collier- Macmillan Canada, Ltd.
132
Lersten, N. R., and Gunn, C. R. (1981). Seed morphology and testa topography in Cicer (Fabaceae: Faboideae). Systematic Botany , 6(3), 223-230.
Linnaeus, C. (1753). Species Plantarum . (Vol. 2. pp. 671-672). London, British Museum (Natural History).
Linskens, H. F., and Jorde, W. (1997). Pollen as food and medicine- A review. Economic Botany , 51(1), 78-87.
Mabberley, D.J. (2008). Mabberley’s Plant-book (3 rd ed., pp. 194). Cambridge, Cambridge University Press.
Marzouk, M.M., Al-Nowaihi, A.M., Kawashty, S. A., and Saleh, N.A. M. (2010). Chemosystematic studies on certain species of the family Brassicaceae (Cruciferae) in Egypt. Biochemical Systematics and Ecology , 38, 680-685.
Mathur, R. (2002). Capparidaceae. In N.P.Singh, D.K. Singh and B. P. Uinyal (Eds.), Flora of Jammu and Kashmir (Vol. 1, pp. 600-604). Calcutta, The Director, Botanical Survey of India.
McClure, D.A. (1957). Seed characters of selected plant families. Iowa State College Journal of Science , 31, 649-682.
Meikle, R. D. (1977). Flora of Cyprus (Vol. 1, pp. 173-176). Kew, The Bentham-Moxon Trust, Royal Botanic Gardens.
MetCalfe, C.R., and Chalk, L. (1950). Anatomy of the Dicotyledons (Vol. 1). Oxford, Clarendon Press.
Metcalfe, C.R., and Chalk, L. (1979). Anatomy of the Dicotyledons, systematic anatomy of the leaf and stem (Vol. 1, 2 nd Ed.). Oxford, Clarendon Press.
Mia, M. D., Ara, H., and Khan, B. (2007). Capparaceae. In : M. Khanum and H. Ara, (Eds.), Flora of Bangladesh (Vol. 57, pp. 7-11). Bangladesh, Bangladesh national herbarium.
133
Mitra, K. (1978). Contribution of the pollen morphology of the family Capparaceae. Bulletin Botanical Survey of India, 17, 7-31.
Nowicke, J. W., and Ridgway, J.E. (1973). Pollen studies in the genus Cordia (Boraginaceae). American Journal of Botany , 60(6), 584-591.
Oliver, D. (1868) Flora of Tropical Africa (Vol. 1, pp. 74-81). Ashford, Kent, L. Reeve & Co. Ltd.
Osman, A.K. (2012). Trichome micromorphology of Egyptian Ballota (Lamiaceae) with emphasis on its systematic implication. Pakistan Journal of Botany , 44(1), 33-46.
Otto, B. (2002). Schriftenreihe fur vegetations kunde. Bundesamt fur Nalussschutz Bonn , 38, 177-196.
Pax. (1891). In A. Engler and K. Prantl, Pflanzenfam. III , (Vol. 2, pp. 220) Leipzig, Verlag Von Wilhelm Engelmann.
Perveen, A., Abid, R., and Fatima, R. (2007). Stomatal types of some dicots within flora of Karachi, Pakistan. Pakistan Journal of Botany , 39(4), 1017-1023.
Perveen, A., and Qaiser, M. (2001). Pollen flora of Pakistan- XXXI Capparidaceae. Turkish Journal of Botany , 25, 389-395.
Perveen, A., and Qaiser, M. (2007). Pollen flora of Pakistan- LIII. Verbenaceae. Pakistan Journal of Botany , 39(3), 663-669.
Perveen, A., and Qaiser, M. (2007). Pollen flora of Pakistan- LIV. Rubiaceae. Pakistan Journal of Botany , 39(4), 999-1015.
Perveen, A., and Qaiser, M. (2007). Pollen flora of Pakistan- LV. Caprifoliaceae. Pakistan Journal of Botany , 39(5), 1393-1401.
Perveen, A., and Qaiser, M. (2007). Pollen flora of Pakistan-LVI. Valerianaceae. Pakistan Journal of Botany , 39(6), 1889-1896.
134
Petiver, J. (1699). Some attempts to prove that herb of the same make or class for the generality, have the like virtue and deny to work the same effects. Philosophical Transactions of the Royal Society , 21, 289-294.
Pope, M.A. (1925). Pollen morphology as an index to plant relationship. I. Morphology of Pollen. Botanical Gazette , 80 (1), 63-73.
Post, G. E. (1932). Flora of Syria, Palestine and Sinai . Natural Science Series no.1, (pp. 132-134). Beirut, Publications of the faculty of Arts and Sciences, American University of Beirut.
Prabhakar, M. (2004). Structure, delimitation, Nomenclature and classification of stomata. Acta Botanica Sinica , 46(2), 242-252.
Prabhakar, M., Kumar, B. K. V., Ramayya, N., and Leelavathi, P. (1985). Structure, distribution and taxonomic significance of trichomes in some Indigofera L. (Fabaceae). Proceedings Plant Science , 95(5), 309- 314.
Punt, W., Hoen, P. P., Blackmore, S., Nilsson, S., Le Thomas, A.(2007). Glossary of pollen and spore terminology . Review of Palaeobotany and Palynology , 143, 1- 81.
Puri, H. S. (1971). Macro and micromorphology of leaf and seed of C. viscosa L. Journal of d’agriculture Tropicale et de Botanique Appliquee , 18(12), 566-571.
Radford, A. E., Dickison, W. C., Massey, J.R., and Ritchie Bell, C. (1974). Vascular Plants Systematics . New York, Evanston, San Francisco, London, Harper & Row.
Raghavan, R. S. (1993). Capparaceae. In B. D. Sharma and N. P. Balakrishnan. Flora of India (Vol. 2, pp. 299-321). Calcutta, Botanical Survey of India.
Refinesque, C. S. (1838). Sylva Tellurina. Trees and shrubs of North America (pp. 110). Philadelphia.
135
Riaz, S., and Abid, R. (2018). Significance of seed micromorphological characters and seed coat elements for the taxonomic delimitation of the genus Cleome L. (Cleomaceae) from Pakistan. Pakistan Journal of Botany , 50(1), 271-277.
Sanchez-Acebo, L. (2005). A phylogenetic study of the new world Cleome (Brassicaceae, Cleomoideae). Annals of Missouri Botanical Garden , 92(2), 179- 201.
Saroop, S., and Kaul, V. (2015). Cleome viscosa: a promising underutilized minor crop. Genetic Resources & Crop Evolution , 62(7), 1121- 1126.
Sarwar, G. R., and Qaiser, M. (2012). A numerical taxonomy of the genus Rosularia (DC.) Stapf. From Pakistan and Kashmir. Pakistan Journal of Botany , 44(1), 349-354.
Shah, G. L., and Kothari, M. J. (1975). Observations on stomata and hairs on vegetative and floral organs in the tribe Trifolieae (family Papilionaceae). Australian Journal of Botany , 23(1), 111-122.
Shaheen, N., Ajab, M., Yasmeen, G., and Hayat, M. Q. (2009). Diversity of foliar trichomes and their systematic relevance in the genus Hibiscus (Malvaceae). International Journal of Agriculture and Biology , 11(3), 279-284.
Sharaf, M., El-Ansari, M.A., and Saleh, N.A.M. (1997). Flavonoids of four Cleome and three Capparis species. Biochemical Systematics and Ecology, 25(2), 161-166.
Smith, P.M. (1976). The chemotaxonomy of plants . United Kingdom, Edward Arnold (Publishers) Ltd.
Sokal, R. R., and Sneath, P. H. A. (1963). Principals of numerical taxonomy . San Francisco, W. H. Freeman and Co.
Stearn, W. T. (1968). Obeservations on a computer aided survey of the Jamaican species of Calumnea and Alloplec tus. In V.H. Heywood (ed). Modern methods in plant taxonomy . London and New York, Academic Press.
136
Stearn, W. T. (1983). Botanical Latin (3 rd ed.). Britain, David & Charles.
Steudel, E. T. (1840). Nomenclator Botanicus (2 nd ed, pp. 382). London, J. G. Cotte.
Stewart, J. L. (1863). Memoranda on the Peshawar valley, chiefly regarding its Flora. Journal of the Asiatic Society of Bangal , 32, 219-266.
Stewart, R. R. (1972). An Annonated Catalogue of the Vascular Plants of West Pakistan and Kashmir (pp. 294-296). Karachi, Department of Botany, University of Karachi.
Theobald, W.L., Krahulik, J.L., and Rollins, R.C. (1979). In C.R. Metcalfe and L. Chalk. Anatomy of the dicotyledons: Systematic anatomy of the leaf and stem (Vol. 1, 2nd Ed.). Oxford, Oxford Science Publications. Clarendron Press.
Thulin, M. (1993). Flora of Somalia (Vol. 1, pp. 53-59). Kew, Royal Botanic Gargen.
Tucker, G. C., and Vanderpool, S. S. (2010). Cleomaceae. In Editorial committee (Eds.), Flora of North America (Vol. 7, pp 199-222). New York, Flora of North America Association.
Verhoeven, R.F., and Venter, H.J.T. (2001). Pollen morphology of the Periplocoidae, Secamonoideae, and Asclepiadoideae (Apocyanaceae). Annals of Missouri Botanical Garden , 88(4), 569-582.
Vicary, N. (1847). Some notes on the botany of Sinde. Journal of the Asiatic Society of Bengal , 16(2), 1152-1168.
Walsh, B. M., and Hoot, B. (2001). Phylogenetic relationship of Capsicum (Solanaceae) using DNA sequences from two noncoding regions: the chloroplast atpB-rbcL spacer region and nuclear waxy Introns. International Journal of Plant Sciences, 162(2): 1409-1418.
Western, A. R. (1989). The Flora of the United Arab Emirates An Introduction (pp. 57- 58). Abu Dhabi, United Arab Emirates University.
137
Whitmore, T. C. (1979). Capparaceae. In H. Hara and L. H. J. Williams. An Enumeration of the Flowering Plants of Nepal (Vol. 2, pp. 46). London, Trustees of British museum (Natural History).
Wight, R., and Arn, G. A. W. (1834). Prodromus Florae Peninsulae Indiae Orientalis . (Vol. 1, pp. 22). London, Parbury Allen & Co.
Wild, H. (1960). Capparidaceae. In A. W. Exell and H. Wild. Flora Zambesiaca (Vol. 1, Part 1, pp. 195-205). London, Crown agents for oversea governments and administrations, 4, Millbank.
Wollenweber, E., Valant-vetschera, k.M., and Roitman, J. N. (2007). Chemodiversity studies on exudate flavonoids of Cleomaceae species (Brassicales). Natural Product Communications, 2(10), 997-1002.
Zafar, M., Ahmed, M., and Khan, M. A. (2007). Palynology of family Asteraceae from flora of Rawalpindi-Pakistan. International Journal of Agriculture and Biology , 9(1), 156-161.
Zavada, M. S., and De Villiers, S.E. (2000). Pollen of the Asteraceae from the Paleocene-Eocene of South Africa. Grana , 39, 39-45.
Zhang, M. G., and Tucker, C. (2008). Cleomaceae. In W. Zhengni, P. H. Raven and H. Deyuan (Eds.), Flora of China (Vol. 7, pp. 429). St. Louis, Missouri Botanical Garden Press.
Zoharay, M. (1966). Flora Palestina : Equisetaceae to Moringaceae (Part 1, pp. 245- 246). Jerusalem, The Israel academy of science and humanities.
138
INDEX
(Listed from systematic treatment only)
Cleome ariana Hedge & Lamond 9
C. ornithpodioides auct. Non Linn. 9
C. iberica Aitch. 9
C. brachycarpa Vahl ex. DC. 12
C. ruta Camb. 12
C. moschata Stocks ex. T. Anders. 12
C. dolichostyla Jafri 19
C. fimbriata vicary 20
C. noeana Boiss 20
C. quinquinervia var. noeana (Boiss.) Parsa 20
C. pentanervia Aitch. 20
C. droserifolia J. L. Stewart 20
C. karachiensis (sp. nov.)S. Riaz, R. Abid & M. Qaiser 24
C. oxypetala Boiss. 27
C. hotsonii Blatt. & Hallb. 27
Cleome pakistanica (Jafri) S. Khatoon & A. Perveen 27
C. heratensis sub. Sp. p akistanica Jafri 27
139
C. rupicola Vicary 30
C. stocksiana Boiss. 30
C. scaposa DC. 34
C. papillosa Stued. 34
C. gracilis Edgew. 34
C. linearis Stocks ex. T. Anders. 34
C. spinosa Jacq. 44
C. viscosa L. 45
C. icosandra L. 45
Polinisia viscosa L. 45
Polinisia icosandra (L.) Wight & Arn. 45
Arivela viscosa Refq. 45
140
Addendum
141