PALYNOLOGICAL STUDY OF CULTIVATED AND NATIVE PALMS FROM PAKISTAN
AND KASHMIR
By
Abid Arzoo Rashid
DEPARTMENT OF BOTANY
UNIVERSITY OF KARACHI
KARACHI-75270
PAKISTAN
2014
1
PALYNOLOGICAL STUDY OF CULTIVATED AND NATIVE PALMS FROM PAKISTAN
AND KASHMIR
By
Abid Arzoo Rashid
Thesis Submitted for the Requirement of the Degree
“Doctor of Philosophy”
DEPARTMENT OF BOTANY
UNIVERSITY OF KARACHI
KARACHI-75270
PAKISTAN
2014
2
PALYNOLOGICAL STUDY OF CULTIVATED AND NATIVE PALMS FROM
PAKISTAN AND KASHMIR
THESIS APPROVED
Supervisor
Prof. Dr. Anjum Perveen ______
External Examiner ______
Date ______
3
CERTIFICATE
TO WHOM IT MAY CONCERN
This is to certify that the thesis entitled “Palynological Study of Cultivated and Native Palms from Pakistan and Kashmir” submitted to the Board of Advanced Studies and Research,
University of Karachi by Mr. Abid Arzoo Rashid, satisfies the requirements for the degree of
Ph.D. in Botany.
______
Research Supervisor,
Prof. Dr. Anjum Perveen
Director
Centre for plant conservation,
Herbarium and Botanic Garden,
University of Karachi,
Karachi. 4
CONTENTS
S. No. Description Pg. No.
1 Abstract i) English 1 ii) Urdu 3 2 General Introduction 4 CHAPTER 1 “Taxonomic of the family Palmae” 8 3 Introduction 8 4 Literature Survey 8 5 Materials and Methods 13 6 Systematic arrangement of the family Palmae 15 i) Description of Palmae Juss. 15 ii) Key to the sub families 16 iii) Description of sub family Arecoidae Griff. 16 iv) Key to the genera 19 v) Description of tribe Areceae Drude 20 vi) Description of genus Areca L. 20 vii) Description of genus Dypsis Naronha ex Mart. 23 viii) Description of genus Ptychosperma Labill. 25 ix) Description of genus Wodyetia A.K. Irvine. 29 x) Description of tribe Cocoseae Griff. 33 xi) Description of genus Cocus L. 33 xii) Description of genus Eleais Jacq. 36 xiii) Description of genus Roystonea O.F. Cook 31 xiv) Description of Sub family Calamoideae Griff. 40
5
xv) Description of genus Calamus L. 40 xvi) Description of Sub family Coryphoideae Griff. 43 xvii) Key to the genera 43 xviii) Description of tribe Borosseae Mart. 45 xix) Description of genus Hyphaene J. Geartn 47 xx) Description of genus Bismarkia Hildebrandt & H. Wendl. 49 xxi) Description of genus Borassus L. 50 xxii) Description of tribe Caryoteae Drude 54 xxiii) Description of genus Caryota L. 54 xxiv) Description of tribe Corypheae Mart. 57 xxv) Description of genus Nannorrhops H. Wendl. 57 xxvi) Description of tribe Trachycarpeae Dransf. et al. 60 xxvii) Description of genus Brahea Mart. ex Endl. 60 xxviii) Description of genus Livistonia R. Br. 61 xxix) Description of genus Pritchardia Seeman & H. Wendl 65 xxx) Description of genus Washingtonia H.Wendl. 66 xxxi) Description of genus Trachycarpus H. Wendl 71 xxxii) Description of genus Raphis L.f. ex Aiton. 72 xxxiii) Description of tribe Phoeniceae Drude 77 xxxiv) Description of genus Phoenix L. 77 xxxv) Description of tribe Sabaleae Mart. 86 xxxvi) Description of genus Sabal Adanson 86 6 Glossary of Taxonomy 88 CHAPTER 2 “Palynology of the Palmae” 93 7 Introduction 93 8 Materials and Methods 98
6
9 General Pollen morphology of the Palmae 100 10 Pollen morphology of the sub family Arecoideae Griff. 101 i. Introduction 101 ii. Observations and Results 103 iii. Discussion 113 11 Pollen morphology of the sub family Calamoideae Griff. 120 i. Introduction 120 ii. Observations and Results 121 iii. Discussion 122 12 Pollen morphology of the sub family Coryphoideae Griff. 123 i. Introduction 123 ii. Observations and Results 126 iii. Discussion 144 CHAPTER 3 “Statistical Analysis of the family Palmae” 157 13 Introduction 157 14 Materials and Methods 159 15 Cluster analysis of the family Palmae Juss. 164 16 Cluster analysis of the sub family Arecoideae Griff. 172 17 Cluster analysis of the sub family Coryphoideae Griff. 177 18 General Discussion 180 19 Glossary 187 20 Appendix 196 21 References 198 22 Index of Taxa 218
7
ABSTRACT
The present work gives a detail account of pollen morphology of family Palmae including its taxonomic revision.The taxonomic studies follow the recent classification of
Dransfield et al., 2005 & 2008, which based on phylogenetic studies. A total of 28 species representing 21 genera distributed in 3 subfamilies and 10 tribes are known from
Pakistan and Kashmir. All the taxa valid or synonyms have been typified with the help of literatures. The detail morphological descriptions of the studied taxa including the artificial generic and specific keys are also given. A description of the range of pollen morphological variation by using Light Microscope (LM) and Scanning Electron
Microscope (SEM) have been carried out.
Considerable pollen variations have been found within the family with regard to size, shape, aperture type and exine pattern. However, a large number of species have monosulcate pollen (i.e., 25 out of 27) show the highest percentage (92.5%). Whereas, disulcate and trichotomosulcate pollen is present in a single species each viz., Calamus tenuis Roxb. and Elaeis guineensis Jacq.,respectively. The shape as seen in polar view is frequently elliptical, but in some cases rounded-triangular or circular pollen is also found.
Pollen shows a wide range in the size, the longest axis of pollen ranging from 15- 60 µm, averaging 30- 50 µm. These palm species also reveals a great diversity in exine pattern such as punctate, reticulate, vermiculate, verrucose, spinose, regulate and, perforate
(ranging from fine to coarse, and with sparse to dense distribution) or combination of these types have also been found, but the most predominant pattern being the reticulate type (terminology according to Erdtman).Muri more or less flat or rounded, simple baculate to multibaculate.
8
Pollen morphological characters are potentially informative in the systematics of monocotyledons including Palmae, both above and below the family level. The qualitative pollen characters (such asaperture, exine pattern and shape) are found to be taxonomically important as compared to the quantitative characters (such as size and exine thickness of pollen). On the basis of the combination of pollen characters, the following pollen types have been recognized such as Areca -type, Dypsis- type, Cocus- type, Elaies-type, Calamus-type, Bismarkia-type, Borassus-type, Brahea-type and
Phoenix-type. Key to the genera based on pollen characters and pollen types are also given. In addition, photographs of represented plants and microphotographs of pollen are incorporated to show the diagnostic features and convienience of identification.
The data obtained from the palynological studies have been numerically analyzed to find out the interrelationships among all the taxa of the family Palmae.
9
GENERAL INTRODUCTION
Palms form a large and diverse family of iconic monocotyledon have about c.2400 species belonging toc.187 genera distributed in five subfamilies viz., Arecoideae,
Calamoideae, Ceraxyloideae, Coryphoideae, Nyphoideae (Dransfield et al.,2005 and
2008; Govaerts& Dransfield, 2005) between 40 N and 45 S in both the Old World and
New World (Mahabalé, 1967). Around half of all palm species (c.1200 species and 57 genera) occurs in tropical Asia (extending from the India to the Suleman Island) with almost 1000 species of 50 genera (about 42% of the total family) found in the Malaysia only (Dransfieldet al.,2008).
The family Palmae is represented by 16 genera and 18 species from Pakistan and
Kashmir.Of these, 2 wildspecies viz., Phoenix sylvestris L., and Nannorhops ritichiana
(Griff.) Aitch.,are foundalmost all the entire part of our region (Kamal, 1984).The members of the family Palmae are widespread throughout the tropical and sub tropical regions of the world mostly found in rain forest, and few taxa are distributed in warm temperate regions. Ranging from minute forest floor species to giant canopy trees and even climbers, arecoid palms often play a prominent role in determining forest composition (Peters et al., 2004) and biotic interactions (Galetti et al., 2006).
Palms, the “Princes” of the Plant Kingdom constitutea unique family of monocotyledons which differ from other families in having an arborescent habit and a dense crown of leaves at the top (Mahabalé, 1967). Remarkably, not a single morphological feature can be recognized as distinctive to differentiate the family Palmae from the other monocotyledonous families. However, theimportant morphological
10 characters used for the delimitation of the subfamilies of Palmae are mainly nature of the leaf (i.e., leaves palmate, pinnate and bipinnate, and whether induplicate or reduplicate), the inflorescence and its bracts, arrangement of flower, floral structure and fruit.
This royal family contains a number of economically important plants, among theseArecasp. (betal palm), Cocus sp. (Coconut), Calamussp. (Rattan palm), Elaeissp.
(Oil palm), Phoenixsp. (Date palm), Raphiasp. (Raffia palm) and Salaccasp. (Salak) are considered as the chief genera of the family Palmae. The versatility of Palms in the hand of man is astonishing. Houses, baskets, mats, hammocks, cradles, quivers, blowpipes, bows, starch, wine, fruit beverages, flour, oil, ornaments, loincloths, medicines, magic, perfumes all are derived from palms (Moore,1973).
Thus Palms act as a key stone species among the monocots and ranking third after grasses and legumes. An indication of the importance of Palms in ancient time is that they are mention more than 30 times in Bible and at least 22 times in the Quran. Date palm (Phoenix dactylifera L.) is the oldest cultivated tree crop. Coconut (Cocus nucifera
L.)is symbolic of the multiple uses i.e. almost every part of the entire body is utilized such as roots extraction is used as mouthwash; stem is used for pillar, as a fuel and a good source of Charcoal; leaves are used for making mat, baskets; hard endosperm of the seed is edible, it can be dried to produce Khopra and the liquid endosperm or Coconut water is portable; mesocarpis used for matting (Menon and Pandalai,1958). Furthermore, vegetable fat obtained from the dried endosperm of fruit, is used for cooking and cosmetic. Another most important non-food palm product is Rattan (Calamus sp.), used as a structural material for furniture. Moreover, the Palm have considerable aesthetic value are used in magic and folk-medicine (Tomlinson, 1979). Now-a-day, palms
11 especially, Coconut, remains a symbol of the stereotypical tropical island paradise. Palms become visible on the flags and seals of many countries where they are native such as
Haiti, Saudi Arabia, Florida and SouthCarolina. Many members of the Palms such as
Livistonia chinensis(China palm), Roystonea regia (Bottle palm), Caryota urens (Wine palm or Fish tail palm), Sabal minor (Bush palmetto) are universally knownfor their majestic look and elegant shapes and have commercial horticultural importance too.
Palms are highly distinctive at the family level but within the family their morphological diversity is probably greater than that of any other monocotyledonous family (Uhl & Dransfield, 1987).At the level of pollen morphology the same diversity is also found such as shape and size of pollen, ornamentation and ultra structure of exine, form, number and arrangement of pollen aperture, all show a wide range of variation.Due to the pollen diversity the extensive workhas been carried out on the palynological studies of the family Palmaeby various workers such as Mahabalé (1967), Thanikaimoni
(1966, 1970a, 1970b), Sowunmi (1968, 1972), Kedves (1980), Ferguson (1986),
Ferguson et al. (1987, 1988), Dransfield et al. (1990), Harley (1990), Harley and Baker
(2001) and Harley and Dransfield, (2003).
There is no detailed study on the pollen morphology of the family Palmae especially from Pakistan and Kashmir.Only the taxonomic treatment of Stewart
(1972)and Malik (1984)are available from study area which are also almost 30 to 40 years old Table(1.1)Since then a lot of nomenclatural changes have taken place and a large number of taxa have been accumulated within the family Palmae.
Therefore, it seemed desirable to conduct detailed taxonomic revision of the family Palmae. The other purposeisto illustrate the wide range of pollen diversity
12 reported in the family Palmae by using techniques of Light Microscopy (LM) and
Electron Microscopy (SEM), and in order to establish their availability for further taxonomic workand also to determine the taxonomicrole of palynology of the family
Palmae from Pakistan and Kashmir.
13
Chapter 1
TAXONOMY OF THE FAMILY PALMAE
INTRODUCTION
The purpose of the taxonomic revision is to find out the nomenclature changes and to introduce aspects of the relationships among the members of the family Palmae, and place them in aglobal and regional perspective. The most recent classificationsystem for the Palmae(Arecaceae), as presented byDransfield et al. 2008, is followed for the taxonomic revision of palm taxa from the area under consideration (Table 1.3) Fig (1.2).
The Palm flora is represented by 186 genera and 27 tribes distributed in five subfamilies, of these, Pakistan has representative of 27 genera belonging to 10 tribes and three subfamilies viz., Arecoideae, Calamoideae and Coryphoideae (Table 1.2&1.3).
LITERATURE SURVEY
The Palmae was described by Jussieu (1789) nom. cons. Arecaceae Schultz (1832) nom.con.alt. Although Linnaeus (1753,1754) distinguishd the palm as a separate group on the basis of the types of leaves and reproductive organ. He reported 10 species of true palms and his view regards to the family Palmae is still accepted. However, the detail taxonomic literature on the palms begins with the work of C.F.P. von Martius (1853) followed by the monographic series of Beccari (1908, 1918, 1924 & 1933) and then
Corner (1966). During the last decades the classification of Palmae has been treated differently in the various system of classification by various workers such as Griffth
(1850), Wendland (1865),Drude (1889), Blatter (1926), Lawrence (1951), Hutchinson
(1959) , Moore (1960), Whitmore (1973) and Tomlinson (1961).Moreover, the family
14
Palmae was taxonomically revised by diversity of authors from time to time on the world or regional basisand almost all treated the family Palmae with a different concept such as
Beccari and Hooker (1883) in their “Flora of British India” presented the first detailed account of the family Palmae from South Asia and included 34 genera belonging to 6 tribes such as Areceae (further divided into 5 subtribe), Phoeniceae, Corypheae,
Lepidocarpae, Borasseae and Cocoineae. Out of 34 genera only 11 are found in study area.
Drude (1889) divided the family Palmae into 5 subfamilies (Coryphinae,
Borassinae, Lepidocaryinae, Ceroxylinae and Phytelephantinae) and 7 tribes (Phoeniceae,
Sabaleae, Borasseae, Mauritieae, Metroxyleae, Arecineae and Cocoineae).
Moore (1960) divided the Palm flora in to 8 subfamilies (Coryphoideae,
Borasoideae, Caryotoideae, Cocoideae, Arecoideae, Phytelephanotoideae and
Nypoideae).
Stewart (1972) in “An Annotated Catalogue of the Vascular Plants of West
Pakistan and Kashmir” reported only 8 generaof the family Palmae from the area under consideration.However, in the present study two species recognized by Stewart are reduced to synonym (viz., Phoenix humilis Royle now treated as the synonym of Phoenix loureiri (Kunth).The other species placed (viz., Trachycarpusexcelsa(Thunb.) H. Wendl. now treated under different genus viz.,Rhapis excelsa (Thunb.) Henry ex Rehder.
Moore (1973a) provided a detailed systematic treatment of the family Palmae based on field knowledge and morphological studies including the relationship among taxa.
Malik (1984) in“Flora of Pakistan” treated the family Palmae and reported 18
15 species belonging to 16 genera from Pakistan and Kashmir including 2 wild species.
Dransfield and Uhl (1998) in “Families and Genera of Vascular Plants:
Monocotyledons” classified the family Palmae into 6 subfamilies, 14 tribes, 36 subtribes and 189 genera. They included all the studied genera of the family Palmae which are also reported from Pakistan and Kashmir. However, they treated the following two genera viz., ChrysalidocarpusWendeland Neodypsis Baillon, under the synonymy of the genus
Dypsis Noronha ex Mart.
Dowe (2010) in “Australian Palms: biogeography, ecology and systematic” included 60 species belonging to 21 genera representing 10 tribes and 5 subfamilies.
Dransfield et al.,(2005) in “Genera Plamarum” revised the systematic of the family Palmae based on the molecular data and classify palms into 5 subfamilies, 26 tribes and 28 subtribes representing 187 genera included 17 unplaced genera.
Though as an evident from the above discussed taxonomic literature, we have quite a lot of monographs and classification but there is a need of better one.For this instance, recent advances in molecular systematic have improve our understanding of the phylogenetic relationships of many angiosperm groups, including monocotyledons
(Chase et al., 1995a,b; 2000; Baker et al., 2001). With the improvement of data analysis techniques and molecular systematic interest on the taxonomic studies of the family
Palmae has significantly increased.
The Chronology of the taxonomic recognition of the taxa representing the family
Palmae from Pakistan and Kashmir is given in Table-1.1.
16
Table1.1.Chronology of the taxonomic recognition of the Palm flora from Pakistan andKashmir
No. of taxa Year Author(s) Source or publication Subfamily Tribes Genera Species
An Annonated Catalogue of Vascular 1972 R.R. Stewart 00 00 06 08 Plants of West Pakistan And Kashmir
1984 Kamal 00 00 16 18 PalmaeFlora of Pakistan
2014 Present author 03 10 21 28 Unpublished thesis
17
Table 1.2.Distribution of taxa within the subfamilies of the family Palmae from the world and Pakistan
WORLD PAKISTAN SUB FAMILIES Tribes Genus Species Tribes Genus Species
Arecoideae 14 113 1247 03 7 8
Calamoideae 03 21 617 01 01 01
Ceroxyloideae 02 08 42 00 00 00
Coryphoideae 08 44 456 06 12 19
Nypoideae 00 01 01 00 00 00
Total 27 187 2363 10 21 28
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MATERIALS AND METHODS
The present work is based on the study of more than 500 herbarium specimens belonging to the following herbaria viz., KUH and RAW (abbreviated according to
Holmgren et al., 1990).The whole range of variation of each taxon was studied and on the basis, diagnostic keys have been prepared for subfamilies, genera and species.The morphological description of all level of the studied taxa are given including scientific name,vernacular name, habit, distribution, flowering period and their uses. Most of the names whether accepted or synonyms have been typified with the help of original description and type specimens. Each valid name is followed by full reference and its type. Similarly, synonyms are followed by references. The concept of such taxa where the type specimens could not be studied was strictly based on original description.
The distribution of all the taxa, from Pakistan and the world has been traced with the help of herbarium specimens and literature. All the examined specimens have been arranged according to the grid system of Flora of Pakistan (Fig.1.1).
The index of studied taxa including their synonymy is also given.The terminology of the examined taxa has been adopted from Dransfield et al., 1998, Dowe, 2010.
The systematic position of the studied taxa belonging to the family Palmae from
Pakistan and Kashmir is given in Table 1.3.
19
Pakistan and grid system
Fig 1.1 Map of
20
SYSTEMATIC ARRANGEMENT OF THE PAKISTAN PALM
FLORAPALMAE
Palmae Juss., Gen. P1. 37. 1789, nom. cons., Arecacae Schultz Sch. 1832. nom. cons. alt.Plant hermaphrodite, polygamous, monoecious or dioecious, armed or unarmed, hapaxanthic or pleonanthic. Stem creeping, subterranean, erect or climbing, solitary or cluster, unbranched usually dichotomously branched, smooth, covered with remains of leaf base, and prominently ringed with leaf scars. Leaves alternate, spirally arranged; laminapalmate, costpalmate, pinnate, bipinnate, or bifid, simple plicate in bud, rarely acanthophylls, splitting along the adaxial fold (induplicate) or abaxial folds (reduplicate) rarely splitting between the folds; leaflets lanceolate or linear to rhomboid or wedge- shaped, V-shaped (induplicate), Λ-shaped (reduplicate); petiole terete or ridged, glabrous or hairy (or scaly), bearing spines or prickles, hastulae present or absent.Inflorescence axillary, solitary or multiple, interfoliar or intrafoliar, spicate or usually paniculate; peduncular bracts 0-many.Flowers 1 or 2, hermphordite or unisexual, or dimorphic, sessile or pendunculate. Perianth segments 6 (into 2 series), uniseriate with a variable number of lobes, usually clearly differentiated into sepals and petals, free or connate, valvate or imbricate.Stamens 6, free or united, basifixed or dorsifixed, versatile, filaments free or variously connate, epipetalous. Gynoecium free or fused, ovary 1-3, superior, 1-3 chambered having a single ovule in each chambered. Fruit berry or drupe, mostly 1- seeded (rarely 2-10 seeded).
A large family of c.187 genera and 2365 species almost confined to the tropics of both hemisphere (Dransfield et al., 2005, 2008 and Dowe, 2010).
21
From Pakistan and Kashmir the family Palmae is represented by about 21 genera
and 28 species including 2 wild species viz., Phoenix sylvestris (L.) Roxb. and
Nannorophs ritichiana (Griff.) Aitch.
Key to the subfamilies
1+ Ovary and fruit covered in imbricate scales; seed sarcotestal; climbing palm armed,
spines, bristles and/or hooks present on the leaves, inflorescences and climbing
organs ...... 2. Calamoideae
- Ovary and fruit not covered in imbricate scales; seed not sarcotestal; non-climbing,
semi-herbaceous to arborescent palms, with spines and hooks confined to the petiole
margins or unarmed ...... 2
2+ Leaves palmate, costapalmate, imparipinnate or bipinnate; segments induplicate;
spines or teeth on the petiolesof palmate or costapalmate leaves, or unarmed on
impairpinnateor bipinnate leaves...... 3. Coryphoideae
- Leaves paripinnate; segments reduplicate; petiolesunarmed...... 1. Arecoideae
1. SUBFAMILY ARECOIDEAE Griff.
Leaves pinnate, pinnae induplicate or reduplicate.Inflorescence usually
moderately branched, infrequently spicate.Flower monoecious or dioecious; in triads, or
in cincinni, or in acervuli, or rarely solitary.Fruit epicarp usually smooth, infrequently
rough or corky-warted. Seed 1–8, endosperm homogeneous or ruminate, embryo apical to
basal.
22
Table-1.3. List of the examined taxa for the taxonomy and pollen morphology of the family Palmae. Systematic position is according to Dransfield et al., (2005& 2008)
SUBFAMILY TRIBE SUBTRIBE EXAMINED TAXA Arecoideae Griff. (1845) Areceae Drude (1881) Arecinae J.D. Hook. (1883) Areca L. Dypsidinae Becc. (1914) Dypis Naronnha ex Mart. Ptychospermatinae J.D. Hook. (1883) Ptychosperma Labill.
Wodyetia A. K. Irvine
Roystoneae Roystonea Cook
Cocoseae Mart. (1837) Attaleinae Drude (1887) Cocos L. Elaeidinae J.D. Hook (1883) ElaeisJacq. Calamoideae Griff. (1844) Calameae Drude (1881) Calaminae Meisner (1842) Calamus L. Coryphoideae Griff. (1844) Borasseae Mart. (1837) Lataniinae Meisner (1842) Borassus L. Hyphaeninae Becc. (1924) Hyphaene Gaertn. Caryoteae Drude (1877) CaryotaL.
Bismarkia Hildebr. & H. Wendl.
Corypheae Mart. (1837) Coryphinae J. Drans. & N.W.Uhl (1986) Nannorrhops H. Wendl.
Trachycarpeae Dransfield et al., Livistonae Saakov (1954) Brahea Mart .ex Endl. Livistonia R. Brown Pritchardia Seemann & H. Wendl. Washingtonia H. Wendl Thrinacinae Becc. (1907) Trachycarpus H. Wendl. Rhapis L. f. ex Aiton Sabaleae Mart ex Dumort (1829) Sabal Adanson Phoeniceae J. Presl. (1846) Phoenix L.
23
Fig. 1.2 Phylogeny (simplified) of Arecaceae based on morphological and molecular data (Baker & al. 2009).
24
A subfamily c. 114 genera and c. 1250 species distributed in 14 diverse tribes, with
world-wide distribution; in Pakistan with 03 tribes, the Areceae with 03 subtribes 05
genera; the Roystoneaewith a single genus species,and the Cocoseae with two subtribe
and 02 genera (Table 1.4).
Key to the Genera
1+ Fruit 25 cm long or more; endocarp 3-pored...... 2
- Fruit less than 25 cm long; endocarp not 3-pored………………………………….…3
2+ Petiole unarmed; anthers linear, latrorse. Fruit without beak…………...……6.Cocus
- Petiole armed; anthers rectangular, introrse. Fruit beaked…….….……….… 7.Elaeis
3+ Stamens indefinite (60-100) in male flower.Pistillode bottle shape………..….….... 4
- Stamens 6-30 in male flowers. Pistillode not as above ………………………..….... 5
4+ Seeds terete…………………..…………………………...... ……...... ….4. Wodeytia
- Seeds angled or grooved………………….……………………….....3. Ptychosperma
5+ Female flowers larger than male, borne at base of rachillae; peduncular bract absent.
Fruit up to 5 cm long………………………….…………………………...….1.Areca
- Female flowers larger than male, borne at base of rachillae; peduncular bract absent.
Fruit up to 5 cm long…………………..……………………………..…….….……...6
6+ Stem solid, solitary; flowers bisexual; petals connate…………....……....5. Roystonea
- Stem slender, clustered; flowers unisexual; petals distinct…………………..2. Dypsis
25
1. Tribe Areceae Drude
Pleonanthic palm.Leaves reduplicate, pinnate or pinnately ribbed.Inflorescence with a prophyll, peduncular bracts 1- 2.Petals valvate in staminate flowers; finely imbricate in pistllate flowers.Gynoecium pseudonumerous; ovary tricarpellary, triovulate Fruit with apical, lateral or basal stigmatic remain; endocarp without pores.
1. Areca
L., Sp. Pl. 1189. 1753; Gen. Pl. ed. 5. 495. 1754; Becc. & Hook. f. in Hook. f., Fl. Brit.
Ind.6: 405. 1894; Blatter in J. Bomb. Nat. Hist. Soc. 24 (2): 329. 1916; Whitmore,Palms
Mal. 97. 1973; Malik, in Nasir & Ali (eds.) Fl. Pak. 153: 33.1984; Dransfield and Uhl in
Kubitzki (ed.), Fam. Gen. Vasc. Pl. 4: 306-388. 1998; Dowe, Austr. Palm, Biogeog. Eco.
Syst. 305. 2010.
Plant solitary, up to 9 m tall with distinct nodes.Stem erect, with stripes and unarmed.
Leaves up to 2 m long, lamina entire or variously pinnate, leaflets unevenly divided, linear or lanceolate, 30-45 cm long, glabrous, petiole smooth, unarmed, base enlarged into sheath, glabrous, forming a swollen crown shaft.Inflorescences intrafoliar or interfoliar, spicate.Spikes flexuous or straight, branched, shortly peduncled.Spathes boat- like, coriaceous. Flowers monoecious, sessile, creamy white, lemon-scented; lower ones female, c. 13 mm long, 1-3 at spike-base; upper ones male c. 3 mm long, numerous.
Staminate flower more or less symmetrical; stamens 3 (6)-30.Pistillate flowers with distinct sepalsand petals.Ovary monocarpellary, up to 8 mm long, ovule solitary.
Fruitoblong or ovoid, 4 x 2.5 cm, with fibrous mesocarps, orange-red in colour.
26
SUBFAMILY ARECOIDEAE
Table 1.4 Synopsis of the tribes, subtribes and genera from Pakistan and Kashmir
Tribe Areceae Drude (1881)
Subtribe Arecinae J. D. Hook. (1883)
Genus ArecaL. (1753)
Subtribe Dypsidinae Becc. (1914)
Genus Dypsis Naronha ex Mart. (1837)
Subtribe Ptychospermatinae J. D.Hook. (1883)
Genus Ptychosperma Labill. (1808)
Genus WodyetiaA. K. Irvine (1983)
Tribe Cocoeae Mart. (1837)
Subtribe Attaleinae Saakov (1954)
Genus CocusL. (1753)
Subtribe Elaeidinae J.D. Hook. (1883)
Genus ElaeisJacq. (1763)
Tribe Roystoeneae J. Drans. et al (2005)
Genus RoystoneaO. F. Cook (1900)
27
A genus with about 60 species distributed in India, south China through Malaysia to New
Guinea and the Solomon Islands (Dransfield et al., 2005). In Pakistan it is represented by only one cultivated species.
1. Areca catechuL., Sp. Pl.1189.1753; Becc. Hook. f. l. c.; Blatter l.c.; Whitmore, l.c.;
Malik,l. c., Dowe, l.c.
Lectotype: Pinanga Rumphius, Herb. Amb. 1. Pl. 4. 1741 (vide Moore & Dransfield in
Taxon 28 (1, 2 & 3): 67.1979.
Monoecious, slender, single-trunked palm with a prominent crown shaft, up to 20 m tall and 25-40 cm in diameter.Leaves pinnate, 90-135 cm long, consists of 8-12 fronds,leaflets 50-60 in number, lanceolate, 30-45 cm long, glabrous, unarmed, frond base enlarged in to sheath and forming a green crown shaft about 55 x 15 cm. Spadix much branched, branches filiform, panicles arise below the leaves, bracteolate, bracts glabrous, upper part having small, sessile, creamy white, lemon scented flowers.
Staminate flowers minute, deciduous, 3 coriaceous sepals and petals. Stamens 6, arrow- head shaped anthers, pistillode 3-lobed. Pistillate flowers larger than staminate, sepals and petals 3 each, staminodes 6, ovary one-celled bearing a sessile stigma.Fruit fibrous, ovoid drupe, 5-10 x 3-5 cm, orange or red when ripe. One-seeded, globose or ovoid, endosperm ruminate.
Specimens Examined: G-4: Shaheed-e-millat road Karachi, 20ft, with orange fruit, 02-
10-1977, Kamal Akhter Malik 666(KUH); North Nazimabad Karachi, erect, with fruit,
02-10-77, Kamal Akhter Malik s.n. (KUH); North Nazimabad Karachi, tree with orange fruit, 02-03-1978, Kamal Akhter Malik 780 (KUH); Shagufta Khatoon s.n. (KUH).
Distribution: Cultivated in the warm regions of Asia.
28
Ecology:It is confined to the banks of fast-flowing streams on ultramafic rock.
Flowering Period: September-October
Vern: Chalia; English: Betal nut palm
Uses: A root decoction is reputed cure for sore lips; dry leaf petiole serves as excellent convenient splints for fracture. The nuts (hard endosperm of the seed) are used as chewing either raw or cured having soothing, diuretic, digestive, aphrodisiac and narcotic. The burnt nuts are often used as dentifrice.
2.Dypsis
Noronha ex Martius in C. F. P. von Martius et al., Hist. Nat. Palm., 3: 180. 1837;
Dransfield & Beentje, Palms Madag., 122-413.1995; Dransfield and Uhl in Kubitzki
(ed.), Fam. Gen. Vasc. Pl. 4: 306-389. 1998.
Chrysalidocarpus H. Wendl. in Bot. Zeit. 36: 171. 1878; Blatter, J. Bomb. Nat. Hist. Soc.
22 (4): 667.1914; Whitmore, Palms Mal. 47. 1973; Malik,in Nasir & Ali (eds.) Fl. Pak.
153: 10l.1984; Dransfield and Uhl in Kubitzki (ed.), Fam. Gen. Vasc. Pl. 4: 306-389.
1998;Neodypsis Baillon Baill., Bull. Mens. Soc. Linn. Paris 2: 1172. 1893.,Dransfield and Uhl in Kubitzki (ed.), Fam. Gen. Vasc. Pl., 4: 306- 389. 1998.
Acaulescent to erect or climbing, solitary or caespitose.Leaves: leaf bases unarmed, yellowish; crownshaft present or not; leaf blade pinnate undivided, unarmed, reduplicate; segments single or multifold, lanceolate, apices acuminate.Inflorescences protoandrous, axillary below crown of leaves, paniculate, with 3 orders of branching; peduncular bracts caducous, tubular.Flowers unisexual, sessile, in triads of 1 pistillate flower flanked by 2 staminate flowers.Staminate flowers: sepals 3, imbricate, free; petals 3, connate at the
29 base, valvate; stamens 6, free; anthers dorsifixed; pistillode present. Pistillate flowers: sepals and petals 3, imbricate, free; staminodes 6; monocarpellary, ovules 1, stigmas
3.Fruit ellipsoid, exocarp yellow, smooth, mesocarp fleshy, endocarp thin, fibrous.Seeds ovoid, endosperm homogeneous.
1. Dypsis lutescens: (H. Wendl.) Beentge & J. Dransf.
Chrysalidocarpus baronii.Becc.
Chrysalidocarpus lutescens.H. Wendl.
Holotype : Madagascar, Ambila – Lemaitso Dransfield et al., JD 6441 (K).
Isotype(BH,P, TAN).
Leaflets c. 60 cm long and 2-2.5 cm broad. Crown shaft swollen, yellow-green covered with caducous whitish wax and scales.Inflorescence arching, c. 30 cm long, highly branched, main stalk arising from a tubular bract.Flowers in clusters. Staminate flowers: sepals and petals 3; stamens free, exserted, c. 4 mm long, anthers 2-celled, dorsifixed; pistillode much shorter than stamens.Pistillate flowers small, c. 3 mm in diameter; sepals and petals 3, coriaceous.Fruit baccate, dark yellow turning black when mature.
Specimen Examined: G-4: Gulshan-e-IqbalKarachi, 24ft, with fruit, 02-06-2010, Abid
Arzoo Rashid, 456 (KUH); North Nazimabad Karachi, with flowers, 22-07-10, Abid
Arzoo Rashid, 459 (KUH).
Distribution: Native to Madagascar. Represented by asingle cultivated species in our region.
Ecology: Locally common, occurup to 300 m, mainly in swampy areas, on littoral forest or heath vegetation on white sand, and on rock, also found on alluvium at much higher elevations. 30
Flowering Period: May-July
Vern: English: Golden cane palm, Madagascar palm, Butterfly and Feather palm
Uses:Primarily it is used as shrub, hedge or screen in subtropical or tropical landscapes,
extensively used as an indoor 'plant scaping' specimen.
3. Ptychosperma
Labill., Mem. Inst. Par. 9: 252. 1809; Blatter in J. Bomb. Nat. Hist. 23 (3): 527. 1915;
Bailey, St. Cyc. Hort. 3:2856. 1950; Bailey, Man. Cult. Pl. (reprint ed.). 174. 1966;
Whitmore, Palms Mal. 97. 1973; Malik, in Nasir & Ali (eds.) Fl. Pak. 153: 33.1984;
Dransfield and Uhl in Kubitzki (ed.), Fam. Gen. Vasc. Pl. 4: 306-389. 1998; Dowe,
Austr. Palm, Biogeog. Eco. Syst. pp.305. 2010.
Solitary or clustering, pleonanthic, monoecious palms.Stems slender to robust, leaningto erect, unarmed; leafscars prominent to obscure. Leavesparipinnate, leaf bases forming a tubular elongate crown shaft, variously scaly, tomentose or glabrous, eventually splitting opposite the petiole at abscission; petiole short to long, channeled on the adaxial surface, rounded on the abaxial surface; pinnaesingle or variously united, linear to cuneate, reduplicate, erect to horizontal; apex truncate and notched, terminal pair united at the base. Inflorescence intrafoliar, paniculate,branched 2–6 orders, protandrous; bicarinate, fully enclosing the tubularbeaked; peduncle short to long, angled, glabrous, scaly or tomentose; rachillae long, straight to flexuous, fleshy tofibrous. Flowers unisexual, sessile, spirally arrangedtriads of a single pistillate flowers subtended by a pair ofstaminate flowers on either side. Staminate flowersbullet-shaped - ovoid; sepals 3, imbricate, rounded and gibbous, margin fimbriate; petals 3, stiff , valvate, much longer than the sepals; stamens 9–100 or more; filaments short; anthers long, basally
31
sagittate,dorsifixed, versatile; pistillode conical or bottle shaped with along neck, longer
thanthe stamens, apically irregularly lobed, trifid or papillate.Pistillate flowersshorter
than the staminate, conicoid; sepals 3, imbricate, rounded, sometimes slightly gibbous,
margin fimbriate; petals 3, imbricate, pointed, valvate at the apex; staminodes1–6,
dentiform, linear, fused; gynoecium conic-ovoid, mono-trilocular; stigma 3, recurved;
Fruitglobose to ellipsoid, red, orangeor black-purple at maturity; mesocarp fleshy. Seed5-
lobed; endosperm homogeneous or ruminate.
A genus with c. 28 species, centered in New Guinea, the D’Entrecasteaux and Louisiade
archipelagos, but extending west to east from the Moluccas to the Solomon Islands and
south from the Caroline Islands to northeastern Australia. In Pakistan it is represented by
two species.
Key to Species
1+ Stem solitary; pinnae 30–60 on each side of the rachis, up to 84 cm long; endosperm deeply ruminate………………………………………….………1.P. elegans
- Stems clustered; pinnae 15–40 on each side of the rachis, up to 56 cm long;
endosperm homogeneous……………………………………….....2.P. macarthurii
1. Ptychosperma elegans (R.Br.) Blume, Rumphia 2: 118. 1843.
Seaforthia elegans R.Br., Prodr.: 267 1810; Ptychosperma seaforthiaMiq., Fl. Ned. Ind.
3: 21. 1855nom. illeg.; Saguaster elegans (R.Br.) Kuntze, Revis.Gen. Pl. 2: 735. 1891.
Holotype: Australia. Queensland. NorthumberlandIs., 1802, R.Brown s.n.Bennett 5794
BM; isotype K.
32
Stemsolitary, 12 m tall, 8 cm wide; internodes 15 cm long, green to grey. Leaves7–11, c.3 m long, 30–60 pinnae each side of rachis; crownspreading; crown shaft c. 65 cm long, densely whitewoollyand minutely brown scales; ligule absent; petiole 19–35 cm long, minutely lepidote, with brown punctiform scales;pinnae 84 cm long, 8 cm wide,dark green on the adaxial surface, lighter green on theabaxial surface.Inflorescence65–70 cm long, 100–125 cm broad, branched to 4 or 5orders, peduncular bract 50 cm long, 18 cm wide, withscattered light and dark scales toward the apex. Staminate flowers 4.5–6 mm long,2 mm wide in bud,sepals 1.5–2 x 2 mm, imbricate, gibbous,margins smooth, light green; petals 6 x 2 mm, thick, apex curved inward, lightgreen, glabrous; stamens 12–22,
6 mm long; filaments4–5 mm long; anthers c. 1.5 mm long, medifixed,versatile, white; pistillode 7–10 mm long,3-lobed. Pistillate flowers3.5–5 mmlong; sepals 2.5 x 2.5 mm, imbricate,green, glabrous, margins fimbriate; petals 5 x 2.5 mm glabrous; staminodes c.
6, fused into double segments; stigma c. 1 mm long, recurved, white.Fruit globose to ellipsoid9–15 mm long, 8–10 mm diam.; red at maturity.Seed10–11 mm long, 7–7.5 mm diam.,endosperm deeply ruminate.
Specimens Examined: G-4: Ghandi garden Karachi, small tree upto 9 m tall,12-02-
1987; Saood Umer 623 (KUH); Hill Park Karachi, cultivated,white flowers, 22-02-1988,
Saood Umer 645 (KUH).
Distribution:Endemic to Queensland, distributed from larger islands in western Torres
Strait to near Yeppoon and Great Keppel Island. In Pakistan it is cultivated in Karachi.
Ecology:At the elevation from 0-900 m, in coastal forest, shore-line, on windward slopes, inhigh-rainfall areas, littoral forest, sheltered gullies.
Flowering Period: February-May.
33
Vern: English: Solitare palm
Uses:Primarily it is used as a shrub, hedge or screen in subtropical or tropical landscapes.
2.Ptychosperma macarthuri (H.Wendl.) H. Wendl. ex Hook. f., in Kew Report., 55.
1882. (1884) New Guinea; Bailey, St. Cyc. Hort. 3: 2856. 1950; Bailey, Man. Cult. Pl.
(reprint).174.1966; Benthall, Trees Calc.464 (reprint ed.) 1946; Malik in Nasir & Ali
(eds.) Fl. Pak. 153: 33. f. 3 (A-J). 1984; Dowe, Aust. Palm, Biogeog. Eco. Syst. 305.
2010.
Kentia macarthuri H. Wendl. in T. Moore, Florist & Pomologist, 114. 1879.
Type: Probably described from New Guinea
Trunk 2.5-7 m long, uniformly thickened, older part of the stem greyish-white, internodes
8.5-12.5 cm long.Leaves 90-120 cm long, pinnately divided into 20-28 segments on either side; segments smooth, coriaceous, unarmed, 5-7.5 x 25-40 cm, terminal leaflet absent; pinnae toothed apically, like a fish tail; leaf sheaths forming a crown shaft, c. 45 cm long, unarmed; bracts 8.75-12.5 cm long, triangular; petiole 10-15 cm long, smooth, concavo-convex.Inflorescence arising from the node near the base of crown shaft.
Flowers white. Male flowers: sepals 3, much smaller than the petals, imbricate, coriaceous, glabrous, forming cupule; petals 3-lobed, coriaceous, c. 0.7 cm long. Stamens
22, filaments unequal, 3.5-6 mm long, anthers versatile, apex curved.Ovary tricarpellary.Fruit drupe, densely packed on the hanging arching spikes, dark orange, ellipsoid with a small terminal beak, 0.8 cm long.Seed hard.
Specimens Examined: G-4: Cliften Karachi, 7.5 m tall, 06-04-1990,Saoodumer 321
(KUH); Bhadrabad Karachi, cultivated, 12-04-1990, Rubina Saleem 356 (KUH).
34
Distribution: Same as the genus
Ecology:Grows at the level of 0-400 m,in lowland, hillslope rainforests, in monsoon and littoral closed forest and brackish mangroves.
Flowering Period: February – April.
Vern: English: Macarthur palm
Uses: Cultivated as ornamental plant as a shrub, hedge or for landscape.
4.Wodyetia
A. K.Irvine, Principes27: 161.1983;Dransfield and Uhl in Kubitzki (ed.), Fam. Gen.
Vasc. Pl. 4: 306-389. 1998.Dowe, Austr. Palm, Biogeog. Eco. Syst.pp. 305. 2010.
Solitary, moderate, erect, pleonanthic, monoeciouspalms.Stemscolumnar; leafscars congested;internodes short. Leavesparipinnate, plumose; leafbases forming a tubular elongatecrownshaft, eventually splitting opposite the petiole atabscission, white-grey- green toward the petiole; petioleshort, stout, flat to concave on the adaxial surface,rounded on the abaxial surface, densely scaly, withwhitish tomentum and scattered brown scales; pinnaein a singleplane on the rachis, reduplicate, divided longitudinally to base into 2–17 linearsegments, apically truncate and dentate.
Inflorescenceinfrafoliar, paniculate, branched to4 orders, erect to pendulous in fruit, branches divaricate,protandrous; completely enclosing peduncular bracts; peduncular bract, tubular; peduncleshort, flattened. Flowersunisexual, sessile, spirally arrangedin well-spaced triads of a single pistillate flower subtendedby a pair of staminate flowers onthe proximal portion and then with staminate flowersdistally, in pairs or solitary.
Staminate flowerssymmetrical,bullet-shaped in bud; sepals 3, imbricate, rounded,inflated,
35 margins dentate; petals 3, thick, rigid, ovate,valvate; stamens 58–72; filaments short; anthers long, versatile,apically bifid, basally sagittate, dorsifixed.Pistillate flowersovoid; sepals 3,imbricate, margins tattered; petals 3, imbricate,hooded, apically valvate; staminodes 3–6, dentiform; gynoecium conic-ovoid, unilocular, uniovulate;style short; stigma 3 large recurved.Fruit globose-ovoid, orange-red at maturity; mesocarpfleshy, moderately fibrous; 1-seeded.Seedellipsoidal, terete in cross-section.
A genus with only one species distributed in northeastern Queensland, Australia. In
Pakistan it is cultivated in Karachi University Botanical Garden, Centre for Plant
Conservation, Karachi.
1. Wodyetia bifurcata Irvine, Principes27: 165, fig pp. 162–164 s.n. 1983; Dowe, l.c.
Holotype: Australia, Queensland.MelvilleRa., 4 km N of Abbey Peak, 23 Nov. 1981,
A.K.Irvine 2184.
Stem6–15 m tall, 20–25 cm in diameter, enlarged at the base; leaf scars 3 cm wide; internodes 20 cm long. Leaves6–10, 2.6–3.2 mlong, arching, 90–110 pinnae either side of rachis;crownshaft 80–120 cm long; petiole 25–90 x 5.0–5.6 cm; pinnae 70 x 4 cm; proximal. Inflorescence 75–112 cm long, 70–100 cm wide; axis cream to light green; peduncular bract c. 58 x 15 cm, glabrous; peduncle 8–13 cmlong, 4–5 cm wide, 2–2.5 cm thick, green, with scatteredsmall dark flat scales; rachis 105 cm long and 5 cm wideat the base, cream to green, small flatbrown scales.Staminate flowers11 mm long, 4 mm wide; sepals 5–6 x 3–4 mm, cream-green, glabrouswith scattered light-coloured scales; petals
9.8–10 x 5 mm, cream-green, glabrous; stamens 10 mm long; filaments 4 mm long; anther medifixed, versatile, white-cream; pistillode 8–15 mm long.Pistillate flowers 15 x
36
10 mm; sepals 6–7 x 5–7 mm, green; petals 9–11 x 6–8 mm, green.Fruit50–65 mm long, and 27–60 mm wide.Seed32 x 26 mm wide.
Specimens Examined: G-4: P.E.C.H.S Karachi, 10ft high, 20-05-2012; Abid Arzoo
Rashid 387 (KUH); ibid; Botany department Karachi University, erect, white flowers, 01-
08-12,Abid Arzoo Rashid 357 (KUH).
Distribution: Same as the genus.
Ecology:Occurring primarily on the lower slopes, 50–400 m at sea level, scattered in open forest.
Flowering Period: May to August.
Vern: Foxtail Palm
Uses: Widely cultivated as an ornamental plant
5. Roystonea
O.F. Cook, Science, Ser. 2.12:479. 1900., Bailey in Gentes Herbarium 3:341-387. 1935;
Benthall, Trees. Cal. 462. 1946; Bailey, Man.Cult. Pl. (reprint ed.). 172. 1966;Whitmore,
Palms Mal. 103. 1973; Malik, in Nasir & Ali (eds.) Fl. Pak. 153: 33.1984; Zona, Fl.
Neotrop. 71: 1-35.1996; Dransfield and Uhl in Kubitzki (ed.), Fam. Gen.Vasc. 4: 306-
389. 1998.
Solitary.Stem unarmed with prominent rings, swollen at the base. Leaves pinnate, divided into numerous leaflets; leaflets in two or more planes, narrow, pointed; petiole concavo- convex, base of leaf stalk enlarged into a leaf sheath which envelops the top of the stem and forms a green elongated crown shaft. Inflorescence branched to 3-4 orders, up to 60 cm long, enclosed in 2 coriaceous, green, stiff, caducous bracts, the outer small and the
37 inner is long. Flowers unisexual, usually in triads; staminate and pistillate flowers borne on the same inflorescence. Male flower: Sepals and petals 3; stamens exserted, 6-12, anthers elongate, dorsifixed, versatile; pistillode subglobose or trifid. Female flower: sepals and petals 3, united to half way; ovary 1-3 chambered, covered with 6 scaly staminodes;stigma sessile.Fruit red, oblong to rounded, up to 1.25 cm.
A genus with 10 species, distributed from Florida to W. Indies, Central and tropical
South America. In Pakistan is represented by one cultivated species.
1. Roystonea regia (H.B. & K.) O.F. Cook, Bull. Torrey Bot. Cl. 531. 1900. Benthall, l.c.462; McCann, Hundred Beaut. Trees. Ind. ed. 2.114. 1959; Bailey, l.c. 173; Stewart,
Ann. Cat. Vasc. Pl. W. Pak. & Kashmir 33. 1972; Whitmore, l.c. 104; Malik, l.c. 11. f. 2.
A-H.
Oreodoxa regia Kunth, Nov. Gen. Sp. 1: 305. 1816.
Oenocarpus regius (Kunth) Spreng., Syst. Veg. 2: 140.1825.
Type: Humboldt and Bonpland (P).
Stem up to 40 m tall, usually displaying a distinct thickening towards the middle, greyish white, smooth. Leaf crown round, leaves up to 4 m long, drooping, divided into numerous leaflets, pinnae in two rows on either side of rachis, lower pinnae 60-71 x 2-2.5 cm, middle pinnae 70-90 x 3.5-5.0 cm, reduplicate from the base, rachis tapering towards the apex; petiole, unarmed, elongated into a leaf sheath forming a shining greenish,c.120 cm long crown shaft. Inflorescence profusely branched. Male flower much larger than the female, calyx and corolla 3-lobed, corolla thin, larger than the calyx, petals 0.5 x c. 0.2 mm, valvate, joined at the base; stamens 9, exserted, filaments subulate, 0.5 mm long,
38 joined at the base beneath the pistillode; pistillode smaller, anthers dorsifixed, large.Female flowers 3-4 mm long, white; stigma triangular and surrounded by 6 scale- like staminodes.Fruit globose, fleshy 1-1.2 cm long, scar near the base of the fruit.
Specimens Examined:G-4:Zological garden Karachi, tree, 30ft, white flower, 20-04-
1978, Kamal Akhter Malik & Saida 842 (KUH); Zological garden Karachi, cultivated tree, 40ft, with fruit spadix,, 20-04-1978,Kamal Akhter Malik & Saida 43 (KUH).
Distribution: Indigenous to Cuba; cultivatedin Pakistan as ornamental plant.
Ecology:on road and near the Island(Roadside tree in Karachi, Pakistan; vide
R.R.Stewart, 1972)
Flowering Period: Throughout the year.
Vern:English: Royal Palm, Bottle palm.
Uses:It is used as timber, for thatching, hog feed and cultivated as ornamental.
2. Tribe Cocoseae Drude
Pleonanthic, monoecious palm.Leaves pinnate, reduplicate; crown shaft absent.
Inflorescence solitary, interfoliar, branched up to a single order; prophyll fibrous; peduncular bracts present, exceeding the prophyll, woody, enclosing entire inflorescence in bud, splitting at the time of anthesis; staminodes connate; irregularly dentiform in a ring; gynoecium tricarpellary, ovules 3 or many, well devolped endocarp is present, pores
3 or more.
6. Cocus
L. Sp. Pl. 1188. 1753; Gen. Pl. ed.5. 495.1754; Becc. & Hook. f. in Hook. f., Fl. Brit.
Ind.6: 482. 1984; Malik in Nasir & Ali (eds.), Fl. Pak. 153: 7-9. 1984; Dransfield &Uhl 39 in Kubitzki (ed.), Fam. Gen. Vasc. Pl. 4: 306-389. 1998; Dowe,Austr. Palm, Biogeog.
Eco. Syst. 305. 2010.
Solitary, leaning to erect, pleonanthic, monoecious palms.Leaves paripinnate, leaf bases clasping, not forming a crown shaft, fibrous at the margins; pinnae in a single plane on the rachis, sub-opposite to opposite, linear to acuminate, reduplicate, acute to tapered at the apex or bilobed, midrib prominent. Inflorescence interfoliar, paniculate, branched to 1 order, erect to pendulous, peduncular bract shallowly grooved. Staminate flowers sessile, asymmetrical, ovoid in bud; sepals 3 imbricate, triangular; petals 3, distinct, longer than sepals, leathery, valvate; stamens 6, anthers medifixed, basally sagittate, filaments not inflexed, pistillode thin, 3-lobed.Pistillate flowers globose in bud; sepals 3, imbricate; petals 3, longer than the sepals, thick, imbricate; staminodal ring not lobed; gynoecium trilocular, triovulate, ovoid, stigma 3.Fruit 1-seeded, epicarp smooth, mesocarp thick, fibrous, endocarp woody with 3 longitudual ridges.Seed large, endosperm homogenous, hollow and partially filled with fluid.
A monotypic genus native to the arch of Islands between Southern India and Malaysia
(Buckly and Harries 1984).It is widely cultivated throughout the tropics. In Pakistan the coconut palm grows on the coast of Sind and Baluchistan (Stewart, 1972).
1. Cocus nucifera L. Sp. Pl. 1188. 1753; Becc. & Hook. f. l.c.; R. R. Stewart, Ann. Cat.
Vasc. Pl. W. Pak. & Kash. 33. 1972; Malik, l.c.; Dowe, l.c.
Lectotype: ‘Tenga’ in H.A. Rheede’s Hortus Indicus MalabaricusVol. 1, plates 1–4
1678, were chosen by Moore and Dransfield 1979.
40
Stem 40 m tall, 30 cm in breadth, enlarged at the base, smooth with prominent rings; leaf scars 1-7 cm wide, irregular, internodes 15 cm long, irregular, green to grey. Leaves pinnate, 3-4 m long,120-180 cm in diam., erect to horizontal, or below the horizontal, straight to gently arching; petiole unarmed, stout, up to 1.3 m long; pinnae70-110 in number, 130 cm long and 5 cm wide, regularly arranged. Inflorescence 1-2 m long and 60 cm wide, branched to 1 order; peduncular bracts present, up to 90 cm long. Staminate flowers cream yellow, coriaceous,7-15 mm long, 3-9 mm wide in bud, asymmetric; sepals 3 x 5 mm, imbricate, entire; petals 15 x 4 mm, asymmetric, thick with pointed tips; stamens medifixed, versatile; pistillodes trifid, lobes 6 mm long, recurved.
Pistillate flowers ovoid, 20-40 mm in length, 30-35 mm wide in bud, sepals 15-30 mm long, 25-32 mm wide, petals 25-35 mm long, 30-40 mm wide, imbricate, margins smooth; stigma erect to re-curved, 4 mm long, glistening white. Fruit 10-35 cm long, 8-
20 cm in diam, 3 angled, green or yellowish, covered with thick fibrous mass, woody.
Seed globose, 6-25 cm in diam.
Specimens Examined:G-4:Sewage farm, Karachi, 28-02-57, A .A .Qureshis.n.(KUH);
P.E.C.H.S Block 2, Inflorescence up to 2 m long, flowers cream yellow, 12-8-2012,Abid
Arzoo Rashid 17 (KUH); ibid; Department of Botany, University of Karachi, 12-08-2012,
Abid Arzoo Rashid 19 (KUH); Botanical Garden, University of Karachi, flower present, c. 30 m tall, 12-08-2012, Abid Arzoo Rashid 30 (KUH); on way to Kashmir
RoadP.E.C.H.S, leaves pinnate, flower yellowish, fruit present, green, 14-5-2014, Abid
Arzoo Rashid 25 (KUH).
Distribution: same as the genus
Ecology:Hall mark of the tropics and coastal area.
41
Flowering Period: Flowering and fruiting through out the year.
Vern: Naryal; fruit Khopra. English: Coconut palm.
Uses:The most versatile providing edible and industrial oil (also making of soaps and cosmetics) protein rich milk, invigorating water and also valuable source of timber, fibre and matting materials.
7. Elaeis
Jacq., Sel. Strip.Americ. Hist.: 280. 1763; Dransfield and Uhl in Kubitzki (ed.), Fam.
Gen. Vasc. Pl. 4: 306-389. 1998.
Monoecious palm.Stem solitary, erectup to 10 m tall, 30-50 cm in diameter.Leaves pinnate, up to 5 m long; segments 100 or more on either side, slotted in groups and spreading in various planes; petiole armed (spinous or thorny), spines formed from leaflet, midribs at the base of the leaf.Inflorescences in clusters, borne among the leaf bases.Stamens 6,anthers rectangular, introrse.Pistil tricarpellary, stigma linear.Fruits glossy,more than 25 cm long, red to black when ripe, beaked, 1-seeded; endocarp 3- pored.
1. Elaies guineensisJacq., Select. Stirp. Amer. Hist., 280. 1763; Dransfield and Uhl, l.c.
Solitary palm, erect, 4-10 m in height.Leaves pinnately lobed, reaching up to 5 m in adult trees; pinnae liner lanceolate, 1 m long, leaf blade has 100-160 pairs of leaflets with distinct midrib. Flower monoecious borne on different inflorescence. Spadix short andthick; male inflorescence have dense staminate spikelets axon, sepals 3, oblong orlanceolate, imbricate; petals 3, oblong valvate, stamens 6, filament base connate into a jar
42
Fig 1.3 A:Cocus nuciferaL.B: Dypsis lutescens(H.Wendl.)Beetge & J. Dransf.
43 shape. Female inflorescence subcapitate, bracteoles 2, acute or spiny, sepals and petals each 3, ovate or oblong, imbricate, larger than the male, ovary trilocular, ovoid or nearly cylindrical, style short, stigma 3, linear. Fruit ovoid globose or obovoid, 1 seeded.
Specimens Examined:G-4:Botanical Garden, University of Karachi, flowers present,
12-08-2012, Abid Arzoo Rashid 35 (KUH); Gulshan-e-Iqbal, 4-5-2013, Abid Arzoo
Rashid 40 (KUH); ibid.
Distribution: same as the genus
Ecology:Found on watercourses in drier areas and in savannas.
Flowering Period: May to August.
Vern:African oil palm
Uses: the most important commercial oil-producing plant in the tropics, and is used locally as a source of wine, thatch and building materials. Even waste endocarp has been used as road metalling.
Table 1.5 Synopsis of the sub family Calamoideae tribes, subtribes and genera from
Pakistan and Kashmir
Calameae Kunth ex Lecoq & Juillet(1831)
Calaminae Meisn.(1842)
CalamusL. (1753)
Calamus tenuis Roxb.
44
Fig 1.4 A: Elaeis guineensisJacq.B:Wodyetia bifurcateIrvine
45
2. SubfamilyCalamoideae Griff.
Armed.Leaves pinnate, pinnately ribbed, infrequently palmate; pinnae reduplicate.
Inflorescence paniculate or spicate branched, frequently withadnation.Flower
monoecious or dioecious, polygamous; solitary, dyads or in cincinni.Fruit epicarp of
imbricate scales. Seed 1–3, usually sarcotestal; endosperm homogeneous or ruminate,
embryo basalor lateral.
A subfamily with about 21 genera and c. 620 species distributed in 3 tribes, with world-
wide distribution.In Pakistan represented by only a genus of the tribe Calameae, and
subtribe Calaminae.
8. Calamus
L. Sp. Pl. 325.1753; Gen. Pl. ed.5. 152.1754; Becc. & Hook. f. in Hook. f., Fl. Brit. Ind.6:
436. 1984; Hutch., Fl. W. Trop. Afr. 3: 166.1968; Malik in Nasir & Ali (eds.) Fl. Pak.
153: 6-7.1984; Dransfield and Uhl in Kubitzki (ed.), Fam. Gen. Vasc. Pl. 4: 306 -389.
1998; Dowe, Austr. Palm, Biogeog. Eco. Syst. 305. 2010.
Dioecious palm.Plant solitary or clustered, acaulescent, erect or climbing, pleonanthic,
leaf scars prominent. Leaves pinnate, pinnae reduplicate, cirrate or ecirrate, leaflet single-
fold, terminal pair basally joined, entire or rarely praemorse, linear- lanceolate or
rhombic, armed or scaly, climbing whip (flagellum) sometime present. Inflorescence
branched to 2-3 orders.Flower distichous or rarely spiral. Staminate flower solitary,
symmetrical; calyx much shorter than corolla; stamens 6, filaments elongate; pistillode
short.Pistillate flowers much longer than staminate.Gynoecium tricarpellary, covered in
46 reflexed scales, stigma 3, ovule basal. Fruit usually 1-or (2-3) seeded. Seed sarcotestal, rounded, grooved or angled.Endosperm homogemous and horny.
A genus with about 400 species distributed in Africa, South East and South East-Asia,
Malaysia, Melanesia to Fiji and Australia. In Pakistan the genus is represented by only 1 cultivated species.
1. Calamus tenuis Roxb., Fl. Ind. 3: 780.1832; Becc. & Hook, f., l.c. 447; Blatter in J.
Bomb. Nat. Hist. Soc., 25 (3): 392: 1918; Malik, l.c. 7.
Type: Described from Bendhari Bet of the inhabitants of Chittagong.
Stem clustered, climbing up to 20 m long. Leaves pinnate 45-75 cm long; leaflets equidistant, 20-30 on either side, linear – lanceolate, acuminate up to 2 m long; petiole short and stout margined with straight spines; leaf sheath armed with flat spines and bearing whip-like (flagella), sterile inflorescence. Staminate flower small,c. 25 mm longin 3-4 series sepals 3-lobed, coriaceous; petals 3, valvate; stamens 6, dorsifixed.
Pistillate flowers with a sterile male flower in the same bracteole; sepals 3-lobed; petals
3, tubular at the base, valvate; staminodes forming a cup; ovary trilocular covered with retrorse scales.Fruit sub-globose.
Specimens Examined:G-4: Karachi University Botanical Garden, Karachi, 10-1-
2011,Abid Arzoo Rashid 20 (KUH); Gulshan-e-Iqbal, Karachi, white flowers,Abid Arzoo
Rashid s.n. (KUH).
Distribution: Tropical Himalaya, India, Asam, Bangladesh, Bhutan, Cambodia, Laos,
Burma, Thailand, Vietnam, Java and Sumatra
47
Fig 1.5Calamus tenuisRoxb.
48
Ecology:Occur at 33-550 m altitude, plain to foothill slopes, and flood plain of rivers,
including degraded habitats, wet, damp and shaddy area.
Flowering Period: January to March
Vern: English: Barilley cane
Uses: The cane is of high quality for handicrafts and accepted in trade, also used in
making furniture. The shoot is edible. The cultivation of this species for the production of
edible shoots is becoming increasingly widespread in Laos (Belcher 1999, Sengdala &
Evans 1999).
3. SubfamilyCoryphoideaeGriff.
Leaves pinnate, bipinnate, palmate, costapalmate; pinnae and segments induplicate rarely
reduplicate.Inflorescence interfoliar, intrafoliar, apically paniculate or rarely spikate
usually highly branched.Flowers monoecious, dioecious, polygamous; solitary or in
cincinni or in triads, unisexual or hermaphrodite.Fruit smooth or corky-warted.Seed 1–3,
endosperm homogeneous or ruminate, embryo apical to basal.
A subfamily with about 44 genera and c. 450 species belonging to 8 tribes, distributed all
over the world. In Pakistan it is represented by 03 genera and 19 species belonging to 6
tribes viz., Borasseae (3 genera distributed in 2 subtribes), Caryoteae, Coryphoideae,
Phoeniceae and Sabaleae (1 genus each), and Trachycarpeae (6 genera distributed in 02
subtribes) Table 1.3 & 1.6.
Key to the Genera
1+ Leaves palmate or costpalmate, acanthophylls absent…...... 2
- Leaves pinnate or bipinnate, acanthophyll present...…………………..……………5
49
2+ Endocarp thin, crustaceous or cartilaginous.Rachillae with deep pits………...... …3
- Endocarp thick and hard. Rachillae without deep pits……………..…...... …...6
3+ Staminate and pistillate inflorescence are not same; flowers without tufts of hairs. Fruit sessile, endocarp forming pyrenes; stigmatic remains apical ……………...……………………………………..……………....……11. Borassus
- Staminate and pistillate inflorescence is same; flowers not bearing tufts of hairs. Fruit pedicillate, endocarp not forming pyrenes; stigmatic remains basal………………………………………………………..…………….…...... ……4
4+ Seed deeply grooved…………………………………..…………....…10. Bismarkia
- Seed not grooved……………………………………….…….....………9.Hyphaene
5+ Plant dioecious, leaves pinnate; pinnae spiny……………..……….……20. Phoenix
- Plant monoecious, leaves bipinnate; pinnae not spiny….…………….……12. Caryota
6+ Trunk branched. Leaves costpalmate………………….……...……13.Nannorrhops
- Trunk unbranched. Leaves palmate……………………………..….……………….7
7+ Gynoecium mono-tricarpellary, free; stamens 5-25………………...... …………...8
- Gynoecium tricarpellary, fused either by thin style or up to their midregions or entirely fused; stamens 6………………………………………….………..…..….. 9
8+ Stem up to 3 m tall and 5 cm wide; stamen filaments connate, upper 3 filaments adnate to petals………………………………...………...... …19.Rhapis
- Stem up to 25 m tall and 30-75 cm wide; stamen filaments distinct, adnate to petals…………………………………………….………...... 18. Trachycarpus
9+ Leaf petiole spiny at the margin; gynoecium tricarpellary, free at the base, fused by their style only………………………………………………………..………..….10
50
- Leaf petiole smooth at the margin; gynoecium tricarpellary, entirely fused or connate at the base, style free…………………………………………..…..21. Sabal
10+ Sepals distinct, imbricate……………………………………………..…..14.Brahea
- Sepals united, 3-lobed, tubular……………………….…………..…….....……….11
11+ Trunk columnar and massive.Leaf lamina divided 1/3 into bifid single-fold segments. Inflorescence branched up to 3 orders; peduncular bracts flattened and flaring………………………………………………………….……….……...…...12
- Trunk slender and not-massive.Leaf lamina divided into single (multiple) fold segments. Inflorescence branched up to 5 orders; peduncular bracts tubular and not flaring………………………………………………....…….…………15.Livistonia
12+ Petiole unarmed. Inflorescence with long peduncle; flowering branches clustered at the end; corolla forming a deciduous cap at anthesis...... ….……....16.Pritchardia
- Petiole armed. Inflorescence with short peduncle; flowering branches no confined at the apex; corolla not forming a deciduous cap at anthesis……..17.Washingtonia
1 Tribe Borasseae Mart.
Dioecious palm.Leaves palmate or costapalmate, induplicate; staminate or pistillate
rachillae similar or dissimilar.Flowers solitary, usually in cincinni; staminate and
pistillate flowers borne in deep pits.Pistillate flowers single, bibracteolate. Fruit 1-3
seeded; endocarp thick and hard, pore present at the apex.
51
SUBFAMILY CORYPHOIDEAE
Table 1.6 Synopsis of the tribes, subtribes and genera from Pakistan and Kashmir
Tribe Borasseae Mart. (1837)
Subtribe Hyphaeninae Becc. (1924)
Genus BismarkiaHildebrandt & H. Wendl. (1881)
Genus HyphaeneJ. Gaertn. (1788)
Subtribe Lataniinae Meisner (1842)
Genus Borassus L.(1753)
Tribe Caryoteae Drude (1881)
Genus Caryota L.(1753)
Tribe Corypheae Mart. (1837)
Subtribe Coryphinae J. Drans. & N. W. Uhl (1986)
Genus NannorrhopsH. Wendl. (1879)
Tribe TrachycarpeaeDransfield et. al(2005)
Subribe Livistoninae Saakov (1954)
Genus BraheaMart. ex Endl. (1837)
Genus Livistonia R. Brown (1810)
Genus Pritchardia Seemann & H. Wendl. (1862)
Genus WashingtoniaH. Wendl. (1879), nom. cons.; Bailey (1936)
Subtribe Thrinacinae Becc. 1907
Genus Rhapis L.f. ex Aiton (1789)
Genus Trachycarpus H. Wendl. (1863? "1861")
Tribe Phoeniceae Drude (1881)
Genus PhoenixL. (1753)
Tribe Sabaleae Mart. (1837)
Genus SabalAdanson (1763)
52
9.Hyphaene
Gaertn., Fruct. 1: 28.1788; Boiss., Fl. Or. 5: 46.1811; Palm. Borass. 23. 1924; Post, Fl.
Syria, Pales. Sinai, 2: 556. 1932; Tackholm, St. Fl. Egypt, 448. 1956; Hutch., Fl. W.
Trop. Afr. 3:169. 1968; Malik in Nasir & Ali (eds.), Fl. Pak. 153: 14-15. 1984; Dransfield and Uhl in Kubitzki (ed.), Fam. Gen. Vasc. Pl. 4: 306-389. 1998.
Dioecious and Pleonanthic.Stem dichotomously branched, rarely unbranched. Leaves costapalmate; petioles spiny usually with black recurved hooks; adaxial hastula large, abaxial absent, blade divided to 1/3 into single fold bifid segments.Inflorescence intrafolior and branched; staminate inflorescence 1-6 rachillae; pistillate with 1-3.
Flowers small, staminate and pistillate flowers equal in size. Staminate flowers: sepals 3; petal 3, united to form a stalk; stamens 6, filaments free, anthers ovate, bifid at base, dorsifixed. Pistillate flowers: solitary; sepals 3, petals 3; staminode rudimentary, 6-9; stigma 3, sessile, ovary 3-celled.Fruit variable, borne on enlarged pedicel, oblong, sweet.
Seed smooth; endosperm homogenous.
About 10 species found in drier parts of Africa, southwards to Natal, Madagascar, Red
Sea and Gulf of Eilat coasts, coastal Arabia, and West Coast of India. In Pakistan it is represented by only one cultivated species.
1.Hyphaene thebaica (L.) Mart. Hist. Nat. Palm. 3: 225. (ed.I). t. 131-133. 1839; Post, l.c., Tackholm, l.c.; Hutch., l.c.; Malik, l.c.
Corypha thebaica L. Sp. Pl. 1187.1753.
Type: Palma thebaica Pococke, description of the East, and other countries, t: 281, pl.72,
73.1743 (vide Moore & Dransfield, Taxon 28 (1, 2/3): 60. 1979).
53
Dioecious palm, 10-17 m in height. Trunk Y-shaped, c. 90 cm in diameter. Tree is easily recognizable by the dichotomy of its stem forming up to 16 crowns. Bark dark grey.
Leaves 120 x 180 cm, fan shaped, borne in tufts at the ends of branches, leaf blade divided into about 60 cm long segments, entire; petiole 60-100 cm long, sheathing at the base with numerous upwardly curving hooks. Inflorescence same in male and female plant, up to 1.2 m long, with short branches at irregular intervals, 2-3 spikes arising from each branch. Staminate flowers with minute green bracts, sepals 3, tubular at the base, acute, hooded lobes; petals 3, imbricate, obovate and hooded lobes; stamens 6. Pistillate flowers larger than the male, pedicel short, sepals 3, free, triangular, acute and leathery; petals 3, rounded, smaller than sepals, scarious; staminodes 6; ovary globose, trilocular, stigmas 3, sessile. Fruit woody, 6-10 x 6-8 cm, smooth, rectangular to cubical, with rounded edges, shiny brown when ripe, 1- seeded. Seeds 2-3.5 x 3 cm, ivory in colour, truncate obtuse.
Specimens Examined:G-4:Karachi University Botanical Garden, Karachi, dioecious plant, flower present, 10-1-2011,Abid Arzoo Rashid 22 (KUH); ibid; Gulshan-e-Iqbal,
Karachi, cultivated, 14cm high,13-02-2014,Roohi Bano 58(KUH).
Distribution: A native of northern half of Africa. Also widespread in the Sahel and grows from Mauritania and Senegal in the west, through Central Africa, and east to Egypt, Kenya and Tanzania. In Pakistan it is cultivated in the gardens (Karachi).
Ecology:Occurs in coastal habitats.
Flowering Period: March – September
Vern: English: Gingerbread tree (also doom palm), Arabic: Doum palm
54
Uses:Hyphaene is derived from the Greek word 'hyphaino' (web), referring to the fibres from the leaves, which are used for weavingmats, bags, baskets, hats, fans, strainers, bowls, rope, string, nets and coarse textiles. Petiole fibre is used for making sponges and brushes. The roots fibre used for making snares and fishing nets. The wood is used as fuel and for making charcoal. Dry fruits yield a black dye, used for dying leather. Root is used to relieve chest-pain, jaundice, intestinal colic and inguinal hernia. Fruit is eaten against stomach pain and bladder infection.
10.Bismarkia
Hildebrandt & H. Wendl., Bot. Zeit. 39: 90, 93. 1881;Becc., Palmae della Tribe
Borasseae, 1924; Dransfield and Uhl in Kubitzki (ed.), Fam. Gen. Vasc. Pl. 4: 306-389.
1998.
Unarmed.Leaves huge, bright glaucous, silver palmate, held stiffly in the crown; adaxial hastula large; abaxial absent; leaf lamina divided to 1/4 - 1/3 into single fold bifid segments.Inflorescence with 3-7 rachillae; staminate flower 3 in each pit; pistillate flower solitary; stamens 6. Fruit ovoid or bilobed, pedicellate, endocarp flanged endosperm homogenous.
A monotypic genus occurs in drier parts of Madagascar. In Pakistan it is represented by a single species and cultivated as ornamental plants.
1. Bismarkia nobilis Hildebrandt & H. Wendl., Bot. Zeit. 39: 90, 93. 1881.
Medemia nobilis (Hildebrandt & H. Wendl.) Gall. Compt.Rend. Hebd.Gall., Compt..
Séances Acad. Sci. 138: 1120 .1904.
55
Holotype: Madagascar, Beravi Hildebrandt s.n.(FI).
Dioecious palm.Trunk solitary, gray, 30-45 cm in diameter, slightly bulging at the base, reach about 25 m in height but usually get no taller than 12 m in cultivation.
Leavesinduplicate, costapalmate,hastulawedge-shaped, 3 m wide, divided to a third its length into 20 or more stiff, single-folded segments; petioles 2–3 m, slightly armed, covered in a white wax, scales caducous; leaf crown nearly-spherical, 6x 7.5 m.
Inflorescence pendent, interfoliar.Flowers small brown.Fruit drupe, brown when mature, ovoid, 1-seeded.
Specimens Examined:G-4:Karachi University Botanical Garden, Karachi, fruit present brown, 10-03-2010, Abid Arzoo Rashid 03 (KUH); ibid, Abid Arzoo Rashid s.n. (KUH);
Stadium road, Karachi, 17-05-2012, Shaukat 34 (KUH).
Distribution: Same as the genus. In Pakistan it is cultivated in the gardens (Karachi).
Ecology:Found asconspicuous component of savannahs.
Flowering Period: March – September
Vern: English: Bismark palm
Uses:The trunk is usedwhole or split for house construction; leaves for thatching and also planted asornamental plants.
11. Borassus
L. Sp. Pl. 1187. 1753; Gen. Pl. ed.5. 494.1754; Becc. & Hook. f. in Hook. f., Fl. Brit.
Ind.6: 481. 1984; Malik in Nasir & Ali (eds.), Fl. Pak. 153: 5-6. 1894; Dransfield and Uhl in Kubitzki (ed.), Fam. Gen. Vasc. Pl. 4: 306-389. 1998.
56
Dioecious palm.Leaves costapalmate; blade suborbicular to flabellate, divided to ½ into single-fold bifid segments; 90-120 cm across; leaflets induplicate; petiole armed with irregular spines; adaxial hastula large, abxial small. Inflorescence spadix, very long, peduncle sheathed everal bracts. Staminate flowers small, mixed with scaly bracts; with sperianth glumaceous, sepals 3, wedge-shaped, imbricate; petal 3, smaller than the sepals, obvate – spathulate, imbricate.Stamens 6, filaments connate with corolla, anthers large, oblong; pistillodes of 3 bristles. Pistillate flowers much longer, solitary, globose; perianth fleshy, sepals 3, reniform, imbricate; petals 3, convolute; staminodes 6-9; ovary globose, subtrigonous, 3-4 celled, stigma 3, sessile, recurved. Fruit large, sub globose drupe, 15-20 cm in diameter, 1-3 seeded, endocarp internally ridged.Seed bilobed.
Seven species recognized, but probably three or four to be maintained, distributed in
Africa, Madagascar, Northern Arabia, through India and South East-Asia to New Guinea and Australia. In Pakistan, it is represented by one cultivated species.
1. Borassus flabellifer L. Sp. Pl. 1187. 1753; Becc. & Hook. f., l.c.; Malik, l.c.
Borassus flabelliformis Murr. Syst. Ed. 13. 1827; Roxb. Cor. Pl. i.50, t.71, 72; Fl. Ind.
3:790.1832.
Lectotype: Ampana Rheede, Hortus Indicus Malabaricus 1: 13-14. Pl. 10: Carimpana
Rheede. op. cit. 11-12, pl. 9. 1678-1703 (vide Moore & Dransfield in Taxon 28: 60, 62.
1979).
Solitary palm.Stem 20 m tall, grey with well-defined leaf scars. Leaves 150-180 cm long including petiole and sheath; petiole up to 7 cm wide, robust, bright yellow, costa 60-110 cm long; adaxial hastula conspicuous, abaxial hastula rudimentary; leaf blade up to 150
57
Fig. 1.6A: Bismarkia nobilisHildebrandt & H. Wendl.B: Borassus flabelliferL.
58 cm in diameter; leaflets 62-65 in number, 4-10 cm wide, acute.Staminate flower 0.2-0.6 cm long, bracteoles 0.4-0.7 cm long; calyx 0.3 x 0.15 cm; petals lobe 0.1 x 0.1 cm; stamens 6 with very short filaments up to 0.03 cm. Pistillate flowers 3 x 3 cm; bracteoles large, 2 cm in diameter; sepals 1.5 x 2 cm; petals 1 x 1.5 cm. Fruit massive, 8-13 x 7-16.5 cm, yellowish black, ovoid, rounded or flattened at the apex.
Specimens Examined:G-4:P.E.C.H.S Block 2, stem c. 22 m high, flower present, 17-10-
2011,Abid Arzoo Rashid 117 (KUH); Department of Botany, University of Karachi, 12-
03-2012, Shoukat Ali19 (KUH); Botanical Garden, University of Karachi, flower present, c.27 m tall, 12-09-2012, Abid Arzoo Rashid 65 (KUH).
Distribution: South and Southeast Asia, ranges from western India through Indo China to the Lesser Sunda Islands of Indonesia.Also introduced in China, Malaysia and Pakistan
(Whitmore, 1973; Malik, 1984 & Pei, 1991).
Ecology:It occurs between sea level and 800 m, more abundant in low altitude, particularly common in coastal areas with sandy or alluvial soil, commonly grown along the margin of rice paddies, flood plains and river valleys.
Flowering Period: March – September
Vern: Palmyrah palm
Uses: The leaves are used for thatching, weaving. The wood is thought to be termite resistant and also used for construction houses, canoes, fence posts. The fruit, undeveloped endosperm and apical bud (palm cabbage) are consumed and the inflorescences are tapped for the sweet sap.
59
2. Tribe CaryotaeDrude
Palms hapaxanthic and unarmed; inflorescence bisexual or unisexual by reduction of triads. Leaves induplicate, bipinnate or imparipinnate; entire and induplicately ribbed; leaflets primrose. Pistillodeabsent. Gynoecium tricarpellary, 3-chambered, ovules inserted at base. Stigmatic remains present on fruit. Epicarp smooth and undifferentiated, mesocarp fleshy. Seed present with black seed coat.
12.Caryota
L., Sp. Pl.1189.1753; Gen. Pl. ed.5. 495. 1754; Becc. & Hook. f. in Hook. f., Fl. Brit.
Ind.6: 422. 1894; Malik in Nasir & Ali (eds.), Fl. Pak. 153: 7-9. 1984; Dransfield and Uhl in Kubtzki (ed.), Fam. Gen. Vasc. Pl. 4: 306- 389. 1998.; Dowe,Austr. Palm, Biogeog.
Eco. Syst. 305. 2010. p.p.
Hapaxanthic, monoecious palm.Stem slender to robust, leaning or erect, leaf scars prominent.Leaves bipinnate, persistent rarely deciduous; leaflets wedge shaped, induplicate.Inflorescence interfoliar or intrafoliar, rarely spicate, usually paniculate, branched to 1 or 2 orders. Staminate flowers symmetrical, sepals 3, imbricate; petals 3, valvate; stamens 6-(ca.100), anthers linear.Pistillate flowers globular in bud; sepals and petals 3 each, staminodes 0-6; ovary tricarpellary with 1-3 fertile ovules, stigma trilobed.Fruit 1-2 seeded.Seed globose or hemispherical.
A genus with about 12 species occurs in Srilanka, India, Southern China, southwards through SE Asia, Malaysia to northern Australia, Solomon Islands, and Vanvatu. In
Pakistan it is represented by single cultivated species.
60
Fig. 1.7 Caryota urensL.
61
1. Caryota urens L., Sp. Pl.1189.1753; Gen. Pl. ed.5. 495.1754; Becc. & Hook. f. in
Hook. f., l.c.; Parker, Fl. Punj. Haz. Del. 525 (Reprint ed.) 1956; Malik, l.c. 8. f. 1.
Plant with solitary – trunked, annulate, erect, up to 12 m in height and 30 cm wide, widely spaced leaf scars rings cover the trunk, leaf crown up to 6 m tall.Leaves bipinnate triangular in shape, bright – deep green, up to 3.5 in. long; petiole stout, 6 m tall. Pinnae obdeltoid up to 30 cm long with a pointed and a jagged edge.Inflorescence axillary, 3m long, emerge at each leaf node, producing pendent cluster of white, unisexual flowers, peduncle curved, covered with a large bract.Male flowers much larger than the female; calyx 3- lobed, ciliate; corolla 3- lobed, valvate and woody. Stamens 40 -45 in numbers, as long as the petals, anthers acuminate, basifixed, filaments 1 – 1.5 mm long, pistillode absent; in female flower sepals and petals same as in the male flowers; staminodes 3.
Fruit round, 2 cm in diameter, reddish and juicy, 1-2 seeded.
Specimens Examined:G-4:Ghandi garden Karachi, 10ft, 02-10-77, Kamal Akhter , 665
(KUH); Ghandi garden Karachi, 10ft, 02-10-1977, Kamal Akhter, 667
(KUH);N.Nazimabad Sakhi hasan Karachi, cultivated, tree, 35ft, spadix very long about
10ft, drooping buds green, 07-05-1978, Kamal Akhter Malik 844 (KUH).
Distribution: Tropical Asia, Malaya, India, southwards to Ceylon, Burma and
Singapore. In Pakistan it is cultivated rarely as an ornamental plant.
Ecology: Found in monsoon forest.
Flowering Period: May- October
Vern: English: Sago palm
62
Uses: Stems provide sago used for flavouring; wood is used for construction purposes; leaf sheath fibres are extremely tough and yielded for thatch, cordage. The woolly hairs of leaf sheaths, petioles, and rachis is used for tinder or wadding. Inflorescences especially tapped for palm wine or sugar and also cultivated as ornamental plants.
3.Tribe Corypheae Mart.
Leaves induplicately palmate, costapalmate, or undivided; hermaphrodite or if unisexual, only dimorphic flowers borne singly or in cincinni; peduncular bracts present, rachillae bracts well developed; rachillae lacking deep pits; 1-seeded or if more than lobed; endocarp thin, undifferentiated, crustaceous or cartilaginous.
13.Nannorrhops
H. Wendl. in Bot. Zeit. 37: 148. 1879; Hook. f. in Benth. & Hook. f. Gen. Pl. 3: 923,
1833; Becc. & Hook. f. in Hook. f. Fl. Brit. Ind. 6: 429. 1892; Moore, Jr. in Rech. f., Fl.
Iran.146: 1. 1980; Malik, in Fl. Pak. 153: 18-20.1984;Dransfield and Uhl in Kubitzki
(ed.), Fam. Gen. Vasc. Pl. 4: 306-389. 1998.
Shrub, hapaxanthic, hermaphorditic and glabrous palm.Stem erect or dichotomously branched, branching up to 10 ft., unarmed.Leaves costapalmate, sheath splitting, hastula absent; leaf lamina divided into single-fold, segments bifid; petiole short.Inflorescence spadix, interfoliar, up to 4 orders, spathes tubular, pedunculer branch 0-several.Flowers polygamous, in cincinnus of 1-3 (-7), 3-lobed, tubular calyx.Corolla 3-partite, stalk like basally, valvate.Stamens 6.Carpels 3, connate except at base, style short, stigma 3. Fruit globose or ovoid, drupe, 1 – seeded.
63
A monotypic genus of semi desert areas of Middle East, Iran, Afghanistan, Pakistan and northwestern India
1. Nannorrhops ritichiana (Griff.) H. Wendl., in Bot. Zeit. 1879; Aitch., J. Linn. Soc.
19: 187. 1882; Beccc. & Hook. f., l.c..; Parker, For. Fl. Punj. Haz. Del. 524 (reprint ed.)
1956; R.R. Stewart, Ann. Cat Vasc. Pl. Pak. Kashm. 33. 1972; Malik, l. c., 19. f. 5;
Dransfield and Uhl, l.c.
Chamaerops ritichiana Griff in Calc. Jr. Nat. Hist. 5: 342. 1844.
Lectotype: Fruit from plant cultivated in Saharanpur Botanical Garden (Beccari, Ann.
Roy. Bot. Gard. Calcutta. 13: 38.1933).
Shrub like clumping palm, clustering and tufted, growing up to 6 m tall. Leaves fan shaped, 30-60 (130) cm long, coriaceous, leaf lamina 60-90 cm long, divided from the middle or lower into 20-30 rigid leaflets with often interposed fibres, induplicate, each leaflet 30-120 cm long; petiole 6-12 inch, serrulate. Inflorescence produced on a 2-3 m tall stem at the top of the trunk, forming an open panicle. Flowers 3 enclosed in a membranous spathella, one sessile and remaining pedicellate, ebracteate. Sepals 2-3 mm long, membranous forming a trilobed tube; petal 4 x 2 mm, coriaceous. Stamens inserted in the corolla tube, dorsifixed, deeply sagittate.Ovary narrowed into the style.Fruit drupe, globose or ellipsoid, brown, from the size of a pea to a bullet.
Specimens Examined: C-6: Hangu Kohat, Muhammad Mosharaf Bellal 68 (RAW);
Kurram valley, Sadela-Arawali, Y.Nasir & Siddiqi 8168 (RAW); Alizai Parachinar road,
M.A.Siddiqi 4332 (RAW); D-5 Hernal, Mohindar Nath 2423 (RAW); F-4: c.40 miles from Seeta village on way to Kute-ji-Qabar, Keerthar range, c.6000 ft., Kamal Akhtar
64
Malik & Saood Omer 2466 (KUH); c.60 miles from on way to Kute-ji Qabar, Kerther
Range, very common, shrub, 21-03-1983, Kamal. A. Malik, Saood Omer& Abdul Wahid
2466(KUH); G-3: Bela Awaran road, 33 miles before Awaran, sandy clay soil, S. I. Ali,
S.A. Farooqi & Sultan Abedin 1443 (KUH). c.25 km from Awaran, common palm in river bed, fruit reddish on maturity, eaten by locals, 21-09-1986, AbdulGhaffoor & Saood
Omer 1731 (KUH); 830 miles before Awaran, Bela-Awaran Road, Jhal Jao Mekran, , 10-
10-1978, Kamal Akhter Malik & S. Nazimuddin 1031(KUH); c.45 miles from uthal on way to Khuzdar, c.1-1.5m, red fruit present, 04-05-1980, Saood Omer 1153(KUH); c. 25 km from Awaran on way to Mangri Mashke, Awaran Khuzdar road, Rhizomaetous tufted, 2-3 m tall, common in almost all Wadi beds, 08-03-1990, A-Ghafoor &Steve M.
Goodman 4419 (KUH).
Distribution:A native to southwestern Asia, from the southeast of the Arabian
Penninsula east through Iranand Afghanistan to Pakistan and north western India. In
Pakistan Chatai palm is wild in Sind, Balochistan, lower Kurram valley, N.W.F.P.; the
Salt range, W. Punjab, Mekran, Kohat (R.R. Stewart, 1972)
Ecology:A semi desert plantgrowing up to 1600 m, on dry mountainous area, on lime stone.
Flowering Period: July-November
Vern: English: Mazari Palm (Chatai palm, vide R.R. Stewart, 1972)
Uses: In its native habitat, the young leaves, inflorescence and fruits are eaten. Other parts of the palm are used for thatching, fiber for weaving and rope, and for fuel.
65
4. Tribe Trachycarpeae Dransf., et al.
Pleonanthic, hermaphrodite or rarely unisexual.Leaf palmate or costapalmate; leaf sheaths form a mass of brown interwoven fibres; adaxial hastula present, abaxial absent or reduced.Stamens 6.Gynoecium tricarpellary, wedge shaped, distinct at the base, only united by their style; stigma trilobed
14.Brahea
Mart. ex Endl., Gen. Pl.:252. 1837; L. H. Bailey, Gentes Herb. 4 (3):119-125. 1937;
Henderson et al., Field Guide: 54-56.1995; Dransfield and Uhl in Kubitzki (ed.), Fam.
Gen. Vasc. Pl. 4: 306-389. 1998.
Clustered palm.Sheath becoming fibrous, eventually splitting at the base. Leaf lamina divided nearly ½ in to single fold, deeply bifid; segments stiff or flexible; petiole toothed or unarmed. Inflorescence branched to 4 orders; bracts 0-many.Flowers solitaryor in cincinni of 2-3; sepals 3, distinct, imbricate; petals 3, tubular at the base, valvate at the apex; stamens epipetalous, filaments connate in 6-lobed.
A genus with c. 16 species distributed in Baja California, Guadalupe Island, Mexico and
Guatemala. In Pakistan it is represented by single species.
1. Brahea brandegeei (Purpus) H.E.Moore, Baileya 19: 168.1975.
Stem clothed with persistent leaf sheaths. Leaves induplicate, shortly costapalmate, marcescent; sheath becoming fibrous, persistent, petiole concave, flattened, blade nearly orbicular, regularly divided deeply bifid, surfaces glabrous, waxy or covered in caducous, floccose indumentum, midribs prominent, Inflorescences solitary, interfoliar, branched to
4 orders; prophyll 2-keeled, tubular, peduncular bracts 0–several, like the prophyll but
66 rachis much longer than peduncle.Flowers spirally arranged, solitary or in cincinni of 2–
3; sepals 3, distinct, imbricate, margins minutely toothed petals 3, united basally in a tube as long as the sepals, briefly imbricate, valvate apically; stamens 6, filaments connate into a 6-lobed ring, , anthers broadly elliptic to nearly oblong, dorsifixed; carpels 3, united by the styles, ovule basal, erect, Fruit, globose or ovoid, dark blue to black at maturity, basal, stigmatic remains apical; epicarp smooth, mesocarp fleshy, endocarp crustaceous. Seed basally.
Specimens Examined: G-4: Defence phase 2 Karachi, cultivated, Abid Arzoo Rashid
137 (KUH); Shaheed-e-millat road Karachi, flowers yellow, Abid Arzoo Rashid 139
(KUH).
Distribution: same as the genus.
Ecology:Growing at altitudes of up to 1600 m
Flowering Period: July-November
Vern: English:
Uses: The leaves are used for thatch and as a source of fibre. Fruits of some species are edible. Attractive ornamentals for drier areas
15.Livistonia
R. Br., Prodr. 267. 1816; Becc. & Hook. f. in Hook. f.,Fl. Brit. Ind. 6: 434. 1894; Becc., in Ann. Roy. Bot. Gard. Calc. 13: 43-108.1933; Whitmore, Palms Mal., 70. 1973; Malik, in Nasir & Ali (eds.), Fl. Pak. 153: 15-16.1984; Dransfield and Uhl in Kubitzki (ed.),
Fam. Gen. Vasc. Pl. 4: 306- 389. 1998;Dowe, Austr. Palm, Biogeog. Eco. Syst. 305.
2010.
67
Solitary, pleonanthic, hermaphroditic, dioecious or functionally unisexual or polygamomonoecious palm. Stem slender-robust. Leaves circular or fan shaped, palmate or costapalmate, narrowly clasping the stem; petiole unarmed or toothed; leaf lamina of single (multiple) fold segments; segments lanceolate, induplicate, tapered at the apex.
Inflorescence interfoliar, branched to 5 orders; peduncular bracts absent or 1- few. Flower solitary or in cicinni, minute, bisexual; sepals 3, tubular, imbricate, lobed apically; petals
3, tubular, valvate, longer than the sepals.Stamens 6, epipetalous, connate in fleshy ring, anthers didymous, ovoid, medifixed. Ovary tricarpellary, free, wedge – shaped, sigma trilobed, style united. Fruit drupe, globose -ovoid, pyriform or ellipsoidal, 1 – seeded.
Seed hollow at the ventral side.
About 25 species are reported from the world, distributed in Africa and Arabia, to the
Himalayas as a Ryukyu Island through Indo China and Malaysia to New Guinea the
Solomon Island, and Australia. In Pakistan it is represented by one cultivated species.
1. Livistonia chinensis(N. J. Jacquin) R. Br. ex Mart, Hist. Nat. Pal. ed. 1. 3: 240. 1839;
Maheshwari, Fl. Dehli 342. 1963; R.R. Stewart, Ann. Cat Vasc. Pl. Pak. Kashm. 33.
1972; Whitmore, l.c.; Malik, l. c., f. 4.
Latania chinensis N.J. Jacquin, Fragm. Bot. 16. t. 11. f. 1. 1801; Livistonia mauritima
Wall. Ex Voigt. Hort. Suburb. Calc. 641. 1845. nom. nud.
Type: Jacq. Fragm. Bot. 16. t. 11, based on a cultivated palm from Mauritius said to have originated in China.
68
Fig. 1.8 Livistonia chinensis(N. J. Jacquin) R. Br. ex Mart.
69
Plant hermaphrodite, solitary, trunk up to 15 m tall and 20 – 30 cm in diameter; leaf scars obscure, internodes narrow. Leaves 40-60 in a globose -ovoid crown, costapalmate, glabrous; leaf lamina regularly segmented, subcirular - reniform, 120 – 200 cm long and
120-180 cm wide, lamina divided into 50 – 90 segments. Inflorescence unbranched at the base, 100-120 cm long; peduncular bracts absent. Flower in clusters of 4-7, subglobose in bud, 2-2.5 mm long, white-yellowish. Sepals imbricate, 1.5 mm long, rounded; petals triangular, fleshy, about 2.2 x 2 mm. Filament united at the base, anthers 0.5 mm long.
Fruit oval, bluish green, 15- 26 x 9- 18 mm, fleshy.
Specimens Examined:C-6: Peshawar on road side, cultivated, tree, c.6m, spadix 3ft long, inflorescence interfoliar, trunk not smooth, covered with leaf bases, 25-08-1978,
S.Nazimuddin &Sultan Abedin 1200(KUH); G-4: Ghandi garden Karachi, 15ft, with fruit (green), 02-10-1977, Kamal Akhter Malik 663(KUH); Mazar-e- Quaid-e-Azam
Karachi, cultivated, tree, yellowish, 12-03-1978, Kamal Akhter Malik 785 (KUH); ibid;
Lawarence garden Lahore, cultivated, 01-07-1978, Kamal A. Malik 1184(KUH); Karachi
University near Ibrahim mosque,24-07-1999, Shaheen Faizi s.n. (KUH).
Distribution:Southern Japan, Ryukyu Island, Bonin Island. In Pakistan cultivated as ornamental plants.
Ecology:It is commonly cultivated on the plains, growing as pot plants and tubs on veranda (R.R. Stewart, 1972)
Flowering Period: February-March
Vern: English: Chinese Fan Palm
Uses: Leaves are used for making fans and fiber used for weaving and making rope.
70
16.Pritchardia
Seemann & H. Wendl. ex H.Wendl., Bonplandia 10:197.1862., nom. cons.; Becc. &
Rock, Mem. Bernice P. Bishop Mus. 8 (1): 1-77.1921; Hodel, Principes, 24 (22): 25-81-
1980. Dransfield and Uhl in Kubitzki (ed.), Fam. Gen. Vasc. Pl. 4: 306-389.1998.
Plant solitary, unarmed,vary in height, ranging from 20 to 130 ft.trunk columnar, smooth or fibrous, obscurely ringed by leaf scars. Leaves costapalmate; leaf lamina divided 1/3 -
1/2 into single fold bifid segments. Inflorescence solitary or cluster of 2-4, branched to 3 orders; peduncular bracts numerous.Flower solitary; calyx 3-lobed, tubular; corolla tubular at the base, 3-lobed, valvate, forming a deciduous cap; filament basally connate.
A genus with about 38 species, distributed in Fiji, Tonga, Danger Islands and Hawaii.
In Pakistan one species is cultivated. i. Pritchardia beccariana Rock, Bull. Torrey Bot. Club 43: 386 .1916.
Pritchardia beccariana var. giffardiana Becc., Mem. Bern. Pau. Bish. Mus. 8: 59. 1921.
Type: Holotype: Glenwood, Hawaii, Hawaii Rock 10356(BISH!), Isotype: Glenwood,
Hawaii, Hawaii Rock 10356(FI, GH)
Plant 20 m tall; fibers present at proximal margins of petiole; leaf blade nearly flat, divided 1/5-1/4, abaxial surface incompletely covered with scattered lepidia, segment tips stiff; inflorescences composed of 2-4 panicles, shorter than or equaling petioles in flower and fruit, panicles branched to 3 orders, rachillae glabrous to covered with scurfy indumentum in flower, glabrous in fruit; fruits 40 x 30-40 mm, globose to ellipsoid.
71
Specimens Examined:G-4: Botany Department Karachi University, cultivated, 17 m tall, 25-05-2010, Abid Arzoo Rashid 200 (KUH); Gulshan-e-Iqbal Karachi, Abid Arzoo
Rashid 221 (KUH).
Distribution:Hawaii. Same as the genus
Ecology:on slope at the elevation between 300-1300 m, on wet forest.
Flowering Period: February-March
Vern: English:
Uses: Leaves and fibres are used for making fans, and for umbrella.
17.Washingtonia
H. Wendl. in Bot. Zeit. 37:68, 148. 1879, nom. cons.; Hook. f., in Benth. & Hook. f.,
Gen. Pl. 3: 923. 1883; Becc., in Ann. Roy.Bot. Gard. Calc. 13: 307-308. 1933; Bailey in
Gentes Herb. 4:51-82. 1936; Palabin in Komarov, Fl. U.R.S.S. 3:374.1964; Malik, in
Nasir & Ali (eds.), Fl. Pak. 153: 15-16.1984; Dransfield and Uhl in Kubitzki (ed.), Fam.
Gen. Vasc. Pl. 4: 306-389. 1998.
Robust tree. Stem tall, massive, unbranched, upper part is densely covered with dead leaves and petioles. Leaves costapalmate, orbicular, plicate, leaf sheath split below petiole, margin fibrous; leaf lamina deeply divided 1/3 into bifid, single-fold segments, segments pendulous, induplicate; petiole armed, planoconvex, margins with hooked spines. Inflorescence axillary, much branched, branched to 3-4 orders, branches more or less zigzag, slender, bracts flattened and almost woody. Flowers solitary, bisexual, white; calyx cup-shaped, tubular with 3 tattered lobes; corolla tubular for ¼ of its length, lobes
72 valvate, longer than calyx with narrow reflexed lobes; stamens 6, in two series, filaments free, fusiform; carpels 3.Fruit small black, ovoid-oblong with thin pericarp.
A genus with two species, native to the southwestern United States (in southern
California, southwest Arizona, Texas) and northwest Mexico (in northern Baja
California and Sonora). In Pakistan it is represented by 2 cultivated species.
Key to the Species
1+ Petioles of young plants are green and without thorns. Basal sheath of the leaf blade
does not have a bright tawny-colored patch. Leaf crown of mature plant has loose
and open arrangement. Leaflets pendulous and not stiff, cottony threads persist.
Trunk barrel shaped rarely exceeds 60 ft in height………...1.Washingtonia filifera
- Petioles of young plants are brown and distinctly thorny. Basal sheath of the leaf
blade has a bright tawny-colored patch. Leaf crown of mature plant is dense and
compact. Leaflets stiff and cottony threads fall off with age. Trunk slender (slightly
swollen at the base), exceeds 100 ft in height…………….2. Washingtonia robusta
1. Washingtonia filifera (L. Linden) H. Wendl. in Bot. Zeit. 36:61.1879; Bailey, l.c.,
3507. f. 1 E-I; Malik, l.c.
Pritchardia filiferaL. Linden, Catalogue 95:13.1876.
Pritchardia filamentosa Fenzi in Bull. Roy. Bot. Soc. Tosc. Ort. 1: 116.1876;
Washingtonia filamentosa (Fenzi) O. Ktze., Rev. Gen. 2: 737.1891; Neowashingtonia filamentosa (Fenzi) Sudw. Forest Trees Pacif. Slope 199.1908.
Stem robust, up to 24 m tall, 60-120 cm in diameter, trunk massive gray, barrel shaped, ringed with old and dead leaf scars. Leaves costapalmate, each leaf 3-6 ft (0.9-1.8 m) 73 across, spread out to form a dense, loose and open crown; circular in outline, divided from the middle into 70-75 segments; segments glabrous, pendulous, a long thread hanging at the sinus of each segment; costa prominent, hastula at the apex of petiole triangular, whitish, spongy; petiole long, plano-convex, armed along the margin with curved thorns, spines yellow; cotton-like threads on and between the leaflets persist even when the palm is mature. Inflorescence among the foliage, longer than the leaves, declined, bracts tubular at base, sheathing the branches and branchlets, flattened at anthesis. Flowers bisexual, white, subsessile; calyx 3-lobed, campanulate, c. 3 mm long; corolla 3-lobed, 6-7 mm long, papery, lanceolate, acuminate; anthers dorsifixed, versatile, hastate, filaments fusiform. Ovary tri-lobed, stigmas 3, style filiform, exserted. Fruit red- black, c. 8 x 6 mm, oblong or rounded, 1-seeded, c. 0.6 cm in diameter.
Specimens Examined:G-4: Near Feroz sons Ibrahim Chanderigur road Karachi, cultivated, tree, 30ft, white spadix very large ( 8ft ), 20-4-1978, Kamal Akhter Malik,
839 (KUH); ibid; Shahra-e- Faisal, Karachi, 18-4-2011, Abid ArzooRashid 66 (KUH);
Karachi University Campus, Karachi, flower white, bisexual, fruit present, 5-2-2012,Abid
Arzoo Rashid& Adil 84 (KUH).
Distribution: Western United States (southeastern California, Arizona & Baja
California)
Ecology:Found in dried open area.
Flowering Period: February-April
74
Fig 1.9 Washingtonia filifera(L. Linden) H. Wendl.
75
Vern: English: Desert fan palm, Petticoat palm.
Uses: Leaves are used for making fans and fiber used for weaving and making rope.
2. Washingtonia robusta H. Wendl. in Bot. Zeit. 36:61.1879.
Neowashingtonia robusta (H. Wendl.) A. Heller
Stem solitary, tapered, erect, 22 m tall and 80 cm in diameter, gray trunk ringed with closely set leaf scars. Leaves costapalmate, induplicate, split about ½ the leaf blade in to numerous segments with stiff apex and threads (Marginal fibres) swing between segments. Leaf crown of mature plant is dense and compact; petiole split, armed with distinct, curved; adaxial hastula prominent with tattered, paper margins; leaf blade densely covered with white hairs at the lower side. Inflorescence arched, numerous, branched, at least 3 orders, 2.5-3 m in length. Flowers hermaphrodite, fragrant, creamy white, 8 mm long. Fruit bluish- black or brownish-black or black when ripe, spherical to pear shaped, 7-10 mm long, up to 1 cm wide.
Specimens Examined:G-4: Abrahim Chanderigur road T&T office Karachi, cultivated,tree, 6 m, with fruit, 20-5-1978,Kamal Akhter Malik 847 (KUH).
Distribution: Palm native to North America, Mexico and Spain.
Ecology:Desert palms found along streams and valleys.
Flowering Period: June- August
Vern: English: Mexican Fan Palm, Mexican Washingtonia, Skyduster
Uses:A widely used vertical accent for its form and foliage, good for tropical effects; street or lawn tree.
76
18.Trachycarpus
H. Wendl in J. Gay, Bzull. Soc. Bot. Fr. 8:429 (1863?”1861”), Becc. & Hook. f. in Hook. f., F1. Brit. Ind. 6 : 435.1894; Becc. in Ann. Roy. Bot. Gard. Calc. 13: 272-286. 1933;
Malik, in Nasir & Ali (eds.), Fl. Pak. 53: 30.1984; Dransfield and Uhl in Kubitzki (ed.),
Fam. Gen. Vasc. Pl. 4: 306-389. 1998.
Solitary or clustered, acaulescenet or caulescent, unarmed or armed, dioecious or polygamous. Leaves palmate, in terminal crown; leaf lamina fan shaped or circular, single-fold segments, bifid; petiole toothed or unarmed, rachis absent. Inflorescence interfoliar, stout with coriaceous and tomentose branches, peduncular bracts 1-3. Flowers solitary or in clusters of 2-3, similar in both male and female. Sepals 3, connate at the base; petals 3, ovate, imbricate. Stamens 6, filaments free, distinct and fleshy, anthers dorsifixed, staminodes present or absent. Ovary tricarpellary, stigma 3, ovule basal. Fruit kidney shaped to oblong. Seeds reniform, ventrally grooved.
A genus with 6 species, distributed in Himalayas to northern Thailand, Burma, Chinaand
Japan. In Pakistan it is represented by one species.
1. Trachycarpus fortunei (Hook. f.) H. Wendl in J. Gay, Bull. Soc. Bot. Fr. 8:429, 1863
(1861); Malik, l.c.
Chaemaerops fortune Hook. f., in Curtis Bot. Mag. t. 5221. 1860.
Type: Plant grown at Kew, introduced by Mr. Fortune in 1849.
Trunk upto 6 m tall and 40 cm in diameter, covered with black fibres. Leaves c.90 cm across, suborbicular; divided in 30-40 segments, segments linear – lanceolate, pendulous; petiole green as long as the leaf lamina. Inflorescence yellowish green. Flowers in
77 clusters of 2-4; maleflower c.4 x 3 mm in size, female flower same as the male; sepals ovate, acuminate; petals 1/3 longer than the sepals, ovate. Fruit black with bluish bloom,
1.25 cm in diameter.
Specimens Examined:G-4: I. I Chanderigur road Karachi, cultivated, tree, showing inflorescence, with fruit, 20-5-1978 Kamal Akhter Malik 47 (KUH); Amir Khusro road,Abid Arzoo Rashid 77 (KUH).
Distribution: Indigenous to China and Japan,south to northern Burma and northern
India. In Pakistan it is rarely cultivated.
Ecology:growing at altitudes of 100–2400 m (328–7,874 ft) on limestone hills.
Flowering Period:March - June.
Vern: English: Hemp palm, Windmill palm
Uses:Stem and leaf sheath fibres are used for making rope, sacks, brushes and raincoats; seeds are used medicinally and are believed to have anticancer properties and also grown as ornamental plant.
19. Rhapis
L. f. ex W., in Aiton, Hort. Kew, ed. 1. 3:473. 1789; Becc., in Ann. Roy Bot. Gard.
Calc. 13: 243-253.1933; Whitmore, Palm. Mal.103. 1973; Malik, in Nasir & Ali (eds.),
Fl. Pak. 153: 26- 27.1984; Dransfield and Uhl in Kubitzki (ed.), Fam. Gen. Vasc. Pl. 4:
306- 389. 1998.
Lectotype: R. flabelliformis L’Hérit. ex Aiton (illegitimate name) = R. excelsa (Thunb.)
A. Henry ex Rehder). Mart., Hist. Nat. Palm 3: 253. 1838; Kunth, Enum. 1, Pl. 3: 251.
1841; Benth. et Hook.f., Gen. Pl. 3 (2): 930. 1883. 78
Plant dioecious or polygamodioecious and unarmed. Stem reed-like, leaf sheath persistent and fibrous. Leaves palmate or deeply bifid, induplicate, splitted between the folds into leaflets; leaflets toothed, blunt at the tips, varying in number of ribs, midrib absent.
Inflorescence small, axillary. Flower solitary, unisexual. In staminate flower calyx cup- shaped; corolla tubular; stamens 6, filaments elongate, epipetalous; pistillodes minute.
Pistillate and hermaphordite flower same, ovary tricarpellary, free. Fruit berry, 1-3 per flower, one-seeded. Seed thin, stony.
A genus with c.12 species, distributed from Japan, southern China, south through
IndoChina to Peninsula Thailand and northern Sumatra. In Pakistan it is represented by twocultivated species.
Key to the Species
1+ Segments up to 375 mm long with relatively straight sides, apices usually truncate
with regular dentate secondary splitting; inflorescence with rachis pale brown,
glabrous; filaments keeled ………………………………………………1. R. excelsa
- Segments up to 450 mm long with slightly curved sides, apices usually oblique,
secondary splitting with irregular appearance; inflorescence with rachis pale or dark
brown, glabrous or tomentose; filaments terete ……………….…..….2. R. multifida
1.Rhapis excelsa (Thunb.) Henry ex Rehder in J. Arn. Arb. 11:153. 1930.
Chamaerops excelsa Thunb., Fl. Jap.130. 1784.
Trachycarpus excelsus (Thunb.) H.Wendl., Bull. Soc. Bot. France 8: 429 1861; R.R.
Stewart, Ann. Cat Vasc. Pl. Pak. Kashm. 34. 1972; Rhapis flabelliformis L’ Her. ex Ait,
Hort. Kew, 3:473; Becc., 1.c., 244-246.
79
Fig. 1.10 A: Raphis excels(Thunb.)Henry ex Rehder B: Raphis multifidaBurret
80
Lectotype: Japan, C.P. Thunberg 24386 (UPS),photo K.
Clustering palm, 90-120 cm in height and up to 8 cm in diameter, enclosed by the leaf- sheath and fibrous network. Leaves glossy, about 30 cm across, palmately divided usually to the base; leaflet 6-9 in number, drooping, hastula fibrous; petiole c.20 cm long,
4 mm wide; margin often smooth rarely scabrid; segments 1 (4)-13, folds 11-25, 375 mm long, broad, relatively straight sides, tapered at the base and apex.Inflorescence male and female similar in general appearance, branching to 2 or 3 orders; rachis bracts 2-3, rachis
260 mm long, 4–8 mm diam. Flowers unisexual, sessile and yellowish, densely packed on the rachillae. Male flowers globose when young, elongating when mature, 5.2× 3.8 mm; calyx 2.8 mm, lobes 2 mm, usually with a regular margin; corolla sometimes narrowed into a short receptacular-stalk; filaments, shorter row 2.2 mm, longer row 2.5 mm x 0.4 mm, with adaxial keel, triangular in cross section; pistillode sometimes present.
Female flowers 3.6 × 3.2 mm; calyx 2.3 mm; corolla with a receptacular-stalk 1.1 mm; staminodes present.Fruit 8–10 × 8 mm, borne on a short receptacular stalk, epicarp shiny translucent, minutely papillose, with conspicuous black lenticels, globose seed, c. 3 mm across.
Specimens Examined:G-4: Botanic Garden Center for Plant ConservationUniversity of
Karachi Karachi, cultivated, with green, 20-03-2010Abid Arzoo Rashid 132 (KUH);
Botanic Garden Center for Plant Conservation University of Karachi Karachi, cultivated, yellow flower, 20-04-2010 Abid Arzoo Rashid 134 (KUH).
Distribution: China, Yunnan; South Central China, Hainan; South East China,
Guangdon, Fujian, Hong Kong and Japan.
81
Ecology: grow at the elevation of 3080 ft (939 m), near the river valley andwooded mountain side.
Flowering Period:Febuary - April.
Vern: English: Dwarf Ground Rattan, Broad leaf lady palm
Uses: Sticks and canes are made from the stem of this species.
2.Rhapis multifida Burret, Notizbl. Bot. Gart. Mus. Berlin. 13: 588. 1937.
Type: China, Guangxi, Chen Bien, S.P. Ko 56092 (holotype probably SYS or IBSC)
Stems 2.5m tall. Leaf sheath fibers close together; petiole 4 mm wide, smooth; leaf lamina large; segments 14, folds 30, the longest segments 450 mm, narrow (1 or 2 folds), pointed at the apex, relatively curved. Inflorescence male not seen, femalebranching to 2 orders; rachis bracts 3 or 4, large, tubular, overlapping the base of the next rachis bract, dark brown, 560 mm long, 8–10 mm in diam.Female flowers 3–5 mm apart, 4.5 × 3.0 mm; calyx 2 mm, tomentose, lobes 0.8 mm, margin pale; corolla darkly pigmented, with a long receptacular-stalk, 2.5 mm; staminodes present.Fruit 5 mm long 8 mm diam., borne on a receptacular stalk; epicarp shiny translucent papillose, apical region with conspicuous lenticels.
Specimens Examined:G-4: Botanic Garden Center for Plant Conservation University of
Karachi Karachi, cultivated, 20-08-2011 Abid Arzoo Rashid 136 (KUH).
Distribution: South China, West Guangxi, South East Guangdong.
Ecology:grow at 1000–1500 m, shrub in mixed forest on rocky slopes.
Flowering Period: April - June.
82
Vern: English: Lady Palm, Finger Palm,
Uses: For decoration.
5-Tribe Phoeniceae Drude
Dioecious palm.Leaves induplicately pinnate; leaflets acute apically, lower leaflets modified as spines; inflorescence branched up to single prophyll present; flowers single, dimorphic. Endocarp membranous; seed deeply grooved longitudinally.
20.Phoenix
L., Sp. Pl.1188.1753; Gen. Pl. ed.5. 495.1754; Becc. & Hook. f. in Hook. f., Fl. Brit.
Ind.6: 424-428. 1984; Blatter in J. Bomb. Nat. Hist. Soc. 24 (2): 329. 1916; Malik in
Nasir & Ali (eds.) Fl. Pak. 153: 33.1984; Uhl & Dransfield, Genera Palmarum: 214-
217.1987; Dransfieldand Uhl in Kubitzki (ed.), Fam. Gen. Vasc. Pl. 4: 306-389. 1998.
Dwarf to large, solitary or clustered, armed, pleonanthic, dioecious palms. Leaves induplicately pinnate, sheath reddish brown - brown-black, fibrous; petiole absent to very short; proximal rachis with leaflets modified as acanthophylls. Leaflets regularly arranged or fascicled, single-fold, acute - acuminate. Inflorescence interfoliar, branching to one order, male and female flower same, peduncular bracts absent. Flower solitary, yellow-white. In staminate flowers: sepals 3 lobed; petals 3 (rarely 4), valvate; stamens 6
(rarely 3 or 9). In pistillate flowers: sepals connate, 3; petals imbricate, 3, about two times longer than the calyx; staminodes 6; carpels 3, distinct, follicular and ovoid.Fruit ovoid – oblong.Seed elongate, terete or plano-convex in shape.
A genus with about 14 species, distributed from Atlantic Islands through Africa, Cerete,
Middle East, and India to Hong Kong, Taiwan, Phillippnes, Sumtra and Malaya (Barrow,
83
1998). In Pakistan it is represented by 5 species including one wild species i.e. Phoenix sylvestris
Key to the Species
1+ Leaflets with discolorous surfaces; abaxial surface pale with white ramenta and
tannin stained veins……..…………………….…………………....1. P. roebelenii
- Leaflets with concolorous surfaces; abaxial surface without ramenta and tannin
stained veins…………………………………………….…………………..…...... 2
2+ Solitary palms. Pseudopetiole with acanthophylls, clearly conduplicate, crowded at
leaf base. Fruit golden brown when mature………………..………………….…..3
- Clustering palms. Pseudopetiole with spine-like acanthophylls, sparsely arranged.
Fruit variable, deep red to purplish- brown to black when mature…………….....4
3+ Trunk up to 15 m tall and 100 cm in diameter. Leaflets 200 on either side of midrib,
closely and regularly arranged in plane of orientation...... 2. P. canarieensis
- Trunk up to 20 m tall and 30 cm in diameter. Leaflets 100 on either side of midrib,
closely and irregularly arranged in plane of orientation...... 5. P. sylvestris
- Stem up to 30 m tall. Leaves 96 cm long or more. Fruit up to 4 cm long with thick
fleshy mesocarp. Seed pointed at the apex……………..…………3. P. dactylifera
- Stem up to 4 m tall. Leaves 20-55 cm long or more. Fruit up to 1.75 cm long,
mesocarp not as above. Seed rounded at the apex……………….…...4..P. loureirii
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1.Phoenix canarieensis Hort. ex Chabaud in La Provence Agricole et Horticole IIIust.
19: 293-297, fig. 66 – 68 (1882) ; Parker, For. Fl. Punj. Haz. & Dehli. 529. 1921; Blatt.,
Palms Brit. Ind. 41 , f. 4 (1926); Malik, in Nasir & Ali (eds.) Fl. Pak. 153: 25. 1984.
Type: Described from Canary Islands.
Solitarypalm. Stem 15 – 20 m tall, 120 cm in diameter, trunk marked with broad, diamond shaped leaf base scars. Leaves arching, 5 – 6 m long; leaf base 25 – 30 cm wide; leaflets closely and regularly inserted in one plane of orientation, c.200 on each side, c.
25 – 30 cm long; lamina concolorous, with adaxial and abaxial surfaces glabrous.
Inflorescence elongate Staminate flowers crowded along full length of rachillae; c. 4 mm long; calyx an eyen-rimmed cupule, 1.5-2 mm high; petals 6 x 3 mm, rounded, minutely serrate. Pistillate flowers mostly in distal half of rachillae, calyx cupule c. 2.5 mm long; petals c. 3 x 4 mm. Fruit obovoid, 1.5 – 2 x 1.2 cm, ripening from yellow green.Seed ovoid, 15 x 10 mm, with rounded apices.
Specimens Examined: G-4: Gandhi garden Karachi, cultivated, tree, 20-5-1979Kamal
Akhter Malik848 (KUH).
Distribution: Endemic to the Canary Islands. In Pakistan it is rarely cultivated as ornamental plants.
Ecology: Grow up to 600 m in a range of habitat from humid areas just below cloud forest to semi arid areas.
Flowering Period: March- July
Vern: Palmera Canaria
85
Uses: The leaflets are used for making woven products including across for Palm Sunday
Celebrations. Inflorescence and buds are tapped for the sweet sap which as palm honey.
2. Phoenix dactylifera L. Sp. Pl., 1188. 1753; Roxb., Fl. Ind. ed. 2: 786. 1832; Aitch.,
Cat. Pl. Punjab Sindh: 143. 1869; Boiss., Fl. Or. 5: 47. 1882; Becc. & Hook. f., l.c.;
Bamber, Pl. Punj. 45. 1916; Kitamura, Fl. Afgh. 2: 62. 1955; Jafri, Fl. Kar. 77. 1966; R.
R. Stewart, Ann. Cat. Vasc. Pl. W. Pak. & Kashm. 33. 1972; Moore, Fl. Iranica, 146: 4.
1980; Malik, l.c.
Lectotype: Palma hortensis mas et foemina of Kaempf., Amoen. Exot. Fasc. 668, 686, t.
1, 2. 1712 (vide Moore & Dransfield, Taxon 28:64. 1979).
Solitary, or sparsely clustering palm, suckers producing offshoots at the base. Stem c. 30 m tall, leaf sheaths absent, trunk dull brown, marked with diamond shaped leaf scars.
Leaves erect, obliquely vertical in orientation, 3-5 m long; 15-20 cm wide at the base; pseudopetiole 50-100 cm long; acanthophylls sparsely arranged, c.20 cm long; leaflets variously arranged in 1-3 planes, 50-130 on either side of rachis, stiff, 40 x 2 cm.
Inflorescence covered by hard, boat-like bracts.Staminate inflorescence 12-25 cm long, peduncle 60-90 cm long, glabrous. Male flower sessile, white; sepals trilobed, smaller than the petals, fused forming cupule; petals 3, creamy yellow-white, 3-lobed, valvate, rounded apically, margin serrate, 7-10 x 3-5 mm; stamens 6, c. 5 mm long; pistillodes 3, small, scaly. Pistillate inflorescence initially erect, becoming; pendulous when mature,
90-120 cm long, peduncle 45-75 mm long, glabrous, spikelets many. Female flowers mostly in distal half of rachillae, yellow-white, sweet scented; Sepals 3-lobed, 2-3 x 2 mm, basally united, forming cupule; petals 4-5 x 4 mm. Fruit 4-7 x 2-3 cm, ripening a
86 range of colour from green- yellow to orange, red, brown and black; mesocarp sweet, fleshy or dry. Seeds stony, longitudinally grooved.
Specimens Examined:G-4: Ghandi garden, erect, 02-10-1977, Kamal Akhter 664
(KUH); Karachi University campus, cultivated, tree 30 ft, female spadix 2.5 m long, green, 07-04-1978, Kamal Akhter Malik 837(KUH); ibid; G.P.O, Karachi, cultivated, tree
6 m, 20-05-1978, Kamal Akhter Malik 846 (KUH); Jham Peer, 06-03-1998, Rubina
Saleem s.n. (KUH); Malir Karachi, cultivated, tree up to 4 m tall, fruit present, common,
Dr. Munnuwar Rashid 01 (KUH);
Distribution: W. Asia and N. Africa. In Pakistan occur in Punjab, Sind and also cultivated in lower Balochistan and N.W.F.P.
Ecology:found in dry desert areas.
Flowering Period: March- July
Vern: Khajoor, Aseel, Khurma
Uses: The primary use is of course, their nutritious fruits, eaten fresh, dried or processed.
Date seed are used as fodders. Vegetative parts have diverse uses such as for building material (leaves, trunks), fencing (leaves, midribs), thatching (leaves), rope (leaflets & midrib fibres), fuel (all vegetative parts).
3.Phoenix loureiriiKunth,Enum. Pl. 3: 257. 1841; H. E. Moore, Principes 7 (4): 157,
179. 1963b; Moore, Jr. in Rech.f., Fl. Iran. 146: 4. 1980; Malik, l.c.
Phoenix humulis Royle, III. Bot. Him. 394-397. 1840, nom. nud; ex Becc. & Hook. f. in
Hook f., Fl. Brit. Ind. 6:426. 1894;R. R. Stewart, Ann. Cat. Vasc. Pl. W. Pak. & Kashm.
33. 1972; Phoenix ouseleyana Griff., Cat. Jr. Nat. Hist. 5:347.1845.
87
Lectotype: Pierre 4832 (FB.), collected by Harmand from Mount Kuang Repen in
Cambodia.
Solitary or clustering palm. Stem 1-5m, without leaf sheaths, 10- 30 (40) cm in diam., with crowded diamond-shaped, persistent leaf-bases. Leaves 2 m long; pseudopetiole 20
– 40 cm long; acanthophylls c. 15 on each side of rachis; leaflets arrange more than one plane, proximally fascicled in 3- 4 , 130 on each side of rachis. Staminate inflorescences erect; 40 x 7 cm; peduncle c. 15 cm long. Staminate flowers sweet-scented initially, turning musty, calyx cupule 1.5 – 2 mm high; petals yellow to white, oblong. 4 – 6 x 2 –
2.5 mm, undulate apically; anthers yellow to white. Pistillate inflorescences erect, coriaceous, splitting twice either along or between margins, c. 20 x 3 cm; peduncle 1.5 m long. Pistillate flowers with calyx cupule 1.5-2 mm long, yellow; petals orange-pink to yellow, 2 - 2.5 x 3 – 4 mm. Fruit ovoid to obovoid, 9 – 18 x 5 – 9 mm, green to blue- black when ripe. Seed obovoid, 11 – 18 x 6 – 9 mm.
Specimens Examined:C-8:Kotli, 14-08-1953, A. Rashid s.n.(RAW); Nepag Kotli, 21-
09-1954, A. Rashid 26959 (RAW).
Distribution:Sub-Himalayan belt southwards through India eastwards through Indochina to southren China, Taiwan and Batanes and Sabtang Islands of the Philippines
Ecology: occur at the sea level to 1700 m, in open scrublands, mixed deciduous or pine forests, along road sides or raised ground bordering rice paddy.
Flowering Period: January to April
Uses:Leaflets used in making of mats and brooms, sun-dried juvenile leaves are woven as raincoats (Gruezo Fernando 1985). The apical bud is sweet and can be used as vegetable
88
(palm cabbage). Fruit is sweet and eatable.
4.Phoenix roebelenii O’ Brien Gard. Chron., ser. 3, 6: 475. f. 68. 1889.
Clustering palms up to 3 m tall and 10 cm in diameter, without sheath, erect or twisted.
Leaves arching, 1 - 2 m long, pseudopetiole c. 50 cm long; leaf sheath reddish brown, fibrous; acanthophylls arranged single or paired, c.12, leaflets regularly arranged, 25 – 50 cm on each side, linear. Staminate inflorescences pendulous; 30 – 60 cm long. Staminate flowers with calyx a three-pointed cupule, 1.2mm high, yellow white; petals pale yellow white with acuminate apices, 7 – 8 x 2 – 2.5 mm; anthers 3.5 - 4 mm long. Pistillate flowers pale green; calyx a three-pointed cupule, rigid at the apex, striate, 2 – 2.5 mm in length; petals 3.5 x 4 mm, acute, monocarpellary. Fruit obovoid, 13 - 18 x 6 – 7 mm.
Specimens Examined:G-4:Botanic Garden Center For Plant Conservation University of
Karachi, Karachi, cultivated, white flowers 20-03-2012 Abid Arzoo Rashid 256
(KUH);P.E.C.H.S Karachi, cultivated, 03-04-2012Abid Arzoo Rashid 257 (KUH).
Distribution: Northern Laos, Vietnam and southern China. In Pakistan it is cultivated.
Ecology: Closely associated with riverside or on cliff habitats
Flowering Period: February - October
Vern: Thailand palm
Uses:cultivated as an ornamental plant
89
Fig. 1.11 A & B: Phoenix roebelenii O’ Brien
90
5. Phoenixsylvestris(L.) Roxb., Hort. Bengal.: 73. 1814 and Fl. Ind. ed. 2: 787. 1832;
Royle, III. Bot. Himal. Mts.: 397, nom. 1840; Aitch., Cat.Becc. & Hook. f., l.c.; Bamber,
Pl. Punj. 45. 1916; R. R. Stewart, Ann. Cat. Vasc. Pl. W. Pak. & Kashm. 33. 1972; H. E.
Moore & J. Dransf., Taxon, 28 (1, 2 & 3): 67.1979; Malik, l.c.
Elate sylvestris L. Sp. Pl. 1189.1753.
Lectotype:Katou-Indel Rheede, Hort. Malab. 3: 15-16, pl.22-25. 1682 (vide Mart., Hist.
Nat. Palm. 3: 270-273, t. 136. 1849) see Moore & Dransfield, 1979.
Dioecious and solitary palm.Stem 10-20 m tall, leaf sheaths absent, 20-30 cm in diameter, persistent and diamond shaped leaf bases. Leaves 1.5 x 4 m; leaf sheath reddish brown,fibrous; pseudo-petiole 40 - 50 x 3-5 cm, acanthophylls closely inserted, conduplicate,yellow green; leaflets 15-60 cm long, irregularly fasicled, 80-90 on each side, often waxy, sharp, 18-35 x 1-2.5 cm. Staminate flowers white-yellow, musty scented; sepals 3-lobed, 2-2.5 mm long; petals 3-4, obtuse, somewhat hooded, 6-10 mm long; anthers 3-4 mm long. Pistillate flowers creamy-white, 40-50 in numbers, sepals 1-
2.5 mm long; petals 3, 4 x 4-5 mm long. Fruit drupe, obovoid, 15-25 x 12 mm, orange- yellow, mesocarp somewhat fleshy, sweet and edible. Seed woody, obovoid, rounded at the apex, 15-20 x 7-10 mm, longitudinally grooved on either side.
Specimens Examined:G-4:Near Central Mosque Karachi University, on cultivated land, cultivated tree, 04-01-2007, Asifullah Khan 02(KUH); 36 m from Wana on way to
Jandola, 18-05-1978, S.Nazimuddin,S.Abedin & Hameedullah 975(KUH).
Distribution:Indigenous in the Indus Basin., common in India and Pakistan.In Pakistan also cultivated in Sindh and Punjab.
91
Ecology: In the plains to the coast in low- lying waste lands, scrub forest and areas that have been disturbed or seasonal inundation with water, causing water logging.
Flowering Period: January to April
Vern: Khaji, Khajoor
Uses: Sweet sap is tapped from the stem and used fresh or processed in to a dark sugar
(gur or jaggery) or alcoholic toddy (Davis 1972). Leaves are used in weaving products.Fruits are eaten fresh and also processed as jellies and jams.
6. Tribe SabaleaeMart. ex Dumort
Solitary, hermaphroditic, pleonanthic, unarmed.Leaves costapalmate, induplicate.
Inflorescence interfoliar; peduncular bracts present.Gynoecium tricarpellary, fused,stylar canal present.
21.Sabal
Adanson, Families des Plantes 2:495. 1763; Bec., Ann. Roy Bot. Gard. Calc., 13: 287-
297.1933; Parker, For. Fl. Punj. Haz. Delhi (reprint ed.) 526.1956. Malik, in Nasir & Ali
(eds.), Fl. Pak. 29, 1984; Dransfield and Uhl in Kubitzki (ed.), Fam. Gen. Vasc. Pl. 4:
306-389. 1998.f. 80I, 83J – R.
Dwarf or tall.Acaulescent or erect.Leaves in a terminal crown, palmate or costapalmate, adaxial hastula short, abaxial obscure; leaf lamina divided ½ into single fold bifid segments.Inflorescence interfoliar, branching to 4 -5 orders; peduncular bracts numerous.
Flowers bisexual, white or green; calyx and corolla tubular, 3-lobed; stamens6, filaments connate, epipetalous; ovary tricarpellary, ovules basal, stigma crepitate.Fruit black and globose.
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One of the larger coryphoid genus comprised of 16 species confined to the
CentralWestern Hemisphere from Colombia to north-eastern Mexico and the south easternUnited States. In Pakistan a single species is cultivated as ornamental plants.
1. Sabal minor (N.J. Jacq.) Persons, Syn. Pl. 1:339. 1805; Becc. 1.c., Malik, l.c.
Corypha minor N.J. Jacquin, Hort. Bot. Vindob. 3:8. 1776; Sabal minimum Nut. in Amer.
J. Ser. 1.5: 293. 1822; Vilmorins Blumeng., ed. e. Sieb. Voss, 1:1146.1895.
Type: Described from N. America
Acaulescent, dwarf, subterranean palm with rhizome.Leaves green or bluish, 45-105 cm long; leaf lamina divided 2/3 or more at the base, 20-40, entire or shortly 2-cleft leaflets; petiole larger than the leaf lamina, unarmed, smooth.Inflorescence up to 150 cm long, erect. Flower whitish, c.3 mm long; sepals 3-lobed; petals 3-lobed, imbricate; carpels united.Fruit black, glossy, globose, c. 1 cm in diameter.
Specimens Examined:G-4: Clifton Karachi, flower white, 05-1-2012, Abid Arzoo
Rashid 364(KUH); Gulshan Karachi, 11-05-2012,Abid Arzoo Rashid 368 (KUH).
Distribution:Indigenous to S.E. United States of America. In Pakistan it is rarely cultivated.
Ecology:occur in marshy areas.
Flowering Period: February-March
Vern: English: Dwarf sabal, Dwarf palm or Bush palm.
Uses: Leaves are used for making fans and fiber used for weaving and making rope.
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GLOSSARY OF TAXONOMY
Abaxial (abaxially) side of an organ that faces away from the axis bearing it
Acaulescent without stem or stemless
Adaxial (Adaxially) side of an organ that faces the axis bearing it
Axil (Axillary) upper angle of the junction between thepetiole and stem
Bifid divided into two equal parts
Bipinnate doubly pinnate
Bract a modified enclosing leaf
Bracteole a bract on the flower stalk
Caducous Deciduous
Campanulate bell-shaped
Carinate Keeled
Chartaceous Papery
Ciliate fringed with hairs
Cincinnus arranged with each flower arising inthe axil of a bracteole on the stalk of the previous flower
Cirrus (Cirrate) a whip-like extension of the leaf rachisarmed with reflexed spines used as a climbing organ
Clustered (Clustering) basally composed of two to manystems
Concolorous of the same colour
Conic (Conical) cone-like
Connate Joined
Coriaceous Leathery
Costapalmate describing a palmate leaf with a strong ribextending through
94
the lamina
Crownshaft cylinder formed by the leafbases
Crustaceous dry and brittle
Cuneate wedge-shaped andwidest distally
Cupule (Cupular) cup-like
Deflexed curved downwards
Dichotomous fork-like
Dimorphic occurring in two different forms
Dioecious (Dioecy) plants with either staminate or pistillate flowers
Discolorous of different colour
Distal farthest from the point of attachment
Distichous arranged in two rows
Endocarp the innermost layer of the fruit wall
Epipetalous united with or borne on petals
Fimbriate fringed with fine hairs or thread-like segments
Flabellate fan-shaped
Flagellum a whip-like climbing organ armed with reflexed spines adnate to or emergent from the leafsheath
Flexuous wavy, bent alternately in opposite directions
Gibbous pouched, more convex in one place than another
Globose Spherical
Hapaxanthic the process of an individual plant (or stem ofa cluster) flowering once then dying
Hastula a flap of tissue borne at the insertion of thelamina on the petiole of a palmate leaf
95
Hermaphrodite bisexual(hermaphroditic)
Imbricate Overlapping
Imparipinnate a pinnate leaf with a terminal pinna
Indumentum a covering of scales or hairs
Induplicate V-shaped in cross-section
Inflexed bent inwards
Inflorescence collective term for the aggregation offlowers
Infrafoliar borne below the leaves
Interfoliar borne among the leaves
LeafBase portion of leaf proximal to the petiole, clasping the stem or forming a closed tubular crownshaft
LeafScar the remnants or impression of the formerattachment of a leaf on a stem
Medifixed attached at the middle
Membranous thin and translucent
Monoecious plant bearing both staminate and pistillateflowers
Obovoid (Ovoid) egg-shaped
Palmate with segments radiating from a central pointresembling a fan in a leaf
Panicle (Paniculate) Inflorescence with iterative branchingpatterns (racemes) from a central axis
Paripinnate a pinnate leaf without a terminal pinna
Peduncle the proximal unbranched portion of aninflorescence
Peduncular Bract(S) a single bract or multiple bracts onthe main axis of the inflorescence
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Pendulous hanging pendant
Perianth the sepals and petals together
Pinna(e) segment(s) of a pinnate leaf
Pinnate pinnae arranged in a feather-like arrangement ona leaf, i.e. with similar number ofpinnae on each side ofthe rachis
Pinnule the pinnae of a bipinnate leaf
Pistillate bearing a pistil
Pistillode a sterile gynoecium
Pleonanthic the process of an individual plant floweringrepeatedly
Polygamodioecious having bisexual and male flowers onsome plants and bisexual and female flowers on others
Polygamomonoecious having bisexual and unisexualflowers on the same plant
Polygamous having bisexual and unisexual flowers onthe same or different plants
Prophyll the first bract borne on the inflorescence
Pseudomonomerous a gynoecium with one fertile carpeland one locule but with parts of two other carpelspresent
Rachilla(e) the flower-bearing branch of an inflorescence
Rachis the axis beyond the petiole or beyond the peduncleof the leaf or of an inflorescence
Reduplicate inverse V-shaped in cross-section
Reflexed folded or turned back
Sagittate arrow-shaped
Segment a single division of a palmate or costapalmate leaf
Solitary with a single stem,
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Spicate an unbranched inflorescence
Staminate a flower that bears stamens
Staminode vestigial stamens, often reduced to tooth-likeflaps of tissue
Triad in threes, flowers grouped with a central pistillateand two lateral staminate flowers
Trilocular with three chambers, each bearing an ovary orseed in the gynoecium
Triovulate bearing three ovules in the gynoecium, one inthe locule of each carpel
Unisexual flower having either stamens or pistils but not both
Valvate margins of segments meeting neatly without overlapping
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Chapter 2
PALYNOLOGY OF THE FAMILY PALMAE
INTRODUCTION
Pollen analysis is a scientific method that can reveal evidence of past ecological and climate change. Pollen plays a critical role in the studies of climate change during the
Quaternary period (Faegri and Iverson, 1975) and is the only branch of Paleoecology for the late Pleistocene and Holocene. Furthermore, the morphological characters of pollen can be useful in studies of plant taxonomy because many pollen characters influenced by strong selective forces involved in various reproductive process such as pollination, dispersal and germination (Erdtman, 1952; Nowicker and Skvarla, 1979; Stussey 1990;
Moore et al., 1991). Thus pollen morphology plays a key role in the plant systematic and is supportive in the delimitation of closely related taxa.Pollen morphology is not only confined to the plant systematic butthe scientific scope of palynological studies is broad since it encompasses knowledge from many disciplines such as botany
(includinghorticulture and agriculture), geology, ecology, climatology and archaeology, medical studies (aeropalynology), and criminology.
Comparative pollen morphology has been studied for about 150 years beginning with workers such as Mohl (1835) and Hassel (1842).Pollen and anther characters have frequently been regarded as systematically significant in monocotyledons. In relation to plant systematic Wodehouse (1935) has made magnificent contribution in the field of
Palynology. Later Erdtman (1952) gives a detailed account on the pollen of 75 species belonging to 55 genera representing the family Palmae. She reported eurypalynous nature
99 of Palmae and described pollen as monosulcate, disulcate or trichotomosulcate; andalso mentioned thesize of pollen ranging from 20 µ to 80 µ.Zavada (1983) presented a general review of apertures and wall structures in monocot pollen.Linder and Ferguson (1985) discussed the significance of aperture types and exine stratification in Poaceae,
Restionaceae and allied families.Sultan et al. (1994) studied the pollen morphology of 25 monocotyledonous species belonging to 14 genera from Karachi (Pakistan) by using the techniques of LM and SEM. Harley and Zvada (2000) studied the pollen morphology of the monocotyledonous plants and also carried out their cladistic analysis.
The first published description of triporate pollenin Arecaceae is that by Erdtman andSingh (1957) concerning pollen of Sclerosperma. During the last years the pollen morphology of Palmae has received considerable attention by various workers such as
Punt & Wessels Boer (1966a, 1966b), Thanikaimoni (1970a, 1970b), Sowunmi
(1968,1972), Kedves (1980), Ferguson (1986) and Ferguson and Harley (1993).
However, all these covered the pollen morphology of the entire family. Thanikaimoni
(1970a) create his own classification based on pollen morphology and also discussed the geographical distribution. Sowunmi (1972) adopted the classification of Thanikaimoni and modified the classification on the basis of palynological evidences.
The detailed pollen morphological studies of the Palmae have been carried out by
Ferguson (1981) in association with Dransfield and Uhl.Afterward a number of studies have been made by various workers from time to time. David (1927) studied the pollen of
Phoenix dactyliferaL. with reference to its longevity and effect on the fruit. Mahabalé
(1967) adopted and gives a complete account on the pollen morphology of the family
Palmae and also discuss their taxonomic value by follow the classification of Drude
100
(1889) and Moore (1960). He discussed that the number and nature of the colpi are more reliable for the classification and phylogeny as compared to the shape, size and the sporic nature of the pollen.Kedvis (1980) investigated the pollen morphologic characters within the family Palmae. Sharma (1967) investigated the pollen of Calamus rotangand reported
1-2 colpate, sub- spheroidial pollen.Sowunmi (1968) examined the pollen grain of 350 species and discussed diversity of pollen with regard to size, shape, aperture and exine pattern within the family. He found five possible trends in aperture (trichotomocolpate, monoporate, diporate, annulocolpate and dicolpate) and a great variety of exine pattern
(punctuate, negatively reticulate, vermiculate,verrucose, pilate, clavate, spinose, verrucose-reticulate and punctuate-clavate).Tisserat & DE Mason (1982) described the pollen of the genus Phoenix L.by using SEM.Ferguson et al. (1983) described the pollen morphology of the 2 representative of the genus Pinanga viz., P. aristataand P. pilosa by using LM and SEM and found remarkable pollen exine ornamentation i.e., urceolate type that have not been reported previously in the Angiosperms. Straka and Friedrich (1984) investigated the pollen of the three Madagascar Palm genera viz., Medemia, Hyphaene and Ravenea with the help of SEM.Ferguson (1986) studied the pollen morphology of the family Palmae based on SEM and TEM and found there is remarkable variation and parallelism in the pollen of Palmae.Ferguson (1987) examined the pollen of the tribe
Borasseae (Palmae:Coryphoideae) of 7 genera and 14 speciesby using LM, SEM and
TEM. He used pollen size, exine thickness, ornamentation and its stratification for the delimitation of taxa.Mendis et al., (1987) investigated the pollen of 11 species belonging to
8 genera of the subtribe Oncospermatinae (Palmae: Arecoideae:Arecaceae). They describedmonosulcatepollen and also found wide range of variation in the exine thickness,
101 ornamentation and its stratification.Harley (1990) studied the pollen grain of the family
Palmae and discussed that the majority of species have simple tectate or semi-tectate, columellate and monosulcate or trichotomosulcate pollen.Grayum (1991) reviewed anther and tapetum types in the systematic of Arecaceae.Ferguson and Harley (1993) reviewed the pollen morphology of the family Palmae primarly based on electron microscopy.
Ambwani and Kumar (1993) studied the pollen morphology of 9 species coryphoid genus
Licula (Palmae) by using LM and SEM. They recognized pollen morphological variation of different species on the basis of 4 exine pattern. Baker et al. (2000) discussed that apart from monosulcate, trichotomosulcate and equatorial disculcate are also quite common. They found monosulcate pollen is present in all subfamilies except Nypoideae.
On the other hand, trichomosulcate occurs in Coryphoideae, Ceroxyloideae and
Arecoideae whereas disulcate is only confined to the Calameae.Harley and Baker (2001) studied the morphology of pollen aperture in Areceae. They described almost all the species have monosulcate pollen, 17 aperture types and 13 exine types and also discussed the interpretation and use of aperture characters in Cladistic analysis of the
Arecaceae.Harley and Dransfield (2003) studied the triporate pollen in the
Arecaceae.Kumar and Ramaswamy (2003) worked on the development of microsporogenesis of 5 species of Calamus L. (Arecaceae). Sannier et al. (2006) studied variation in the microsporogenesis of monosulcate palm pollen. They observed the following characteristics such as aperture pattern, ontogeny in pollen and described microsporogenesis in 8 species (4 species belong to subfamily Arecoideae and 4 from
Coryphoideae) producing distal monosulcate pollen.
102
Though numbers of palynological studies of the Palmae were conducted as evident from the literature but there are no detailed information on the pollen of several taxa belonging to thePalmsoccurring in Pakistan and Kashmir.
Therefore, the main purpose of the present work is to complete the detail information of the pollen morphological characterand then to correlate the palynological data with the taxonomy of the family Palmae.
103
MATERIALS AND METHODS
The present work is based on 27 species belonging to 21 genera of the family Palmae.
For the study of pollen, the mature, healthyun-dehisced and fresh flower were collected from the field and have been fixed in the 70% alcohol contained in vials, whereas in few cases herbarium specimens were also taken from Centre for plant conservation Karachi
University Herbarium (KUH).The palynological study was based on 10-15 pollen grains per specimens. Details of voucher specimens are given in Appendix 1.Fertile anthers were processed according to the standard acetolysis method outlined by Erdtman (1952,
1960). Anthers were drenched for 12 to 18 hours in glacial acetic acid. After that the soaked material was crushed, then centrifuged at 3000 – 4000 r.p.m and acid was decanted. Add 5 c.c. acetolysed mixtures (Sulphuric acid and acetic anhydride in the ratio of 1:9). The acetolysed mixtures were placed in the water bath up to the boiling point then leave it for cool and re-centrifuged. The material was washed with distilled water and then divided in to two parts. One part was used for the examination of Light
Microscopy and the second part for the Scanning Electron Microscope.
Light Microscopy (LM)
For Light Microscopy pollen were mounted in glycerine jelly stained with 1% safranine.
To melt the glycerine jelly, slide was placed on the hot plate followed by mounting with a cover slip. When slide cooled, sealedby running molten paraffin wax and labeled withrelevant informationwas pasted on the slide. The following pollen characters such as polar length (P), equatorial diameter (E), colpus length, exine thickness, aperture number and aperture type has been examined. The pollen data were analysed statiscally i.e. calculated range, mean and standard error (±) by using MS Excel (Table.2.1-2.3).The
104 measurement is given in the Table-2.1 to 2.3 are based on at least 10-15 grains from each specimens.The photographs of studied pollen were also taken with the Nikon FX-35
Phase Contrast Microscope.
Scanning Electron Microscope (SEM)
For SEM, the sample was suspended in a drop of water and directly mounted on metallic stub using double sided adhesive tape. The stub was left for few hours to evaporate the water. Thenthe samples were coated with gold using Jeol JFC 1100 E ion sputtering device. SEM observation and photographs were carried out on a Jeol Microscope
JSM6380LV.
Terminology
The terminology of the examined data was made accordance to Erdtman (1943, 1952&
1969), Faegri & Iverson (1953), Walker & Doyle (1975) and Punt et al., (1994).
105
GENERAL POLLEN MORPHOLOGY OF PALMS
The present study reveals that the Palmae is a eurypalynous family which shows the pollen diversity within the family such as size, shape, and aperture and exine pattern.
The predominant aperture type is colpus i.e, majority of pollen are monosulcate, occasionally disulcatepollen (in Calamus tenuis L.) or trichotomosulcate pollen
(Elaeisguineensis Jacq.) is also examined. Theaverage size range of palm pollen is 13.0
– 36 µm in lengthand 20 – 78 µm in breadth (such as in Phoenix, the pollen is smalli.e.
13-15 µm and quite large in SabalAdanson and CocusL., i.e., 36- 78 µm).The shape of the grain, as seen in polar view, is frequently elliptical either symmetrical or asymmetrical but sometimes subcirculr or rounded to triangular pollen have also been found. On the other hand, in equatorial view the pollen is kidney- shaped in transverse position and more or less boat shaped in longitudinal position.Aperture more or less equal or somewhat shorter or longer than the longest axis of pollen.The exine of most of the examined species (about 80%) is semi-tectate.Infratectum collumelate. Muri simpli to dupli or tripli to multibaculate.There is a great variety in exine pattern i.e., reticulate, punctate, vermiculate, regulate and in some cases verrucose and spiny exine but the most common pattern found is reticulate type. In most of the species, ornamentation is mottled, perforate or granulate, finely reticulate-granulate. The exine thickness ranges from 1.0 –
3.0 µm.
Important pollen characters of all the studied taxa have been arranged according to the subfamilies (Table 2.1-2.3).
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1 SUB FAMILY ARECOIDEAEGriff.
Introduction
The Arecoideae is the largest and most diverse of the fivesubfamilies recognized in the family Palmae consists of 14 tribes and 14 sub-tribes, from which Areceae is the largest tribe (Dransfield et al., 2008; Baker et al, 2011). About 60% of palm genera (113 out of 186) and 50% of species (c.1300 out of c.2400) representing the subfamily
Arecoideae, extend equally in the America and Southeast Asia to Western Pacific regions. The members of arecoid palms are widespread in the tropics and sub-tropics regions(mainly in rain forest) and, to a lesser extent, inseasonally dry habitats. The monophyly of subfamily Arecoideae as circumscribed in the current classification
(Dransfield et al., 2005, 2008) is strongly supported by a substantial body of phylogenetic evidence (Uhl and Dransfield, 1987; Asmussen et al., 2000, 2006; Asmussen and Chase,
2001; Lewis and Doyle, 2002; Baker et al., 2009).
Morphological characters that define the arecoid clade and distinguish it from other subfamilies include the presence of reduplicate, pinnate and pinnately veined simple leaves, highly differentiated primary inflorescence bracts and flower mostly in triads
(floral/ flower triad is a cluster of three unisexual flowers, comprisinga central female flower flanked by two male flowers). Outside arecoids, triads are only found in tribeCaryoteae (Coryphoideae), which is the main reason for the erroneous placement of
Caryotae within Arecoideae in the earlier classification of Uhl and Dransfield (1987).
The differentiated primary inflorescence bracts of Arecoideae contrast with those of some other subfamilies, which are conspicuous and relatively uniform throughout the main axis of the inflorescence. In arecoids, however, the primary bracts subtending the first-order
107 branches (rachis bracts) are always highly reduced, and well-developed bracts occur only on the peduncle. This feature is shared with subfamily Ceroxyloideae, the sister of
Arecoideae (Asmussen and Chase, 2001; Asmussen et al., 2006; Baker et al., 2009).
Phylogenetically various workers contributions on sub-clades of Arecoideae (Gunn,
2004; Roncal et al., 2005; Loo et al., 2006; Norup et al., 2006; Cuenca and Asmussen-
Lange, 2007; Cuenca et al., 2008, 2009), provided sufficient evidence for the circumscription of monophyletic tribes and sub-tribes in the current classification
(Dransfield et al., 2005, 2008). The subfamily Arecoideae contains various species that are the sources of economically important palm such as oil palm (Elaeis guineensis), coconut (Cocos nucifera), betel nut palm (Areca catechu), peachpalm (Bactris gasipaes) and important species in the global horticultural trade like Dypsis lutescensand Roystonea regia.
The pollen morphology of the subfamily has been studied by Erdtman 1952;
Thanikaimoni 1971;Sowunmi 1972; Kedves 1980 and Ferguson & Harley 1993.
However, no detailed information is available from the area under consideration. In the present work 8 species belonging to 7 genera distributed in 3 tribes (viz., Areceae,
Cocoseae and Roystoneae) have been studied by using Light Microscope (LM) and
Scanning Electron Microscope (SEM).
1. Tribe Areceae
The Areceae is monophyletic in most phylogenetic studies (Asmussen et al. 2006;
Dransfield et al. 2008), but its relationships to other tribes is unresolved. Out of 14 tribes of the subfamily, the Areceae is the largest tribe with c.41 genera (including 10 unplaced genera)distributed in 11 sub-tribes (see Dransfield et al., 2008; Baker et al, 2011). Of
108 these 3 sub-tribes have representative in Pakistan, Arecinae and Dypsidinae with a monotypic genus Areca catechuL. and Dypsis lutescence(H.Wendl.) Dransf. & Beentje., respectively and the subtribe Ptychospermatinae with 2 genera viz., Ptychosperma Labill and WodeytiaIrvine. Comprehensive pollen morphology of the tribe Areceae (subfamily
Arecoideae) including genera has been studied by various workers (Thanikaimoni
1971;Sowunmi, 1968 and 1972; Kedves 1980 and Ferguson & Harley 1993).
Observations and Results
The summary of pollen morphological data of the studied taxa belonging to the sub- family Arecoideae is given in the Table 2.1 where as LM and SEM photographs are given in Figure 2.1-2.3
Pollen description of the tribe Areceae
Pollen monosulcate.Amb (Polar view) elliptical or asymmetric or circular.Size: Pollen
15- 32 µm in length and 30- 45 µm in breadth. Colpus length is 14-32 µm.Aperture equal or shorter than or as long as the length of the longer axis of pollen.Exine (Sexine) tectate or semitectate.Tectum reticulate, perforate or finely perforate-rugulate, without process or spinulose-reticulate, spinules borne on the tectum.Infratectum columellate.Murisimpli- through to multi baculate.
Represented by 4 genera viz.,Areca L., Dypsis Noronha ex Mart., Ptychosperma Labill. and Wodeytia Irvine
Key to the Genera
1+ Tectum perforate or finely perforate-rugulate.Aperture equal in length to the longer
axis of pollen……...…….…Ptychosperma group (3. Ptychosperma &4.Wodeytia)
109
- Tectum not as above. Aperture equal or shorter than or as long as the length of the
longer axis of pollen………………………………………………………..………..2
2+ Tectum finely reticulate. Muri simple baculate to dupli-baculate.…..……....1. Areca
- Tectum spinulose. Muri not as above…………………………..………..….2. Dypsis
1.Areca L.
Pollen monosulcate, elliptic in polar view; boat-like in equatorial view, Exine finely reticulate.
1. Areca catechuL. (Fig. 2.1 A & B; 2.3 A)
Pollen (21.20-) 23.85(-26.50) µm in length and (37.50-) 41.35 (-45.20) µm in breadth, asymmetric.Colpus length is (23.50-) 24.00(-26.50) µm, margin wavy. The aperture is equal or shorter than the longest axis of pollen.Exinetectate or semitectate, 1.3 - 2 µm thick. Tectum 0.8 µm thick, finely reticulate. The muri is 0.6 µm wide, simple baculate.
2. DypsisNoronha ex Mart.
Pollen monosulcate, elliptic to pyriform in polar view; boat-like in equatorial view.Tectum spinulose.
1. Dypsis lutescence(H.Wendl.) Dransf. & Beentje (Fig. 2.1 C & D; 2.3 B)
Pollen (15.50-) 21.0 (-26.50) µm in length and (30.80) -36.70 (-42.60) µm in breadth, usually asymmetric.Colpus length is (15.50-) 20.75 (-26.00) µm. The aperture is shorter than, or as long as, the longest axis of pollen. Exine is usually tectate occasionally semitectate, 1.5-2.5 µm thick.Tectumspinulose or perforate. Muri at reticulum coarsely granulate, spinulose.
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Figure2.1. Scanning Electron Micrographs (SEM) of the pollengrains: Areca catechu: A, pollen grain; B, exine pattern. Dypsis lutescens: C, pollen grain;D, exine pattern. Ptychosperma elegans: E, pollen grain; F, exine pattern. Ptychosperma macarthuri:G, pollen grain; H, exine pattern (scale bar: B, F, H = 1μm; D, G = 2; A= 5μm; C = 10 μm).
111
112
3. Ptychosperma Labill.
Pollen monocolpate, ellipsoidal in polar view occasionally pyriform; boat-like inequatorial view.Tectum reticulate or finely perforate-rugulate.
1. Ptychosperma elegans(R.Br.) Blume (Fig. 2.1 E & F; 2.9 G)
Pollen (22.40-) 24.95 (-27.50) µm in length and (31.60-) 36.40 (-41.20) µm in breadth, asymmetric Colpus length is (21.20-) 24.10 (-27.00) µm. The aperture is about equal in length to the longer axis of pollen.Aperture margin similar or slightly finer.Exine tectate,
1.2-2 µm thick.
2. Ptychosperma macarthuri (H.Wendl.) H. Wendl. ex Hook. f. (Fig. 2.1 G & H; 2.9 H)
Pollen (25.20-) 28.35 (-31.50) µm in length and (39.40-) 41.47 (-43.55) µm in breadth, asymmetric Colpus length is (25.00-) 27.85 (-30.70) µm. The aperture is about equal in length to the longer axis of pollen.Aperture margin similar or slightly finer.Exine tectate,
1.2-2 µm thick.
4. WodeytiaA. K. Irvine
Pollen monosulcate, ellipticin polar view; boat-like in equatorial view.Tectum reticulate or finely perforate-rugulate.
1. Wodeytia bifurcate Irvine(Fig. 2.3 D; 2.8 G & H)
Pollen (24.60-) 28.50 (-32.40) µm in length and (31.50-) 35.85 (-42.20) µm in breadth, slightly asymmetricColpus length is (24.50-) 28.25 (-32.00) µm. The aperture is about equal in length to the longer axis of pollen.Exine tectate, 1.5-2 µm thick.
2. Tribe Cocoseae
The Cocoseae Mart. is one of the 14 tribes of subfamily Arecoideae, with three subtribes and
18 genera (Dransfield et al., 2005, 2008; Dowe, 2010; Baker et al., 2011). Among these, two
113 genera viz., Cocus L. and Elaeis Jacq., of the subtribe Attaleinae and Elaeidinae respectively have been recognized from Pakistan and Kashmir.The members of this tribe are widely distributed in the Neotropics, Madagascar, Africa and S. America.
A striking synapmorphy of the tribe Cocoseae is the presence of three or more “eyes” or the germinating pores on the hardened seed endosperm. Other diagnostic characters included a 1- branched (rarely spicate) inflorescence, an inconspicuous prophyll, a prominent or often woody peduncular bracts that spilts abaxially with respect to the main axis, imbricate petals of the pistillate flowers and triovulate gymnosperms (Uhl & Dransfield, 1987).
Cocoseae Mart. is monophyletic, and is variously sister to the Reinhaditeae, Roystoneae and the Oranieae in the available molecular evidences (Dransfield et al. 2008).
Few literatures are available on the pollen morphology of the tribe Cocoseae Mart. and their representatives (Thanikaimoni, 1967; Sowunmi, 1968, 1972; Kedves 1980). The present research has been carried out to study the pollen of 2 monotypic genera viz., Cocus nucifera L. and Elaeis guineensis Jacq. by using LM and SEM and also discuss its systemic significance.
Pollen description of the tribe Cocoseae
Pollen elliptical to more less circular or oblate- triangular in apertural view, usually asymmetric.Size: Pollen (23.20-) 26.80 (-30.40) µm in length and (44.20-) 54.60
(-65.00) µm in breadth.Aperture monosulcate or trichotomosulcate.Colpus 18 - 65 µm in length, margin wavy or broad and psilate.Exine tectate or semi tectate, up to 2.0 µm thick. Tectum 0.6 µm thick, punctuate or finely perforate or reticulate to vermiculate.Infratectumcolumellate with wide or very narrow, irregular spaces between the columella (Table 2.1).
Represented by 2 genera viz., Cocus L. and Elaeis Jacq.
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Key to the Genera 1+ Pollen monocolpate, up to 65 µm long…………………..……...…………...1. Cocus
- Pollen trichotomocolpate, up to 37 µm long………………………….….… 2. Elaeis
5.Cocus L.
Pollen monosulcate,elliptical to more or less circular in polar view. Tectum punctate or
perforate.
1. Cocus nucifera L. (Fig. 2.2 C & D ; 2.3 F)
Pollen (24.60-) 28.50 (-32.40) µm in length and (44.20-) 52.15(-60.10) µm in breadth,
frequently elongate, usually asymmetric.Colpus (23.20-) 26.50 (-29.80) µm in length,
margin wavy.Exine tectate or semi tectate, 1.5 µm thick, undulating, segments with
exinous infoldings tubercles. Tectum 0.6 µm thick, perforation rugulate supported by a
finely but densely baculate layer.
6.Elaeis Jacq.
Pollen trichotomosulcate,triangular in polar view.Tectum reticulate to vermiculate.
2. Elaeis guineensis Jacq. (Fig. 2.2 E & F ; 2.3 G)
Pollen (31.60-) 34.30 (-37.00) µmin length and (16.64-) 21.16 (-25.68) µm in breadth,
radio-symmetrical with concave sides.Colpus (31.60-) 34.30 (-37.00) µm in length
Aperture margin wavy, broad and psilate, not always reaching the periphery of the
pollen.Exine tectate or semitectate, 1.1 µm thick.Tectum 0.6 µm thick.Bacula
present.
3. Tribe Roystoneae Drans., et al.
The Roystoneae Drans., et al.is one of the 14 tribes of subfamily Arecoideae, with a
single genus RoystoneaO. F. Cook. ((Dransfield et al., 2005, 2008).The placement
115 of RoystoneaO. F. Cook., within the Arecoideae is uncertain. Thephylogenetic studies failed to determine the taxonomic position of the genus within the Arecoideae. As of
2008, there appear to be no molecular phylogenetic studies of Roystonea O. F. Cook.
The Royal Palms, spectacular majestic solitary pinnate-leaved palms from the
Caribbean islands and neighbouring parts of North, Central and South America.The genus is distinct due to the presence conspicuous crownshaft; inflorescence branched to at least 4 orders with rather stiff spreading rachillae.A long, green, caducous, peduncular bract protects inflorescence, and when it splits and abscisesfrom the peduncle, it falls off together with a multitude of “free trichomes” of farinaceous aspect,a unique feature in the world of palms (Zona 1996). Like the other members of the palm the genus also have economic value such as the seeds are used as a source of oil and forlivestock feed; leaves are used for thatching; wood for construction; roots are diuretic in nature and for the treatment of diabetes.Of the ten existing Roystonea species, only one is known from
Pakistan viz., Roystonearegia (H.B. & K.) O. F. Cook.
Thanikaimoni 1971 and Sowunmi 1972 describe the pollen of the subtribe
Roystoneinae (tribe Roystoneae) as monosulcate, elliptic or triangular in apertural view, tectm is tectate-perfrorate or semitectate and foveolate to reticulate. However, no detailed information is available from the area under consideration.In the present work pollen of
Roystonea O. F. Cook has been studied by using Light Microscope (LM) and Scanning
Electron Microscope (SEM).
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Figure 2.2. Scanning Electron Micrographs (SEM) of the pollengrains:Roystonea regia: A, pollen grain; B, exine pattern. Cocus nucifera: C, pollen grain;D, exine pattern. Elaeis guineensis:E,pollen grain;
F, exine pattern Caryota urens: G, pollen grain; H, exine pattern (scalebar: B,
D, F, H = 1μm; C = 2 μm; A, G = 5μm;E = 10μm).
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118
Pollen description of the tribe Roystoneae
1. RoystoneaO. F. Cook
Pollen monosulcate, ellipticin polar view; boat-like in equatorial view.Tectum punctate to vermiculate.
1. Roystonearegia(H.B. & K.) O.F. Cook (Fig. 2.2 A & B; 2.3 F)
Pollen (40.30-) 46.25 (-52.20) µm in length and (39.15.-) 44.20 (-49.25) µm in breadth, with slight or obvious asymmetric.Colpus length is (39.50-) 45.45 (-51.40) µm. The aperture is about equal in length to the longer axis of pollen.Exine tectate, 1.5µm thick.
Discussion of the subfamily Arecoideae
This is the largest subfamily within the Palmae and has awide range of pollen. It is very heterogenous morphologically (Moore,1961) and palynologically (Erdtman, 1952;
Sowunmi, 1972), but uniform anatomically (Tomlinsin, 1961). The Arecoideae was found to be heterogenous with regard to aperture forms such as monocolpate, monoporate, annuporate, trichotomocolpate, and triporate pollen (Sowunmi, 1972). In the present work two types of apertures are found viz., monosulcate and trichotomosulcate.
Out of 8 species, monosulcate pollen is present in 7 speciesrepresenting three tribes viz.,
Arecaceae (Areca catechu L., Dypsis lutescenceNoronha ex Mart.,Ptychosperma Labill and Wodeytia Irvine); Cocoseae (Cocus nuciferaL.) and Roystoneae (Roystonearegia
(H.B. & K.) O. F. Cook). The remaining one species i.e., Elaeis guineensisJacq., havetrichotomocolpate pollen. Similarly, there is a variety in an exine pattern such as pollen finely reticulate;spinulose or perforate;perforate or finely perforate- rugulatepunctate or finely perforate, minutely sub-reticulate to vermiculate.
Palynologically, the subfamily is found to be eurypalynous and our findings are in
119 accordance withErdtman, 1952; Thanikaimoni 1971; Sowunmi, 1968, 1972; Kedves,
1980; Harley, 1990; Faegri & Harley, 1993 and Dransfield et al., 2008. On the basis of the aperture type and exine pattern six pollen types are recognized in this sub family.
These are Elaeis type (Elaeis guineensisJacq.), Cocustype(Cocus nuciferaL.), Roystonea type (Roystonearegia (H.B. & K.) O. F. Cook), Dypsis type (Dypsis lutescenceNoronha ex Mart.), Areca type (Areca catechu L.) and Ptychosperma type (Ptychosperma Labill and Wodeytia Irvine). Present results are in accordance with Sowunmi (1972) and Faegri and Harley (1993), also represented these genera under the different groups.
Key to the Pollen Types
1+ Pollen trichotomosulcate…………………………………………...…...…Elaeis type
- Pollen monosulcate…………………...………………….…….…….……..………..2
2+ Pollen more than 45 µm in length.Tectum punctate to vermiculate……..…...... 3
- Pollen up to 45 µm in length.Tectum not as above…….………...…….……………4
3+ Pollen up to 60 µm in length.Tectum punctate …………………….....….Cocus type
- Pollen up to 50 µm in length. Tectum punctate to vermiculate…..….Roystonea type
4+ Pollen spinulose, spinules borne on the tectum…………………….….…Dypsis type
- Pollen not as above…..…………………………………………………..…...……….…….5
5+ Tectum finely reticulate…………………………………………..….……Areca type
- Tectum perforate or finely perforate-rugulate. ………………...... Ptychosperma type
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Pollen Morphology and Taxonomy
Tribe Areceae
In this tribe pollen morphology of four genera belonging to 3 subtribes viz.,
Arecinae (Areca L.), Dypsidinae (Dypsis Noronnah ex Mart.) and Ptychospermatinae
(Ptychosperma Labill and Wodyetia Irvine) have been studied. The exine pattern of the pollen classified these genera into 3 pollen types. The aperture type is same in all the studied genera i.e.,pollen monosulcate and elliptical in polar view. Ptychosperma Labill and Wodyetia Irvine, distinctive by having perforated or finely perforate-rugulate tectum but cannot be distinguished from each other by their pollen morphology. Ferguson &
Harley (1993) described the pollen of Wodyetia is tectate to perforate or finely perforate to rugulate. Similarly, in the genus ArecaL., tectum is finely reticulate which also been observed byThanikamoni (1971), Sowunmi (1968, 1972), Kedves (1980), (Dransfield &
Uhl, 1998), Dransfield et al., (2005,2008). Pollen morphology is significantly helpful at generic and subtribal level. The genus DypsisNoronnah ex Mart is characterized by spinulose pollen, spinules present on the tectum.
Tribe Cocoseae:
The tribe is represented by the following two genera such as Cocus L. (Cocus nuciferaL.) and Elaeis Jacq. (Elaeis guineensis Jacq.). Palynologically, both the genera can be easily distinguished on the basis of aperture and exine pattern. The former genus is distinguished by monocolpate pollen with a punctate tectum whereas, the later genus havetrichotomosulcate pollen with reticulate-vermiculate tectum.Various workers such as
Mahabalé (1967), Thanikamoni (1971), Sowunmi (1968, 1972), Kedves (1980) and
Harley (1990) have placed these two genera in to two different groups.
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It is also known that between these genera there is an extraordinary distinction in morphological characters. The genus Elaeis Jacq.,is distinguished by the presence of unarmed petiole; bisexual inflorescence; beaked fruit and endocarp with apical pore. On the other hand, Cocus L., is charcterised by having armed petiole, unisexual inflorescence, unbeaked fruit and endocarp with basal pore. Thus these genera fall into two subtribes viz., Attaleinae (Cocus L.) and Elaeidinae (Elaeis Jacq.) that show the genera appear to be closely related butdefinitelydistinct.
Thus foregoing discussion clear that morphological variation in the pollen can also be considered as a feature of taxonomic value.
TribeRoystoneae
This tribe is represented by a single species viz., Roystonearegia (H.B. & K.) O.
F. Cook. This specie is charctarized by punctate to vermiculate tectum.
Thanikaimoni(1970 a) and Sowunmi(1972) also described the pollen as monosulcate, elliptical in polar view, tectum punctate to vermiculate to negatively reticulate.
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Figure 2.3. Light Micrographs (LM) of the pollen grains at 100 x:A, Areca catechu: pollen grain. B,Dypsis lutescens:pollen grain. C,
Trachycarpusfortune:pollen grain.D,Wodyetia bifurcata: pollen grain.
E,Roystonea regia:pollen grain. F, Cocusnucifera: pollen grain. G,Elaeis guineensis: pollen grain. H, Caryota urens: pollen grain. I, Calamus tenuis:pollen grain.(Scale Bar = 10μm)
123
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Table 2.1 General Pollen Characters of the Sub Family Arecoideae
Exine Length Breadth Colpus length Name of taxa Tribe Aperture thickness Tectum (µm) (µm) (µm) (µm)
Areca catechu Areceae 21.20 (23.85) 26.50 37.50 (41.35) 45.20 20.50 (23.50) 26.50 Monosulcate 1.3 - 2 finely reticulate ± 0.595 ±0.876 ± 0.639
Dypsis lutescence Areceae 15.50 (21.0) 26.50 30.80 (36.70) 42.60 15.50 (20.75) 26.00 Monosulcate 1.5 - 2.5 spinulose or ± 1.220 ±1.25 ± 1.170 perforate
Ptychosperma elegans Areceae 23.20 (25.85) 28.50 35.60 (39.90) 44.20 21.00 (23.70) 26.40 Monosulcate 1.2 - 2 reticulate or ± 0.593 ±0.830 ± 0.650 finely perforate- rugulate P. macarthuri Areceae 25.20 (28.35) 31.50 39.40 (41.47) 43.55 25.00 (27.85) 30.70 Monosulcate 1.2 - 2 reticulate or ± 0.721 ±0.521 ± 0.592 finely perforate- rugulate Wodyetiabifurcata Areceae 24.60(28.50) 32.40 31.50 (35.85) 40.20 24.50 (28.25) 32.00 Monosulcate 1.5 - 2 reticulate or ± 0.784 ±1.083 ± 0.967 finely perforate- rugulate Cocus nucifera Cocoseae 23.20 (26.80) 30.40 44.20 (52.15) 60.10 23.20(26.50) 29.80 Monosulcate 1.5 punctate ± 0.842 ±1.734 ± 0.722
Elaeis guineensis Cocoseae 31.60 (34.30) 37.0 16.64 (21.16) 25.68 25.20 (29.90) 34.60 Trichotomosulcate 1.1 reticulate - ± 0.632 ±0.907 ± 0.834 vermiculate
Roystonea regia Roystoneae 40.30 (46.25) 52.20 39.15 (44.20) 49.25 39.50 (45.45) 51.40 Monosulcate 1.5 punctate to ± 1.284 ±1.002 ± 1.269 vermiculate
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2 SUB FAMILY CALAMOIDEAEGriff.
Introduction
The subfamily Calamoideae is monophyletic and resolves the most phylogenetic studies as sister to all other palms (Uhl et al., 1995; Baker et al., 1999, 2000 a, 2000 b
&2000 c; Asmussen et al., 2000; Asmussen et al., & Chase, 2001; Lewis & Doyle, 2001;
Hahn 2002 a; Asmussen et al., 2006; Dransfield et al., 2008). A subfamily of 21 genera and c. 620 species falls in to three tribes viz., Calameae, Lepidocaryeae and Eugissoneae
(Fig. 6.1),distributed globally whereas highest diversity occurs in westMalesia.
The subfamily Calamoideae includes the largest and wide spread palm genus,
CalamusL., commonly known as Rattan or Cane palm, represented by c. 400 species. The species is harvested for the canes which are used for making furniture, cordage, basketry and various handicrafts. In Pakistan, only a single genus with one species is present i.e.,
Calamus tenuis Roxb., belonging to the tribe Calameae and subtribe Calaminae.
1. Tribe Calameae
On the basis of molecular evidences, the tribe Calameae is also resolved asmonophyletic (Baker et al. 2000 a; 2000 b; Asmussen et al., 2006; Dransfield et al.,
2008). Morphologically, the tribe is easily distinguished due to the presence of following characters such as erect, acaulescent or climbing; Leaves reduplicate, pinnate; usually hapaxanthic, hermaphordite, monoceious or dioecious palms. The tribe consists of six subfamilies, of which the subfamily Calaminae with the monotypic genus Calamus L. has representative in Pakistan Calamus tenuis Roxb .
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As far as palynological studies are concerned various studies are available such as
Sharma (1967) investigated the pollen of Calamus rotang and reported 1 – 2 colpate (2 sino colpate), sub-spheroidal pollen. Thainkaimoni (1971) in his monograph of palm, studied the pollen morphology of many species of Calamus L. with the help of Light
Microscope (viz., C. huegelianus, C. pseudotenuis, C. rotang, C. thwaitesil and C
.travancoricus. Sowunmi (1968 and 1972) has also studied the pollen morphology of the
Palmae. She reported dicolpate pollen in the genus Calamus L., which distinct the genus from all the palm species. Kedves (1980) investigated the pollen of C. andamanicus and
C. palustris.
Pollen morphological studies of the CalamusL. were scanty and most of these work mainly based on LM observations. Therefore, the present investigation is carried out with LM and SEM along with its taxonomic significance.
Pollen description of the tribe Calameae Drude
1. CalamusL.
Pollen disulcate,ellipsoidal or circular in polar view.Tectum finely coarsely reticulate.
1. Calamus tenuisRoxb. (Fig. 2.3 I; 2.4 A & B)
Pollen (22.50-) 25.75 (-29.00)μm in length and (23.50-) 28.00 (-33.00)μm in breadth, asymmetrical and isopolar.Colpus (21.50-) 24.00 (27.50-) 휇m in length.Exine tectate or semitectate, 1.2 – 2.0 휇m thick. Tectum 0.2휇m, veruccate, sometime with reduce excrescences. Muri simple, baculate, bacula indistinct (Table 2.2)
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Discussion of the subfamily Clamoideae
The Calamus type is distinct from all other palm species (with the exception of
Pinanga javana) in having dicolpate grains. It is significantto note that the genera
belonging to the tribe Calameae (sensu Hutchinson), of which all genera in the Calamus
subgroup form a part, are distinguished from all other palms by the hard, shining and
imbricate scales on their fruits; pollen-morphologically, too, they are distinct in having
disulcate pollen which show the remarkable association between gross morphology and
pollen morphology.
The Calamoideae is the only subfamily where disulcate pollen occur (Ferguson &
Harley, 1992). Within the subfamily, pollen of a single species viz., Calamus
tenuisRoxb., of the tribe Calameae is examined. This species is mainly characterized by
the presence of disulcate pollen (Fig. 2.3 I; 2.4 A & B). This exclusive pollen feature
remains distinct cane palm from rest of the members of the family Palmae by having
monosulcate or trichotomosulcate. The present results are in accordance with previous
reports like Sharma (1967), Thanikaimoni (1971), Sowunmi (1968 and 1972), Kedves
(1980) and Ferguson & Harley (1993), Dransfield & Uhl (1998) and Dransfield et al.,
(2005, 2010). Other feature such as pollen elliptic or circular in polar view and
semitectate exine with coarsely reticulate tectum are also in accordance with Ferguson &
Harley, 1993.
Table 2.2 General pollen characters of the sub family Calamoideae
Exinethic Length Breadth Colpus length Name of taxa Aperture kness Tectum (µm) (µm) (µm) (µm) finely 22.50 23.50(28.00) 33.00 21.50(24.0) 27.50 coarsely Calamus urens Disulcate 0.8-1.0 (25.75)29.0±0.797 ±0.970 ±0.997 reticulate- vermucate
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3.SUBFAMILY CORYPHOIDEAE Griff.
Introduction
The Coryphoideae is monophyletic and resolves in most phylogenetic studies as sister to a clade including the Arecoideae and the Ceroxyloideae (Hahn 2002b; Asmussen et al. 2006; Dransfield et al. 2008) and distributed globally, with centre of diversity in the
New World and Asia/Malaysia. The subfamily can be easily differentiated by the presence of combination of characters such as leaves palmate or costapalmate, rarely entire, induplicate, rarely reduplicate (when flowers is apocarpous), or mixed induplicate- reduplicate, or pinnate.Flowers solitary or clustered; never arranged in triads of one central pistillate flower and two lateral staminate flowers. The combinations of these characters segregate the Coryphoideae from other sub families. The subfamily comprised of about 456 species and 40 genera belonging to the eight tribes (Fig. 8.1). Out of which six tribes, the Borasseae (with 3 genera), Caryoteae, Corypheae, Sabaleae (with a single genus), Phoeniceae (a genus with 5 species) and Trachycarpeae (with 6 genera), have been reported from Pakistan (Table 1.2 &1.3).
The pollen morphology of the tribe Coryphoideae has been studied comprehensively with light microscopy by Thanikaimoni (1970 a), Sowunmi (1972).
Various workers such as Ferguson et al., (1987) studied the pollen of the tribe Borasseae in detail with the help of Scanning Electron Microscope. Dransfield et al., (1990) gives an account on the pollen of the Coryphoid palms including the SEM microphotographs.
Ferguson and Harley (1993) examined the pollen of the tribe Coryphoideae in detail with electron microscopy. On the other hand, not a single report on the pollen morphology of
129 the Coryphoid Palm is available from the area under consideration. Therefore, the present work has been carried out from our region.
Key to the Genera
1+ Pollen with supratectal process. Exine verrucose and up to 3 µm thick...……...... 2
- Pollen without supratectal process.Exine not as above and up to 2 µm thick……...3
2+ Pollen up to 50 µm in length.Collumella more than 0.5 µm, dense and narrowed towards tectum…………………………………………………….…..…..Borassus
- Pollen up to 40 µm in length.Collumella less than 0.5 µm, less dense and not narrowed towards tectum……………………………………………...... Hyphaene
3+ Exine regulate or scrobiculate, up to 1 µm thick…………………….…Bismarkia
- Exine reticulate or perforate up to 2 µm thick………………..…………….………4
4+ Length of pollen is less than 20 µm…………………………………………….….5
- Length of pollen is more than 20 µm………………………………..…………….10
5+ Pollen simple baculate……………………………………………………..……….6
- Pollen dupli to multibaculate……………………………………………………….7
6+ Pollen less than 20 µm in length and up to 35 µm in breadth……..……...... Brahea
- Pollen more than 20 µm in length and up to 29 µm in breadth……...……..Phoenix
7+ Pollen up to 47 µm in breadth.Sexine duplibaculate…………………..……..Rhapis
- Pollenup to 33 µm in breadth.Sexine tri-multibaculate………………………..…..8
8+ Pollen multibaculate and more than 35 µm in breadth………………………..Sabal
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- Pollen tribaculate and less than 35 µm in breadth..………………………..……….9
9+ Pollen more than 20 µm in length. Exine perforate to reticulate.…………Livistonia
- Pollen less than 20 µm in length. Exine densly reticulate……….….Trachycarpeae
10+ Pollen reticulate………………………………………………….……Nannorrhops
- Pollen densly perforate-reticulate……...….Group I (Pritchardia & Washingtonia)
1.Tribe Borasseae
The Borassoid Palms represent a well circumscribed apparently monophyletic group of dioecious palms with induplicately palmate or costpalmate leaves (Uhl &
Dransfield, 1987) and by flowers dimorphic, the staminate and sometimes also the pistillate flowers borne in deep pits on the rachillae (Dransfield et al., 1990, Dransfield &
Uhl, 1998). The phylogenetic studies by using morphological and molecular datasets have determined the tribe Borasseae as a monophyletic group (Uhl et al., 1995; Barrow,
1996).The tribe is divisible into two apparently natural subtribes on the basis of gross morphological characters viz., subtribe Lataniinae Meisner and subtribe Hyphaeninae
Beccari. The former subtribecomprised of 4 genera viz., Borassus L., Borassodendron
Becc., Latania Comm. Ex Juss. and Lodoicea Comm. and easily keyed out on the basis of their reproductive features such as pistillate flowers are sessile and pits absent on the rachillae (the axis of a spikelets). Whereas, the later subtribe is also comprised of 4 genera viz., Bismarkia Hildebr. & H. Wendl, Hyphaene Geartn., Medemia WÜrrtemb. ex
H. Wendl and Satranala Dransf. & Beentje. In thissubtribe, the pistillate flowers are pedunculate and pits are present on both theflowers.
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As far as the morphological study of pollen is concerned various reports are available such as Thanikaimoni (1970), Sowunmi (1972), Kedves (1980) and Ferguson
&Harley (1992) discussed the pollen of the tribe Borasseae as they give general overview of the pollen morphology of the family Palmae. Ferguson et al. (1986) gives a detailed account on the pollen morphology of the tribe Borasseae by using LM and SEM. The present study gives a detailed examination of pollen morphology of the taxa belonging to the tribe Borasseae (subfamily Coryphoideae) from Pakistan and Kashmir.
Observations and Results
The summary of pollen morphological data of the studied taxa belonging to the sub-family Coryphoideae is given in the Table 2.3 where as LM and SEM photographs are given in Figure 2.4-2.7.
Pollendescription of the tribe Borasseae
Pollen monosulcate, usually elliptical or subcircular in apertural view.Size:
Pollen(26-) 29 (-32) µm in length and(27-) 38.5 (-50)µm in breadth. Aperture membrane is smooth, thin and usually narrow, inconspicious aperture margin. Aperture length is almost to the size of longest axis in polar view (i.e., 32 - 81 µm). Colpus 33.50 (40.75)
48.00µm in length.Exine tectate or semitectate, 1.0 – 6.0 µm thick, supratectal process present or absent. The exine ornamention in general may be verrucose, rugulate or reticulate, or finely perforate with supratectal gemmae.
It is represented by 3 genera viz.,Bismarkia Hildebr. & H. Wendl, Borassus L. and Hyphaene Gaertner
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Key to the Genera
1+ Supratectal gemmae present…………………………..……………………………2
- Spratectal gemmae absent……………………………….……..…....…1. Bismarkia
2+ Pollen more than 45 µm long. Columella 0.6 µm high, dense. Tectum 0.8 µm thick ...……...……………………………………..……………….…………..2. Borassus
- Pollen less than 45 µm long. Columella 0.3 µm high, less dense. Tectum 0.4 µm
thick …………………………...... …..……….…………...…….……...3. Hyphaene
1.Bismarkia Hildebr. & H. Wendl.
Pollen monosulcate, elliptical to more or less circular in polar view.Tectum regulate,
supratectal gemmae absent.
Bismarkia nobilis Hildebr. & H. Wendl. (Fig. 2.4 E & F; 2.9 A)
Pollen (42.50-) 45.95 (-49.40) µm in length and (26.00-) 28.80 (-31.60) µm in breadth,
asymmetrical.Aperture equal or somewhat shorter than the length.Colpus (41.0-) 44.80
(48.60-) 휇m in length.Exine tectate, 1.0 µm thickTectum 0.8 µm thick, perforate.
2.Borassus L.
Pollen monosulcate, elliptical in polar view.Tectum verrucose, supratectal gemmae
present.
2. Borassus flabellifer L. (Fig. 2.4 G & H; 2.9 C)
Pollen (39.00-) 45.00 (-51.00) µm in length and (33.00-) 36.50 (-40.00) µm in
breadth.Aperture long (equal or somewhat shorter than the length).Colpus (37.5-) 43.65
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(49.8-) 휇m in length.Exine tectate, 2.0 – 3.0 µm thick including gemmae.Tectum 0.8 µm thick including gemmae.
3. Hyphaene Gaertner
Pollen monosulcate, elliptical to sub circular in polar view.Tectum verrucose, supratectal gemmae present.
1. Hyphaene thebaica(L.) Mart. (Fig. 2.4 C & D; 2.9 B)
Pollen (34.00-) 36.00 (-40.00) µm in length and (31.00-) 34.30 (- 38.00) µm in breadth.Aperture equal or somewhat shorter than the length.Colpus (33.50-) 36.25 (-
39.00) 휇m in length.Exine tectate, 2.0 - 3.0 µm thick including gemmae.Tectum 0.4 µm thick including gemmae, perforate.
2. Tribe Caryoteae
The Caryoteae is monophyletic and resolves in most phylogenetic studies as sister to a clade including the Corypheae and Borasseae (Bayton 2007; Asmussen et al. 2006;
Dransfield et al. 2008). Previously, the tribe is treated under the subfamily Arecoideae due to the presenceof remarkable similarity in the inflorescence including triad flowers or triad deravatives in the inflorescence (Dransfield & Uhl, 1998). The phylogenetic studies clear the status of the tribe Caryoteae by placing it under the tribe Coryphoideae instead of Arecoideae (Uhl et al., 1995; Baker et al., 1999; Asmussen et al., 2000; Asmussen &
Chase, 2001; Hahnn, 2002a, b; Dransfield et al., 2005; Dowe, 2010).ThisOld World tribe immediately identifiable by the presence of induplicate; pinnate and
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Figure 2.4. Scanning Electron Micrographs (SEM) of the pollengrains:Calamus tenuis:
A, pollen grain; B, exine patternHyphaene thebaica: C, pollen grain;D, exine pattern.
Bismarkia nobilis: E, pollen grain; F, exine pattern. Borassus flabellifer:G, pollen grain;
H, exine pattern (scale bar: B, D = 2 μm; C, F = 5μm; A, E = 10 μm).
135
136 bipinnate-leaved taxa, in three genera, Caryota, Arenga and Wallichia. The genus is highly supported to a clade of Wallichia and Arenga (Bayton 2005, Asmussen et al.,
2006). Among these genera, Caryota L. (with 12 species), is one of the most distinct genera of palms with its concolorous bipinnate leaves, ruminate endosperm and bisexual inflorescence with triads of flower. In Pakistan the genus have a single representative i.e.,
Caryota urens L.
The pollen of the tribe as well as the species has been described by Thanikaimoni
(1970), Sowunmi (1972), Kedves (1980), Ferguson et al. (1986)and Ferguson & Harley
(1992). However, not a single study has been made on the pollen morphology of the tribe
Caryoteae therefore; the present work has been carried out by using LM and SEM.
Represented by a single genusviz., Caryota L.
Pollen Description of the Tribe Caryoteae
1. Caryota L.
Pollen monosulcate, elliptical or circular in polar view.Tectum finely clavate or less frequently spinose.
1. Caryota urens L. (Fig. 2.2 G & H; 2.3 H)
Pollen (13.50-) 20.35 (-27.20) µm in length and (16.20-) 25.40 (-34.60) µm in breadth, generally small, asymmetric.Aperture equal in length to the longer axis of pollen.Colpus
(13.00-) 19.70 (26.40-) 휇m in length.Exine intectate, 2-2.5 µm thick.Tectum densly clavate-baculate, less frequently spinose or with protuberances; spines attached upper surface of foot layer.
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3.Tribe Corypheae
The tribe Corypheae is resolved as monophyletic in most phylogenetic studies, and as sister to the Borasseae (Bayton 2007; Asmussen et al. 2006; Dransfield et al.
2008).
The tribe Coryphaea is distinguished by the induplicate, palmate or costapalmate leaves; by genera being hermaphrotide, polygamous, or rarely dioecious and the absence of pits on the rachillae. The Coryphineae, a heterogenous subtribe of 4 Old World genera
(viz., Chuniphoenix Burr., Corypha L., Kerridoxa Dransf. and Nannorrhops H. Wendl.).
Of which the genus Nannorrhops H. Wendl., with a single species Nannorrhops ritichiana (Griff.) Aitchi., has representative in Pakistan, is a source of fibre for weaving and rope making.The members of the Coryphineae is distinct by the presence of basally fused carpels with free or connate styles but often with separate styler canals. The phylogenetic studies have resolved Chuniphoenix Burr., Kerridoxa Dransf. and
Nannorrhops H. Wendl., in a monophyletic group with Corypha L. usually in a sister position and the subtribe is further resolved as related to the Borasseae.
The pollen morphology of the tribe including the genus Nannorrhops H. Wendl., has been studied quite extensively with the help of light microscope (LM) by
Thanikaimoni (1970 a), Sowunmi (1972), Kedves (1980), Ferguson & Harley (1993),
Dransfield et al., (1990); Dransfield & Uhl (1998).
The purpose of present study is to provide the detailed account of the pollen characters by using light microscope (LM) and Scanning Electron Microscope (SEM) and to determine its systematic significance.
138
Represented by a single genus viz., Nannorrhops H. Wendl.
Pollen Description of the Tribe Corypheae
1. Nannorrhops H. Wendl.
Pollen monosulcate, ellipsoidal or sub-circular in polar view, Tectum reticulate.
1. Nannorrhops ritichiana(Griff.) Aitc. (Fig. 2.5 A & B; 2.9 D)
Pollen (21.20-) 29.85 (-38.50) µm in length and (28.30-) 31.40 (- 34.50) µm in breadth, usually slightly asymmetric.Colpus (20.66-) 29.30 (-37.40) in length.Aperture margin psilate or scabrate.Exine tectate, 1.5 µm thick.Infratectum columellate.Muri simpli-dupli baculate.
1. Tribe Phoeniceae
The tribe Phoeniceae is one of the eight tribes representing the subfamily
Coryphoideae, with the single genusPhoenix L.with about 14 species (Govaerts and
Dransfield, 2005; Henderson et al., 2006; WCSP, 2013)distributed from Canary Island east across the Northern and Southern Africa in to the Middle East. from South Europe and South East Asia from Turkey east to Southern China and Malaysia. In Pakistan the monotypic genus Phoenix L. is represented by 5 species among these speciesPhoenix sylvestris(L.) Roxb., is the wild.
The tribe is distinguished amongCoryphoideae, being the only genuswith pinnate leaves. Furthermore, a combination of characters also helpful to distinguished Phoenix
L., such as dioecious,apocarpioc flowers and induplicate leaves with basal leaflets modified as spines.
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Figure 2.5. Scanning Electron Micrographs (SEM) of the pollen grains:Nannorrhops ritichiana: A, pollen grain; B, exine pattern.Brahea brandeegie: C, pollen grain;D, exine pattern. Livistoniachinensis: E, pollen grain; F, exine pattern. Trachycarpus fortunei:G,pollen grain; H, exine pattern (scale bar: B, F, H =1μm; D, = 2μm; A, E,
G = 5μm; C=10μm).
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141
At the molecular level, the genus also appears highly divergent from other palms but
it remains hardly classifiable in Cladistic analysis (Asmussen and Chase 2001; Hahn
2002; Baker et al., 2009).Human use of palms is as old as human civilization itself,
starting with the cultivation of the date palm.The importance of palms is mentioned more
than 30 times in the Bible and at least 22 times in the Quran.The Phoenix dactylifera
(date palm) is cultivated mainly for its fruit and also provides a favorable environment for
the cultivation of other species such as olives, figs, vegetables and in Asianagro systems
(Tengbergand Newton, 2007).The sap of P. sylvestris (L.)Roxb.is boiled to produce a
sweet juice (Newton et al., 2013). P.canarrensis Hort. ex Chabaud is used for
ornamentation. The pollen morphology of cultivated plants has attained a great deal of
attention in recent years due to its effective application in interpretation of the taxonomy
and to find out the inter-relationships of cultivated taxa.
There are several palynological studies on species belonging to the genus Phoenix L.
such as Erdtman (1952), Mahabale (1967) and Sowunmi (1968, 1972), Kedves (1980),
Dransfield et al., (1990).This is the first attempt to analyze the detailedpalynological
studies of the genus Phoenix L. to examined the pollen morphological variations of
differen species from Pakistan and Kashmir.
PollenDescription of the tribe Phoeniceae
1. PhoenixL.
Pollen monosulcate elliptic and oblong or sub rounded in polar view, transversely kidney shaped and boat-shaped longitudinally.Tectum finely reticulate, perforate.
1. Phoenix dactyliferaL.(Fig. 2.6A&B; 2.10 C)
Pollen(15.60-) 18.05(-20.50) µm in length and (22.10-) 24.65(-27.20) µm in
142 breadth.Colpus(15.20-) 17.60(-20.00) µm in length.The aperture more or lesssame length as long axis and located at the distal pole of pollen.Muri being simplibaculate.Exine tectate 1.0 µmthick. .
2.Phoenix loureiriiKunth(Fig. 2.6 C & D; 2.10 D)
Pollen(15.70-) 18.75 (-21.05) µm in length and (21.05-) 23.67 (-26.30) µm in breadth.asymmetric sometime symmetric, isopolar to heteropolar.Colpus(20.40-) 22.95(-
25.50) µm in length.The aperture more or less same length as long axis and located at the distal pole of pollen.Muri being simpli baculate.Exine tectate 1.1 µm thick.
3.Phoenix roebelenii O’Brien(Fig. 2.6 E & F; 2.10 E)
Pollen(20.30-) 24.50 (-28.70) µm in length and (21.20-) 24.42 (-27.65) µm in breadth.Asymmetric sometime symmetric, isopolar.Colpus(20.00-) 23.85(-27.10) µm in length. The aperture more or less same length as long axis and located at the distal pole of pollen.Muri being simpli baculate.Exine tectate and thickness ranges from 1.0 µm thick.perforation ranging from fine to coarse with sparse to dense distribution.
4.Phoenix sylvestris (L.) Roxb. (Fig. 2.6 G & H; 2.10 F)
Pollen(14.50-) 17.75 (-21.00) µm in length and (22.10-) 24.95 (-27.80) µm in breadth, asymmetric.Colpus(14.10-) 17.35(-20.60) µm in length.The aperture more or less same length as long axis and located at the distal pole ofpollen.Muribaculate. Exine tectate 1.0
µm thick.
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Figure 2.6. Scanning Electron Micrographs (SEM) of the pollen grains:Phoenixdactylifera: A, pollen grain; B, exine pattern. Phoenixloureiri:
C, pollen grain;D, exine pattern. Phoenix roebelenii: E, pollen grain; F, exine pattern. Phoenix sylvestris:G, pollen grain; H, exine pattern(scale bar: B, D,
F, H = 1μm; A, E = 2 μm; C, G = 5 μm).
144
145
5. Tribe Sabaleae
The Sabaleae is one of the 6 tribes of subfamily Coryphoideae, represented by a single New World genus Sabal Adanson with 16 species (Dransfield & Uhl, 1987; Zona,
1990), widely distributed in and around the Carribean region in Bermuda, South eastern
United States, Mexico, Central America, and through northern Columbia and Venezuela to Trinidad. The presence of costapalmate leaves with smooth and non thorny petiole and long arching costa that gives the leaf a three dimension shape, and fused carpels through out the stylar region with a single stylar canal distinguished the tribe Sabaleae from the other tribes.Sabal Adanson, a sole genus is used for making brooms, locally used as a source of thatching and also cultivated as ornamental plants. In Pakistan the genus is represented by a single species viz., Sabal minor.
Observation and Results
The pollen morphological data of the studied taxon is given in Table 2.3 and LM and
SEM micrograph is given in Fig. 2.5 & 2.6.
Pollen description of the tribe Sabaleae
1.Sabal Adanson
Pollen monosulcate, elliptic in apertural view. Tectum finely reticulate.
1. Sabal minor (N.J. Jacq.) Persons (Fig. 2.7 G & H; 2.10 B)
Pollen (18.50-) 19.95 (-20.80) μm in length and (27.60-) 33.70(-40.00) μm in breadth. asymmetric.Colpus (17.90-)19.05(-20.20) μm in length.Exine tectate, up to 2 µm thick.
Infratectum columellate. Muri simple multibaculate.
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6.Tribe Trachycarpeae
The Trachycarpeae (previously known as Livistoneae) is the most diverse and commontribe among the coryphoid palm of Pakistan, comprised of 6 genera and 10 species divided in two subtribes,the Livistoninae and the Rhapidinae. The tribe is resolved as monophyletic in most studies (Asmussen et al. 2006; Dransfield et al.
2008).Both these subtribes previously treated under the tribe Coryphoideae (see
Dransfield and Uhl, 1998) but at present these subtribes has been placed under the tribe
Trachycarpeae (Dransfield et al., 2008 and Dowe, 2010). The subtribe Livistoninae is represented by five genera, of which 4 genera (viz., Brahea Mart. ex Endl., Livistona R.
Br., PritchardiaLabill and WashingtoniaH. Wendl) has representative in Pakistan.The subtribe Rhapidinae occurs in south-eastAsia and in the Mediterranean region, and is represented by 6 genera, of which only 2 Old World genera (viz., Rhapis L. and
Trachycarpus H. Wendl) has representative in study area(Table. 1.3). Both these subtribes can be distinguished on the basis of the degree of fusion of the carpels. For instance, in Livistoninae gynoecium tricarpellary, connate; carpels only free at the base but fused through out their styles. On the other hand,subtribe Rhapidinae hasapocarpous gynoecium (carpels distinct).
Palynological studies of the members belonging to the tribe Trachycarpeae has been carried quite extensively with Light Microscopy (by Thanikaimoni, 1971; Sowunmi,
1968; 1972) and with LM, SEM and TEM (by Kedves, 1980; Dransfield et al., 1990;
Ferguson and Harley, 1993). The present work reports the first descriptive information on the pollen morphological study of eight taxa of the tribe Trachycarpeae by using LM and
SEM from study area.
147
Observation and Results
The pollen morphological data of the studied taxon is given in Table 2.3 and LM and
SEM micrograph is given in Fig. 2.5 & 2.6.
Pollen description of the tribe Trachycarpeae
Pollen monosulcate, elliptic to subcircular in polar view.Size: Pollen(14.50-) 16.55(-
18.60)μm in length and (20.00-) 27.35 (-34.20)μm in breadth.Aperture moreor less equal in length to the longest axis, aperture membrane thin, smooth.Colpus (14.50-) 16.25(-
18.00) μm in length. Exine tectate, up to 1.5-2 µm thick.Tectum finely or densly reticulate, perforateto reticulate.Muri simple baculate to dupli or tripli to multibaculate.
Represented by 6 genera viz., Brahea Mart. ex Endl., Livistona R. Br., Pritchardia Labill and WashingtoniaH. Wendl, Rhapis L. and Trachycarpus H. Wendl.
1.Brahea Mart. ex Endl.
Pollen monosulcate, elliptic in polar view.Tectum perforate.
1. Brahea brandegeei(Purpus) H.E.Moore (Fig. 2.5 C & D; 2.9 E)
Pollen(14.50-) 16.55(-18.60)μm in length and (20.00-) 27.35(-34.20)μm in breadth, asymmetric.Colpus (14.50-) 16.25(-18.00) μm in length.Exine tectate, up to 2 µm thick.
Infratectum collumellate.Muri simple multi-baculate.
2. LivistoniaR. Br.
Pollen monosulcate, elliptic in apertural view.Tectum finely perforate to reticulate.
1. Livistonia chinensis (N. J. Jacquin) R. Br. ex Mart. (Fig. 2.5 E & F; 2.9 F)
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Pollen(15.10-) 19.70(-24.30)μm in length and (23.68-)28.94(-34.20)μm in breadth,asymmetric.Colpus (15.10-) 19.55 (-24.00)μm in length.Exine tectate, up to 2 µm thick. Infratectum collumellate.Muri simple multibaculate.
3.PritchradiaLabill
Pollen monosulcate, elliptic in polar view.Tectum densly perforate to reticulate.
1. Pritchardia beccariana Rock (Fig. 2.8 E & F; 2.10 I)
Pollen(19.50-) 21.00 (-22.60) μm in length and (25.20-) 30.45 (-35.70)μm in breadth, asymmetrical.Colpus(19.00-) 20.55 (-22.10)μm in length.Exine tectate, up to 2.2 µm thick.Infratectum collumellate.Muri simple multibaculate.
4.WashingtoniaH. Wendl.
Pollen monosulcate, elliptic in polar view.Tectum finely reticulate.
1. Washingtonia filifera (L. Linden) H. Wendl. (Fig. 2.7 A – D; 2.9 I)
Pollen(32.89-) 36.18 (-39.47)μm in length and (30.26-) 32.89 (-35.52) μm in breadth,
Colpus (32.60-) 35.75 (-38.90)μm in length.Usually asymmetrical.Exine tectate, up to 1.1
µm thick. Infratectum collumellate. Muri simple multibaculate.
2. Washingtonia robusta H. Wendl.(Fig. 2.7 E & F; 2.10 A)
Pollen(34.62-) 37.24 (-39.86)μm in length and (30.20-) 33.90 (-37.60) μm in breadth,
Colpus (33.25-) 36.32 (-39.40)μm in length.Exine tectate, up to 1.2 µm thick. Infratectum collumellate Muri simple multibaculate.
5.Raphis L.
Pollen monosulcate, elliptic in polar view.Tectum finely reticulate.
1. Rhapis excelsa (Thunb.) Henry ex Rehder (Fig. 2.8 A & B; 2.10 G) 149
Pollen (14.30-) 16.65 (-19.00) μm in length and (21.80-) 30.50 (-38.30) μm in breadth, asymmetric.Colpus (14.10-) 15.80 (-17.50)μm in length.Exine tectate, up to 2 µm thick.
Infratectum columellate.Muri simple multibaculate.
2. Rhapis multifida Burret (Fig., 2.8 C & D; 2.10 H)
Pollen(-17.50) 19.95(-22.40) μm in length and (27.20-) 32.92 (-38.65) μm in breadth, asymmetric.Colpus (17.20-) 19.50 (-21.80)μm in length.Exine tectate, up to 2 µm thick.
Infratectum columellate.Muri simple multibaculate.
6.Trachycarpus H. Wendl.
Pollen monosulcate, elliptic in polar view.Tectum finely reticulate.
1. Trachycarpus fortunei (Hook. f.) H. Wendl. (Fig. 2.3 C ; 2.5 G & H)
Pollen (14.80-) 17.20 (-19.60) μm in length and (19.40-) 25.30 (-31.00)μm in breadth.Colpus (14.60-) 17.10 (-19.60)μm in length.Exine tectate, 1.8 µm thick.
Infratectum columellate Muri simple multi-baculate.
Discussion of the subfamily Coryphoideae
All the species studied in this subfamily have remarkably similar and uniform pollen i.e., monosulcate and elliptical in polar view. Thanikaimoni (1970) described that all the coryphoid genera have similarity in their pollen morphological characters. Sowumni
(1972) mentioned coryphoid members in three subfamilies viz., Borissoideae (Borassus
L., Bismarkia and Hyphaene Gaertner), Coryphoideae (Brahea, Livistonia,Nannorhops,
Pritchardia, Rhapis, Sabal, TrachycarpusH.Wendl. and WashingtoniaH.Wendl.) and
Phoenicoideae (Phoenix L.). She divided Borrisoideae in to two groups on the basis of exine pattern whereas Coryphoideae and Phoenicoidae described as homogenous
150 subfamilies. Similarly,Dransfield et al., (1990) included all the members of the coryphoid palms in the tribe Corypheae excluding the genus Phoenix L. They reported that the pollen of tribes Corypheae and Phoeniceae show more similarity in their pollen characters.The results are more or less in accordance with the present findings. However, the pollen of the coryphoid palm share same aperture type while the differences are found in the pollen size, exine thickness, pattern and the number of baculae. On the basis of these pollen characters, the members of the subfamily Coryphoideae has been divided in to 4 pollen types such as Bismarkia-type, Borassus-type, Brahea-type and Lavistonia type.
Key to the pollen types
1+ Exine verrucose, up to 3µm thick, supra tectal gemmae present……... Borassus type
- Exine not as above, up to 2 µm thick, supra tectal gemmae absent…….……………2
2+ Exine rugulate; up to 1 µm thick; pollen more than 30 µm in length..Bismarkia type
- Exine not as above; pollen less than 30 µm in length…………..…….………………3
3+ Exine mostly perforate……………………………………...... …….……Brahea-type
- Exine reticulate……………………………………………..…...…..……..Sabal-type
Tribe Borasseae
In this tribe pollen morphology of the 3 genera are studied which falls in to 2 pollen types on the basis of exine features viz., Borassus-type and Bismarkia-type.The two taxa i.e., Borassus flabellifer L. (subtribe Lataniinae) and Hyphene thebaica(L.)
Mart. (subtribe Hyphaeninae) falls in the former pollen type because they show more
151 similarity in their pollen characters such as shape, aperture, exine and particularly striking in the supratectal process only a difference has been found in their pollen size. It is interesting to observe that in terms of gross morphological characters both these taxa are not closely related, as they belong to different subtribes of the Borasseae. On the other hand, later pollen type is represented by a single species i.e., Bismarkia nobilisin which exine is rugulate, up to 1.0 µm thick where as in the other pollen type exine is verrucose up to 1.0 µm thick. Similartype of pollen in the Borossoid palm have also been examined by Thanikaimoni (1970), Sowunmi (1972), Ferguson and Harley (1983) and Dransfield et al., (1990).
Tribe Caryoteae
Within this tribe a single species i.e., Caryota urens L. is characterized by reticulate exine and belonging to the Sabal-type.The pollen is up to 25 µm in breadth which differ the species from the other members of this pollen type.
Tribe Corypheae
Within this tribe a single species i.e., Nannorhops ritichiana(Griff.) H.Wendleis examined. This species is characterized by reticulate exine and belonging to the Sabal- type.The pollen is up to 34µm in breadth which differ this from the other members of this pollen type.
Tribe Phoeniceae
A tribe of monotypic genus represented by 5 species, of which pollen of 4 species has been studied viz., Phoenix dactylifera L., Phoenix loureirii Kunth, Phoenix sylvestris
(L.) Roxb &Phoenix roebelenii O’Brien. The pollen of this tribe is almost similar to that
152 of other coryphoid members i.e., monosulcate, elliptic in polar view. Nevertheless, the pollen of Phoenix L. is distinguishable by the combination of characters such as small accordance with earlier workerssuch as Sowunmi, 1972; Dransfield et al., 1990; Harley and Hall, 1991; Ferguson and Harley, 1993.
Tribe Sabaleae
In this tribe a single taxa viz., Sabal minor(N. J. Jacq.) Persons is examined. It is characterized by reticulate exine thus belonging to Sabal-type.
Tribe Trachycarpeae
In this tribe pollen of 8 species belonging to 6 generas are examined. All these members can be clearly delimited on the basis of exine pattern and represented in two different pollen types.For instance, Brahea brandegeei (Purpus) H. E. Moore, Livistonia chinensis(N. J. Jacquin) R. Br. ex Mart., Pritchardia beccariana Rock, have perforate exine and belong to Brahea-type. Wheras in the remaing taxa viz., Rhapis multifida
BurretRhapis excelsa (Thunb.) Henry ex Rehder,Trachycarpus fortunei (Hook. f.) H
Wendl., Washingtonia filifera (L. Linden) H. Wendl.Washingtonia robusta H. Wendl., exine is reticulate and represented by Sabal-type.
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Figure 2.7. Scanning Electron Micrographs (SEM) of the pollen grains:Washingtonia filifera: A, C pollen grain; B, D exine pattern.
Washingtoniarobusta: E, pollen grain; F, exine pattern. Sabalminor:G, pollen grain; H, exine pattern (scale bar: D, H = 1 μm; B = 2 μm; A, C, E = 10 μm).
154
155
Figure 2.8. Scanning Electron Micrographs (SEM) of the pollen grains:
Rhapis excelsa: A, pollen grain; B, exine pattern. Rhapis multifida:C,pollen grain;D, exine pattern. PritchardiabeccarianaE, pollen grain; F, exine pattern.Wodyetia bifurcata G, pollen grain; H, exine Pattern (scale bar: B, D
= 1 μm; A, C = 5μm).
156
157
Figure 2.9 Light Micrographs (LM) of the pollen grains at 100 x:A,
Bismarkia nobilis: pollen grain. B, Hyphaene thebaica:pollen grain. C,
Borassus flabellifer:pollen grain. D,Nannorrhops ritichiana:pollen grain.
E,Brahea brandeegie:pollen grain.F, Livistoniachinensis: pollen grain.
G,Ptychosperma elegans: pollen grain. H, Ptychospermamacarthuri:pollen grain. I, Washingtoniafilifera:pollen grain.(Scale Bar = 10μm).
158
159
Figure 2.10 Light Micrographs (LM) of the pollen grains at 100x:
A,Washingtoniarobusta:pollen grain.B,Sabalminor:pollen grain.C,
Phoenixdactylifera:pollen grain. D,Phoenix loureiri:pollen grain.
E,Phoenixroebelenii:pollen grain.F, Phoenixsylvestris: pollen grain. G,Rhapis excelsa:pollen grain. H, Rhapismultifida: pollen grain. I, Pritchardia beccariana:pollen grain. (Scale Bar = 10μm).
160
161
Table 2.3 General Pollen Characters of the Sub Family Coryphoideae
Exine Colpus length Name of taxa Tribe Length(µm) Breadth(µm) Aperture thickness Tectum (µm) (µm) Bismarkia nobilis Borasseae 42.50 (45.95 ) 49.40 26.00 (28.80) 31.60 41.00(44.80)48.60 monosulcate 1.0 rugulate or reticulate perforate ± 0.769 ± 0.627 ± 0.798 Borassus flabellifera Borasseae 39.00(45.00) 51.00 33.00 (36.50) 40.00 37.50(43.65)49.80 monosulcate 2.0-3.0 reticulate - verrucose ± 1.437 ± 0.755 ± 1.419 Hyphaene thebaica Borasseae 34.00(36.00) 40.00 31.00 (34.30) 38.00 33.50 (36.25) 39.00 monosulcate 2.0 reticulate - verrucose ± 0.672 ± 0.996 ± 0.566 Caryota urens Caryoteae 13.50(20.35) 27.20 16.20(25.40) 34.60 13.00(19.70)26.40 monosulcate 2 – 2.5 finely and densly ± 1.772 ± 2.361 ± 1.534 clavate-baculate Nannorrhops ritichiana Corypheae 21.20(29.85)38.50 28.30 (31.4) 34.50 20.66(29.03) 37.40 monosulcate 1.5 reticulate or foveolate - ± 1.899 ± 0.643 ± 1.896 reticulate Brahea brandegeei Trachycarpeae 14.50 (16.55) 18.60 20.00 (27.35) 34.20 14.50 (16.25) 18.00 monosulcate 1.0-1.5 reticulate ± 0.502 ± 1.541 ± 0.430 Livistonia chinensis Trachycarpeae 15.10(19.70) 24.30 23.68(28.94)34.20 15.10(19.55)24.00 monosulcate 1.0 perforate - reticulate ± 1.052 ± 1.049 ± 0.944 Pitrichardia beccariana Trachycarpeae 19.50 (21.05) 22.60 25.20 (30.45) 35.70 19.00 (20.55) 22.10 monosulcate 2.2 perorate- reticulate ± 0.365 ± 1.068 ± 0.361 Washingtonia filifera Trachycarpeae 32.89(36.18) 39.47 30.26(32.89)35.52 32.60 (35.75) 38.90 monosulcate 1.1 reticulate ± 0.747 ± 0.569 ± 0.714 Washingtonia robusta Trachycarpeae 34.62(37.24) 39.86 30.20(33.90)37.60 33.25(36.32)39.40 monosulcate 1.2 reticulate ± 0.622 ± 0.741 ± 0.846 Rhapis excelsa Trachycarpeae 14.30 (16.65) 19.00 21.80 (30.50) 38.30 14.10 (15.80) 17.50 monosulcate 1.9 reticulate ± 1.35 ± 1.75 ± 1.22 Rhapis multifida Trachycarpeae 17.50 (19.95) 22.40 27.20 (32.92) 38.65 17.20 (19.50) 21.80 monosulcate 2 reticulate ± 0.762 ± 1.251 ± 0.499 Trachycarpus fortunei Trachycarpeae 14.80 (17.20)19.60 19.40 (25.30) 31.00 14.60 (17.10) 19.60 monosulcate 1.8 coarsely reticulate ± 0.560 ± 1.389 ± 0.523 Phoenix dactylifera Phoeniceae 15.60 (18.05) 20.50 22.10 (24.65)27.20 15.20(17.60) 20.00 monosulcate 1.0 reticulate ± 0.720 ± 0.688 ± 0.679 Phoenix loureirii Phoeniceae 15.70 (18.75)21.05 21.05 (23.67)26.30 20.40(22.95) 25.50 monosulcate 1.10 reticulate ± 0.723 ± 0.620 ± 0.641 Phoenix roebelenii Phoeniceae 20.30(24.50)28.70 21.20(24.42)27.65 20.00(23.85) 27.10 monosulcate 1.0 reticulate. ± 0.955 ± 0.657 ± 0.780 Phoenix sylvestris Phoeniceae 14.50(17.75) 21.00 22.10 (24.95) 27.80 14.10(17.35) 20.60 monosulcate 1.0 reticulate ± 0.325 ± 0.425 ± 0.360 Sabal minor Sabaleae 18.50 (19.95) 20.80 27.60 (33.70) 40.00 17.90(19.05) 20.20 monosulcate 2.0 reticulate ± 0.269 ± 1.04 ±0.248
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Chapter 3
STATISTICAL ANALYSIS BASED ON PALYNOLOGY
INTRODUCTION
Numerical taxonomy or Phenetics is a classification system in biological systematic (Sokal & Sneath, 1963). The modern concept of numerical taxonomy had their beginning for the contributions of Sneath (1957), Michener & Sokal (1957) and Sokal &
Michener (1958) which culminated in the publication of “Principles of Numerical
Taxonomy” (Sokal & Sneath, 1963).This was later elaborated by the same authors (Sneat cladeh & Sokal, 1973). Numerical taxonomy aims to develop methods that are objective, explict and repeatable, both in the evolution of taxonomic relationships and in the erection of taxa (Sokal & Sneath, 1962).Currently statistical or multivariate analysis are frequently using in different disciplines to explain the quantitative attributes. In
Taxonomy it uses to classify and ordinate different plant species on the basis of similarities or dissimilarities.The major aspire of classification is to summarize quantitative data in order to understand the relationship between groups of species
(Greig-Smith, 1983; McCune and Grace, 2002).
In the first cladistic analysis of the family Palmae based on morphological and chloroplast DNA (cp DNA) restriction site data. Uhl et al., (1995) identified the
Livistonneae as part of clade that also included the Thrinacinae and was therefore paraphyletic.There are several studies of the utilization of numerical analysis in solving taxonomic problems and to find out inter relationship among different taxa viz.,
163
During the last 20 years numerous phylogentic studies of the Palm have been published at various taxonomic levels to maintain its taxonomic stability and to recognize morphologically diagnosable entities (Uhl et al., 1995; Asmussen, 1999; Baker et al.,
1999 b; Pintaaud, 1999; Asmussen et al., 2000; Baker et al., 2000; Asmussen & Chase,
2001; Hahn, 2002a, b; Lewis, 2002; Baker and Loo, 2004; Gunn, 2004; Bayton, 2007;
Dransfield et al., 2005; Asmussen et al., 2006). However, advance data sources are elucidate ambiguities that remain in the systematic of the family Palmae.
The aim of the present study is to classify taxa on the basis of pollen characters and to quantify the species relationships belonging to the family Palmae based on numerical taxonomic techniques.
PHYLOGENETIC RELATIONSHIPS OF PALMS FROM PAKISTAN
The family Palmae is placed in the order, Arecales, whichresolves in most recent phylogenetic studies as sister tothree other orders, namely the Commelinales, the
Poalesand the Zingiberales, all of which are included in thebroadly termed ‘commelinid monocots’ (Chase et al. 2000;Soltis et al. 2005).The Palm family (Arecaceae-Palmae) is a monophyletic group in all higher level phylogenetic studies of monocotyledonous plants (Chase et al., 2000, Asmussen & Chase, 2001; Govaerts & Dransfield, 2005;
Dransfield et al., 2008).
There are five major lines of evolution in the Palmae (Asmussen et al. 2006;
Dransfield et al. 2008) and representatives of three lines occur within the Pakistan palmflora. The major lines of evolution are classified systematically as five subfamilies
(Fig. 1.2; Table- 1.3) and are further divided into tribes and subtribes. Of the 14 tribes in the diverse subfamily Arecoideae, only three, Areceae, Cocoseae and Roystoneae, occur
164 in Pakistan. Within the tribe Areceae, out of 11 subtribes, three have representative genera in study area whereas the other two tribes are represented by a single genus.
Similarly, the Coryphoideae, with eight tribes globally, has representatives of six tribes
(viz., Borasseae, Caryoteae, Corypheae, Phoeniceae, Sabaleae and Trachycarpeae) in the area under considerations. The Calamoideae is represented, by a monotypic genusCalamus. The remaining two subfamilies viz.,Ceroxyloideaeand Nypoideae have no representatives in Pakistan (Table-1.2).
MATERIALS AND METHODS
In the present work more than 500 herbarium specimens of 27 taxa has been studied based on pollen characters. The morphological characters of used in the study were based on personal observation of the material and were given equal weight. The qualitative characters were recorded in binary state i.e., in terms of 1 and 2.Where as for quantitative characters their minimum, maximum and average values were used. While in cases of presence or absence, certain characters were recorded as 1 and 0 respectively.
The characters and character state used for performing hierarchical clustering and are listed in tables 3.1 – 3.6.
Agglomerative cluster analysis was performed choosing the Euclidean distance as the resemblance function and Ward’s method for a group linkage method (McCune & Grace,
2002) so as to expose the group structure in the species on the basis of different pollen characters. The computations were performed using the computer program PC-ORD (version
6.0) (McCune and Grace, 2002; Peck, 2010). Different pollen characters of plant species belong to three sub families (Arecoiadeae and Coryphoideae)of Palmeae are use in data matrix. Sub family Calamoideae are not use to run cluster analysis due to small sample size.
165
Table 3.1 List of characters, scored for cluster analysis for the taxa of the family Palmae in Table 3.2.
Character description
1. Length (µm)
2. Breadth (µm)
3. Exine Thickness (µm)
4. Aperture number
Aperture type
5. Sulcate: Absent (0), Present (1)
6. Porate: Absent (0), Present (1)
7. Zonosulcate: Absent (0), Present (1)
Aperture Position
8. Polar: Absent (0), Present (1)
9. Equatorial: Absent (0), Present (1)
10. Merdonial: Absent (0), Present (1)
Symmetry
11. Symmetric: Absent (0), Present (1)
12. Asymmetric: Absent (0), Present (1)
Shape
13. Elliptic (narrow or broad):Absent (0), Present (1)
14. Triangular: Absent (0), Present (1)
15. Oblong: Absent (0), Present (1)
16. Subrounded or circular: Absent (0), Present (1)
166
Tectum
17. Tectate including semitectate: Absent (0), Present (1)
18. Intectate: Absent (0), Present (1)
19. Supratectal process (gemmae): Absent (0), Present (1)
Exine pattern
20. Reticulate: (including sub-reticulate): Absent (0), Present (1)
21. Perforate: Absent (0), Present (1)
22. Punctate: Absent (0), Present (1)
23. Spinulose: Absent (0), Present (1)
24. Rugulate: Absent (0), Present (1)
25. Verrucose: Absent (0), Present (1)
Sexine Pattern
26. Muri simpli- baculate: Absent (0), Present (1)
27. Muri simpli-duplibaculate: Absent (0), Present (1)
28. Muri simpli-triplibaculate: Absent (0), Present (1)
29. Muri simpli-multibaculate: Absent (0), Present (1)
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Table 3.2 Data matrix ofthe taxa representing family Palmae scored for 29 characters present in the table 3.1
Name of taxa 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 27 28 29
Areca 23.85 41.35 1.6 1 1 0 0 1 0 0 1 1 1 0 0 1 1 0 0 1 0 0 0 0 0 1 0 0 0 Dypsis 21.0 36.70 2 1 1 0 0 1 0 0 0 1 1 0 0 0 1 0 0 1 0 0 1 0 0 0 0 0 0 Ptychosperma 28.85 41.41 1.6 1 1 0 0 1 0 0 0 1 1 0 1 0 1 0 0 1 1 0 0 1 0 0 0 0 0 Wodyetia 23.85 41.35 1.75 1 1 0 0 1 0 0 0 1 1 0 1 0 1 0 0 1 1 0 0 1 0 0 0 0 0 Cocus 26.80 54.60 1.5 1/3 1 0 0 1 0 0 0 1 1 1 0 0 1 0 0 0 0 1 0 0 0 1 0 0 0 Elaeis 34.30 21.16 1.1 3 1 0 0 1 0 0 0 1 0 1 0 0 1 0 0 1 0 0 0 1 0 0 0 0 0 Roystonea 46.25 44.20 1.5 1 1 0 0 1 0 0 0 1 1 0 0 0 1 0 0 0 0 1 0 1 0 0 0 0 0 Calamus 25.75 28.00 1.8 2 1 0 0 1 0 0 0 1 1 0 0 0 1 0 0 1 0 0 0 0 0 0 1 0 0 Bismarkia 40 28 1.0 1 1 0 0 1 0 0 0 1 1 0 0 0 1 0 0 1 1 0 0 1 0 1 0 0 0 Borassus 40.8 36.5 2.5 1 1 0 0 1 0 0 0 1 1 0 0 1 1 0 1 1 1 0 0 1 1 1 0 0 0 Hyphaene 36 34.3 2.0 1 1 0 0 1 0 0 0 1 1 0 0 1 1 0 1 1 1 0 0 1 1 1 0 0 0 Caryota 20.3 25.4 2.0 1 1 0 0 1 0 0 0 1 1 0 1 0 1 0 0 1 0 0 0 0 0 1 0 0 0 Nannorrhops 29.8 31.4 1.5 1 1 0 0 1 0 0 0 1 1 0 0 1 1 0 0 1 0 0 0 1 0 0 1 0 0 Brahea 16.6 27.3 1.2 1 1 0 0 1 0 0 1 1 1 0 0 1 1 0 0 1 1 0 0 0 0 1 0 0 0 Livistonia 19.7 28.9 1.9 1 1 0 0 1 0 0 0 1 1 0 0 0 1 0 0 1 1 0 0 0 0 0 0 1 0 Pritchardia 21.0 30.4 2.2 1 1 0 0 1 0 0 0 1 1 0 0 0 1 0 0 1 1 0 0 0 0 0 1 0 0 Washingtonia 36.7 33.4 1.5 1 1 0 0 1 0 0 0 1 1 0 1 0 1 0 0 1 0 0 0 0 1 0 1 0 0 Trachycarpus 17.2 25.31 1.8 1 1 0 0 1 0 0 0 1 1 0 0 1 1 0 0 1 0 0 0 0 0 0 0 1 0 Rhapis 18.85 30.22 1.9 1 1 0 0 1 0 0 0 1 1 0 0 1 1 0 0 1 0 0 0 0 0 0 1 0 0 Phoenix 22.22 24.45 1 1 1 0 0 1 0 0 1 1 1 0 0 1 1 0 0 1 1 0 0 0 1 1 0 0 0 Sabal 19.9 33.7 2.0 1 1 0 0 1 0 0 0 1 1 0 1 0 1 0 0 1 0 0 0 1 1 0 0 0 1
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Fig. 3.1 Dendrogram obtained by Ward’s cluster analysis, showing three groups of species belong to family Palmae, separated on the basis of different pollen characteristics
Key to abbreviations: Ac = Areca, Wo = Wodyetia,Pt = Ptychosperma,Dy = Dypsis, Ry = Roystonea,Co = Cocus, El = Elaeis,Ca = Calamus,Nr = Nannorrhops,Cr = Caryota,Ph = Phoenix, Br = Brahea,Tr = Trachycarpus,Lv = Livistonia, Pr = Pritchardia, Rh = Rhapis,Sb = Sabal,Bi = Bismarkia, Bo= Borassus, Hy = Hyphaene,and Ws = Washingtonia
169
CLUSTER ANALYSIS OF THE FAMILY PALMAE
Pollen morphological data fully support the taxonomicdelimitation of palm flora at various levels.The main dichotomy divides all the studied taxa into three broad group viz., group-1, group-2 and group-3. The details of characters and characters state are listed in Table-3.1 and 3.2
Group-1
This group accommodates six genera belonging to three different tribes of the subfamily Arecoideae viz., ArecaL., DypsisNoronnha ex Mart., Ptychosperma Labill.and
Wodeytia A.K. Irvine. (tribe Areceae), CocusL. (tribe Cocoseae) and Roystonea O.F.
Cook (tribe Roystoneae). These genera linked together due to the almost same pollen size, aperture and combination of exine pattern. However, the genus Cocus L., does not form cluster with any genera mainly due to the difference in the pollen size (i.e., the longest axis up to 60-70 µm) and exine pattern (punctate) whereas in the other genera the longest axis of the pollen is up to 50µm and exine pattern is reticulate, spinulose, regulate and vermiculate.Furthermore, ArecaL.,Ptychosperma Labill.and Wodeytia A.K. Irvine show close association with each other as they share more pollen morphological characters. The grouping among these genera is only due to slight difference in pollen size and exine features (Table 3.2, Fig. 3.1 & 3.2). The other genera DypsisNoronnha ex
Mart.,and Roystonea O.F. Cook forms a common clade and distinct from other due to the exine pattern. In the former genus exine is spinulose where as in the later exine is vermiculate and punctate. In this group pollen morphology show marked variation and therefore divided into different pollen types such as Areca-type, Dypsis-type,
Ptychosperma-type, Cocus-type and Roystonea-type. Sowunmi (1972) also placed these
170 genera in to different groups on the basis of exine pattern. In terms of gross morphology, the present results fully support the placement of the above said genera into different tribes and subtribes.
Group-2
This is the largest and diverse group accommodating 10 genera of three subfamilies viz., Arecoideae (1 genus), Calamoideae (1 genus) and Coryphoideae (8 genera) as they have more or less same pollen size (i.e., in range of 20 -30 µm). The dendrogram based on pollen morphology further divided the group-2 into two distinct subgroupsviz., group-2a and group-2b (Fig. 3.1).
Group 2a
This diversesubgroup comprises 3 genera representing the different subfamilies viz., ElaiesJacq. (Arecoideae),Calamus L.(Calamoideae) and Nannorrhops H. Wendl.
(Coryphoideae). The genus ElaiesJacq., of the subfamily Arecoideae is not only different from the member of this group but also different from the entire palm flora due to thepresence of trichotomosulcate pollenwhereas in the remaining genera the pollen is monosulcate excluding the genus Calamus L. On the other hand, in the cluster analysis of the subfamily Arecoideae (Fig., 3.2), the genus seemsElaiesJacq., to be partially linked with Cocus L. and thus divided into two groups on the basis of pollen shape, aperture and exine pattern (Table 2.1). Similarly, in terms of gross morphology, these two genera belong to the same tribe Cocoseae but placed in different subtribes (Table 1.3 & 1.4). The genus Calamus L. (subfamily Calamoideae)is represented by only one species and different from all the studied palm taxa due to the presence of disulcate pollen (Table
2.2). Therefore, both these genera do not show any affiliation with other genera and form
171 a separate clade. Thanikaimoni (1971), Sowunmi (1968, 1972), Ferguson and Harley
(1993) also treated these genera in separate groups on the basis of pollen morphological characters. The remaining genus Nannorrhops H.Wendl., of the subfamily coryphoideaepartially linked with Calamus L. due to only the same pollen size otherwise the genus is also different from the above said genus due to the presence of monosulcate pollen.
The present result clearly indicates that all these genera are not closely related with each other in terms of both pollen morphological as well as gross morphological characters.
Hence, the pollen morphology plays a significant role at the generic or even at the higher level (i.e., subfamily and tribes).
Group 2b
This subgroup is represented by 7 genera of three different tribes (subfamily
Coryphoideae) such as Phoenix L. (Phoeniceae),Caryota L. (Caryotaae), Brahea Mart. ex
Endl.,Livistonia R.Br., Pritchardia Seeman and H.Wendl., RhapisL. and TrachycarpusH.
Wendl. (Trachycarpeae).All these genera share almost all the pollen characters but differ in pollen size, exine and sexine features (Table 2.3, 3.6; Fig. 3.1, 3.3).The following two genera Phoenix L. and Caryota L., shows linkage as they share all the pollen morphological characters excluding the exine pattern (Table 3.6; Fig.3.3). While, grouping of the subfamily Coryphoideae both these genera seems to be linked. The other
3 members viz., LivistoniaR.Br.,PritchardiaSeeman and H.Wendl.and Rhaphis L., appear in the same group as they show more or less similarity in pollen morphological characters. In the present studies, the first two genera have been placed in the same pollen type (i.e., Brahea-type) as they have perforate pollen and in the cluster analysis of the
172 subfamily Coryphoideae they also seems to be more linked. Whereas, the genus Rhapis
L., is different from these on the basis of reticulate exine.The remaining two genera viz.,
BraheaMart. ex Endl. and Trachycarpus H. Wendl., appear in the same clade as they have same pollen size but their exine pattern is different. In the former genus exine is densely perforate whereas in the later exine is finely reticulate.Although, in terms of gross morphology they are not closely related, as they belonging to two different tribes.
Sowunmi (1972) placed all the coryphoid palms in one group and mentioned the homogenous nature of pollen i.e no pollen type have been recognized. Similarly,
Dransfield et al., (1993) also describe that all the coryphoid palm have uniform pollen.
As far as gross morphological characters are concerned they are almost closely related excluding the genera Phoenix L. (Phoeniceae) andCaryota L. (Caryotaae), as indicated by their inclusion of different subtribes.
Group 3
The group includes 6 taxa viz., Bismarkia Hildebrandt and H. Wendl., Borassus
L.,Hyphaene J. Gaertn., Sabal Adanson,WashingtoniaH.Wendl., of the subfamily
Coryphoideae. The results of cluster analysis show that the genus BismarkiaHildebrandt and H. Wendl. does not form cluster with any species but appear to be linked with
Hyphaene L. However, these two genera representing the same subtribe Hyphaenineae
(tribe Borosseae) as they have similar small, globose, pedicellate female flowers and globule fruit with basal stigmatic remains (Dransfield & Uhl, 1998;Dransfield et.al,
2008), but palynologically both are different mainly on the basis of exine feature thus treated in to two distinct pollen-types i.e., Bismarkia-type and Borassus-type. On the other hand, the genus Borassus L. shows close association with Hyphaene Geaertner, as
173 they share similar pollen shape, aperture and even the exine pattern. Otherwise, they are not closely related, indicated by their inclusion in separate subtribes such as Borassus
L.(Latannineae) and Hyphaene Gaertner (Hyphaeninae)of the Borossoid palm as the former genus have large and sessile female, symmetric fruits with apical stigmatic remains where as in the later genus the flower and fruits are different. Thus this relationship among these three genera clearly shows the lack of association between pollen morphology and gross morphology. The present findings are in accordance with some earlier reports such as Thanikaimoni, 1970a; Sowunmi, 1972; Ferguson, 1986;
Ferguson et al., 1987.
The genus WashingtoniaH. Wendl., of the tribe Coryphoideae is placed in basal position and show association due to sharing more or less same pollen characters (pollen aperture type and size).The remaining genus Sabal Adanson of the tribe Sabaleae is quite distinct from the other members of this group and does not form any clade but moderately linked as they share same pollen characters in some extant except the exine features.
The present observations clearly indicate that pollen morphological characters play a significant role for the delimitation of taxa at all the levels (subfamily, tribes, subtribes and generic) except the specific level.
174
Table 3.3 List of characters, scored for cluster analysis for the taxa of the subfamily Arecoideae in Table 3.4.
Character description 1. Length (µm) 2. Breadth (µm) 3. Exine Thickness (µm) 4. Aperture number Aperture type 5. Sulcate: Absent (0), Present (1) 6. Porate: Absent (0), Present (1) 7. Zonosulcate: Absent (0), Present (1) Aperture Position 8. Polar: Absent (0), Present (1) 9 Equatorial: Absent (0), Present (1) 10. Merdonial: Absent (0), Present (1) Symmetry 11. Symmetric: Absent (0), Present (1) 12. Asymmetric: Absent (0), Present (1) Tectum 13. Tectate including semitectate: Absent (0), Present (1) 14. Intectate: Absent (0), Present (1) 15. Supratectal process: Absent (0), Present (1) Exine pattern 16. Reticulate (including sub-reticulate): Absent (0), Present (1) 17. Perforate 18. Punctate: Absent (0), Present (1) 19. Vermiculate:Absent (0), Present (1) 20. Rugulate:Absent (0), Present (1) 21. Spinulose:Absent (0), Present (1)
175
Table 3.4 Data matrix of the taxa representing subfamily Arecoideae scored for 21 characters present in the table 3.3
Name of taxa 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21
Areca catechu. 23.85 41.35 1.65 1 1 0 0 1 0 0 0 1 1 0 0 1 0 0 0 0 0
Dypsis lutescence 21.0 36.70 2 1 1 0 0 1 0 0 0 1 1 0 0 0 1 0 0 0 1
Ptychosperma elegans 23.85 41.35 1.6 1 1 0 0 1 0 0 0 1 1 0 0 1 1 0 0 1 0
P. macarthuri 33.85 41.47 1.6 1 1 0 0 1 0 0 0 1 1 0 0 1 1 0 0 1 0
Wodyetiabifurcata 23.85 41.35 1.75 1 1 0 0 1 0 0 0 1 1 0 0 1 1 0 0 1 0
Cocus nucifera 26.80 54.60 1.5 1 1 0 0 1 0 0 0 1 1 0 0 0 0 1 0 0 0
Elaeis guineensis 34.30 21.16 1.1 3 1 0 0 1 0 0 0 1 1 0 0 1 0 0 1 0 0
Roystonea regia 46.25 44.20 1.5 1 1 0 0 1 0 0 0 1 1 0 0 0 0 1 1 0 0
176
Fig. 3.2 Dendrogram obtained by Ward’s cluster analysis, showing two groups of species belong to family Arecoiadeae, separated on the basis of different pollen characteristics.
Key to abbreviations: Ac = Areca catechu, Dl = Dypsis lutescence, Pe = Ptychosperma elegans, Pm = Ptychospermamacarthuri, Wb = Wodyetiabifurcata, Cn = Cocus nucifera,Eg = Elaeis guineensis, Rr = Roystonea regia.
177
CLUSTER ANALYSIS OF THE SUBFAMILY ARECOIDEAE
Cluster analysis of the 8 OTUs of the subfamily Arecoideaeclearly indicates the presence of two main group’sviz., Group-1 and Group -2 (Fig. 3.2). Details of characters and character state are listed in Table 3.3 and 3.4. Pollen morphological data support the taxonomic delimitation of Arecoid palm at various levels. For instance, Group-1 and
Group-2 is mainly distinguished on the basis frui
Group-1: This group includes the following species such as Areca catechu L., Dypsis lutescence(H.Wendl.) Dransf. & Beentje, Ptychosperma elegans(R.Br.)
BlumeandPtychosperma macarthuri (H.Wendl.) H. Wendl. ex Hook. f. and Wodeytia bifurcate Irvine.
Among these, Wodeytia bifurcata Irvine appears to be closely linked with Areca catechu L., whereas both the species of Ptychosperma Labill falls in between these two taxa as they share almost same pollen characters i.e., aperture type and exine pattern. The differences are found in their pollen size and also in the exine pattern. For instance, in
Areca catechu L. exine is finely reticulate while in Ptychosperma Labill and Wodeytia
Irvine, exine is reticulate or perforate to rugulate (Table 2.1& 3.4).
On the otherhand, Dypsis lutescence(H.Wendl.) Dransf. & Beentje does not form cluster to any species of the Arecoideae due to the difference in their pollen size and exine pattern but partially linked with the other species of Group-1 as it share same pollen aperture(Table 2.1& 3.4).
Group-2: This group comprised of 3 species viz., Cocus nucifera L., Elaeis guineensis
Jacq. and Roystonearegia (H.B. & K.) Cook.
178
The cluster analysis clearly indicate thatElaeis guineensis Jacq., is quite distinct from other taxa of the Group-2 and other members of the Arecoideae mainly due to aperture type, pollen shape and exine pattern.The other two species viz., Cocus nucifera
L., and Roystonearegia (H.B. & K.) Cook.,forms a cluster as they share same aperture type and show differences in their pollen size and exine pattern (Table 2.3) and thus classified in to two groups.
The cluster analysis of the subfamily Arecoideae clearly support the grouping based on pollen morphological data. Similarly, various workers such as Mahabale (1967),
Thanikaimoni (1971), Sowunmi (1968, 1972), Kedves (1980), Ferguson (1986),
Ferguson and Harley (1993), Dransfield et al., (1998, 2008) separate arecoid palms mainly on the basis of pollen shape, aperture type and exine ornamentation.
The present result clearly shows that species of the subfamily Arecoideae or even the species representing the other genera of the entire family have remarkably similar pollen.Thus the pollen morphological data usually support the delimation of taxa at all the levels tribal, subtribal and generic level except the specific level.
179
Table 3.5. List of characters, scored for cluster analysis for the taxa of the subfamily Corphoideaeae in table 3.6.
Character description
1. Length (µm)
2. Breadth (µm)
3. Exine Thickness (µm)
4. Aperture number
Symmetry
11. Symmetric: Absent (0), Present (1)
12. Asymmetric: Absent (0), Present (1)
Tectum
13. Tectate including semitectate: Absent (0), Present (1)
14. Intectate: Absent (0), Present (1)
15. Supratectal process (gemmae): Absent (0), Present (1)
Exine pattern
16. Reticulate: (including sub-reticulate): Absent (0), Present (1)
17. Verrucose:Absent (0), Present (1)
18. Perforate:Absent (0), Present (1)
19. Punctate or foeveolate: Absent (0), Present (1)
20. Rugulate:Absent (0), Present (1)
Sexine Pattern
21. Muri simpli- baculate:Absent (0), Present (1)
22. Muri simpli-duplibaculate:Absent (0), Present (1)
23. Muri simpli-triplibaculate:Absent (0), Present (1)
24. Muri simpli-multibaculate:Absent (0), Present (1)
180
Table 3.6 Data matrix of the taxa representing subfamily Coryphoideae scored for 24 characters present in the table 3.5
Name of taxa 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 Bismarkia nobilis 40 28 1.0 1 1 0 0 1 0 0 0 1 1 0 0 1 0 0 0 0 1 0 0 0
Borassus flabellifer 40.8 36.5 2.5 1 1 0 0 1 0 0 0 1 1 0 1 1 1 0 0 0 1 0 0 0 Hyphaene thebaica 36 34.3 2.0 1 1 0 0 1 0 0 0 1 1 0 1 1 1 0 0 0 1 0 0 0 Caryota urens 20.3 25.4 2.0 1 1 0 0 1 0 0 0 1 1 0 0 1 0 0 0 0 1 0 0 0 Nannorrhops ritichiana 29.8 31.4 1.5 1 1 0 0 1 0 0 0 1 1 0 0 1 0 0 0 0 0 1 0 0 Brahea brandegeei 16.6 27.3 1.2 1 1 0 0 1 0 0 0 1 1 0 0 1 0 1 0 0 1 0 0 0 Livistonia chinensis 19.7 28.9 1.9 1 1 0 0 1 0 0 0 1 1 0 0 1 0 1 0 0 0 0 1 0 Pritchardia beccariana 21.0 30.4 2.2 1 1 0 0 1 0 0 0 1 1 0 0 1 0 1 0 0 0 1 0 0 Washingtonia filifera 36.2 32.9 1.1 1 1 0 0 1 0 0 0 1 1 0 0 1 0 0 0 0 0 1 0 0
Washingtonia robusta 37.2 33.9 1.2 1 1 0 0 1 0 0 0 1 1 0 0 1 0 0 0 0 0 1 0 0 Trachycarpus fortunei 17.2 25.31 1.8 1 1 0 0 1 0 0 0 1 1 0 0 1 0 0 0 0 0 0 1 0 Rhapis excelsa 16.6 30.0 1.9 1 1 0 0 1 0 0 0 1 1 0 0 1 0 0 0 0 0 0 0 0
Phoenix.dactylifera 21.8 24.6 1.0 1 1 0 0 1 0 0 0 1 1 0 0 1 0 1 0 0 1 0 0 0 P. loureirii 18.7 23.6 1.1 1 1 0 0 1 0 0 0 1 1 0 0 1 0 1 0 0 1 0 0 0 P. sylvestris 23.9 25.2 1.0 1 1 0 0 1 0 0 0 1 1 0 0 1 0 1 0 0 1 0 0 0
P .robelli 24.5 24.4 1.0 1 1 0 0 1 0 0 0 1 1 0 0 1 0 1 0 0 1 0 0 0 Sabal minor 19.9 33.7 2.0 1 1 0 0 1 0 0 0 1 1 0 0 1 0 0 0 0 0 0 0 1
181
Fig. 3.3 Dendrogram obtained by Ward’s cluster analysis, showing two groups of species belong to sub family Coryphoideae, separated on the basis of different pollen characteristics.
Key to abbreviations: Bn = Bismarkia nobilis, Bf = Borassus flabellifer, Ht = Hyphaene thebaica, Cu = Caryota urens, Nr = Nannorrhops ritichiana, Bb = Brahea brandegeei, Lc = Livistonia chinensis, Pb = Pritchardia beccariana, Wf = Washingtonia filiformis, Wr = Washingtonia robusta, Tf = Trachycarpus fortune, Re = Rhapis excels, Pd = Phoenix dactylifera, Pl = P. loureirii, Ps = P. sylvestris, Pr = P .robelli and Sm = Sabal minor.
182
CLUSTER ANALYSIS OF SUBFAMILY CORYPHOIDEAE
The dendogram of the 17 OTUs based on pollen morphological characters clearly indicates the existence of two major groups mainly based on the size of pollen
(Fig 3.3). In the group-1, pollen is more than 30 µm in length whereas in the group-2, pollen length is less than 30 µm. Details of characters and character state are given in
Table 3.5 and 3.6.
GROUP-1
The first group includes 6 taxa viz., Bismarkia nobilisHildebrandt & H. Wendl.,
Borassus flabelliferL.,Hyphaene thebaica (L.) Mart., Nannorrhops ritichiana (Griff.) H.
Wendl., Washingtoniafilifera (L. Linden) H. Wendl. and Washingtonia robusta H.
Wendl. Among these species, Bismarkia nobilisHildebrandt & H. Wendl., partially linked withHyphaene thebaica (L.) Mart., as they sharelittle pollen morphological characters (Table 2.1; Fig. 3.3). However, they have been treated in the subtribe
Hyphaeninae of the Borossoid palm because of strong gross morphological characters.
The present grouping is accordance with Dransfield & Uhl, (1998), Ferguson et. al,
(1987), Dransfield et.al, (2008). It is interesting to note that the pollen of Hyphaene thebaica (L.) Mart., is remarkably similar to Borassus flabelliferL.,in shape, aperture and even the exine pattern as compare to Bismarkia nobilisHildebrandt & H. Wendl.
(Thanikaimoni, 1970a; Sowunmi, 1972; Ferguson, 1986; Ferguson et al., 1987).The cluster also shows the close association of both these genera (Table 2.1; Fig. 3.3).
Nevertheless, in terms of gross morphological characters they are not closely related, as indicated by their inclusion of different subtribes. The species of the genus
WashingtoniaH. Wendl., is placed in basal position and show slight association due to
183 sharing more or less same pollen characters (aperture type and size).The remaining species Nannorrhops ritichiana (Griff.) H. Wendl., is quite distinct from the other members of this group and does not form any clade but moderately linked as they share same pollen characters in some extant.
GROUP-2
This group is characterized by the combination of pollen characters such as pollen less than 30 µm in length, monosulcate, mostly elliptic in polar view and accommodates 11 taxa which are further divided in to two subgroups viz., group 2a and group 2b (Fig. 3.3).
GROUP 2a
This subgroup is distinct from the subgroup 2b mainly on the basis of pollen size, exine and sexine pattern. The following two genera Phoenix L. (with 4 species) and
Caryota L. (with a single species) representing this group and show linkage as they share all the pollen morphological characters excluding the exine pattern (Table 3.6;
Fig.3.3). Furthermore, all the taxa representing the genus Phoenix L. arise from the common point which proves strong affinities between them as they share more gross morphological and pollen morphological characters. The presentfindings are also supported by the previous findings of Sowunmi (1972) as she described the genus
Phoenix L.,is homogenous on the basis of pollen characters.
GROUP 2b
Out of 6 species representing this subgroup, Brahea brandegeei (Purpus)
H.E.Moore, Rhapis excelsa (Thunb.) Henry ex Rehderand Trachycarpus fortunei(Hook.
184 f.) H. Wendl., show close linkage and occur in similar clade whereas Livistonia chinensis (N.J. Jacq.) R. Br. ex Mart., Pritchardia beccariana Rock and Sabal minor
(N.J. Jacq.) Persons, also appear in the same group. Both these clades share almost all the pollen characters but differ in the exine and sexine features and slight difference have also been found in their pollen size (Table 2.3, 3.6; Fig. 3.3). Sowunmi (1972) placed all the coryphoid palms in one group and mentioned the homogenous nature of pollen i.e no pollen type have been recognized. Similarly, Dransfield et al., (1993) also describe that all the coryphoid palm have uniform pollen. As far as gross morphological characters are concerned they are almost closely related excluding the genus Sabal
Adanson, as indicated by their inclusion of different subtribes such as the tribe Sabaleae.
185
GENERAL DISCUSSION
The present comparative study is the first attempt to provide detail information on the pollen morphology including systematic revision of the family Palmae from Pakistan and Kashmir.Previously,Stewart (1972) in his “Annotated Catalogue of Vascular Plants of
W. Pakistan and Kashmir” reported only 8 genera of the family Palmae from the area under consideration. However, in the present study two species recognized by R.R. Stewart are reduced to synonym (viz., Phoenix humilis Royle now treated as the synonym of Phoenix loureii) and the other species placed under different genus (viz., Trachycarpusexcelsa
(Thunb.) H. Wendl. now treated under Rhapis excelsa (Thunb.) Henry ex
Rehder.Furthermore,Malik (1984) reported 18 species belonging to 16 genera from Pakistan and Kashmir. However, in the present study a total of 27 species belonging to 21 genera distributed in 10 tribes and 3 subfamilies of Palmae are included from study area.Among these subfamilies, the Coryphoideae is represented by 13 genera and 19 species distributed in 6 tribes followed by the subfamiliy Arecoideae i.e., represented by 7 genera and 8 species distributed in 3 tribes and the last subfamily Clamoideae is represented by a single genus with one species of the tibe Calameae (Table 1.1 & 1.3).
Palms are extremely distinctive at the family level but, inside the family, theirdiversity in the morphological characters is probably greater than among the monocotyledonous family (Uhl & Dransfield, 1987).At the microscopic level this diversity is also found in the pollen morphology by different workers such asErdtman (1952),
Sowunmi (1968, 1972), Thanikaimoni (1966, 1970), Kedves (1980); Dransfield et al.,
(1990), Ferguson (1986), Ferguson & Harley (1993), Harley (1990, 1999a), Harley & Baker
(2001), Harley & Dransfield (2003). The basic and most common pollen type in palms, as
186 elsewhere in the monocotyledons (Dahlgran & Clifford, 1982), is monosulcate (i.e., with one aperture in the shape of sulcus), located at the distal pole of the pollen. The simple tectate, monosulcate pollen present in all the subfamilies except Nypoideae (Harley 1990,
Harley & Baker, 2001). In the subfamily Calamoideae monosulcate pollen is rare or deficient whereas the remaining subfamilies show a high percentage of monosulcy type of pollen.
The present study also observeddiversity in pollen morphological characters of the family Palmae, particularly in the subfamily Arecoideae (Table 2.1). In general, the size of pollen ranges from 15 to 50 µm in length and 20 to 60 (-70) µm in breadth. The polar outline of pollen is frequently elliptical, sometimes rounded triangular, or circular. The three types of aperture have been found viz., monosulcate, disulcate and trichotomosulcate, the monosulcate is common among the palms.A vast variety have been examined in exine pattern such as punctate, reticulate, vermiculate, verrucose, pilate, spinulose, or combination of some of these patterns – however, the highest proportion is the reticulate type. Generally, the species of the following genera such as, Phoenix L., Washingtonia Irvine, Rhapis,
Ptychosperma Seemann & H. Wendl. ex H.Wendl.have similar aperture and exine pattern.These species are hardly classified on the basis of their pollen size or they are placed in the same group. However, within the studied genera they may have different pollen characters and can be recognized easily mainly on the basis aperture and exine pattern.The subfamily Arecoideae shows a great diversity in the aperture type (i.e., monosulcate and trichotomosulcate) and exine pattern (finely reticulate, punctate, spinulose, rugulate and vermiculate) followed by the subfamily Coryphoideae have The unigeneric tribes viz.,
Phoenicoideae and Roystoneae each have uniform pollen types.
187
DISCUSSION ON SIGNIFICANT POLLEN CHARACTERS
A palynological study of 27 palm taxa reveal the diversity of pollen in the family
Palmae with regard to shape, size, exine pattern, aperture form and number (Table 2.1-
2.3; Fig 2.1-2.10). Many workers such as Sowunmi (1972), Ferguson et al., (1983,
1987); Dransfield and Uhl (1998),Furness and Rudall (2001); Dransfield et al., (2008),
Ambwani and Kumar (2009), Dowe (2010), Rashid and Perveen (2014) have usedvarious pollen charactersto find out additional micro-morphological characters and also to strengthn the taxonomic status of the taxa representing the family
Palmae.However, the following pollen characters proved quite important of the palm flora.
Size: Thestudied taxa show a wide range in the pollen size of palmae. For instance, in
Phoenix L., pollen are small (15 x 26 µm) but SabalAdanson and CocusL. have quite large pollen i.e., 20 x 40 µm and 30 x 60µm respectively.The present results are in accordance with Mahabalé (1967), Sowunmi (1972) and Dransfield et al., (1990) results.
Furthermore, the size of the longest axis of pollen ranging from 18-60 µm, averaging 30-
50 µm. Kupriankova (1948), Erdtman (1952), Mahabalé (1967), Sowunmi (1968 &
1972) and Harley (1990) also observed the more or less same result (i.e., the longest axis of ranging from 18µ-60µ (-80µ), rarely 50µ or averaging30µ to 50µ).
Shape: In monosulcate pollen the shape as seen in polar view, is frequently elliptical
(either symmetrical or asymmetrical) or subcircular but in one species pollen is triangular viz., Elaeis guineensis Jacq.Whereas, in equatorial view the pollen is kidney- shaped in transverse position and more or less boat or lemon shaped in longitudinal
188 position (2.1-2.10).Punt and Wessels Boer (1966a) considered that the pollen shape depended on the aperture type. Thus monocolpate pollen is oblong or oval (usually asymmetric) in polar axis and kidney-shaped in equatorial view and trichotomocolpate grains were said to be regularly triangular. Howerver, the results obtain from the present study regarding the shapes of pollen agree with earlier investigations such asCranwell
(1953),Nair and Sharma (1963),Mahabalé (1967), Sowunmi (1968 & 1972), Kedves
(1980), Ferguson (1986), Harley (1990), Dransfield et al., (1990, 2008).
Aperture:The predominant aperture type of pollen is colpus. The colpi, as examined in the present study is usually long i.e., they extended from one pole to another, and coincided with the longest axis of pollen.Most of the species have monosulcate pollen
(i.e., 25 out of 27) show the highest percentage (92.5%).While disulcate and trichotomosulcate pollen is found in Calamus tenuis Roxb. and Elaeis guineensis
Jacq.,respectively. A few investigators had recorded the occurrence of monocolpate pollen in the Palmae (Gassner, 1941; Erdtman, 1952; Punt and Wessels Boer, 1966a, b;
Thanikaimoni, 1970; Mahabalé, 1967; Sowunmi, 1968 & 1972; Kedves, 1980;
Ferguson, 1986; Harley, 1990; Dransfield & Uhl, 1987; Dransfield et al., 1990, 2008;
Harley & Baker, 2001; William & Zona, 2006).This monosulcate aperture is considered to be a primitive character pointed out by Wodehouse (1935, 1936), Kuprianova (1948),
Meeuse (1965)Thanikaimoni (1966) and traces its origin to the Palaeozoic
Cordaitales.On the other hand, occurrence of trichotomosulcate pollen in the family
Palmae has been observed by Erdtman, 1952;Cranwell, 1953; Nair and Sharma, 1963;
Punt and Wessels Boer, 1966a and Thanikaimoni, 1966.
189
Exine Pattern: The palm flora of study area show a great diversity in exine pattern such as reticulate, punctate (in Cocus nucifera L. and RoystoneaO.F.Cook), rugulate (in
Bismarkia nobilis, Wodeytia A. K. Irvine, PtychospermaLabill), vermiculate (in Elaies
Jacq. and RoystoneaO.F.Cook), spinose (in Dypsis sp. and Caryota urens L.) verrucose- reticulate (in Borassus flabellifer L. and Hyphaene thebeica) or perforate to reticulate (in
Brahea Mart ex Endl., Livistonia R. Brown, PritchardiaSeemann & H. Wendl. ex
H.Wendl, Phoenix L.).The most predominant pattern being the reticulate type i.e., about
81.5%(Table 2.1-2.3).Kuprianova (1948) gave only three types of exine pattern (viz., tubercular, reticulate and negatively reticulate) in the family Palmae. The muri is supported by one to many rows of bacula. Erdtman (1952) gave reticulate and tegillate type provided with warts, spines or spinules. Cranwell (1953) mention only two types of exine i.e., reticulate and spiny. Punt and Wessels Boer (1966a) also reported two type viz., perforated (punctatum) and vermiculate tectum.Thanikaimoni (1966) studied the exine pattern of 608 species and the main pattern found by him were reticulate or foveolate, scabrate, spinulose, and fossulate (vermiculate). Sowunmi (1972) investigated the pollen of 350 species and found 11.5% species were intectate, but with sexinous process (i.e., exine spinulose), c. 82.5% species had semitectate exine (i.e., reticulate, vermiculate or negatively reticulate) and 6% of the species was pertectate (i.e., finely perforated). In most of the cases the present findings are in accordance with the earlier reports as discussed above (also see table 2.1 -2.3; Fig. 2.1-2.10).
Exine thickness: According to present results, exine thickness rangingfrom 1.0 to 3.0
µm.A highest proportion of the range between 1.0-2.0µm (i.e., about 24 species out of
27 having 88.88%).
190
191
COMPARISIONS OF POLLEN WITHIN THE SUBFAMILIES
The family Palmae is represented by 5 species viz., Arecoideae, Calamoideae,
Coryphoideae, Ceroxyloideae and Nyphoideae (Table 1.2; Fig., 1.2), of which first three subfamilies has representatives in study area.
The pollen of Arecoideae is probably the most varied, although there are many genera with the generalized monosulcate type (Harley, 1990) but sometimes associated with trichotomosulcate pollen grain (a triradiate variant of monosulcate) as recorded in the literature (Harley, 1999a). Similarly, in the present study out of 8 taxa, the 7 species have monosulcate and a single taxonviz., Elaeis Jacq.,is represented by trichotomosulcate pollen. Some earlier workers such as Thanikaimoni, 1970a; Sowunmi,
1972; Ferguson, 1986 have found trichotomosulcate grains in some others Arecoid also,
Pinanga, Elaeis, Bactris, Astrocaryum, and Acrocomia.
The pollen morphology of Coryphoideae is fairly uniformrepresented by monosulcate, perforate or finely reticulate pollen excluding the Borossoid palm,a much generalized type similar to that found in some other monocotyledonous families. The monosulcate pollen type with perforate or reticulate tectum occurs throughout the entire family (Harley, 1990) and is considered primitive in view of its appearance in the early
Crateceous fossil records (Doyle, 1973). Moreover, many of the pollen representing the subfamily Coryphoideae are remarkably uniform to early Cretaceous pollen grain illustrated by Walker and Walker (1985; 1986). On the other hand, more variation have been found in the pollen of the tribe Borassease (Sowunmi, 1972; Ferguson et. al., 1986) as compare to the other tribes such as Caryotae, Corypheae, Phoeniceae, Trachycarpeae and Sabaleae.
192
Within the subfamily Calamoideae, a single species viz., Calamus tenuis L.is known from Pakistan. The pollen of Calamus L. is distinct from all other palm flora by having disculcate pollen. Similarly, in terms of gross morphological characters the genus is not closely related with any other palm member by the presence of stiff, shiny and imbricate scales on their fruits. However,the pollen literature shows a huge range of variation within the subfamily Calamoideae in pollen characters such as ornamentation, apertures, and in exine stratification. For example in the Calamoideae there is a intectate gemmate and spinose pollen in Salacca, Daemonorops and Korthalsia as well as tectate psilate or sparsely perforate tectate pollen in the two former genera (Ferguson, 1986).
Extended sulcate, dicolpate, and diporate pollen occurs in Calamus,Salacca,
Daemonorops,and Korthalsia (Thanikaimoni, 1970a; Frederiksen et al., 1985; Ferguson,
1986). Supratectal spines and Supratectal gemmae occurs in the Retispatha and
Calamus. Mauritia, Mauritiella, and Lepidocaryum have spinose pollen, the spines being characteristically sunk into the foot layer (Sowunmi, 1972; Ferguson, 1986).
Although, the present investigations clearly indicates that pollen morphology of the family Palmae fully support the delimitation of generic level or even at the higher level also. However, at the specific level pollen data are not helpful because species of the representative genus have uniform pollen. In some cases, pollen morphology shows a remarkable correlation with gross morphology (as in the subfamily Calamoideae) on the other hand they also show a lack of correlation as in the tribe Cocoseae (Cocus L. and Elaeis Jacq.) and Borosseae (Bismarkia Hildebrandt. & H. Wendl.; Borassus L. and
Hyphaene J. Gaertn.).
193
GLOSSARY OF POLLEN
Amb The outline of pollen grain or spore seen in polar
view(Erdtman, 1952)
Aperture A specialized region of the sporoderm, that is thinner than the
remainder of the sporoderm and generally differs
inornamentation and/or in structure(Erdtman, 1947)
Apocolpium A region at the pole of a zonocolpate pollen grain delimited by
lines connecting the apices of the colpi(Erdtman, 1952)
Apolar Describing pollen and spores without distinct polarity(Erdtman,
1952)
Areola A feature of ornamentation in which the sexine/ectexine is
composed of circular or polygonal areas separated bygrooves
which form a negative reticulum(Erdtman,1947)
Atectate Describing pollen grains that have an exine with little or no
internal structure(Walker and Doyle, 1975)
Baculum A cylindrical, free standing exine element more than 1μm in
length and less than this in diameter(Potonié, 1934)
Brochus A brochus consists of one lumen of a reticulum and half of the
width of the adjacent muri(Erdtman, 1952)
194
Colpus An elongated, aperture with a length/breadth ratio greater than
2(Erdtman, 1943)
Columella A rod-like element of the sexine/ectexine, either supporting a
tectum (Iversen and Troels-Smith, 1950)
Compound aperture An aperture with two or more components that are situated in
more than one aperture wall layer(Erdtman, 1969)
Dicolpate, dicolporate Pollen grains with two ectocolpi, two compound or diporate apertures two pores(Iversen and Troels-Smith, 1950)
Distal face That part of a palynomorph that faces outwards the centre of the
tetrad, between equator and distal pole (Erdtman, 1952)
Distal pole The centre of the surface of the distal face(Erdtman, 1952)
Disulcate Describing pollen grains with sulci arranged in pairsequatorial
disulcate: with opposing, equatorially arranged sulci and distal
disulcate with paired sulci lying parallel to the long axis of the
pollen grain on the distal face (Harley, 1998)
Dyad Term for two microspores (pollen grains or spores) united as a
dispersal unit.
Ectexine The outer part of the exine, which stains positively with basic
fuchsin in optical microscopy and has higher electron density in
conventionally prepared TEM sections(sensu Fægri, 1956)
195
Ectoaperture An aperture in the outer layer of the sporoderm(Van Campo,
1958)
Ectoaperture An aperture in the inner layer of the sporoderm, often the inner
aperture of a compound aperture (Van Campo, 1958)
Equator The dividing line between the distal and proximal faces of a
pollen grain or spore (Wodehouse, 1935)
Equatorial diameter A line, lying in the equatorial plane, perpendicular to the polar
axis and passing through it(Erdtman, 1943)
Equatorial outline General description of the equator when a pollen grain is seen
in polar view.
Equatorial plane The plane perpendicular to the polar axis and lying midway
between the poles(Fægri and Iversen, 1950)
Equatorial view The view of a pollen grain or spore where the equatorialplane is
directed towards the observer (Erdtman, 1943)
Etectate Describing pollen grains interpreted as having lost their
tectumduring evolutionary development.
Eu- A prefix for true.
Eurypalynous Describing plant taxa characterized by possession of a great
diversity of palynomorphs(Erdtman, 1952)
196
Exine The outer layer of the wall of a palynomorph, which is highly
resistant to strong acids and bases,and is composed primarily of
sporopollenin(Fritzsche, 1837)
Foot layer The inner layer of the ectexine(Fægri, 1956)
Foveola A feature of ornamentation consisting of more or less rounded
depressions or lumina more than 1μm in diameter (Erdtman,
1952)
Furrow A common word for an elongate aperture. Examples: colpus,
sulcus.
Granulum A very small and rounded element of thesexine/ectexine that is
less than 1μm in all directions (sensu Erdtman,1952)
Heteropolar Describing pollen or spores in which the distal and proximal
faces of the exine are different either in shape, ornamentation or
apertural system(Erdtman, 1952)
In- A prefix used to emphasis the absence of a feature.
Inaperturate Describing a pollen grain or spore without apertures (Iversen
and Troels-Smith, 1950)
Intectate Describing pollen grains without a tectum, but with
sculpturing(Iversen and Troels-Smith, 1950)
197
Intine The innermost of the major layers of the pollen grain wall
underlying the exine and bordering the surface of the cytoplasm
(Fritzsche, 1837)
Isopolar Describing a pollen grain or spore in which the proximal and
distal faces of the exine are alike(Erdtman, 1947)
Lumen The space enclosed by the muri (Potonié, 1934)
Meridional Describing longitudinal features on the surface of a pollen grain
or spore which run along line perpendicular to the
equator(Fægri and Iversen, 1950)
Meso- A prefix meaning middle
Mesocolpium The area of a pollen grain surface delimited by lines between
the apices of adjacent colpi or the margins of adjacent
pores(Erdtman, 1952)
Mono- A prefix for one
Monoaperturate Describing a pollen grain or spore with a single aperture.
Examples: monocolpate (Iversen and Troels-Smith, 1950),
monoporate (Iversen and Troels-Smith, 1950) monosulcate
(Erdtman, 1952).
Murus (pl. Muri) A ridge that is part of the ornamentation and, for example,
separatesthe lumina in a reticulate pollen grain or the striae in
198
striate pollen (Erdtman, 1943)
Nexine The inner, non-sculptured part of the exine which lies below
the sexine(Erdtman, 1952)
Oblate Describing the shape of a pollen grain or spore in which the
polar axis is shorter than the equatorial diameter (Erdtman, 1943)
Oblate spheroidal Describing the shape of a pollen grain or spore in which the
ratio between the polar axis and the equatorial diameter is 0.88-
1.00(Erdtman, 1952)
Operculum (pl. A distinctly delimited sexine, ectexine structure which covers opercula) part of anectoapertureand which is completely isolated from the
rest of the sexine (Wodehouse, 1935)
Ornamentation A general term that is useful for describing the organization
offeatures(Potonié, 1934)
P/E ratio The ratio of the length of the polar axis (P) to the
equatorial diameter (E)(Erdtman, 1943)
Pilum (pl. pila) A sexine element, usually standing directly on the nexine,
consistingof a rod-like part (columella) and a swollen apical
part (Erdtman, 1952)
Polar axis The straight line between the distal and proximal poles of a
pollen grain or spore (Wodehouse, 1935)
199
Polar area index (PAI) (Iversen and Troels-Smith, 1950)
Polar view A view of a pollen grain or spore in which the polar axis
is directed towards the observer(Erdtman, 1943)
Pole Either of the two extremities of the polar axis (Wodehouse,
1935)
Pollen The microgametophyte of seed plants, developed from
the microspore (Linnaeus, 1751)
Pollen class An artificial grouping of pollen grains that share a distinctive
character, or suite of characters. Such classes are useful in
identification keys and may be subdivided into more restrictive
categories, pollen types and pollen groups (Fægri and Iversen,
1950)
Pore A general term, applied in palynology to a circular orelliptic
aperture with a length/breadth ratio less than 2(Jackson, 1928;
Wodehouse, 1935)
Punctum A rounded or elongate tectal perforation, less than 1μm in
length ordiameter(Erdtman, 1952)
Reticulum A network-like pattern consisting of lumina or other spaces
wider than µm bordered by elements narrower than the lumina
(Praglowski and Punt, 1973)
200
Semitectum A partially discontinuous tectum in which the tectal
perforations are equal to or wider than the muri and usually
larger than 1μm in diameter(Fægri and Iversen, 1964)
Sexine The outer, sculptured layer of the exine, which lies above the
nexine(Erdtman, 1952)
Shape classes (pl) Categories of pollen and spore shape based on the relations
between polar axis (P) and equatorial diameter (E) (Erdtman,
1943)
Spheroidal Describing the shape of a pollen grain or spore in which the
polar axis and the equatorial diameter are approximately
equal(Erdtman, 1943)
Spine A general word, applied in palynology to long and tapering
pointed elements, exceeding 1μm(Erdtman, 1952)
Spinule Small spines, less than 3μm in length(Erdtman, 1952)
Sporoderm The entire wall of a pollen grain or spore (Bischoff, 1833)
Stenopalynous Describing plant taxa characterized by only a slight variation in
their palynomorphs (Erdtman, 1952)
Sulculus An elongated latitudinal ectoaperture not situated at a pole
(Erdtman, 1952)
201
Sulcus An elongated latitudinal ectoaperture situated at the distalor
proximal pole of a pollen grain (Erdtman, 1952)
Tectum The layer of sexine, which forms a roof over the columellae,
granules or other infratectal elements (Fægri and Iversen, 1950)
Tetrad A general term for a group of four united pollen grains or
spores,either as a dispersal unit or as a developmental stage.
Trichotomocolpate Describing a pollen grain with a three-armed colpus(Erdtman,
1945 a)
Trichotomosulcate Describing a pollen grain with a three-armed sulcus (Erdtman,
1952)
Ulcus A rounded ectoaperture situated at the distal or proximal pole of
a pollen grain(Erdtman, 1952)
Vermiculate A general descriptive term used to describe winding features
(Kosanke, 1950; Harris, 1955)
Verruca A wart-like sexine element, more than 1μm wide, that is
broader than it is high and is not constricted at the base (Iversen
and Troels-Smith, 1950)
Verrucose Synonym of verrucate (Erdtman, 1969)
202
APPENDIX-I
Areca catechu Shagufta s.n.(KUH); Kamal Akhtaer 666, 780(KUH); Kamal Akhtar s.n. (KUH).
Bismarkia nobilis Abid A. Rashid 03 (KUH);Abid A. Rashid s.n. (KUH);Shaukat 34 (KUH).
Borassus flabellifera Abid A. Rashid 117 (KUH); Abid A. Rashid 65 (KUH).
Brahea Abid A. Rashid 137 (KUH); Abid A. Rashid 139 (KUH).
Calamus tenuis Abid A. Rashid 20 (KUH); Abid A. Rashid s.n. (KUH).
Caryota urens Kamal Akhtar 665 (KUH). Kamal Akhtar 667 (KUH). Kamal Akhtar 844 (KUH).
Cocus nucifera A. A. Qureshi s.n. (KUH); Abid A. Rashid 17 (KUH); Abid A. Rashid 19 (KUH);Abid A. Rashid 30 (KUH).
Dypis lutescens Abid A. Rashid 456 (KUH); Abid A. Rashid 459 (KUH).
Elaeis guineensis Abid A. Rashid 35 (KUH); Abid A. Rashid 40 (KUH).
Hyphaena thebaica Abid A. Rashid 22 (KUH); Roohi Bano 58 (KUH).
Livistonia chinensis Kamal Akhtar 663 (KUH); Kamal Akhtar 785 (KUH); S. Nazimuddin & Sultan Abedin 1200 (KUH).
Nannorhops ritichiana Abdul Ghaffoor & Saood Umer1731 (KUH); Abdul Ghaffoor & Steve M. Goodman 4419 (KUH); S. I. Ali, S. A. Farooqi & Sultan Abedin 1443 (KUH).
Phoenix dactylifera Kamal Akhtar 664 (KUH); Kamal Akhtar 837
203
(KUH);Kamal Akhtar 846 (KUH).
Phoenix loureirii A. Rashid s.n. (RAW); A. Rashid 26959 (RAW).
Phoenix robelenii Abid A. Rashid 256 (KUH); Abid A. Rashid 257 (KUH).
Phoenix sylvestris Asifullah Khan 02 (KUH);S. Nazimuddin, S.Abedin & Hameedullah 975 (KUH).
Ptychosperma elegans Saood Omer623 (KUH); Saood Omer 645 (KUH).
Ptychosperma macarthuri Rubina Saleem 356 (KUH); Saood Umer 321 (KUH).
Rhapis excelsa Abid A. Rashid 132 (KUH); Abid A. Rashid134 (KUH).
Rhapis multifida Abid A. Rashid 136 (KUH).
Roystonea regia Abid A. Rashid 16 (KUH);Abid A. Rashid &Adil 89, 33 (KUH); Abid A. Rashid & Shaukat 110 (KUH).
Sabal minor Abid A. Rasheed 364 (KUH); Abid A. Rasheed 368 (KUH).
Trachycarpus fortunei Kamal Akhter 47 (KUH);Abid A. Rasheed 77 (KUH)
Washingtonia filifera Abid A. Rashid 66 (KUH); Abid A. Rashid 84 (KUH); Kamal Akhtar 839 (KUH).
Washingtonia robusta Kamal Akhtar 847 (KUH).
Wodeytia Abid A. Rasheed 357 (KUH); Abid A. Rasheed 387 (KUH).
204
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INDEX OF TAXA
Areca L. 20
Areca catechuL. 22
Bismarkia Hildebrandt & H. Wendl. 49
Bismarkia nobilis (Hildebrandt & H. Wendl.) 49
Medemia nobilis (Hildebrandt & H. Wendl.) Gall. 49
Borassus L. 50
Borassus flabellifer L. 51
Borassus flabelliformis Murr. 51
Brahea Mart. ex Endl. 60
Brahea brandegeei (Purpus) H. E. Moore 60
Calamus L. 40
Calamus tenuis Roxb. 41
Cocos L. 33
Cocus nucifera L. 34
Caryota L. 54
Caryota urensL. 56
Dypsis Noronha ex Mart. 23
ChrysalidocarpusH. Wendl. 23
Neodypsis Baillon 23
Dypsis lutescens (H.Wendl.) 24
Chrysalidocarpus baronii Becc. 24
Chrysalidocarpus lutescens H.Wendl. 24
225
Elaeis Jacq. 36
Elaeis guineensisJacq. 36
Hyphaene J. Gaertn. 47
Hyphaene thebaica (L.) Mart. 47
Corypha thebaica L. 47
Livistonia R.Br. 51
Livistonia chinensis(N. J. Jacquin) R. Br. ex Mart. 62
Latania chinensis N.J. Jacquin 62
Livistonia mauritima Wall. Ex Voigt. 62
Nannorrhops H. Wendl. 57
Nannorrhops ritichiana (Griff.) H. Wendl 58
Chamaerops ritichiana Griff 58
Phoenix L. 77
Phoenix canarieensis Hort. ex Chabaud 79
Phoenix dactylifera L. 80
Phoenix loureirii Kunth 81
Phoenix humilis Royle. 81
Phoenix ouseleyanaGriff. 81
Phoenix roebelenii O Brien Gard. 83
Phoenix sylvestris L.(Roxb). 85
Elate sylvestris L. 85
Pritchardia Seemann and H. Wendl. ex H.Wendl 65
Pritchardia beccariana Rock 65
226
Pritchardia beccariana varr giffardiana Becc. 65
Ptychosperma Labill 25
Ptychosperma elegans(R.Br.) Blume 26
Seaforthia elegans R.Br. 26
Saguaster elegans R.Br. 26
Ptychosperma macarthuri (H.Wendl.) H. Wendl. ex Hook.f. 28
Kentia macarthuri H. Wendl. 28
Raphis L. 72
Rhapis excelsa (Thunb.) Henry ex Rehder 73
Chamaerops excelsa Thunb. 73
Trachycarpus excelsus(Thunb.) H. Wendl. 73
Rhapis flabelliformis L’ Her. ex Ait. 73
Rhapis multifida Burret 76
Roystonea O.F.Cook 31
Roystonea regia (H.B. & K.) O.F. Cook 32
Oreodoxa regia Kunth, Nov. 32
Oenocarpus regius (Kunth) Spreng. 32
Sabal Adanson 86
Sabal minor (N.J. Jacq.) Persons 87
Sabal minimum Nut 87
Corypha minor N.J. Jacq. 87
Trachycarpus H. Wendl. 71
Trachycarpus fortunei (Hook. f.) H. Wendl 71
227
Chaemaerops fortune Hook. 72
Washingtonia H. Wendl. 66
Washingtonia filifera (L. Linden) H. Wendl. 67
Pritchardia filiferaL. Linden 67
Pritchardia filamentosa Fenzi 67
Washingtonia filamentosa (Fenzi) O. Ktze. 67
Neowashingtonia filamentosa (Fenzi) Sudw. 67
Washingtonia robusta H. Wendl. 70
Neowashingtonia robusta (H. Wendl.) A. Heller 70
Wodyetia A.K.Irvine 29
Wodyetia bifurcate A.K.Irvine 30
228