i. leaf anatomy as an aid in the classification of living gymnosperms
TRANSCRIPT
I. Leaf Anatomy as an aid in the Classificationof Living Gymnosperms
Introduction
The gymnosperms are of scientific Importance to Indian Forestry, since they are well represented in the Flora of India, particularly in the Himalaya where there are extensive regions of coniferous forests from the stand-point of resin, yield and timber production which together constitute a potential source of revenue (Raizada and Sahni, 1958). The first time we have a glimpse of them in the fossil records, the remains can be conveniently divided into two broad groups, namely, the Cycadophyta and Conifero- phyta. The compound leaf is an outstanding feature of the Cycadophyta. The former group included relatively small plants with unbranched stems and pinnate leaves, thick cortex, thin wood and large pith whereas the latter group had large trees with profusely branched stems and simple leaves, thin cortex, thick wood and small pith.
From the phylogenetic view-point, the gymnosperms are much more ancient than the anglosperms. The gymnosperms represent the most primitive type of seedplants (Datta, 1973). They were abundant In thelj^te Paleozoic and apparently had evolved from psilop-
sldan stock in the Late Devonian, perhaps independently of the ferns and fern-allies. The Cycadofilicales and Cordaitales made their greatest display before the Triassic. The Paleozoic plants gave rise to the wealth of forms Including the Bennettitaies and Cycadales which literally covered the landscape during the Meso-
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zoic. The time of maximum development of the ancient gymnosperms coincided almost exactly with the age of giant dinosaurs. It is believed that young leaves and seeds of the extinct Cycads were consumed by certain hervivorous dinosaurs (Chamberlain, 1934),
The extant gymnosperms, numbering about 70 genera with 725 species and belonging to the Coniferales, Cycadales and Gnetales,
are considered to be remnants of an once-large and diversified group (Datta, 1973). Despite the loss of much splendour, they now comprise a major forest type of primarily temperate regions of both southern and northern hemispheres. A case in point is the taiga (coniferous-spruce, fir forest) of North America and Eurasia which has surpassed the geographical extent of any vegetation in the world. The recent cycads symbolise veritable living fossils and are confined to limited areas of the tropics or subtropics. Ginkgo biloba is one of the wonders of ,the plant world," It is, in the words of Seward (1938), "cne of a small company of living plants which Illustrates continuity and exceptional power of endurance in a changing environment'*. It exists in the wild state in south-eastern China and as an ornamental plant in many parts of the temperate zone now-a-days. The,living members of the Gnetales contain many puzzling.features of anatomy, organography and reproduction. As their evolutionary status is a speculative topic, they form a somewhat isolated group.
Though the anatomical characters of vegetative parts have long been extensively used in the study of cycads, conifers and
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the other vascular groups (Durrell, 1915; Lamb, 1923$ Poole, 1923$ Harlow, 1931$ Marco, 1931 & 1939$ Sutherland, 1933$ Bannan, 1934$ Fulling, 1934$ Peirce, 1935$ Abbe and Crafts, 1939$ Florin, 1940, 1951, 1958 & 1963$ Cross, 1941$ Johnson, 1943$ Orr, 1944$ Buchholz and Gray, 1948$ Johnson and Thomas, 1963$ Konar, 1963$ Griffith, 1971$ Rao, 1972$ Ziepski, 1972; Alfieri and Evert, 1973$ Behnke and Paliwal, 1973$ Ghouse, 1973 & 1974$ Grill, 1973$ denOuter and Toes, 1974$ Litvintseva, 1974$ Miller, 1974$ Paliwal et al., 1974$ Eremin, 1975$ Gaussen and Waltz, 1975$ Kausik, 1975 & 1976$ Kausik and Bhattacharya, 1977$ Lotova, 1975$ Baig and Tranqullini, 1976), there is no elaborate effort to classify living gymnosperms on the basis of their leaf anatomy. Keeping this object in view, thepresent work was designed to evolve a simplethe easy identification of the Indian specimens based on the microscopic study of the readily-available leaf material. Moreover, an attempt has been to present a concise account of the anatomical features of foliage leaves of the different families, genera and
■ species, as information is hard to get from the relevant literature regarding most of the Indian forms. This, it is hoped, will provide an easy means to identify gymnospermous plants by leaf specimens alone, since very often complete material is not forthcoming and makes identification a difficult task. An explanatory note on anatomical features selected has been provided$ this is indicative of the uses to which such features can be put.
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Materials and Methods
In the present study, leaf material of the following species were collected from Forest Besearch Institute (Dehra Dun), Lloyd Botanic Garden (Darjeeling) and Indian Botanic Garden (Howrah)s
Cycadaceae
GinkgoaceaeAraucariaceae
s 1) Cyca3 rumphii Miq.2) £• revoluta Thunb.3) Encephalartos villosus__Lem.4) Zamia, angustifolia Facq.
: 5) Ginkgo biloba Linn.8 6) Agathis loranthifolia Salisb.
7) Araucaria bldwillii Hook.8) A. columnaris Hook.9) A. cunnlnghamii Sweet.
Cupressaeeae 8 10)11)12)13)14)15)16) 17)
. 18)
Cephalotaxaeeae % 19)20)
Gallitris robusta B. Br,Chamaecyparls pisifera Sieb. & Zucc. Cupressus cashmeriana Boyle ex Carriers C, torulosa D. Don Juniperus recurva Buch.-Ham.J. wallichiana Hook.Thu.ia occidentalis Linn.T. orientalis Linn.Tfau.1opsi3 dolabrata Sieb. & Zucc.
Cephalotaxus drupacea Sieb. & Zucc.C. griffithii Hook.
10 -
Pinaceae
Podoearpaceae
TaxaceaeTaxodiaceae
s 21) Abies spectabilis Spach.22) Cedrus libani Laudon23) Picea smithlana Boiss24) . Pinus densiflora Sieb. & Zucc.25) P. gerardlana Wall, ex Lamb26) P, kesiva, Royle27) P. roxburghii Sarg.28) £,' sylvestris Linn.29) P. wallichiana A. B. Jacks30) Tsuga dumosa Eiehl.
s‘ 31) Podocarpus macrophyllus D. Don32) P. neriifolius D. Don33) P, sinensis Teijsm. & Binn.
8 34) Taxus baccata Linn.: 35) CrvPtomeria .laponica D. Don
36) Cunninghamia sinensis R, Br.37) Metasequoia glyptostroboides Hu et Cheng38) Taxodlum distichua Rich.
A number of leaves from each source of collection were used to avoid any discrepancy, in the. observations originating fromnatural variation. Unless otherwise stated, only fully matured leaves were selected for sectioning. In each case, the sections were chosen from various regions of the same leaf to depict anoverall picture of its anatomical characters. Free-hand sectionswere made and stained with Bismarck Brown and Safranin and Fast-
- IX -green. Finally, the sections were mounted in Canada balsam.
Herbarium specimens were revived by boiling in water until they came to their natural form and size. Some of the dry specimens were preserved in 10% glycerine at room temperature for 2 days, while most of the fresh specimens were fixed in FAA before sectioning.
In studying the stomata! type and stomatal frequency, a. somewhat different procedure was adopted. Those leaf materials, fixed in FAA overnight, were washed in tap water. Then cut pieces from the middle and from two ends of a leaf were boiled in 95% alcohol for 15-30 min. Again, the same material was boiled in a mixture of 1:1 5% NaOH and 95% ethyl alcohol for additional 15-20 min. Next, the material was washed in tap water and finally rinsed with distilled water. Later on, the washed material was kept in 4% sodium hypochlorite solution for a single night or two nights. Those materials, provided with thick cuticle, took at least two nights to be completely cleaned. Finally, these materials weremounted and observed under the compound microscope (Olympus^__brand). _ ]
vCl.
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Fig. I. Fig. II.
Fig. Ill,
Fig. IV.
Fig. V.
Fig. VI.
Fig. VII.
Fig. VIII,
Key to tissues of living gymnosperms.Diagrams of the transverse sections of the leaves of Cycads j (1) Cycas rumphii: (2) C, revoluta:(3) Bncenhalartos villosus: (4) Zamla floridana.
Diagrams of the transverse sections of the leaves of ginkgos and conifers i (5) Ginkgo bllobat (6) Asathis loranthifolias (7) Araucaria bidwillii;(8) A. columnaris: (9) A. cunningha.mil.
Diagrams of the transverse sections of the leaves of conifers j (10) Callltris robustat (11) Chamae- cyparis pisiferas (12) Cupressus cashmeriana;(13) C. torulosa: (14) Juniperus recurva.
Diagrams of the transverse sections of the leaves of conifers s (15) Juniperus wallichiana: (16) Thuja occidentaliss (17) T. orientalis: (18) Thu.lopsis dolabrata
Diagrams of the transverse sections of the leaves of conifers i (19) Cephalotaaius drupacea:(20) C. griffithil;
Diagramms of the transverse sections of the leaves of conifers s (21) Abies spectabilist (22) Cedrus libani; (23) Picea smithiana: (24) Pinus densi. flora? (25) P. gerardiana.
Diagrams of the transverse sections of the leaves of conifers s (26) P. kesiyat (27) P. roxburghii:
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(28) P. sylvestrls: (29) P. wallichianas (30) Tsuga dumosa.
Fig. IX. Diagrams of the transverse sections of the leaves of conifers s (31) Podocarpus macrophvllus : (32)P- neriifolius: (33) P. sinensis: (34) Tax us baccata.
Fig. X. Diagrams of the transverse sections of the leaves of conifers : (35) Cryptomeria iaponicat (36) Cunnlnghamia sinensis: (37) Metasequoia glypto- stroboides: (38) Taxodium distichum.
Fig. XI, Camera lucida drawings of stomatal apparatus s(1) Cycas rumphli - Paracytic; (2) C. revoluta - Actinocytic; (3) Encephalartos vlllosus - Hypocytic; (4) Zamla angustlfolia - Paracytic.
Fig. XII. Camera lucida drawings of stomata! apparatus j(5) Ginkgo biloba - Anomocytic; (6) Agathis loran-thlfolia - Staurocytic; (7) Araucaria bidwillii - Anisocytie; (8) J\,. columnaris - Anisocytie; (9) JL. cunnlnghamil - Anomocytlc.
Fig. XIII. Camera lucida drawings of stomatal apparatus jCallltris robusta - Paracytic5 (11) Chamaecyparis
pisifera - Anomocytlc; (12) Cupressus cashmeriana - Anomocytic; (13) C. torulosa - Anomocytlc; (14) Juniperus recurva - Anomocytlc.
Fig. XIV. Camera lucida drawings of stomatal apparatus s (15) J. wallichiana - Anomocytic; (16) Thu.la occldentalis - Anomocytlc and Paracytic; (17)Thu.la orientails - Actinocytic; (18) Thu.1opsl3 dolabrata - Anomocytic.
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Fig. XV. Camera lucida drawings of stomatal apparatus s (19) Cenhalotaxus drupacea - Hexacytic; (20) C. griffithii - Paracytic.
Fig. XVI, Camera lucida drawings of stomatal apparatus 5 (21) Abies spectabilis - Pericytic; (22) Cedrus libani - Tetracytic; (23) Picea smith!ana • Tetracytic; (24) Pinus densiflora - Graminaceous;(25) P. gerardiana - Semiactinocytic.
Fig. XVII, Camera lucida drawings of stomatal apparatus 5(26) Pinus kesiva - Paracytic; (27) P. roxburghiiGraminaceous; (28) P. recurva - Staurocytic and Paracytic (complex type); (29) P. wallichiana - Paracytic; (30) Tsuea dumosa- Paracytic.
Fig.XVIII. Camera lucida drawings of stomatal apparatus 5(31) Podocarpus macrophvllus - Staurocytic;(32) P. neriifoliU3 - Paracytic; (33) P. sinensisParacytic; (34) Taxus baccata - Paracytic and Hexacytic.
nFig, X3X. Camera lucida drawings of stomatal apparatus s"
(35) Cryptomeria .laponica - Actinocytic or Cyclo- cytic; (36) Cunninghamia sinensis - Anomocytic;(37) Metaseauoia glyptostroboides • Anomocytic;(38) Taxodlum distichum - Anomocytic.
S = subsidiary cell; g = guard cell; a = epidermal cell; us = undifferentiated subsidiary cell; st = storm.
Fiq. I
KEY TO TISSUES
* EZS- EPIDERMIS
'yWtk~ SCLERENCHYMA
? miH- PALISADE
r~-h- SPONGY PARENCHYMA
£y~ PLICATE PARENCHYMA
f- XYLEM>/ PHLOEM
RESIN DUCT
TRANSFUSION TISSUE
ENDODERMIS
PERI CYCLE
AIRSPACE
STOMATA
©0— ALBUMINOUS CELL
CO— SCLEROTIC CELL
ooo— PARENCHYMA
;'YV>y\
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Fig. nr
Fig, 12
Fiq.3E
Fiq.
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Fig.IK
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Fiq.301
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Fiq. 22:
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Fi« WIT
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5
Explanation of Anatomical Characters
General formThe conifers hear simple leaves having various forms, e. g.
acicular, linear, scaly or subulate (Laubenfels, 1953; Esau, 1965; Foster and Gifford, 1974), Impending on the leaf-form, they assume various shapes in cross sections. These are laterally winged, vertically winged, semi-lunar, semi-circular or triangular.
EpidermisVariations in epidermal structure have been put to extensive
uses for the purpose of constructing phylogeny and taxonomy in a variety of taxa including the gymnosperms (Florin, 1940, 1951 & 1963; Tateoka, 1957; Davies, 1959; Metcalfe, 1960). The Indian forms are no exception to this rule, since the epidermis of these plants vary in their wall structure and composition.
The cuticle may be thick and thin and lumen broad or narrow.
StomataThe mode of stomatal development and their spatial relation
ships have been taken into account while dealing with the classification and phylogeny of both angiosperms and gymnosperms (Florin 1940, 1951 & 1953; Metcalfe and Chalk, 1950; Pant, 1965). In the present work, the distribution, frequency and type of stomata have been utilised in separating the various genera and species of cycads and conifers.
35 -
Distribution, frequency and type of stomata
In general, the stomata are situated in parallel rows, running largely on either side of the midrib and forming vertical bands. However, they are located without any regular arrangement in Cedrus. Crvptomeria. Cupressus and Thuja. The stomata are amphistomatic, I. e. distributed on all sides as in Callitris. Cedrus. Crvptomeria. Plcea and Pinus. whereas the stomata are hypostomatic, i. e. usually confined to abaxial-epidermis as in all cycads, Ginkgo. Abies. Cephalotaxus. Cunninghamia. Metaseauoia. Podocarpus. Taxodium. Taxus and Tsuga and rarely hyperstomatic, i. e. confined to the adaxial epidermis as in Chamaecyparis. Cupressus. Thu.1a and Thu.lopsis. In the same genus, it is amphistomatic in Juniperus recurva and hypostomatic in J. walllchiana.
The stomata may be entirely absent in the lower epidermis, e. g. Cupressus cashmeriana. Thu.ia and Thu.lopsis. The stomata are lacking in the upper epidermis as in all cycads, Ginkgo. Abies, Podocarpus. Metaseauoia. Taxus. Tsuga and Pinus densiflora. In the rest of genera and species studied, the stomata are present in lower epidermis mainly.
The stomatal type differs considerably in the Cycadaeeaa and Pinaceae. The stomata is anomocytie in Ginkgo. Juniperus and Taxodiaceae (except Crvptomeria where it Is actinocytic). The stomatal type is paracytic In the Cupressaceae, Podocarpus and Tsuga, However, Thuja occidentalis has a complex stomatal type, being both anomocytie and paracytic.
36
HypodermisThe development and distribution of hypodermal cells vary
in different gymnosperms, This tissue is entirely absent in Ginkgo. Callitris. Cenhalotaxus. Metaseauoia. Taxodium. Taxus and Tsuga, The hypodermis is both less developed and well developed in Cycas. Cupressus. Pinus and Podocarpus. This tissue is less developed in Agathis and Chamaecvparis. It is well developed in Cedrus. Cryptomeria. Cunninghamii. Juniper us. Picea, Thu.ia and Thu.lopsis.
In Araucaria bidwillii. Pinus gerardiana and Podocarpus sinensis, there are distinct bundles of sclerified cells. In the remaining gymnosperms, where hypodermis is present, the bundles are uniformly distributed in one or more layers.
Me sophy11
In most gymnosperms studied, the mesophyll is differentiated into palisade and spongy parenchyma. The tissue is less differentiated in Zamla and Cryptomeria. In the same genus, it is undifferentiated in Thu.ia occidentalis and less differentiated in Zamia and Cryptomeria. In the same genus, it is undifferentiated in Thuja occidentalis and less differentiated in T. orientalis. Undifferentiated mesophyll Is observed in Araucaria columnaris and A. cunninghamii as well as -Chamaecvparis. While the mesophyll cells are undifferentiated in Cedrus, Picea and Pinusf they are sometimes unique in possessing internal ridges on the walls which
- 37 -project into the cell lumen (plicate parenchyma). In these forms, masophyll cells are either vertically aligned or arranged in horizontal layers demarcated from each other by intercellular spaces.
Besin ductsThey are absent in Encenhalartos. Zamia and Taxus. In the
same genus, they are absent in Cycas revoluta and present in C. rumphii. Single resin duct is seen in Chamaecyparis. Crypto- meria, Cupressus. Juniperus. Taxodlum and Tsuga. There are more than two resin ducts in Callitris robusta as well as in Podocarpus macrophvlla and just two in P, nerlifolius. Two resin ducts are noteworthy in Abies. Araucaria. Cedrus and Picea. In the rest of the gymnosperms, more than two resin ducts are observed. Epithelial cells are associated with resin ducts and the number of such cells vary from 6-7 in Pinus kesiva to 33 in Abies specta- bilis and 37 in Cryptomeria japonica.
Though the taxonomic value of resin ducts is indisputable (Phillips, 1948; Ghouse, 1969), their position and number vary in the members of the same family, e. g. Araucariaceae, Cupressa- ceae, Pinaceae and Podoearpaceae and even in the same species, e. g. Pinus roxburghii. As to their number, the resin ducts do not always run continuously from one end of the leaf to the other and rather form elongated cysts or sacs (Marco, 1939).Endodermis
The endodermis is ill-defined or indistinct in the Cycada-
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ceae, Araucariaceae and Cephalotaxaceae as well as Charoaecyparis. Cupressus. Juniperus. Meta sequoia. Podocarpus. Taxodiua, Thujop- sis and Tsuga. It is slightly distinct in Thu.la and is even more distinct in Callitris robusta. The endodermal layer is well differentiated in most members of the Pinaceae and certain other gymnosperms. The cells of this layer are thick-walled, sometimes containing starch and exhibit prominent casparian strips particularly when they are young (Strasburger, 1891; Esau, 1965).
Transfusion tissueThe ontogeny, the evolution and function of the transfusion
tissue is controversial (Abbema, 1934), However, this special tissue system is universally present in gymnosperms (Griffith, 1957; Esau, 1965) and varies in arrangement and quantity in different species (Gathy, 1954; Lederer, 1955). In view of spatial relationships of the transfusion tissue with the vascular tissues and variations in constituent elements, this tissue system has been profitably used in the classification of gymnosperilous leaves (Buchholz and Gray, 1948).
The transfusion tissue occurs all round the vascular bundle or bundles in Bneenhalartos. Zamia. Ginkgo. Agathis, Araucaria. Pinus, Plcea and Chamaecyparis. This tissue system has been noted to occur in Cycas. Abies. Callitris. Cedrus. Cephalotaxus, Crypto- meria. Cunninghamii. Metasequoia. Tax us and Taxodium to the left and right of the vascular bundles. In the same genus, the trans-
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fusion tissue occupies lateral position in Podocarpus macrophvlla and P. neriifolius and is all round the vascular bundle in P. sinensis. The transfusion tissue is both lateral to and all round the vascular bundle in Juniper us wallichiana and Thu.la orien tails. While the transfusion tissue is lateral to vascular bundle in the two species of Cupressus, it is specialised in forming wing-like extensions towards the leaf margin in,C. torulosa. ,
Accessory transfusion tissue
An additional tissue system, which is composed of non-living tracheidal cells and horizontally-elongated elements of parenchyma, runs from the vascular area to the wings. It has been identified as accessory transfusion’tissue'by Griffith (1957). Among gymnos- perms studied, this tissue is found in the species of Cycas and Podocarpus. As a matter of fact, the taxonomy of Podocarpus has been revised on the basis of this tissue (Buchholz and Gray, 1948).
Vascular bundles
Though cambial activity is common in the leaves of gymnos- perms in general and in the conifers in particular (Strasburger, 1891; Esau, 1960), their value in . leaf taxonomy has not been ascertained.
In general, vascular bundles remain either one or two and rarely more in the coniferous leaves. In cross section, the phloem is situated towards the abaxial side and xylem towards the adaxial side. In all cases, the xylem is endarch. As the number and posi-
. 40tion of vascular strands remain constant at the mature stage, this'character has been employed in separating the forms of the Cycadaceae as well as the species of Abies, Araucaria. Chamae- cyParis and Tsuga* Another interesting point is bundle-sheath extension, being present in Zamla and absent in Cycas and Encephalartos. '
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Concise Account of Individual Family and Genera
Cycadaceae
Chamberlain (1919) recognised nine genera which he accommodated in the single family Cycadaceae under the order Cyea- dales. According to Pilger and Melchior (1954), the Cycadaceae is divided into five subfamilies s Cycadioideae, Stangeroideae, Bowenioideae, Dioonoideae, Zamioide&e. Johnson (1959) added the genus Lepidozamia and set up three separate families of cycads t the Cycadaceae and Stangeriaceae,.each represented by a single genus and the Zamiaceae which includes the rest of the eight genera. The only genus found wild in India is Cycas. though Encephalartos and Zamia are sometimes observed in cultivation in Botanic Gardens (Raizada, and Sahni, 1958),
Anatomically, the members of the Cycadaceae (Fig. II &Plates 1-4) studied possess thick-walled epidermis, well-developed hypodermis, indistinct endodermis and largely no resin ducts.While accessory transfusion tissue is present in Cycas, both Bncephalartos and Zamia are devoid of them. Eneephalartos is characterised by the presence of hypocytic stomata and the absence of bundle-sheath extension. However, the stomata is paracytic in Zamia where the bundle-sheath extension is present.
The species differentiation of Cycas. merely on the basis of anatomical characters, is easy (Table 1).
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GinkeoaceaeMi monotypie genus on which the family Glnkgoaceae is foun
ded, differing from the Coniferales, hy its wide flattened leaves, and by fertilization effected by motile spermatozoids1* (Raizada and Sahai, 1958),
Ginkgo blloba is widely cultivated as a park specimen or street tree in many temperate regions of the world, (Foster and Gifford, 1974). One of the remarkable morphological features of this species is the foliage leaf which consists of a petiole and a fan-rshaped dichotomously veined lamina.
Anatomically, the lamina (Fig, III, Plate 5 & Table 1) bears undulated thin-walled epidermis with distinct endodermis and plenty of resin ducts.
Araucariaceae
Evergreen, dioecious or monoecious trees. Branches whorled. Leaves coriaceous, spirally arranged, scaly or subulate, broad- ovate. (Agathis) to stiff (Araucaria). Micrpsporophylls with 5-20 pendant microsporangia; ovuliferous scale fused with its bracts and producing a single median ovule; pollen-grains without wings.
Of the two genera, Araucaria is more commonly grown in India. A. bidwillii. A. columnaris and A. cunninghamii have been introduced in gardens and parks of eastern India (Datta, 1973).
Anatomically, the leaves of the Araucariaceae (Fig. Ill &
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Plates 6-9) are characterised by lateral or vertical wings* meso- phyll with air spaces* transfusion tissue all round the vascular bundle which are largely more than one* resin ducts not placed below the vascular bundle.
The leaf anatomy of this family has not been worked out by Ghouse and Yunus (1972). The separation of the two genera on anatomical basis is possible. This statement also holds good for the delimitation of the species of Araucaria (Table 1).
C ephalotaxaceae
Shrubs or trees, mostly dioecious. Branches opposite or whorled. Leaves alternate, spirally arranged, shining green above. Microsporophylls with 3-8 microsporangia; ovulate cones with decussately arranged bracts, each bract subtending 2 ovules* pollen-grains without wings.
This is a northern family and is represented by a single genus (Cephalotaxus) which is restricted to eastern Asia. There are six species out of which two occur in India. C. griffithii and £• ntannii are 3mall trees of the evergreen forests of eastern Himalaya (Datta, 1973). Since G. mannil was not available, C. drunacea (an introduced species) was studied here.
The leaves of Cephalotaxus (Fig. VI & Plates 19-20) are endowed with bifacial structure* abaxial stomata; absence of hypodermis; mesophyll differentiated into adaxial palisade and
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abaxial spongy parenchyma; ill-defined endodermis; single resin duct lying below the phloem; single vascular bundle and the transfusion tissue confined only to lateral side of the vascular bundle. All these characters were also recognised by Ghouse and Yunus (1972). While these authors found straight-walled epidermis with wide lumen, the epidermal cells appear to be more undulating with narrow lumen in C. griffithii and less undulating with broad lumen in C. drupacea. It appears possible to separate the two species of Cephalotaxus on anatomical basis (Table 1).
The leaves of Cephalotaxus closely resemble those of Taxus. but are easily distinguished by the presence of a large resin duct on the lower side between the epidermis and the midrib.
Cephalotaxus can be separated from Tsuga by the presence of a well-defined midrib and adaxial keel-like structure.
Cupressaceae
.Evergreen, dioecious or monoecious, shrubs or trees. Leaves opposite or whorled, acicular or scaly, iicrosporophylls with 3-6 or more microsporangia; ovulate cones woody or somewhat fleshy at maturity; ovuliferous scale fused with its bracts; ovules invariably erect and usually 3-20 per scale; pollen-grains without wing's.
In this family, there are about 23 genera and 148 species unequally divided between the northern and southern hemispheres.
- 45 -
Cupressus and Juniper us are indigenous to India, while Callitris. Chamaeevparls and Thu.jopsis have been introduced in this country,
Anatomically, the leaves of the Cupressaceae (Fig. IV & Plates 10-18) are triangular and characterised by vertical or lateral wings; compact or lax mesophyll; transfusion tissue lateral to and all round the vascular bundle, resin ducts 1-3, placed below the vascular bundle and towards the margin. It is possible to separate the six genera on anatomical basis.
Callitris s This genus is a native of Australia and Tasmania. Here the leaves are in whorls of three and those of the adult stage closely elapsing the shoot except at the triangular scalelike tip.
The stomata is amphistomatic and paracytie; the mesophyll is compact; the transfusion tissue Is lateral to the single vascular bundle (Table 1).
ChamaecvParis s This genus is found in North America, Japan and Formosa. Here the stomata is hyperstomatlc and anomocytic; hypodermls Is less developed (Table 1).
Cupressus 5 This genus has 20 species, out of which only two from India has been worked out. C, cashmeriana Is reported to have migrated to India through Tibet and is widely cultivated In this country. C. torulosa occurs in the Himalaya In the outer ranges from Chamba to Nepal and has been introduced in the Nilgiris where It is getting naturalised (Eaizada and Sahni,1958).
— 46 —
In C. cashing riana. the twigs are somewhat flattened with the leaves being blue-green and not appressed. In C. torulosa, the twigs, are roundish and leaves truncate with appressed tips. While C. cashmerlana varies in length from 1.5 to 2.0 mm, C. torulosa is about 1.5 mm long.
Anatomically, the leaves of Cunressus (Table 1) are characterised by triangular surface and thick-walled epidermis5 stomata are hyperstomatic and anomocytic; single-layered hypo- dermisj differentiated mesophyll$ one resin ducts ill-defined endodermis; single vascular bundle and the transfusion tissue lateral to the vascular bundle, forming wing-like extensions.
Although Ghouse and Yunus (1972) were not able to separate the species of Cupressus on anatomical basis, this has been done in the present study.
Juninerus $ This is an alpine conifer, .enjoying' a continuous distribution in the northern hemisphere alongja_br.oad' belt. There are about 70 species (Sporne, 1965); out of this number, six are reported from India (Datta, 1973). All the (Indian'species are — found in the inner valleys and higher ranges above the tree-limit
?
In Juninerus the leaves are acicular, scaly or subulate. They are arranged in four rows or borne in whorls, sometimes opposite and decussate or closely Imbricate. They are dull grey in colour, with a concave upper surface.
Anatomically, the leaves of Juninerus (Table 1) are ehara-
- 47 -o '
cterised by thick or thin-walled epidermis bearing anomocytic stomata; differentiated mesophyll; ill-defined endodermis; single but large resin duct located below the single vascular bundle which is flanked on either side with transfusion tissue. On the basis of anatomical features, the two species of Juniperus can be separated.
On the basis of morphological and anatomical features, the leaves of Juniperus resemble other members of the Cupressaceae. However, the leaves of Juniperus can be separated from Cupressus and Thuja by the large single resin duct and saucer-shaped outline in cross section.
Thuja s This is a northern. conifer of discontinuous distri- bution. There are about five species (Sporne, 1965) among which T. orientales is widely cultivated.
jr
f ***"
The leaves of Thuja are small, scaly, overlapping and in four ranks. Both T. occidentalss and T. orientales are laterally winged, the former with pointed apex and the latter with blunt -— apex.
Anatomically, the leaves of Thuja (Table 1) are characterised by thick-walled epidermis; anomocytic to paracytlc stomata; well- developed hypodermis of a single layer; undifferentiated or slightly differentiated mesophyll; one or more resin ducts with the same number of epithelial cells; slightly distinct endodermis; single vascular bundle flanked with transfusion tissue on either
— 48 —
side, varying in quantity and distribution in the same leaf from the base to the apex. On the basis of anatomical characters, the two species of Juniperus can be separated.
Thujopsis s This is a monotypic genus, occurring in Japan (Datta, 1973). According to Dallimore and Jackson (1948), wThu.1opsis differs mainly from Thu.1a by its larger leaves which
t 1are silvery on the undersurface, the lateral ones, axe-shaped, curved and pointed, the facial ones narrow and blunt, and by its rounder fewer-scaled cones with 3-5 seeds on each fertile scale against 2-3 seeds in Thuja”,
Anatomically, the leaves of Thujopsis (Table 1) are characterised by thick-walled epidermis bearing hyperstomatic and anomoeytic stomata* single-layered well-developed hypodermis dis- eontinuously distributed* lax differentiated mesophyll* ill- defined endodermis* single resin duct below the single vascular bundle which is transversely placed with the palisade parenchyma.
Thujopsis is closely related,,to Callltris. In both of them, the mesophyll is fully differentiated. While mesophyll Is compact in Callitris. it is |axjin Thujopsis. Moreover, the transfusion tissue is lateral to the vascular bundle in Callitris arid all round the vascular bundle in Thujopsis.
PinaceaeMonoecious trees. Leaves spirally arranged, linear or acl-
cular. Microsporophylls with 2 microsporangia* ovuliferous scale
- 49 -
free from the bract? ovules 2 per scale? pollen-grains mostly
winged.
The Pinaceae is mainly a northern family, being composed of
about 10 .genera and 200 species (Florin, 1963? Sporne, 1965).. In
India, there are about six genera of this family and all of them,
the leaves are largely acieular except in Abies and Tsuga where
they are linear.and somewhat flat.
Anatomically,.the leaves of the Pinaceae (Figs..VII-VIII &
Plates 21-30) have mostly thick-walled epidermis? well-developed
or less-developed hypodermis? lax mesophyll? resin-ducts 2-9 and
transfusion tissue all round the vascular bundles. The five genera
of the family can be separated on the basis of anatomical chara
cters.1Abies 5 This is a genus well distributed in the Himalaya ;
(Raizada and Sahni, 1958). The leaves are variously arranged on
shoots and those on the lateral shoots pectinate or spreading all
round or imbricate pointing outwards. They persist for 3-6 years
and generally bear 2 white bands, 6 cm one on either side of the
midrib running parallel to each other. Abies can be separated
from all other conifers by the disc-like leaf-scars and erect
cones which break-up as soon as the seeds are ripe.
Anatomically, the leaves of Abies (Table 1) are characteised
by 2 resin ducts, located one at the end of each wing slightly
towards the abaxlal side. In this respect, the genus differs from
Tsuga'and Cunninghamia. Besides, the leaves possess thick-walled
- 50 -
epidermis;, the stomata is hypostomatic and pericytie, the hypo-
dermis is well-developed and the mesophyll differentiated; trans
fusion tissue is lateral to the double vascular strands.
Cedrus s This genus is represented by a solitary species in
India, namely C. libani which is widely distributed in the wes- ,
tern Himalaya, from Afghanistan to Garhwal at 1200-3000 m elevation
(Raizada and Sahni, 1958). It has leaves of dark-green colour with
sharply pointed apex.and persisting for 3-6 years. The length of
the leaves ranges from 2.5 to 3.8 cm.
Anatomically, the leaves of Cedrus (Table 1) have thick-
walled epidermis; the stomata Is amphistomatlc and tetracytie. The
hypodermis is well-developed and the mesophyll undifferentiated, ^ containing^pl^^t^ parenchyma; there are 2 resin ducts placed
laterally, one on either side of the vascular bundle but more
towards the abaxlal side; the endodermis is distinct and the trans
fusion. tissue Is lateral to the double vascular strands.
Picea i In India, there are two sjpecies of Picea — P.
smlthiana and P. spinulosa. Whereas the latter is confined to the
Eastern Himalaya, the former occurs in Central and Western Himalaya.
Borne on normal shoots, the leaves of Picea persist for 6-8
years. They are acicular and vertically winged. They vary in len
gth from 2.5 to 3.8 m. Picea can be distinguished from other coni
fers by peg-like leaf bases on the twigs. They are similar to the
- 51 -
leaves of Acmonvle. Cryntomeria and Dacrydium in containing vertically placed wing-like structures.
Anatomically, the leaves of Picea (Table 1) are characterised by ado-abaxially extended keels which form vertically-placed wings; thick-walled epidermal cells with amphistomatic and tetra- cytic stomata; hypodermis of one or two layers; undifferentiated but horizontally aligned non-plicate mesophy11 cells; the double sac-like discontinuous resin ducts; well developed endodermis; double vascular strands and the transfusion tissues all round the vascular tissues.
Pinus j This is the largest and most important genus of the Pinaceae. About 90 species of Pinus are known, out of which five are found in the Indian subcontinent. Eecently, a Chinese species (P. armandi) has been reported from North-west frontier of Assam.
Morphologically, Pinus is characterised by long and dwarf shoots, the latter bearing the foliage in clusters terminally.In each cluster, there are 3 needles in P. gerardiana. P. keslva and P. roxburghli; 5 needles in P. wallichiana and 2 in P. merkusii.
Among conifer leaves, the anatomy of pine needles has been studied in great details (Strasburger, 1891; Harlow, 1931; Sutherland, 1933; Huber, 1947; Konar, 1963). Depending on the number of needles per cluster, pine needles assume various shapes in cross- section, namely circular, semi-circular, triangular and terete
- 52 -
(Dolivo, 1948). Pine needles (Table 1) show cuticularised epidermis with thickened walls encircling the almost obliterated lumen. Although the stomata is amphistomatic, the type varies from species to species. The hypodermis is less developed or well developed, occurring in many layers or In patches. The mesophyll cells are undifferentiated but peculiar in possessing plicate parenchyma (Esau, 1960); cells are arranged in horizontal layers and separated from one another by intercellular spaces. The number of resin ducts differ from species to species and sometimes in the same species (Ghouse and Yunus, 1972). But all of them possess two basic ducts, situated lateral to the vascular bundles but slightly towards the abaxial side and the other ducts vary in distribution and number. The endodermis may be either distinct or not distinct. The transfusion tissue occurs all round the vascular tissues. While P. wallichlana shows a single vascular bundle, other species contain 2 vascular bundles. Although the five species of Pinus resemble each other, they can be separated on the basis of anatomical characters.
Tsuea s Although the genus contains 10 species, it is represented by two species in the Indian region. T. dumosa is mainly distributed in the Central and Eastern Himalaya at an elevation of 1800-3300 m. T. yunnanensis is mainly found in the mountainous areas of Western Szechuen and Yunnan.
Unlike Pinus. Tsuga does not show dimorphic branches. The leaves are flat, dark green above, usually two-ranked with two
- 53 -
whitish hands below. In appearance and arrangement, the leaves of Tsuga are similar to those of Abies and Pseudotsuga.
Anatomically, the leaves of Tsuga (Table 1) are characterised by a bifacial structure without distinct midrib; the upper epidermis is thick-walled without any undulations and lower epidermis with sinous cells bearing stomata; there Is the absence of any hypodermal development; mesophyll is differentiated and without plicate parenchyma; single resin duct is situated right below the phloem; endodermis is ill-defined and the transfusion tissue distributed all round the vascular bundle.
Tsuga differs from Cenhalotaxus in lacking well-defined midrib and adaxial keel-like structure. Tsuga can be separated from Cunninghamia in not having any hypodermis and from Taxus in having a resin duct. However, Tsuga can be related to the other three genera by dint of other anatomical attributes,
Podocarnaceae
Evergreen shrubs or large trees, usually dioecious. Leaves linear, lanceolate or elliptic. Microsporophylls with 2 micro- sporangia; megasporangiate stroblli cone-like or greatly modified; ovule erect or reflexed; pollen-grains with 2 or 3 wings.
The Podocarpaceae is a southern family, having 7 extant genera and 150 species. Though some of them are found north of the equator, they are thought to have reached there only recently
(Florin, 1963). Podocarpus is the principal genus with about 125 species (Ghouse and Yunus, 1972). Podocarpus is the only- natural conifer of Peninsular India and only surviving indigenous conifer of this country.
The leaf anatomy of the Podocarpus has been studied in details (Orr, 1944; Buchholz and Gray, 1948; Griffith, 1957; Florin, 1958; Gray, 1958), The leaves of Indian species (Fig.IX & Plates 31-33) are characterised by thick-walled epidermis; hypostomatic, paracytic and staurocytic stomata; discontinuous hypodermis; resin ducts one or more; absence of well defined endodermis; single vascular strand; primary and accessory transfusion tissues; the former constituting lateral wings to the vascular elements and the latter extending to the extremities of the wings. The separation of species on anatomical basis seeds to be possible in the case of Podocarpus (Table 1).
TaxaceaeEvergreen shrubs or trees, dioecious. Leaves acicular or
linear, spirally arranged. Microsporophylls peltate with 2-8 microsporangia; ovuliferous branch with a single terminal ovule, entirely or partly enclosed at maturity by a fleshy aril; pollen- grains without wings.
The Taxaceae is a northern family embracing 5 extant genera and about 20 species. In India, this family is represented by Taxus. T. baccata prevails In moist shady regions above 1800 m
55 -
and all along the Himalaya (Datta, 1973).
The leaves of the Indian taxads are flat, linear and arranged in apparent distichous order. They vary in length from 2.5 to 3.8 cm. They are single-veined, having a well-defined midrib.
Anatomically, the leaves of Tax us (Fig, IK & Plate 34) are characterised by thick-walled epidermis with hypostomatic stomata; absence of both hypodermis and resin ducts; differentiated mesophyll and transfusion tissue lateral to the single median vascular bundle.
Although the leaves of Taxus closely resemble those of Cenhalotaxus and Tsuga, they can be easily separated from the rest of the other genera studied by the lack of resin ducts (Table 1).
Taxodiaceae :
Evergreen or deciduous trees, monoecious. Leaves opposite or spirally arranged, aeieular or linear. MIcrosporophylls with 2-9 microsporangia; ovullferous scale fused with each bract and ' bearing 2-9 ovules; pollen-grains without wings.
This is an northern family, having relict distribution and historical significance. This family comprises 10 genera and 15 species (seven of which are monotypic). There is no indigenous species but Cryptomeria. Cunninehamia. Metasequoia and Taxodium are exotics. They are forming natural forests wherever they
- 56 -
have been introduced in India,
Anatomically, the members of the family (Fig. X & Plates 35-38) are characterised by one or more resin ducts and a single vascular bundle supported^with transfusion tissue; accessory transfusion tissue is entirely absent here. The four genera studied can be separated by means of anatomical characters.
Cryptomeria s This is a monotyplc genus which Is restricted to China and Japan and got the entry by introduction and is rapidly spreading In the Eastern Himalaya (Troup, 1921).
The, leaves are awl-shaped, quadrangular with decurrent bases and spirally arranged in five ranks.
Anatomically, the leaves of Cryptomeria are characterised by vertically extended wings; thick-walled epidermis; stomata amphistomatic and actinocytic; one to two-layered hypodermis; slightly differentiated mesophy11; lonely resin duct placed below the single vascular bundle which is flanked on either side with transfusion tissue.
Cunninghamia : This is a Chinese genus which have been introduced in Dehra Dun and adjacent places.
The leaves of Cunninghamia are lanceolate, spirally arranged, distichous, with wide base and spiny apex, glaucous green in colour (Ghouse and Yunu3, 1972). Anatomically, the leaves of Cunninghamia are characterised by laterally extended wings;
- 57 -
thick-walled epidermis with harrow lumen; stomata hypostomatic and anomocytic; one- to" four-layered well-developed hypodermis; differentiated mesophyll; resin dhcts more than one; transfusion tissue' lateral to the single vascular bundle.
letasequoia s “Because Metasequoia had long been known only from its fossil record, uncovered in various parts of western North America and Asia, the discovery in 1944 by Chinese botanists of living specimens of the genus in Szechuan Province, China, was of exceptional scientific Importance*1 (Foster and Gifford, 1974). The morphology and relationships of the living species with the other extant genera of the family have been discussed by Sterling (1949). Like living cyeads and Ginkgo biloba, Metaseauoia is in reality a living fossil. Further studies on its structure and reproduction should help in discoveries of evolutionary interest.
Unlike other three genera in the family, the leaves are opposite in Metaseauoia.
Anatomically, the leaves of Metaseauoia are characterised by laterally extended wings; thin-walled epidermis, containing hypostomatic and anomocytic stomata; absence of both hypodermis and accessory transfusion tissue; differentiated mesophyll; ill- defined endodermis; transfusion tissue lateral to the single vascular bundle.
Taxodium * This genus contains two species, known from North American (Datta., 1973). Taxodium dlstichum is a semi-deciduous
- 58 -
'tree which has been planted at the Forest Research Institute,Dehra Dun and is a highly prized ornamental (Raizada and Sahni, 1958). "On well drained soil the trunks taper from base to apex but in swamps the submerged roots form ’mnees*. which are woody / structure several feet high and hollow1*.
The leaves of Taxodium- are spirally arranged on the branch- lets, but they appear in two ranks on the deciduous shoots due to twist near the base. They are scaly and shorter on the persistent branches and are delicate green in autumn.
Anatomically, the leaves of Taxodium are characterised by laterally extended wings; thin-walled epidermis, containing hypo- stomatic and anomocytic stomata; absence of both hypodermls and accessory transfusion tissue; differentiated mesophyll; resin duct one.; ill-defined endodermis and transfusion tissue lateral to the single vascular bundle.
Table
1 5 A
nato
miea
l da
ta for t
he c
lassificat
ion
of li
ving I
ndian
gymnosperms
based
on the m
icroscopic
characters of
the f
oliage l
eaves
Zamia angustifolla
Late
rall
y winged
Thick
Narrow
Hypostomatic
Upper
surface 0
Lower
surface
7079*4
Paracy
tic
Well
dev
eloped
One
layer
Disc
onti
nuous
Less d
ifferentiated
Spongy p
arenchyma
loosely
arra
nged e
nd p
rovided
with
air s
pace
0 0No
t distinct
All
roun
d V.B,
Not
developed
Ence
phal
arto
s vlllosus
Late
rall
y wi
nged
Thic
kNarrow
»
Hypost
omatic
Upper
surface
0Lower
surface
4719*6
Hypocytic
Well
developed
One
layer
Discon
tinuous
Diff
eren
tiat
ed
Spongy p
arenchyma
compac
tly ar
rang
ed
0 0i
Not distinct
All
roun
d V.B*
Not
develo
ped
Two
- 6-11
Not
distinct
Latera
l to V*B.
Well
developed
eye A
D AC! EA
E Cycas
rum' ph
ii
Late
rall
y win g
ed
Thic
kBr
oad
Hypost
omatic
Uppe
r surface
0 Lower
surface
6961*6
Paracytic
Less d
eveloped
One
layer
Discon
tinuous
Diff
eren
tiat
ed
Cvca
s revoluta
Late
rall
y wi
nged
Thick
Narrow
Hypostomatic
Upper
surface
0 Lower
surface 68
31*0
Actinocytic
Well
developed
One/two
layers .
xDiscontinuous
Diff
eren
tiat
ed
0 -
0 .
Not
distinct
Late
ral
to V.B.
Well
developed
I. G
eneral f
orm in T.S
II. Epidermis
Cuticle
Lumen
III. S
tomata (
Fig. XI)
Dist
ribu
tion
Fr
eque
ncy
Type
IV.
Hypodermis
V. M
esop
hy11
VI*
Besin
ducts
Epithelial
cells
VII. E
ndodermis
VIII.
Transfusion
tissue
3X.
Accy.
transfusion
tissue
XVascular b
undle
One
" Two
More t
han
two
More t
han
two
Bundle s
heath
Bundle s
heath)
Bundle s
heath
Bundle s
heath
extension
absent
extension
absent
extension ab
sent
extension
present
I,II.
III.
IV.V.
VI.
VII.VIII.
IX.
X.
- 60 -
Table 1 (Contd.)
GINKGOACEAE
General form in T.S.Epidermis
CuticleLumen
Stomata (Fig. XII) Distribution Frequency
Type
Hypodermis Mesophy11
Besin ducts Epithelial cells
EndodermisTransfusion tissueAccy. transfusion tissueVascular bundle
Ginkgo biloba
Laterally winged
Thin (undulated)Narrow
HypostomaticUpper epidermis 0 Lower epidermis 11799.0Anomocytic
0Differentiated but; palisade and spongy Irregular
More than two 11-13
DistinctAll round V.B.Not developed
More than two
(Contd...,)
Tabl
e 1| (Co
ntd.
)
ABA
UCid
ilAC
EAE
• • •
•)
One
Mor
e than
two (3
)M
ore th
an t^
'oM
ore th
an tw
oIX
, Vascu
lar b
undl
e
Not
dist
inct
All r
ound
V.B
,
10Tw
o
Not
diff
eren
tiate
d,
with
prom
inen
t air
spac
es
/A. cu
nnin
gham
ml
Ver
tical
ly wi
nged
Thic
kN
arro
w ’ •
Am
phis
tom
atic
U
pper
surf
ace 4
968
Low
er su
rfac
e 746
2 A
nom
ocyt
icy
Wel
l deve
lope
d O
ne/tw
o laye
rs
Dis
cont
inuo
us
Ver
tical
ly wi
nged
Thic
kN
arro
w
Am
phis
tom
atic
U
pper
surf
ace 5
216,
4 Lo
wer
surf
ace 6
085.
8 A
nisp
cytic
Less
deve
lope
d O
ne lay
e'r
Dis
cont
inuo
usN
ot di
ffer
entia
ted,
le
ss ai
r spac
e
One
- 18
Not
dist
inct
5 A
ll rou
nd V
.B.
A. /
col
umna
r!s
Thic
kN
arro
w
Hyp
osto
mat
i<j
Upp
er 3ur
fa< !
e 310
5.0
Low
er 3u
rfa<
JQ 50
92.0
Ani
socy
ticW
ell de
velo
] >ed
In bu
ndle
s D
isco
ntin
uot is
Diff
eren
tia^
:ed
i j
Mor
e than
tv ro
8-12
Not
dist
inct
:
All r
ound
vi B
.
Lnge
d
Ldw
illi
Late
rally
wA
rauc
aria
, bA
gath
ls lo
rant
hifo
lla
Late
rally
win
ged
Thin
Bro
ad
Hyp
o3to
mat
ic
Upp
er su
rfac
e 0
Low
er sur
face
7700
.4
Stau
rocy
ticLe
ss de
velo
ped
One
/two la
yers
D
isco
ntin
uous
Diff
eren
tiate
d
Mor
e than
two
11
Not
dist
inct
All r
ound
V.B
.
V. Mes
ophy
ll . ’
VI. Be
sin d
ucts
Epith
elia
l cel
lsV
II. End
oder
mis
VII
I. Trans
fusi
on tis
sue
I, Gene
ral fo
rm in
T.S,
II
. Epide
rmis
C
utic
le
Lum
en
III. Sto
mat
a (Fig
, XII
) D
istri
butio
n Fr
eque
ncy
Type
IV • Hyp
oder
mis
i
(Con
td
CEPHAL
OTAX A(E AE
(Con
td....)
C. g
riffithil
Late
rall
y wi
nged
More u
ndulated
(homogenous)
Thic
kNarrow
Hypo
stom
atic
’Up
per
surface
0 Lo
wer
surface
12644,2
Para
cyti
cAbsent
Diff er
enti
ated
One
13-16
Not
distin
ct
Late
ral
to .V.B,
One
$
Late
rall
y winged
Less u
ndulated (
heterogenous)
Thin
Broa
d
Hypostomatic
Upper
surface
0 Lower
surface
12420
Hexacytic
Absent
Diff
eren
tiated
One
10 Not
distinct
Lateral
to V.B,
One
I, Ge
neral
form
in
T.S,
II. E
pidermis
Cuticle
Lumen
III. St
omata
(Fig. XV)
Distribu
tion
Frequenc
y
Type
IV. H
ypodermis
V. Me
sophyll
VI. R
esin
ducts
Epit
heli
al c
ells
VII. En
dodermls
VIII. T
ransfusion t
issue
IX• V
ascular
bundle
Cephalotaxus d
rupacea
CUPR] ISSACEAE
(Contd
...)
Thick
Broad
Hyperstomatic
Upper
surface
3477,6
Lower
surface
12047.4
Anomocytic
Well
developed
One
layer
Disc
ontinuous
. Differentiated, l
ax
One
(at the
concave
side)
26 Not
distinct
Lateral
to V.B.
OneTriangular b
ut a
pex
knotched or
concave
Thick
Broad
Hype
rsto
mati
c but
at t
wo
latera
l sides an
d not
at
the
upper
surface
Upper
surface
12792,6
Lowe
r surface
0
Anom
ocyt
icLe
ss d
eveloped
One
layer
Cont
inuo
us ,
Diff
eren
tiat
ed wit
h (2-3)
layers o
f palisade
tissue,
lax 5
One
(just
below
phloem)
23 Not
distin
ct
Late
ral
to V.B.
OneTriang
ular
C. t
orulosa
i •
Cupr
essu
s cashmeriana
One
(at th
i leaf
margin) |
7-8
Not
distin« j
i
All ro
und Vi
B.
Two
Chamaecypa: 'is pi
slfe
ra
Triangular
Thic
kBroad
Hyper s
toma' ;lc
Upper
surfi ice 24
964.
2 Lower
surfj ijce 2
0865.6
Anomoc
ytic
Less d
evel< >ped
One
layer
Discontinue >us
Not
differ^ intiated
Diff
eren
tiat
ed wit
h a
little a
baxlal p
ali
sade
More t
han
two
(3),
one
belo
w V.B.
19-22
Dist
inct
Late
ral
to V.B.
One
Uppe
r surface
7700.4
Lowe
r surface
8445.6
Para
cyti
c
Late
rall
y wi
nged
Thin
Broa
d
Amphistomatic
I. Ge
neral
form i
n T.S,
II. E
pidermis
Cuticle
Lume
nIII. St
omata
(Fig. XIII)
Distribu
tion
Freq
uenc
y
Type
IV. H
ypodermia
V. Me
sophyll
VI. R
esin
ducts
Epit
heli
al c
ells
VII. En
dodermis
VIII. T
ransfusi
on t
issue
IX. V
ascular
bundle
Call
itri
a robusta
Table
! /Contd.)
Jun
iper
us
reeu
rva
I.
Gen
eral
for
m i
n T
.S.
· L
ate
rall
y w
inge
d,
bu
t ab
axia
l su
rtac
e co
n.
cave
II
. E
pid
erm
is
Cu
ticl
e T
hic
k (
un
du
late
d)
Lum
an
Nar
row
II
I.
Sto
mat
a (F
igs.
XII
I-X
IV)
Dis
trib
uti
on
A
mph
isto
mat
ic
Fre
quen
cy
Typ
e
IV.
Hyp
qder
mis
v.
Mas
ophy
ll
VI.
R
esin
du
cts
Ep
ith
eli
al ~ells
VII
. ~
End
oder
mis
VII
I.
Tra
nsf
usi
on
tis
sue
IX.
Vas
cula
r b
un
dle
Upp
er
surf
ace
5216
L
ower
su
rfac
e 1
16
74
.8
Ano
moc
ytj,a
. W
ell-
dev
elo
ped
O
ne
lay
er
Dis
con
tin
uo
us
Differentiated~
wit
h
lax
pal
isad
e ti
ssu
e
..t
...... -
,~
-'>
o~e
(at
the
leaf
mar
gin)
16
Not
d
isti
nct
Late
ral
to V
.B.
one
'·
'~
-6
4 -
Tab
le 1
. (C
on
td.)
CUPR
ESSA
CEA
E (C
on
td.)
l.• w
alli
chia
na
Late
rall
y w
inge
d
Th
in
Bro
ad
Hyp
o st
om
atic
Upp
er
surf
ace
0 L
ower
su
rfac
e :8
3349
.6
Ano
moc
ytic
Wel
l-d
evel
op
ed
One
la
yer
C
onti
nuou
s
Dif
fere
nti
ate
d1
aba
xia
l p
alis
ade
t.~sso
cia
ted
wit
h c
o~1pact
spon
gy p
aren
cbjr
ma.
i
One
(a
t th
e le
i1,f
mar
gin)
'
25
.
Not
dis
tin
ct
Late
ral
to V
.Be.
and
all
rou
nd V
.B.
One
,. T
hu
jop
sisf
do
lob
rata
Late
rall
y w
inge
d
Th
ick
N
arro
w
Hyp
er s
tom
atic
Th
uja
occid
en
tali
s
Late
rall
y w
inge
d,
wit
h p
oin
ted
ape
x
Thi
ck
Nar
row
Hyp
er s
tom
atic
Upp
er
surf
ace
10
30
8.6
U
pper
su
rfac
e 12
420
Low
er
surf
ace
0 L
ower
su
rfac
e 0
Ano
moc
ytic
Wel
l-d
evel
op
ed
One
la
yer
D
isco
nti
nu
ou
s
Dif
fere
nti
ate
d,
wit
h
one
lay
er o
f .p
ali
sa
de t
issu
e,
lax
One
(j
ust
abo
ve t
he
phlo
em)
13
Not
d
isti
nct
(Com
plex
ty
pe)
A
nom
ocyt
ic &
Pa.
ra.
cy
tic
Wel
l-d
evel
op
ed
One
la
yer
D
isco
nti
nu
ou
s
Un
dif
fere
nti
ated
, la
x
'One
/tw
o (b
ig s
ize,
just
at
the t
ip)
24
Sli
gh
tly
dis
tin
ct
All
rou
nd V
.B.
Late
ral
to·v
.B.
(V.B
. tr
an
svers
ely
p
lace
d w
ith
pal
isad
e ti
ssu
e)
One
O
ne
!·
ori
en
tali
s
Late
rall
y w
inge
d,
wit
h b
lun
t ap
ex
Th
ick
N
arro
w
Hy
per
sto
mat
ie
(at
two
late
ral
sid
es)
Upp
er
surf
ace
23
47
3.8
L
ower
su
rfac
e 0
Act
ino
cyti
c
Wel
l-d
evel
op
ed
One
/tw
o la
yers
C
onti
nuou
s
Sli
gh
tly
dif
fere
n
tiate
d,
wit
h l
ax
spon
gy t
issu
e
Mor
e th
an t
wo
(3)
{at
th
e le
af
mar
gin)
24
Sli
gh
tly
dis
tin
ct
Late
ral
to V
.B.
and
all
ro
un
d V
.B.
One
(Con
td •
•• )
Abie
s SPectabiliS
Cadr
us d
eoda
m var. H
bani
Pldea
sralthlana
Plnu
s de
nsiflora
Distin
ctAlt ro
und V.
B Two
(Contd.,...)
Semicircular
Thic
k , '
Broa
d
Amphis
tomatic
Uppe
r surface
0 ,
Lower
surface
€706
.8
Graminaceous
Well
-dev
eloped
■One l
ayer
Disc
onti
nuous
Distin
ctAll round V*
B Two
Vert
ical
ly win
ged
Thick
Broad
Amphis
tomatic
Upper
surface 44
71.2
Lover
surface
12047.4
Tetr
acyt
icWe
ll-d
evel
oped
One/tw
o layers
Disc
onti
nuou
s
Late
rall
y winged
4
Thic
kNarrow
Amphis
tomatic
Uppe
r surface 6682.6
Lowe
r surface 73
27.8
Tetrac
ytic
Well-developed
One/tw
o layers
i
Cont
inuo
us j
Not
differentiated
i
Plicat
e parenchyma p
resent
l
Two
21 Distinct
Latera
l to V.Ijf
Two
Late
rall
y wi
nged
Thin
Narrow
Hypo
stoa
atic
Upper
surface
0 Lo
wer
surface
0439.2
> Per
icyt
icWe
ll-d
evel
oped
On
e/tw
o layers ‘
Disc
onti
nuou
s
Diff
eren
tiat
ed
Plicate
pare
nchy
ma absent
Two
33 Distinct
Late
ral
to V.B.
Two
Type
IV• H
ypoderals
V. M
e3ophyll
VI.
Basi
n ducts
Epit
heli
al c
ells
VII*
Endoder»is
VIII.
Transfusion
tissue
IX. V
ascu
lar bundle
I. G
eneral f
orm in
T.S.
. II. E
pidermis
Cuticle
Lume
nIII. S
tomata (
Figs. XVI-XVII)
Distributi
on
Freq
uenc
y
plic
ate
pare
nchy
ma abs
ent
Plicat
e parenchyma p
resent
Two
More t
han
two
(9)
9 e
10-11
Not differ
entiated
Not
differ
entiated
I.
Gen
eral
for
m i
n T
,S •.
II,
Epi
derm
is
Cu
ticl
e Lu
men
III,
S
tom
ata
(Fig
. X
VII
) D
istr
ibu
tio
n
Fre
quen
cy
Typ
e
IV.
Hyp
oder
mis
v.
Mes
ophy
ll
VI.
R
esin
du
cts
Ep
ith
eli
al
cell
s
VII
. E
ndod
erm
is
VII
I.
Tra
nsf
usi
on
tis
sue
IX.
Vas
cula
r bu
ndle
Pin
us
wal
lieh
ian
a
Tri
ang
ula
r
Thi
n N
arro
w
Am
phis
tom
at1c
U
pper
s~tace
64
58
,4
Low
er
surf
ace
53
40
.6
Par
acy
tic
Wel
l-de
velo
ped
One
to
tw
o la
yer
s C
onti
nuou
s
Not
d
iffe
ren
tiate
d
Pli
ca 'lie
-,. p
aren
chym
a p~esent
Two
14
-15
Dis
tin
ct
All
rou
nd V
.B.
One
',
·,·,
• 66
-
Tab
le ;
L (C
ontd
,}
PlN
ACE
J ~
(Con
td.)
!·
gera
rd1a
na
Sem
i-lu
nar
Thi
ck
Nar
row
Am
phis
tom
a.tic
U
pper
su
rfac
e 4
71
9,6
L
ower
su
rfac
e 6
33
4,2
S
emia
ctin
ocyt
j.c
ltle
ll-d
evel
opea
In
Bun
dles
D
isco
nti
nu
ou
s
Not
differenti~ted.
Pli
cate
par
enc:
uym
a p
rese
nt
Mor
e th
an 2
(3
)':
8 Not
d
isti
nct
All
rou
nd V
.B.·
Two
.. P
, ke
si:v
a.
-I
Tri
ang
ula
r,
uppe
r la
yer
sli
gh
tly
cu
rved
Thi
n N
arro
w
P.
!'OX
bUT.
ghii
Tri
ang
ula
r,
uppe
r la
yer
sl
ign
tly
conve~
Thi
n N
arro
w
Am
phis
tom
atic
A
mph
isto
ma.
tic
Upp
er
surf
ace
77
00
,4 U
pper
su
rfac
e 4
47
1.2
L
ower
su
rfac
e 93
15
Low
er
surf
ace
83
21
.4
Par
acy
tic
Gra
min
aceo
us
.f.
sylv
est
ris
Cre
scen
t-sh
aped
. L
ower
la
yer
sl
igh
tly
kn
otch
ed
. T
hin
Nar
row
Am
ph1s
torn
atic
U
pper
su
rfac
e 6
45
8
Low
er
surf
ace
1055
7 S
ta.u
rocy
tic
and
Par
acy
tic
(Com
plex
ty
pe)
Les
s-de
velo
ped
One
to
tw
o la
yer
s D
isco
nti
nu
ou
s
Les
s-de
velo
ped
Les
s-de
velo
ped
Not
d
iffe
ren
tiate
d
Pli
cate
par
ench
yma
pre
sen
t
Two
6-7
Dis
tin
ct
All
rou
nd V
.B.
Two
One
to
th
ree
lay
ers
One
la
yer
C
onti
nuou
s
Not
d
iffe
ren
tiate
d
Pli
cate
par
enph
yma.
p
rese
nt·
.·
·
Mor
e th
an t
wo
(3-4
) 8 D
isti
nct
All
rou
nd Y
.B,
Two
Dis
con
tin
uo
us
Not
differ~ntiated
Pli
cate
par
ench
yma
pre
sen
t
Mor
e th
an t
wo
(6)
14
Not
d
isti
nct
All
rou
nd V
.B.
Two
(Con
td •
••• )
- 67 -Table 1 (Contd.)PINACEAE (Contd.)
'Tsuga dumosa
I. General form In T.S. Laterally wingedII, Epidermis
Cuticle ThinLumen Broad
III. StomataDistribution HypostomaticFrequency Upper surface 0
Lower surface 15276,6Type Paracytic
IV. Hypodermis AbsentV. Me sophyll Differentiated, plicate
parenchyma absent
VI. Eesin ducts OneEpithelial cells 14
VII. Endodermis Not distinctVIII. Transfusion tissue All round V.B.
IX. Vascular bundle Two
(Contd..,)
PODO
CARP
ACEA
E
(Con
td....)
Hypo
stom
atic
Uppe
r surface
0 Lower
surface
19251.0
Paracytic
Well
-dev
elop
ed
In b
undles
Discon
tinuous
Differentiated,
with
abaxia
l palisade
One
9 Not
distinct
All ro
und
V.B.
Well
-dev
elop
ed
OneLate
rall
y winged
Thin
Broa
d
Late
rall
y winged
Thic
kBr
oad
*
Hypo
stom
atic
Up
per
surface
0 Lo
wer
surface
8445.6
Para
cyti
cLess d
evel
oped
One
layer
Disc
onti
nuou
sDi
ffer
enti
ated
Two
7 Not
distin
ct
Late
ral
to V.B,
Well
-dev
elop
ed
One
P. s
inensis
P* n
eriifollus
Podo
carn
us m
acro
nhvl
la
Late
rall
y wi
nged
Thick
Broad
Hypostomatic
Upper
surface
0 Lower
surface
12171.6
Staurocytic
Less
developed
One/
two
layers
Disc
onti
nuou
sDiff er
entiated
More t
han
two
(3)
18-19
Less d
istinct
Late
ral
to V.B.
Well
-dev
elop
ed
V. Me
sophyll
VI. R
esin
ducts
Epithelial
cells
VII. En
dodermis
VIII. T
ransfusion t
issue
IX.
Accy.
transfusion
tissue
X.I. Ge
neral
form
in T.S,
II, E
pidermis
Cuticle
Lume
nIII. St
omata
(Fig. XVIII)
Distributi
on
Frequency
Type
IV. H
ypod
ermi
s
Table
1 (Contd.)
Vascular
bundle
One
69
Table 1 (Contd.)TAXACEAE
Tax us baccata
I. General form in T.S.II. Epidermis
Cuticle Lumen
III. Stomata (Fig. XVIII) Distribution Frequency
Type
Laterally winged
ThickBroad
Hypos tom ticUpper surface 12171.6Lower surface 0Paracytie and Hexacytic
IV. HypodermisV. Mesophyll
VI. Resin ductsEpithelial cells
0Differentiated0
VII. EndodermisVIII. Transfusion tissue
IX, Vascular bundle
Distinct Lateral to V.B. One
(Contd....)
Differentiated
One
15 Not
distinct
Late
ral to V.B.
Absent
OneLaterally winged
Thin
* 'Br
oad
Hypostomatic
Upper
surface
7948.8
Lower
surface
9315.0
Anomocytic
Absent
Latera
lly wi
nged
Thin
Broad
»
Hypost
omatic
Upper
surface
0 Lower
surface
21238.2
Anomocytic t
Absent
Diff
eren
tiat
edMore t
han two
(3)
10 Not distinct 8
Lateral
to V.B.
Absent
OneMetase
quoia
glypto
3troboides Tax
odiu
m distichum
- 70
-Table
1 (Contd.)
TAXOD E
ACEAE
Cunn
ingh
amia s
inensis
Late
rall
y wi
nged
Thin
Narrow
Hypost
omatic
Upper
surface
13786.2
Lower
surface
2496
4.2
Anomocytic
We 11
- de v
e lo ptfd
One/four l
ayers
Discontinuous
Differentiated |
More t
han
twc> (3
)27 Distinct
Latera
l to V,B.
Absent
One
Cryp
teaw
ria
japonica
Vert
ical
ly winged
Thic
kBr
oad
Amphistomatic
Uppe
r surface
9563.4
Lower
surface
16394.4
Actinocytic
Well
-dev
elop
ed
One/two
layers
Continuous
Slightly differentiated
One
37 Distinct
Late
ral
to V.B.
Absent
One
V. Me
sophyll
VI. B
esin
ducts
Epit
heli
al c
ells
VII. En
doderm
lsVIII. T
ransfusi
on t
issue
IX.
Accy.
transfusion
tissue
X.
Vascular b
undle
I, Ge
neral
form
in
T.S,
II. E
pidermis
Cuticle
Lumen
III, St
omata
(Fig. XIX)
Dist
ribu
tion
Fr
eque
ncy
Type
XV. H
ypodermis
PLATE-1
Plate 1. Cycas rumphii Plate 2. C. revoluta
f
(Left : mesophy11 tissue rights vascular tissue)
PLATE-3
Plate 3* Encenhalartog villo3U3 Plate 4* Zamia florldanaCleft t mesonhyll tissue; Right t vascular tlarae )
)
PLATE-7
*>
t
i
i
i
flat# 1# Ginkgo bilob* flat# l« ittthli loranthlfolla Flat# f • Araucaria bMwlllll(Left t mesophy 11 tl*#tsaf Blgbt « vascular tissue)
fist# S* kxmmmrlm columnar Isflat* f * ummsaria curm Inehamll(Left § mesophy 11 tiesue§Right t vascular tissue)
flat* 20. Callitrli robust*f lat* 11. ChjHaattyMris ptslfay* Plate *21. Cupreaeas eaahaertaa*(Left i ae Sophyll tissue; light, t vnscalar tissue )
PLATE-14
Flat® 13* Cupressas torulosa Flat® 14* JTaalparug yistam(Left • mesonhyll tissue; lifMt t msealAr tissma )
PLATE-I5:
Flat* US. Junit>eru3 wallichianfi Fluts li. Thmia oeeidentales(Left t raesophyll tissue; Bight § vascular tissue)
{
PLATE-18
- 78 -
Plate 17* Thuja orientalia Plate 18. Thu.1opsi3 dolabmta(Left : mesophyll tissue; Eight s vascular tissue)
PLATE-19
PLATE-20
Plate 19. Cenhalotaxus - Plate 20. C. griffithil
(Left s mesophyll tissue Right ; vascular tissue)
- SO -
Plate 21. Abies special) ill a Plate 22. Cedrua libani Plate 23. Picea saithiana(Left s mesophyll tissue; Eight t vascular tissue )
PLATE-24
PLATE-25
81
Plate 24. Pinus densiflora Plate 25. P. gerardlana(Left s raesophyll tissue; Right i vascular tissue)
PLATE-26
PLATE-28
- 82 -
Plate 26. Plrius keslya Plate 27. P. roxburghii Plate 28. P • sylvestris
(Left s mesophyll tissue; Eight 5 vaseular tissue)
PLATE-29
- 83 -
Plate 29. Pinus wallichiana Plate 30, Tsuga dumosa
(Left 5 mesophyll tissue; Eight * vascular tissue)
Plate 31. Podocarpus macrophvlla£ Plate 32. P. neriifolius
(Left 5 mesophyll tissue;Right 5 vascular’ tissue )
PLATE-34
85 -
Plate 33. Podocarpus sinensis Plate 34. Taxus baecata
(Left t mesophyll tissue; Right : vascular tissue )
■» 86 ■*
Plate 35. Cryptomeria japonica Plate 36. Cunnlnghamia sinensis(Left s mesophyll tissue; Eight i vascular tissue )
- 87
Plate 37. Plate 38.
(LeftRight
Metasequoia glyptostroboides Taxodium distiehums mesophyll tissue; t vascular tissue )
\
- 88 -Key to the families
A. Cuticle thin, undulated; hypodermis absent; endodermls distinct; resin ducts more than 2 .. Ginkgoaeeae
AA. Cuticle thick, not undulated; hypodermis present; endodermis not distinct; resinducts 0-2 .. Cycadaceae
A. Resin duct absent. Hypodermis absent Taxaceae
AA. Resin ducts present,B, Vascular bundles 1,
C. Accessory transfusion tissue present; hypodermis present, discon-
1 tinuous; cuticle not undulatedCC. Accessory transfusion tissue
absent; hypodermis absent; cuticle undulated
BB. Vascular bundles 1-C. Vertically winged or laterally
winged.D. Transfusion tissue lateral
to vascular bundle and one of the resin ducts lying below the vascular bundle; mesophy11 with no air space
DD. Transfusion tissue all round vascular bundle; resin duct lying not below the vascular bundle; mesophyll with airspaces .. Araucariaceae
Podocarpaceae
Cephalotaxa-ceae
Taxodiaceae
CC. Triangular; vertically or laterally winged.
- 89 -
D, Resin ducts 2-9$ plicate parenchyma present in some; mesophyll not lax
DD. Resin ducts 1-3; no plicate parenchyma; mesophyll lax
CYCADACEAl
A, Vascular bundles 1 or 2; accy. transfusion tissue present; resin ducts 0-2
AA. Vascular bundles more than 2; accy. transfusion tissue absent; resin ducts 0.B. Bundle-sheath extension absent;
stomatal type hypocyticBB. Bundle-sheath extension present;
stomatal type paracytic
Eej^to^^e^Sgecies
A. Resin ducts 0-2;’ transfusion tissue lateral to vascular bundle.B. Vascular bundles l; dome-shaped
stomata presentBB. Vascular bundles 2; dome-shaped
stomata absentAA. Resin ducts *0; transfusion tissue all round
vascular bundle.B. Spongy parenchyma compactly arranged
with no air space; bundle-sheath extension absent; stomatal type hypocytic
Pinaceae
Cupressaceae
Cvcas
Encenhalartos
Zamia
Cvcas revoluta
C. rumphii
Encephalartosvillosus
- 90 -
BB. Spongy parenchyma loosely arranged with air spaces; bundle-sheath extension present; stomatal type paracytic
ARAUCARIACEAE Key to the Genera
A. Hypodermis of one to two layers; lumen of epidermal cell broad
AA. Hypodermis of one layer or in bundles; lumen of epidermal cell narrow
Species
A. Resin ducts 1 or 2; vertically winged.B. Resin ducts 1; vascular bundles 3;
hypodermis of one layer, less developed; mesophyll with less air spaces
BB. Resin ducts 2; vascular bundles 1; hypodermis of one to two layers, well developed; mesophyll with prominent air spaces
AA. Resin ducts more than 2; laterally winged.B. Hypodermis well developed in bundles;
lumen of epidermal cell narrow ..BB. Hypodermis less developed in one or
two layers; lumen of epidermal cell broad ..
Zamiaangustifolia
Agathls
Araucaria
Araucaria columnaris
A.cunninghamii
A. bidwillli
Agathlsloranthifolia
- 91 -CEPHALOTAXACEAEfegBjtoatth0a-Sgecl§s
A. Epidermal wall less undulated; cellsheterogenous; lumen broad; cuticle thin; stomatal type hexaeytie; epithelial cells 10.
AA. Epidermal wall more undulated, cellshomogeneous; lumen.narrow; cuticle thick; stomatal type paracytic; epithelial
, cells 13-16.
CUPRESSACEAE Key to the Genera
A. Mesophyll fully differentiated.B. Mesophyll compact
BB. Mesophyll lax.C, Transfusion tissues all round
vascular bundleCC. Transfusion tissue lateral to
vascular bundle.D. Triangular or triangular with
concave surface; transfusion tissue forms wing-like structure towards leaf margin
DD. Laterally winged; transfusion tissue forms no wing-like structure
Cephalotaxusdrupacea
C. griffithii
Callitris
Thu.lopsis
Cupressus
Junineru3AA. Mesophyll undifferentiated or slightly
differentiated.
- 92 -
B. Endodermis not-distinct; hypodermis less developed; lumen of epidermal cell broad; -transfusion tissue allround vascular bundle •• Chamaecyparis
BB. Endodermis slightly distinct; hypo- dermis well-developed; lumen of epidermal cell narrow; transfusion tissue lateral to vascular bundle .. Thu.ia
Key to the SpeciesI» «J"J LE«-Im|»LLLg>
A. Eesin ducts 1.B. Hypodermis continuous.
C. Hypodermis well developed; resin duct at the leaf margin; laterally winged;nypostomatic, anomocytic .. Juniperus
wallichlana
CG. Hypodermis less developed.Differentiated with two to three layers of-palisade tissue, /Lax; resin ducts just below thephloem t .. Cupressus
ca shiner iana
BB. Hypodermis discontinuous.C. Besin ducts at the leaf margin
D. Vascular bundles 2; meso- phyll undifferentiated; epithelial cells 7-8 Chamaecyparis
plsifera• •
- 93 -
DD. Vascular bundles 1; mesophyll differentiated,E. Hyperstomatie, paracytie;
resin ducts at the concave region of leaf margin; epithelial cells 26; transfusion tissue forms wing-like structures towards the leaf margin; triangular with concave surface ., Cunressus
torulosaEE, Amphistomatie, anomocytic;
resin ducts at the apical region of leaf margin; epithelial cells 16; transfusion tissue forms no • wing-like structures; surface laterally winged Juniuerus
recurvaCC. Resin ducts just above the phloem.
Differentiated with one layer ofpalisade tissue, lax; vascularbundle transversely placed withpalisade tissue .. Thujonsis
dolabrataAA. Resin ducts more than 1,
B, Mesophyll undifferentiated or slightly differentiated,G, Hypodermis of one layer, discon
tinuous; stomatal type complex; resin ducts 1-2 at the region of leaf margin •. Thuja
OccidentallsCC. Hypodermis of one to two layers,
continuous; stomatal type actlno- cytic; resin ducts 3 at the leaf margin ., Thuja
oriental!s
- 94 -
BB. Mesophy 11 differentiated.- .Amphistomatic; hypodermis absent; cuticle thin; mesophyll differentiated with little palisade tissue1; endodermis distinct
PINACEAB ; ^ytotheG|Mr|
A. Mesophyll not differentiated.B. Plicate parenchyma present.
C. Semicircular, triangular other;than laterally winged; transfusion . , tissue all round vascular bundle ..
CC., Laterally winged; transfusion !tissue lateral to vascular bundle ..
BB. Plicate parenchyma absent; verticallywinged ..
AA, Mesophyll differentiated.B. Vascular bundles i;‘ hypodermis absent;
resin ducts l,1 *’
BB. Vascular bundle- 2; hypodermis present;resin ducts 2 .,
I ’ V
122. Jo^J^^SpeciesA. Mesophyll differentiated.
B. Basin ducts 1; epithelial cells 14; -endodermis not distinct; hypodermis absent ..
Callitrisrobusta
Pinu3
Cedrus
Picea
Tsuga
Abies
Tsuga dumosa
- 95 -
AA.
BB* Resin duets 2; epithelial cells 33; endodermis distinct; hypodermis present
Mesophy11 not differentiated.
B. Plicate parenchyma present.C. Resin ducts 2.
D. Hypodermis continuous,1, Laterally winged; cuticle
thick; transfusion tissue lateral to vascular bundle; epithelial cells 21 ..
EE. Triangular; cuticle thin; transfusion tissue all round vascular bundle; epithelial cells 14-15
DD. Hypodermis discontinuous.Triangular; upper layer slightly curved; hypodermis less developed; transfusion tissue all round vascular bundle; epithelial cells 6-7 ..
CC. Resin ducts more than 2.D. Endodermis distinct.
1. Cuticle thick; semicircular; hypodermis of one layer,well developed resin ducts 9; epithelial cells 10-11 ..
EE. Cuticle thin; triangular, upper layer slightly convex; hypodermis of one to three layers; less developed; resin ducts 3-4; epithelial cells 8 ^ ..
Abies3Pectabilis
Cedrus libani
Plnus"‘wallichiana
P. kesiva
P. densiflora
P. roxburghii
- 96 -, tDD, Endodermis not distinct,
E, Cuticle thick; semilunar;- • hypodermis in bundles, well developed; resin ducts 4, epithelial cells 8 .,
EE, Cuticle thin; crescentshaped; hypodermis or one- layer, less developed; resin ducts 6; epithelial cells 14 . •
BB. Plicate parenchyma absent; vertically winged
' PODOCARPACEAE Kev to the Species
A, Resin ducts 1; hypodermis well developed; cuticle thin"; hypodermis in bundles; transfusion tissue all round vascular bundles, mesophyll differentiated with abaxlal palisade; stomatal frequency 19251,0
AA. Resin ducts more than 1; hypodermis less developed, *B, Resin ducts 2, epithelial cell 7;
hypodermis of one layer; stomatal frequency 8445,6
BB, Resin ducts 3, epithelial cells 18-10; hypodermis of one to two layers; sto- matal frequency 12171,6
P. gerardiana
P. svlvestris
Picea morinda
Podocarnussinensis
P. neriifolius
P. macrophvlla
97 -TAXODIACEAE
Key to the Genera
A, Basin ductr 1.B. Vertically winged; amphistomatie
BB. Laterally winged; hypostoraatieAA. Besin ducts more than 1.
B. Hypodermis present; endodermis distinct
BB• Hypodermis absent; endodermis not distinct
CryptomeriaTaxodium
Cunninghamia
Metasequoia
Key to the Spe„cie,s
A. Besin ducti lvB. Mesophyll slightly differentiated;,
cuticle thick; amphistomatie; actino- cytic; hypodermis one to two layers
BB. Mesophyll differentiated; cuticle thin; hypostomatlc; anomocytic; hypodermis 0
AA. Besin ducts more than 1,B. Hypodermis of one to four layers;
epidermis distinct; epithelial cells27; lumen of epidermal cell narrow .. Cunninghamia
sinensis
BB. Hypodermis absent; endodermis notdistinct; epithelial cells 10; lumen
, of epidermal cell broad .. Metasequoiaglyptostro-boides
Cryptomeria.laponica
Taxodiumdistichum
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