7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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International Research Journal of Plant Science (ISSN 2141-5447) Vol 4(6) pp 168-191 June 2013Available online httpwwwinteresjournalsorgIRJPSCopyright copy 2013 International Research Journals

Full Length Research Paper

Anatomical studies on the genus Euphorbia L SaudiArabia (Subgenera Triucalli Ermophyton Esula andChamaesyce )

Aldhebiani A1 and Jury S2

1Department of Biology King Abdulaziz University Jeddah Saudi Arabia

2Centre for Plant Diversity and Systematics University of Reading

Corresponding Author Email aaldhebianikauedusa

Abstract

The genus Euphorbia is the largest in Saudi Arabia even though no anatomical study has been doneintensively In this study the epidermis the stomata and the venation patterns have been investigatedThe shape of the epidermal cell in Euphorbia species in Saudi Arabia varies polygonal rectangularundulate or elongated Moreover the cell shape relies on the cell location on a leaf ie the middleregion the margin the apex or above the vein Furthermore in some cases both leaf surfaces have thesame cell shape but more often they are unlike Hairs are generally simple unbranched and with awarty ornamentation on their surface Papillae occur only in one species E hypericifolia The mostcommon stomata type is anomocytic while the rare type is actinocytic recorded only in E helioscopia Stomata of more than one type (have been encountered on the same leaf surface as in E scordiifolia and E hirta Venation patterns vary from one-veined three-veined to those with four or more veins

Keywords Euphorbia Saudi Arabia stomata venation epidermal cells anatomical characters

INTRODUCTION

Euphorbiaceae the spurge family one of the majorflowering plant families with 334 genera grouped in 52tribes and 5 subfamilies is considered as the sixthlargest family of Angiospermae In Saudi ArabiaEuphorbiaceae is represented by 15 genera (AndrachneL Flueggea Willd Phyllanthus L Clutia L ChrozophoraNeck ex Juss Ricinus L Mercurialis L ErythrococcaL Micrococca Benth Acalypha L Tragia LDalechampia L Jatropha L Croton L and Euphorbia L)(Chaudhary 2001) Among them Euphorbia is the

largest and varies from herbs to shrubs and trees andfrom succulent to non-succulent plants Species arescattered all over the country but the succulent taxamostly occur in the South West Region According toGovaerts et al (2000) the genus Euphorbia L is thethird largest genus in the flowering plants (afterAstragalus [Fabaceae] and Psychotria [Rubiaceae]) withabout 2000 species distributed worldwide both in Oldand New Worlds and mainly in the tropical subtropicaland warm temperate regions In addition to the widedistributional range Euphorbia L has various life formsas annual or perennial herbs shrubs or trees The

species can be prostrate or erect monoecious odioecious and succulent or non-succulent Most of thesucculent Euphorbia species are endemic to Africa

Even though the genus Euphorbia is the largest inSaudi Arabia no extensive study on the anatomy of thegenus has been done

This work has been done to revise the genusEuphorbia in Saudi Arabia in three parts morophologicalanatomical and phytochemical studies Thephytochemistry study has been published in a separate

paperIn the Euphorbiaceae anatomical characters havebeen found significant since the work of Pax (1884) Inhis work Pax found the importance of laticifers as acharacter to define natural groups In addition he usedthe type of laticifers (articulate or non-articulate) phloemcharacters and trichome type as the major anatomicafeatures to support his redivison of the majosuprageneric taxa in the family The first significananatomical survey of the family was provided bySolereder (1899) who revised the groups of Radlkofer(1870) and added many observations by himself Then

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Gaucher (1902) tried to omit all the work of the Germanschool by publishing an independent anatomical surveyof the family His work was criticized by Solereder(1908) Yet it was valuable in providing a comparativereview of anatomical characters which was arranged bytribe Later in a series of studies Mahlberg (1973 1974

1975 1982) and Mahlberg et al (1987) showed thesystematic importance of laticifers in Euphorbia especially with regard to the starch grains producedFurthermore the vascular anatomy of petioles has beenstudied in cross section by Dehay (1935) who found aconsiderable variation in stellar configurations Howeverit seems unlikely that this feature will be of general valueMeanwhile Miller and Webster (1962) have useddifferences in petiolar steles to separate Cridoscolus fromJatropha Then Dehgan (1982) could prove that thepetiolar stellar dissection were significant at the sectionaland sub-sectional level in some genera such asJatropha However in a considerable number of taxastipules have become reduced or are early deciduousFor example many species lack stipules whereas theyare large and remarkable in others In Euphorbia thepresence or absence of stipules is a diagnostic characterfor some sections and subgenera (Webster 1994a)

Anatomical Characters in Euphorbia

Latex

The white latex is a useful distinguishing character in thegenus Euphorbia L It is distributed throughout the plantin a series of tubes derived from either single cells (non-

articulated laticifers) or articulated laticifers formed by thefusion of several cells The value of laticifer types as ataxonomic marker in systematic comparisons betweenand within families has been established by Carlquist(1961)The laticifer system of the mature Euphorbia plantwas explained by Gaucher (1898 1902) In additionRosowski (1968) studied the branched non-articulatedlaticifers system in mature tissue of the internode andnode in transition from the node to and throughout themature leaf of Esupina Raf He found that the latexsystem in the E supina stem is restricted to the cortexand does not break through the leaf gaps In the leaflamina he found that the widest laticifers are in

association with the vascular system These laticifersbegin to branch from the base of the leaf and continuethroughout certain areas in the mesophyll Moreoverthey are associated with the phloem and may send somebranches at right angles to the vein and between thebundle-sheath cells

Epidermis

Epidermal cells may vary greatly in size shape and

Aldhebiani and Jury 169

outline from species to species especially when seenin a surface view Generally in dicotyledons epidermacells have irregular shapes and sizes Sometimes theshape of the cells in both leaf surfaces are similar butmore often they are different The costal cells usuallydiffer from those intercostal regions they tend to be

elongated in the direction of the veins Some marginacells develop unicellular or multicellular prickles Moreoveranticlinal walls can be either very thin and hardly visible fromthe surface or they may range through degrees of thicknessto be very thick (Kakkar and Paliwal 1972)

One of the important epidermal features for altaxonomists is the hairs They can be glandular or non-glandular and can be divided depending on thecomponent number of cells and degree of branching Theoccurrence of distinctive types of hair can be a valuablecharacter to recognize a whole family Moreover hairsare more useful in determination at the level of genus ospecies In other words variations in size and densityshould be accepted in the differentiation of closely related

genera or species after a comprehensive investigation ofa wide range of material (Metcalfe and Chalk 1950)

In a study of 150 species of Euphorbia Kakkar andPaliwal (1974) described the epidermal cells in Euphorbiaas circular trapezoidal rectangular or polygonal inoutline Additionally they found that the shape may varydepending upon the location of the epidermal cell on theleaf ie the middle region margin apex or above thevein Some xeromorphic species exhibit a waxy coveringwhich takes on many crystalline forms Hairs were alsorecorded in the epidermis of Euphorbia in some speciesIn addition the papillae occurred in the surface view asrounded structures in the centre of cell lumen

Stomata

A stoma consists of the stomatal aperture and the pair oguard cells that form it Stomata usually tend to be on thelower surface only But in some cases this distributionvaries from species to species and depends on whethethe plant is a xerophyte or a mesophyte They might besuperficial or sunken Stomata sometimes aresurrounded by specialized epidermal cells which arecalled subsidiary cells These subsidiaries differ fromunmodified epidermal cells in shape size and staining

properties (Baranova 1992 Metcalfe andamp Chalk 1950Stace 1965) On the other hand the arrangement osubsidiary cells where present is of the greatest interesto the taxonomist This variation is used to define thedifferent types of stomata Occasionally species haveseveral types of stomata on one leaf while some haveonly one type for the species (Stace 1984) In additionVan Cotthem (1973) pointed out that thosemorphological stomata types can provide not onlydiagnostic characters but also very valuable taxonomicones or even phylogenetic clues Metcalfe and Chalk

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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170 Int Res J Plant Sci

(1950) had established some terms to replace therepresentative lsquofamilyrsquo name proposed by Vesque(1889) Anomocytic was substituted for theranunculaceous type anisocytic replaced cruciferousdiacytic the caryophyllaceous and finally paracytic for therubiaceous The tetracytic type which can be found in

most of the monocotyledons was added by Metcalfe(1960) Later Stace (1965) proposed the term cyclocyticfor the narrow ring of four or more subsidiary cellssurrounding the stomata Metcalfe and Chalk (1950)have named and defined the actinocytic type as stomatasurrounded by a circle of radiating cells Three moretypes were introduced by Van Cotthem (1970)hexacytic epicytic and hemiparacytic And someintermediate types were added by Payne (1970) whodescribed the helicocytic and allelocytic types in relationto mesogenous forms of anisocytic paracytic and diacyticpatterns Stace (1989) lists 35 types of stomata invascular plants Closely related families are distinguishedby the presence of a specific type of stomata such asAcanthaceae and Scrophulariaceae separated by thepresence of diacytic stomata in the former as againstanomocytic in the latter Moreover some stomatal types aredistinctive of certain families for example Ranuculaceaehas the anomocytic type Brassicaceae the anisocyticCaryophyllaceae diacytic Rubiaceae paracytic and finallyPoaceae has the graminaceous type (Singh 2004)

According to Metcalfe and Chalk (1950) the maturestomata of Euphorbiaceae are anomocytic paracytic andanisocytic They are usually confined to the lower leafsurface more rarely on both surfaces of the laminaParacytic stomata were reported by Tognini (1897)which are mesogenous in development in E variegata

and Ricinus communis On the other hand according toRaju and Rao (1977) stomata in the Euphorbiaceaeshow considerable variation They found that the woodytaxa have predominantly the paracytic stomata typewhile the anisocytic stomata are characteristic of theherbaceous Phyllanthoideae Moreover they indicatedthat Chamaesyce has a high percentage of anomocyticstomata (Raju and Rao 1987) Finally a considerablediversity of stomatal types were found in Euphorbia byKakkar and Paliwal (1974) They reported that thecommon type of stomata in Euphorbia species is theanomocytic even though stomata of paracytic andanisocytic have been also observed

Venation

Some features that give the leaf its structure are forinstance leaf shape margin type and venation patternsThese together form the leaf architecture as discussed byHickey (1979) A more sophisticated knowledge of leafarchitecture has allowed a start in discriminatingphylogenetic trends from leaves (Hickey 1973 Roth-Nebelsick et al 2001) Due to its importance especiallyfor systematic classification attention has been paid

largely to the architectural properties of leaf venation(Koumlhler 1993) The careful description of venation incooperation with studies of other leaf anatomy details canprovide valuable taxonomic evidence The works of Pray(1955a 1955b) suggest that the venation can be useful ina comparative analysis Moreover Wagner (1979) has

proved that the vein patterns were significant forclassification at the level of species genus and family inferns Of course this is more noticed and observed inangiosperms especially dicotyledons where thehierarchical network patterns of the veins are much moreadvanced and complicated

Leaf venation has been neglected for a long time inthe Euphorbiaceae until the work of Levin (1986a1986b 1986c) Levin developed a new insight intorelationships within subfamily Phyllanthoideae Nodetailed studies have been provided for othesubfamilies except the work of Sehgal and Paliwa(1974) on the tribe Euphorbieae In part II of their studyon the leaf anatomy of Euphorbia Sehgal and Paliwadesignated the venation in petiolate leaves as uni- bi-and tri-veined according to the number of strandsentering the petiole or lamina base They noticed that inthe 150 species under investigation the majority bearleaves belonging to the tri-veined category Moreoverthey divided the latter into ornamented andunornamented veins When the veins are surrounded bya parenchymatous sheath it is called ornamented Andwhen this sheath is absent it is unornamented Besidessometimes trachedial nodules are encountered at theapices and in the serrations of the leaf Small veinsusually form networks in the lamina These networksmay vary in size and shape and subdivide the area o

the mesophyll The thinnest branches of the bundlesbounded by the smallest areas or regions are calledareoles These areoles usually contain blind veinendings The degree of branching of these vein endingsvaries in the leaves of different species Therefore thiscan be a useful identification tool

MATERIAL AND METHODS

Herbarium specimens were obtained from Kew (KewGarden herbarium UK) and RIY (National Herbarium inRiyadh) Since leaves are early deciduous in succulen

species most of the herbarium sheets or spirit collectionsof these species have no leaves As a result thesespecies were omitted from this anatomy study except fothe stick-like Euphorbias E balsamifera subspadenensis E schimperi ldquoE aff schimperirdquo ldquoE affconsobrinardquo and E cuneata Only mature leaves wereused for studying the epidermis stomata and venation

Epidermal peel

This method was used to observe stomata epidermal

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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cells and hairs1- Leaves were rehydrated using polyoxyethylenesorbitan monolaurate (Tween20) in water and heated for30 minutes Then washed by water and conserved in70 ethanol for future work2- Both surfaces of the leaf were scraped and

peeled off under a dissection microscope with the aid offine forceps and a razor blade to remove loose cells3- Samples were bleached by Jefferyrsquos Solution orthe bleaching agent Vortex for 10-20 minutes dependingon leaf thickness Sometimes this was overnight4- Samples were washed mounted and observedunder a light microscope5- Photographs were taken using a Leitz Diaphlanpolarizing microscope or a Reichert Polyvar 2Microscope

Staining by alucin blue was tried but withoutsuccess Samples became too dense or obscure due tothe presence of tannins and other compounds in theherbarium specimens Therefore some samples were leftwithout staining

Leaf clearing

The following method was adopted from Radford et al(1974) with some modifications for use on herbariummaterials1- The leaf was placed in a Petri dish and coveredwith 5 Sodium Hydroxide (NaOH) The dish waswrapped in cling film leaving a small gap in one area forventilation2- This was microwaved for five seconds on

medium power3- The Petri dishes were left on a hot plate at 35-37degrees centigrade for a week or until the leaf wastransparent The NaOH was changed twice a day foreach sample4- When the leaf had cleared sufficiently it waswashed in water and covered in 90 bleaching agentVortex for ten minutes to one hour depending on thereaction of the sample5- The leaf was washed again in water6- Samples were dehydrated in ethanol series andstained as followinga) 30 ethanol 2 minutes

b) 50 ethanol 2 minutesc) 1 safranin in 50 ethanol 4 minutesd) Wash in water three timese) 30 ethanol 2 minutes repeated 3 timesf) 50 ethanol 2 minutes repeated twiceg) 70 ethanol 2 minutesh) 90 ethanol 2 minutesi) 100 ethanol 2 minutes

j) Absolute ethanol histoclear 11 2 minutesk) Histoclear 2 minutes

Aldhebiani and Jury 171

l) Mounted in 100 glycerol and sealed with Candabalsamm) Observed under light microscopen) Photographs were taken using a Leitz Diaphlanpolarizing microscope or Reichert Polyvar 2 Microscope

Scanning Electron Microscopy

1- Dry specimens were mounted surface up onscanning electron microscope stubs using Bostik No1adhesive2- Stubs were sputter coated for 2-3 minutes with agold palladium alloy using an Edwards sputter coater togive a coating about 15-20 nm thick3- Samples were examined in FEI Quanta FEG 600Environmental Scanning Electron Microscope (ESEM) inhigh vacuum SEM mode4- Photographs were taken using the computerizeddigital system of the microscope

RESULTS

Epidermis

Results of the epidermal characters in Euphorbia speciesunder investigation are summarized in Table 1

Cell shape

Depending upon the shape of the cell wall and

considering the results from Kakkar and Paliwal (1974)the shape of the epidermal cell varies polygonalrectangular undulate or elongated Moreover the celshape relies on the cell location on a leaf ie the middleregion the margin the apex or above the veinFurthermore in some cases both leaf surfaces have thesame cell shape but more often they are unlike Thiscombination of cell shape characters gives each speciesits unique leaf surface appearance

The normal straight cell walls in both adaxial andabaxial surfaces are shown in the following taxa Eschimperi E cuneata E helioscopia E dracunculoidesand E balsamifera subsp adenensis see Figure 1 In

addition ldquoE aff schimperirdquo and ldquoE aff consobrinardquo havealso the same straight cell wall on both surfacesMeanwhile E grossheimii E retusa and E pirottae haveelongated cells on both leaf surfaces (Figure 2 a and b)On the other hand the plicate cell walls on both adaxiaand abaxial surfaces occur in E chamaepeplus Ehypericifolia and E peplus (Figure 2 c and d) but thewalls are more folded on the lower surface In somespecies the cell wall on the adaxial surface is straightwhilst it is plicate on the abaxial such as E hirta E

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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172 Int Res J Plant Sci

Table 1 Epidermal characters of Euphorbia taxa in Saudi Arabia

Ab abaxial + PresentAd adaxial ++ the plicate folded wall is morefully developed in the lower surface than in the upper one

+ hardly observable by LM only

Subgenus

Cell shape (cell wall) Hairs

Straight Elongated Plicate Unicellular MultSpecies Ad Ab Ad Ab Ad Ab Ad Ab Ad T

i r u c al l i

E balsamifera subspadenensis + + +

E schimperi + +

ldquoEaff schimperirdquo + +

ldquoE aff consobrinardquo + +

E cuneata + +

ErmophytonE acalyphoides + + +

E pirottae + + +

E helioscopia + + E

s ul a

E grossheimii+ +

E retusa + +

E dracunculoides + +

E falcate + +

E peplus + ++

E chamaepeplus

E schimperiana +

+ ++

+ C h am a e s y c e

E Arabica + +

E serpens + +

E scordiifolia + + +

E granulate + + + + +

E inaequilatera + +

E hirta + + +

E hypericifolia + ++

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 173

983106

983108

983109

983105

983107

983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

Figure 1 Normal irregular epidermal cells in both adaxial and abaxial surfaces of the leaf (LM)(A) E balsamifera subsp adenensis (B) E schimperi (C) E cuneata (D) E falcata (E) Abaxial in E dracunculoides

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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174 Int Res J Plant Sci

983105

983106

983107

983108

983106 983106

983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

Figure 2 Error No text of specified style in document Both adaxial and abaxial surfaceshave the same shape of epidermal cells (LM) Elongated in A ampB plicate in CampD (A) Egrossheimii (B) E pirottae (C) E chamaepeplus and (D) E peplus

schimperiana E helioscopia E inaequilatera Earabica E granulata E serpens and E scordiifolia see Figure 3 and Figure 4 This last group of species belongto subgenus Chamaesyce except for E helioscopia and E schimperiana

Hairs

Hairs on the leaf epidermis can be both unicellular and

multicellular and may appear only on one surface or bothFurthermore hairs in the Euphorbia species undeinvestigation are generally simple unbranched and with awarty ornamentation on their surface Papillae appear inESEM as outgrowths whereas in the surface view theygive the impression of being rounded structures at thecentre of the cell Prominent papillae occur only in onespecies of Euphorbia in Saudi Arabia this species is Ehypericifolia (Figure 5 and 6) whereas in E arabica theywere poorly distinguishable using LM

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 175

983105983140983137983160983145983137983148

983107

983108

983105

983106

983105983138983137983160983145983137983148

Figure 3 Epidermal cells vary on leaf surfaces adaxial surface has irregular cells whilethe abaxial surface has plicate (LM)(A) E schimperiana (B) E arabica (C) E serpens (D) E scordiifolia

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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176 Int Res J Plant Sci

983105983140983137983160983145983137983148

983107

983105

983106

983105983138983137983160983145983137983148

Figure 4 Epidermal cell shape varies on the adaxial andthe abaxial surfaces straight cell walls in the formerwhile folded plicate walls occur on the latter (LM) (A) Egranulata (B) E inaequilatera and (C) E helioscopia

983112983141983149983145983152983137983154983137983139983161983156983145983139 983155983156983151983149983137983156983137

983151983150 983138983151983156983144 983155983157983154983142983137983139983141983155

Figure 5 E hypericifolia (LM)Above photo the adaxial surface with irregular epidermal cells andvery clear papillae as a circle in the middle of the cellLower photo the abaxial surface with plicate cells and papillae ahair base is obvious in the right side of the picture

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 177

983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

983120983137983152983145983148983148983137983141

983117983157983148983156983145983139983141983148983148983157983148983137983154 983144983137983145983154983155 983145983150 983137983138983137983160983145983137983148 983155983157983154983142983137983139983141983086

Figure 6 E hypericifolia (ESEM) papillae on the adaxial and abaxial surfacesmulticellular hairs are on the abaxial surface of the leaf only

Figure7 Multicellular hairs on adaxial and abaxial surfaces of E hirta

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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178 Int Res J Plant Sci

Figure 8 Hairs wart ornamentation on hair surfaces of (A) E scordiifolia and (B) E pirottae (C) Short hairs onabaxial surface of E balsamifera subsp adenensis

In E balsamifera subsp adenensis the hairs are shortand unicellular (Figure 8C) and they are located mostly atthe leaf margins and the mid-vein area on the abaxialsurface only

Hairs on the leaf of E pirottae are present on theabaxial surface only and are both unicellular andmulticellular Euphorbia granulata is similar to E pirottae in having both unicellular and multicellular hairs but herethey occur on both surfaces of the leaf (Figure 8B)

On the other hand in E scordiifolia E hirta and Ehypericifolia the hairs are multicellular and on bothsurfaces in the first two species and on the abaxial only inthe latter (Figure 7) Moreover E hypericifolia and Ehirta have long hairs when compared with the otherspecies (Figure 6)

No hairs were observed and leaves were glabrous inthe following species E serpens E arabica E peplusE inaequilatera E falcata E helioscopia E cuneata Eretusa E grossheimii E schimperiana E schimperiE chamaepeplus and the new variety of E granulata

Wax

The cuticle layer and wax in the Euphorbia species undeinvestigation vary from smooth to densely cover in waxscales This may be a thick smooth layer of wax whichsometimes has upright scales In some species only thecell surface has a smooth wax layer whereas the walbetween cells has upright scales of wax as in Eschimperiana especially around the stomata pores

(Figure 9) In contrast the anticlinal walls of somespecies have a smooth layer of wax whereas the celsurface is densely covered with upright wax flakes or withsome stellate wax flakes in other species

Stomata

Stomata were observed through LM and ESEM In thespecies under investigation stomata commonly appeaon both leaf surfaces (amphistomatic leaves) excepin E cuneata where the stomata are located only on the

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 179

983109

983105

983108983107

983106

Figure 9 Wax formation in Euphorbia species in Saudi Arabia (A+B) smooth andthick in E peplus and E schimperi (C+D) smooth on top of cell while formingupright scale flakes in between especially around stomata as in E schimperiana smooth in anticlinal walls (E) upright scale flakes in E balsamifera

abaxial surface (hypostomatic leaves) The presence andposition of subsidiary cells are variable in Euphorbia

species Subsidiary cells were found to be of unequalsize and shape whether the stoma was anisocyticparacytic actinocytic or tetracytic While some specieslack them around the stomata others have more thanone arrangement of subsidiary cells

The most common stomata type is anomocytic whilethe rare type is actinocytic recorded only in Ehelioscopia Moreover sometimes stomata of more thanone type (up to three or four types) have beenencountered on the same leaf surface as in E scordiifolia (three types on the abaxial surface) and E hirta (five

types on the abaxial and three on the adaxial) Thedistribution and type of stomata are shown in Table 2

In addition in some species such as E falcata and Ehelioscopia the stomata are deeply sunken andrepresented only by cuticular ledges surrounding theminute pores as seen in ESEM

Examples of species lacking subsidiary cells aroundstomata are E peplus E chamaepeplus E granulataE dracunculoides and E acalyphoides Therefore theyall have anomocytic stomata

On the other hand E balsamifera subsp adenensishas the mononcytic type on both leaf surfaces andanisocytic and paracytic only on the abaxial surface In

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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180 Int Res J Plant Sci

Table 2 Stomata type in Euphorbia taxa in Saudi Arabia

Ab abaxial surface + stomata presentAd adaxial surface

addition E schimperiana has anomocytic andtetracytic on the abaxial only In contrast Ecuneata has three types of stomata anomocyticparacytic and anisocytic only on the lower surfaceAs well ldquoE aff consobrinardquo ldquoE aff schimperirdquo

and E schimperi have paracytic and anisocyticonly in the abaxial on the first two and in both leafsurfaces on the latter Therefore ldquoE affconsobrinardquo and ldquoE aff schimperirdquo are the same taxa Then comparing to E schimperi results

from ansubspec

Eupsimilar and an

Subgenus

Stomata

Anomocytic Hemiparacytic Paracytic Anisocytic T

Species Ad Ab Ad Ab Ad Ab Ad Ab AT i r u c al l i

E balsamifera subsp adenensis + + +

E schimperi + + + +

ldquoEsp aff consobrinardquo + +

ldquoE sp aff schimperirdquo + +

E cuneata + + +

ErmophytonE acalyphoides + +

E pirottae + + + +

E helioscopia + + E s ul a

E grossheimii + + + + +

E retusa + + + + +

E dracunculoides +

E falcate + + E peplus +

E chamaepeplus + +

E schimperiana + C h am a e s y c e

E Arabica + + + + + + +

E serpens + + + +

E scordiifolia + + + + + + +

E granulate + + + +

E inaequilatera + + + +

E hirta + + + + + + +

E hypericifolia + + + + + +

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Table3 Venation type in Euphorbia taxa in Saudi Arabia

Subgenus Species

Venation

Number of veins atthe leaf base

Margins Sheathed

1 2 3 4 or more Open Closed Rounded T i r u c al l i

Ebalsamiferasubspadenensis

+ + +

E schimperi + + +

E aff Schimperi + + +

E cuneata + + +

ErmophytonE acalyphoides + + +

E pirottae + + E s ul a

E grossheimii + +

E retusa + + +

E dracunculoides + + +

E falcate + + +

E peplus + +

E chamaepeplus + +

E schimperiana

E helioscopia

+

+

+

+ C h am a e s y c e

E Arabica + + +

E serpens + + +

E scordiifolia + + +

E granulate + + + +

E inaequilatera + + +

E hirta + + + E hypericifolia + + +

Poinsettia E cyathophora + +

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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182 Int Res J Plant Sci

Figure 10 Sunken stomata on the abaxial surface1) Deeply sunken (A) E helioscopia (B) E falcata (C) E schimperiana 2) Slightly sunken (D) E peplus (E) E pirottae

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 183

983105983138983137983160983145983137983148

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983105983139983156983145983150983151983139983161983156983145983139

983105983140983137983160983145983137983148

Figure 11 Stomata types in Euphorbia taxa in Saudi Arabia (A) E balsamifera subsp adenensis (B) E schimperi (C) E cuneata and (D) E grossheimii

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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184 Int Res J Plant Sci

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983105983138983137983160983145983137983148983105983140983137983160983145983137983148

Figure 12 Stomata types in Euphorbia taxa (A) E falcata (B) E peplus (C) E chamaepeplus and (D) Eschimperiana

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 185

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Figure 13 Figure 13 Stomata types in Euphorbia taxa in Saudi Arabia (A) E arabica (B) E serpens (C)E scordiifolia and (D) E helioscopia

distribution between the leaf surfaces (see Figure 13)While E hirta has the most types E helioscopia Edracunculoides and E acalyphoides have the least

Regarding the position of the stomata on the surfacelevel of the epidermis a variable degree of sunkenesshas been recorded Using the ESEM the stomata appear

deeply sunken in some species whilst in others they areonly slightly sunken with the outer rims of the subsidiarycells surrounding the aperture The deeply and slightlysunken species are recorded in Table 2 AdditionallyFigure 10 shows the sunken stomata in different speciesunder the ESEM

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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186 Int Res J Plant Sci

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983107

983106

983105

983105983140983137983160983145983137983148983105983138983137983160983145983137983148

Figure 14 Stomata types in Euphorbia taxa in Saudi Arabia (A) E hirta (B) E hypericifolia (C) E pirottae and (D)Espaff schimperi

Venation

The venation patterns in the Euphorbia species underinvestigation vary from one-veined three-veined to thosewith four or more veins The predominant type is three-veined However in E balsamifera subsp Adenensis

there are seven middle parallel veins three middle thickand four thinner lateral ones (Figure 15 A and B)Whereas in E pirottae there are three main middleparallel veins and two thinner laterals (Figure 15 C) Incontrast E chamaepeplus E schimperiana and Earabica have only one thick main midrib coming from the

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 187

983105

983106

983107

Figure 15 Figure 15 Venation patterns at base of leaves invariants with more than three veins (A amp B) E balsamiferasubsp adenensis and (C) E pirottae

petiole then branched from the side into two lateral veinsat the point when the leaf starts widening (Figure 16)Each of these veins branches again into secondary veinsat the leaf margin

The rest of the species have just three veins Egrossheimii E dracunculoides E falcata E peplus Eserpens E scordiifolia E granulata E inaequilateraE helioscopia E hirta E hypericifolia and E

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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188 Int Res J Plant Sci

Figure 16 Venation patterns with one main mid vein at the leaf base in (A) E arabica and (B) E chamaepeplus

983106983105

Figure 17 Venation patterns with three-veined leaves in (A) E falcata and (B) E granulata

cyathophora All the three veins come from the petioleand start branching into secondary veins both toward themargin and the mid vein direction (Figure 17 B)

The vein endings at the leaf margin vary from closed

to open There is a relationship between the closedmargins and the sheathed veins (midrib and secondaryveins that are surrounded by parenchyma tissue) Whenthe veins are ornamented (sheathed) the margins areclosed This can be seen in subgenus Chamaesyce inspecies such as E serpens E scordiifolia E granulataE inaequilatera E hirta E hypericifolia and E arabica Additionally these species have the three-vein venation

On the other hand when the veins are not sheathedthe veins ends are open at the leaf margin as in Ebalsamifera subsp adenensis E schimperi E cuneataE acalyphoides E grossheimii E dracunculoides E

falcata E peplus E chamaepeplus E schimperiana Ehelioscopia E pirottae ldquoE aff schimperirdquo and Ecyathophora

CONCLUSIONS

Anatomical characters are seen here to be very useful inEuphorbia classification and identification The resultsobtained from anatomical studies were useful to groupsome species and identify others

Depending on the venation type the Euphorbiaspecies under investigation are divided anatomically intothree main groups see Figure 18

Group one veins are sheathed epidermal cellsdifferentiated on adaxial and abaxial surfaces Group two

983105 983106

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Figure 18 Hierarchial relation of Euphorbia taxa in Saudi Arabia based on anatomical charac

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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190 Int Res J Plant Sci

veins are not sheathed and epidermal cells are alike inboth surfaces And the third group includes species thatdo not belong to any of the previous groups The firstgroup can be further divided into three-veined and one-veined while the second group into three-veined and fouror more veins

In addition E grossheimii E retusa and E pirottae are related by being three-veined open at the marginswith rounded to oval tracheas and elongated cells bothon adaxial and abaxial surfaces However in E retusa the epidermal cells have straight cell walls on the adaxialsurface They are also similar in having hemiparacyticand paracytic types of stomata either on the adaxial orthe abaxial surface

On the other hand species with two differentepidermal cells on the leaf surfaces usually have three-sheathed veins with closed margins and elongatedtrachea such as E serpens E scordiifolia E granulata E inaequilatera E hirta and E hypericifolia which belongto subgenus Chamaesyce according to (Carter andSmith 1988)

In the case of E granulata which has beendescribed as having three different variants E granulata var granulata (both leaf surfaces hairy) E granulata varglabrata (adaxial glabrous and abaxial hairy) and aglabrous variant (the whole plant glabrous) Anatomicalwork has shown especially by using the ESEM that theleaves are either hairy or glabrous on both surfaces Sowhen the leaf is hairy both surfaces have differences inhair distribution Usually the lower surface is more hairythan upper surface (E granulata var glabrata ) butsometimes they are evenly distributed (Egranulata vargranulata ) In addition hairs on both surfaces are

multicellular In contrast when the leaf is glabrous bothleaf surfaces are glabrous and the whole plant isglabrous (suggested new variety) They were similar inhaving the same venation type (three-veined)

Finally ldquoE aff schimperi ldquohas some similarity with E schimperi in having the same open margin not theornamented venation type As well they both have thesame type of stomata paracytic and anisocytic but theydiffer in their distribution on abaxial and adaxial Thusthis may support the suggestion considering ldquoE affschimperi rdquo as a new subspecies of E schimperi

REFERENCES

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Carlquist S (1961) Comparative Plant Anatomy Holt Rinehart ampWinston New York

Carter S Smith AR (1988) Euphorbiaceae (Part2) In Polhill ed Floraof Tropical East Africa Rotterdam Brookfield AA Balkema 409-597

Chaudhary SA (2001) Flora of The Kingdom of Saudi Arabia Vol 3Ministry of Agriculture and Water Riyadh

Dehay C (1935) Lagraveppareil Libero-ligneux foliare des EuphorbiaceaeAnnales des Sciences Naturelles Botanique 10 147-190

Dehgan B (1982) Comparative anatomy of the petiole and infragenericrelationships in Jatropha (Euphorbiaceae) American Journal oBotany 69 1283-1295

Gaucher L (1898) Ē tude anatomique du genre Euphorbia LPKlincksieck Paris

Gaucher L (1902) Recherches anatomiques sur les EuphorbiaceacuteesAnnales des Sciences Naturelles Botanique 15 161-309

Gaucher L (1902) Recherches anatomiques sur les Euphorbiaceacutees

Annales des Sciences Naturelles Botanique 15 161-309Govaerts R Fordin D Radcliffe-Smith A (2000) World Checklist and

Bibliography of Euphorbiaceae The Royal Botanic Gardens KewHickey LJ (1973) Classification of the architecture of dicotyledonous

leaves American Journal of Botany 60 17-33Hickey LJ (1979) A revised classification of the architecture o

dicotyledonous leaves In Metcalfe CR and Chalk L eds Anatomy oDicotyledons Systematic Anatomy of the leaf and stem 2nd edClarendon Press Oxford 25-39

Kakkar L Paliwal GS (1972) Epidermis in Euphorbia Indian ScienceCongress Association Proceedings 59 326-327

Kakkar L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 5 Epidermis Proceedings of the Indian National ScienceAcademy Part B Biological Sciences 40 55-67

Koumlhler E (1993) Blattvervatur-Muster der Buxaceae Dumortierr undSimmondsiaceae Van Tieghem Feddes Repertorium 104 145-167

Levin GA (1986a) Systematic foliar morphology of Phylanthoideae(Euphorbiaceae) II Phenetic analysis Annals of Missouri BotanicaGarden 73 86-98

Levin GA (1986b) Systematic foliar morphology of Phylanthoideae IICladistic analysis Systematic Botany 11 515-530

Levin GA (1986c) Systematic foliar morphology of Phyllanthoideae(Euphorbiaceae) I conspectus Annals of Missouri Botanical Garden73 29-85

Mahlberg P (1973) Histochemistry and Scanning Electron Microscopyof Starch Grains from Latex of Euphorbia-Terracina and EuphorbiaTirucalli Proceedings of the Indiana Academy of Science 82 132

Mahlberg PG (1974) Scanning Microscopic Comparison of StarchGrains from the Latex of Euphorbia -Spp Proceedings of the IndianaAcademy of Science 83 83-84

Mahlberg PG (1975) Evolution of the Laticifer in Euphorbia asInterpreted from Starch Grain Morphology American Journal oBotany 62 577-583

Mahlberg PG (1982) Comparative Morphology of Starch Grains inLatex from Varieties of Poinsettia Euphorbia-PulcherrimEuphorbiaceae Botanical Gazette 143 206-209

Mahlberg PG Davis DG Galitz DS Manners GD (1987) Laticifers andthe Classification of Euphorbia the Chemotaxonomy of EuphorbiaEsula L Botanical Journal of the Linnean Society 94 165-180

Metcalfe C Chalk L (1950) Anatomy of the Dicotyledons At theClarendon Press Oxford

Miller K Webster GL (1962) Systematic position of Cnidoscolus andJatropha Brittonia 14 174-180

Pax F (1884) Die Anatomie der Euphorbiaceen in ihrer Beziehung zumSystem derselben Botanische Jahrbuumlcher fuumlr SystematikPflanzengeschichte und Pflanzengeographie Leipzig 5 384-421

Payne WW (1970) Helicocytic and allelocytic stomata unrecognizedpatters in the Dicotyledonae Am J Bot57 140-147

Pray TR (1955a) Foliar venation of angiosperm II Histogenesis of thevenation of Liriodendron Am J Bot 42 18-27

Pray TR 1955b Foliar venation of angiosperms III Pattern andhistology of the venation of Hosta Am J Bot 42 11-18

Radford AE Dickison WC Massey JR Bell CR (1974) Vascular PlanSystematics Harper amp Row Publishers London New York

Radlkofer L (1870) Ueber Pausandra ein neues EuphorbiaceenGenus Flora 53 81-95

Raju VS Rao PN (1977) Variation in the structure and development ofoliar stomata in the Euphorbiaceae Botanical Journal of the LinneanSociety 75 69-97

Raju VS Rao PN (1987) The Taxonomic Use of the Basic StomataType in the Generic Delimitation of Chamaesyce EuphorbiaceaeFeddes Repertorium 98 137-142

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2424

Rosowski JR (1968) Laticifer Morphology in the Mature Stem and Leafof Euphorbia supina Botanical Gazette 129 113-120

Roth-Nebelsick A Uhl D Mosbrugger V Kerp H (2001) Evolution andfunction of leaf venation architecture a review Annals of Botany 87553-566

Sehgal L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 2 Venation Patterns Botanical Journal of the Linnean Society 68 173-208

Singh G (2004) Plant Systematics An Integrated Approach SciencePublishers Enfield

Solereder H (1899) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Solereder H (1908) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Aldhebiani and Jury 191

Stace CA (1965) Cuticular studies as an aid to plant taxonomy Bulletinof the British Museum Natural History Botany 4 3-78

Stace CA (1989) Plant Taxonomy and Biosystematics Hodder ampStoughton London

Tognini F (1897) Contribuzione allo studio della organogeniacomparata degli stomi Institut de Botanique University de Geneve 41-42

Van Cotthem WR (1970) A classification of stomatal types Botanica

Journal of Linnean Society 63 235-246Wagner W (1979) Reticulate veins in the systematics of modern ferns

Taxon 1 87-95Webster GL (1994a) Classification of the Euphorbiaceae Annals of the

Missouri Botanical Garden 81 3-32

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Gaucher (1902) tried to omit all the work of the Germanschool by publishing an independent anatomical surveyof the family His work was criticized by Solereder(1908) Yet it was valuable in providing a comparativereview of anatomical characters which was arranged bytribe Later in a series of studies Mahlberg (1973 1974

1975 1982) and Mahlberg et al (1987) showed thesystematic importance of laticifers in Euphorbia especially with regard to the starch grains producedFurthermore the vascular anatomy of petioles has beenstudied in cross section by Dehay (1935) who found aconsiderable variation in stellar configurations Howeverit seems unlikely that this feature will be of general valueMeanwhile Miller and Webster (1962) have useddifferences in petiolar steles to separate Cridoscolus fromJatropha Then Dehgan (1982) could prove that thepetiolar stellar dissection were significant at the sectionaland sub-sectional level in some genera such asJatropha However in a considerable number of taxastipules have become reduced or are early deciduousFor example many species lack stipules whereas theyare large and remarkable in others In Euphorbia thepresence or absence of stipules is a diagnostic characterfor some sections and subgenera (Webster 1994a)

Anatomical Characters in Euphorbia

Latex

The white latex is a useful distinguishing character in thegenus Euphorbia L It is distributed throughout the plantin a series of tubes derived from either single cells (non-

articulated laticifers) or articulated laticifers formed by thefusion of several cells The value of laticifer types as ataxonomic marker in systematic comparisons betweenand within families has been established by Carlquist(1961)The laticifer system of the mature Euphorbia plantwas explained by Gaucher (1898 1902) In additionRosowski (1968) studied the branched non-articulatedlaticifers system in mature tissue of the internode andnode in transition from the node to and throughout themature leaf of Esupina Raf He found that the latexsystem in the E supina stem is restricted to the cortexand does not break through the leaf gaps In the leaflamina he found that the widest laticifers are in

association with the vascular system These laticifersbegin to branch from the base of the leaf and continuethroughout certain areas in the mesophyll Moreoverthey are associated with the phloem and may send somebranches at right angles to the vein and between thebundle-sheath cells

Epidermis

Epidermal cells may vary greatly in size shape and

Aldhebiani and Jury 169

outline from species to species especially when seenin a surface view Generally in dicotyledons epidermacells have irregular shapes and sizes Sometimes theshape of the cells in both leaf surfaces are similar butmore often they are different The costal cells usuallydiffer from those intercostal regions they tend to be

elongated in the direction of the veins Some marginacells develop unicellular or multicellular prickles Moreoveranticlinal walls can be either very thin and hardly visible fromthe surface or they may range through degrees of thicknessto be very thick (Kakkar and Paliwal 1972)

One of the important epidermal features for altaxonomists is the hairs They can be glandular or non-glandular and can be divided depending on thecomponent number of cells and degree of branching Theoccurrence of distinctive types of hair can be a valuablecharacter to recognize a whole family Moreover hairsare more useful in determination at the level of genus ospecies In other words variations in size and densityshould be accepted in the differentiation of closely related

genera or species after a comprehensive investigation ofa wide range of material (Metcalfe and Chalk 1950)

In a study of 150 species of Euphorbia Kakkar andPaliwal (1974) described the epidermal cells in Euphorbiaas circular trapezoidal rectangular or polygonal inoutline Additionally they found that the shape may varydepending upon the location of the epidermal cell on theleaf ie the middle region margin apex or above thevein Some xeromorphic species exhibit a waxy coveringwhich takes on many crystalline forms Hairs were alsorecorded in the epidermis of Euphorbia in some speciesIn addition the papillae occurred in the surface view asrounded structures in the centre of cell lumen

Stomata

A stoma consists of the stomatal aperture and the pair oguard cells that form it Stomata usually tend to be on thelower surface only But in some cases this distributionvaries from species to species and depends on whethethe plant is a xerophyte or a mesophyte They might besuperficial or sunken Stomata sometimes aresurrounded by specialized epidermal cells which arecalled subsidiary cells These subsidiaries differ fromunmodified epidermal cells in shape size and staining

properties (Baranova 1992 Metcalfe andamp Chalk 1950Stace 1965) On the other hand the arrangement osubsidiary cells where present is of the greatest interesto the taxonomist This variation is used to define thedifferent types of stomata Occasionally species haveseveral types of stomata on one leaf while some haveonly one type for the species (Stace 1984) In additionVan Cotthem (1973) pointed out that thosemorphological stomata types can provide not onlydiagnostic characters but also very valuable taxonomicones or even phylogenetic clues Metcalfe and Chalk

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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170 Int Res J Plant Sci

(1950) had established some terms to replace therepresentative lsquofamilyrsquo name proposed by Vesque(1889) Anomocytic was substituted for theranunculaceous type anisocytic replaced cruciferousdiacytic the caryophyllaceous and finally paracytic for therubiaceous The tetracytic type which can be found in

most of the monocotyledons was added by Metcalfe(1960) Later Stace (1965) proposed the term cyclocyticfor the narrow ring of four or more subsidiary cellssurrounding the stomata Metcalfe and Chalk (1950)have named and defined the actinocytic type as stomatasurrounded by a circle of radiating cells Three moretypes were introduced by Van Cotthem (1970)hexacytic epicytic and hemiparacytic And someintermediate types were added by Payne (1970) whodescribed the helicocytic and allelocytic types in relationto mesogenous forms of anisocytic paracytic and diacyticpatterns Stace (1989) lists 35 types of stomata invascular plants Closely related families are distinguishedby the presence of a specific type of stomata such asAcanthaceae and Scrophulariaceae separated by thepresence of diacytic stomata in the former as againstanomocytic in the latter Moreover some stomatal types aredistinctive of certain families for example Ranuculaceaehas the anomocytic type Brassicaceae the anisocyticCaryophyllaceae diacytic Rubiaceae paracytic and finallyPoaceae has the graminaceous type (Singh 2004)

According to Metcalfe and Chalk (1950) the maturestomata of Euphorbiaceae are anomocytic paracytic andanisocytic They are usually confined to the lower leafsurface more rarely on both surfaces of the laminaParacytic stomata were reported by Tognini (1897)which are mesogenous in development in E variegata

and Ricinus communis On the other hand according toRaju and Rao (1977) stomata in the Euphorbiaceaeshow considerable variation They found that the woodytaxa have predominantly the paracytic stomata typewhile the anisocytic stomata are characteristic of theherbaceous Phyllanthoideae Moreover they indicatedthat Chamaesyce has a high percentage of anomocyticstomata (Raju and Rao 1987) Finally a considerablediversity of stomatal types were found in Euphorbia byKakkar and Paliwal (1974) They reported that thecommon type of stomata in Euphorbia species is theanomocytic even though stomata of paracytic andanisocytic have been also observed

Venation

Some features that give the leaf its structure are forinstance leaf shape margin type and venation patternsThese together form the leaf architecture as discussed byHickey (1979) A more sophisticated knowledge of leafarchitecture has allowed a start in discriminatingphylogenetic trends from leaves (Hickey 1973 Roth-Nebelsick et al 2001) Due to its importance especiallyfor systematic classification attention has been paid

largely to the architectural properties of leaf venation(Koumlhler 1993) The careful description of venation incooperation with studies of other leaf anatomy details canprovide valuable taxonomic evidence The works of Pray(1955a 1955b) suggest that the venation can be useful ina comparative analysis Moreover Wagner (1979) has

proved that the vein patterns were significant forclassification at the level of species genus and family inferns Of course this is more noticed and observed inangiosperms especially dicotyledons where thehierarchical network patterns of the veins are much moreadvanced and complicated

Leaf venation has been neglected for a long time inthe Euphorbiaceae until the work of Levin (1986a1986b 1986c) Levin developed a new insight intorelationships within subfamily Phyllanthoideae Nodetailed studies have been provided for othesubfamilies except the work of Sehgal and Paliwa(1974) on the tribe Euphorbieae In part II of their studyon the leaf anatomy of Euphorbia Sehgal and Paliwadesignated the venation in petiolate leaves as uni- bi-and tri-veined according to the number of strandsentering the petiole or lamina base They noticed that inthe 150 species under investigation the majority bearleaves belonging to the tri-veined category Moreoverthey divided the latter into ornamented andunornamented veins When the veins are surrounded bya parenchymatous sheath it is called ornamented Andwhen this sheath is absent it is unornamented Besidessometimes trachedial nodules are encountered at theapices and in the serrations of the leaf Small veinsusually form networks in the lamina These networksmay vary in size and shape and subdivide the area o

the mesophyll The thinnest branches of the bundlesbounded by the smallest areas or regions are calledareoles These areoles usually contain blind veinendings The degree of branching of these vein endingsvaries in the leaves of different species Therefore thiscan be a useful identification tool

MATERIAL AND METHODS

Herbarium specimens were obtained from Kew (KewGarden herbarium UK) and RIY (National Herbarium inRiyadh) Since leaves are early deciduous in succulen

species most of the herbarium sheets or spirit collectionsof these species have no leaves As a result thesespecies were omitted from this anatomy study except fothe stick-like Euphorbias E balsamifera subspadenensis E schimperi ldquoE aff schimperirdquo ldquoE affconsobrinardquo and E cuneata Only mature leaves wereused for studying the epidermis stomata and venation

Epidermal peel

This method was used to observe stomata epidermal

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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cells and hairs1- Leaves were rehydrated using polyoxyethylenesorbitan monolaurate (Tween20) in water and heated for30 minutes Then washed by water and conserved in70 ethanol for future work2- Both surfaces of the leaf were scraped and

peeled off under a dissection microscope with the aid offine forceps and a razor blade to remove loose cells3- Samples were bleached by Jefferyrsquos Solution orthe bleaching agent Vortex for 10-20 minutes dependingon leaf thickness Sometimes this was overnight4- Samples were washed mounted and observedunder a light microscope5- Photographs were taken using a Leitz Diaphlanpolarizing microscope or a Reichert Polyvar 2Microscope

Staining by alucin blue was tried but withoutsuccess Samples became too dense or obscure due tothe presence of tannins and other compounds in theherbarium specimens Therefore some samples were leftwithout staining

Leaf clearing

The following method was adopted from Radford et al(1974) with some modifications for use on herbariummaterials1- The leaf was placed in a Petri dish and coveredwith 5 Sodium Hydroxide (NaOH) The dish waswrapped in cling film leaving a small gap in one area forventilation2- This was microwaved for five seconds on

medium power3- The Petri dishes were left on a hot plate at 35-37degrees centigrade for a week or until the leaf wastransparent The NaOH was changed twice a day foreach sample4- When the leaf had cleared sufficiently it waswashed in water and covered in 90 bleaching agentVortex for ten minutes to one hour depending on thereaction of the sample5- The leaf was washed again in water6- Samples were dehydrated in ethanol series andstained as followinga) 30 ethanol 2 minutes

b) 50 ethanol 2 minutesc) 1 safranin in 50 ethanol 4 minutesd) Wash in water three timese) 30 ethanol 2 minutes repeated 3 timesf) 50 ethanol 2 minutes repeated twiceg) 70 ethanol 2 minutesh) 90 ethanol 2 minutesi) 100 ethanol 2 minutes

j) Absolute ethanol histoclear 11 2 minutesk) Histoclear 2 minutes

Aldhebiani and Jury 171

l) Mounted in 100 glycerol and sealed with Candabalsamm) Observed under light microscopen) Photographs were taken using a Leitz Diaphlanpolarizing microscope or Reichert Polyvar 2 Microscope

Scanning Electron Microscopy

1- Dry specimens were mounted surface up onscanning electron microscope stubs using Bostik No1adhesive2- Stubs were sputter coated for 2-3 minutes with agold palladium alloy using an Edwards sputter coater togive a coating about 15-20 nm thick3- Samples were examined in FEI Quanta FEG 600Environmental Scanning Electron Microscope (ESEM) inhigh vacuum SEM mode4- Photographs were taken using the computerizeddigital system of the microscope

RESULTS

Epidermis

Results of the epidermal characters in Euphorbia speciesunder investigation are summarized in Table 1

Cell shape

Depending upon the shape of the cell wall and

considering the results from Kakkar and Paliwal (1974)the shape of the epidermal cell varies polygonalrectangular undulate or elongated Moreover the celshape relies on the cell location on a leaf ie the middleregion the margin the apex or above the veinFurthermore in some cases both leaf surfaces have thesame cell shape but more often they are unlike Thiscombination of cell shape characters gives each speciesits unique leaf surface appearance

The normal straight cell walls in both adaxial andabaxial surfaces are shown in the following taxa Eschimperi E cuneata E helioscopia E dracunculoidesand E balsamifera subsp adenensis see Figure 1 In

addition ldquoE aff schimperirdquo and ldquoE aff consobrinardquo havealso the same straight cell wall on both surfacesMeanwhile E grossheimii E retusa and E pirottae haveelongated cells on both leaf surfaces (Figure 2 a and b)On the other hand the plicate cell walls on both adaxiaand abaxial surfaces occur in E chamaepeplus Ehypericifolia and E peplus (Figure 2 c and d) but thewalls are more folded on the lower surface In somespecies the cell wall on the adaxial surface is straightwhilst it is plicate on the abaxial such as E hirta E

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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172 Int Res J Plant Sci

Table 1 Epidermal characters of Euphorbia taxa in Saudi Arabia

Ab abaxial + PresentAd adaxial ++ the plicate folded wall is morefully developed in the lower surface than in the upper one

+ hardly observable by LM only

Subgenus

Cell shape (cell wall) Hairs

Straight Elongated Plicate Unicellular MultSpecies Ad Ab Ad Ab Ad Ab Ad Ab Ad T

i r u c al l i

E balsamifera subspadenensis + + +

E schimperi + +

ldquoEaff schimperirdquo + +

ldquoE aff consobrinardquo + +

E cuneata + +

ErmophytonE acalyphoides + + +

E pirottae + + +

E helioscopia + + E

s ul a

E grossheimii+ +

E retusa + +

E dracunculoides + +

E falcate + +

E peplus + ++

E chamaepeplus

E schimperiana +

+ ++

+ C h am a e s y c e

E Arabica + +

E serpens + +

E scordiifolia + + +

E granulate + + + + +

E inaequilatera + +

E hirta + + +

E hypericifolia + ++

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 173

983106

983108

983109

983105

983107

983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

Figure 1 Normal irregular epidermal cells in both adaxial and abaxial surfaces of the leaf (LM)(A) E balsamifera subsp adenensis (B) E schimperi (C) E cuneata (D) E falcata (E) Abaxial in E dracunculoides

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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174 Int Res J Plant Sci

983105

983106

983107

983108

983106 983106

983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

Figure 2 Error No text of specified style in document Both adaxial and abaxial surfaceshave the same shape of epidermal cells (LM) Elongated in A ampB plicate in CampD (A) Egrossheimii (B) E pirottae (C) E chamaepeplus and (D) E peplus

schimperiana E helioscopia E inaequilatera Earabica E granulata E serpens and E scordiifolia see Figure 3 and Figure 4 This last group of species belongto subgenus Chamaesyce except for E helioscopia and E schimperiana

Hairs

Hairs on the leaf epidermis can be both unicellular and

multicellular and may appear only on one surface or bothFurthermore hairs in the Euphorbia species undeinvestigation are generally simple unbranched and with awarty ornamentation on their surface Papillae appear inESEM as outgrowths whereas in the surface view theygive the impression of being rounded structures at thecentre of the cell Prominent papillae occur only in onespecies of Euphorbia in Saudi Arabia this species is Ehypericifolia (Figure 5 and 6) whereas in E arabica theywere poorly distinguishable using LM

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 175

983105983140983137983160983145983137983148

983107

983108

983105

983106

983105983138983137983160983145983137983148

Figure 3 Epidermal cells vary on leaf surfaces adaxial surface has irregular cells whilethe abaxial surface has plicate (LM)(A) E schimperiana (B) E arabica (C) E serpens (D) E scordiifolia

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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176 Int Res J Plant Sci

983105983140983137983160983145983137983148

983107

983105

983106

983105983138983137983160983145983137983148

Figure 4 Epidermal cell shape varies on the adaxial andthe abaxial surfaces straight cell walls in the formerwhile folded plicate walls occur on the latter (LM) (A) Egranulata (B) E inaequilatera and (C) E helioscopia

983112983141983149983145983152983137983154983137983139983161983156983145983139 983155983156983151983149983137983156983137

983151983150 983138983151983156983144 983155983157983154983142983137983139983141983155

Figure 5 E hypericifolia (LM)Above photo the adaxial surface with irregular epidermal cells andvery clear papillae as a circle in the middle of the cellLower photo the abaxial surface with plicate cells and papillae ahair base is obvious in the right side of the picture

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 177

983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

983120983137983152983145983148983148983137983141

983117983157983148983156983145983139983141983148983148983157983148983137983154 983144983137983145983154983155 983145983150 983137983138983137983160983145983137983148 983155983157983154983142983137983139983141983086

Figure 6 E hypericifolia (ESEM) papillae on the adaxial and abaxial surfacesmulticellular hairs are on the abaxial surface of the leaf only

Figure7 Multicellular hairs on adaxial and abaxial surfaces of E hirta

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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178 Int Res J Plant Sci

Figure 8 Hairs wart ornamentation on hair surfaces of (A) E scordiifolia and (B) E pirottae (C) Short hairs onabaxial surface of E balsamifera subsp adenensis

In E balsamifera subsp adenensis the hairs are shortand unicellular (Figure 8C) and they are located mostly atthe leaf margins and the mid-vein area on the abaxialsurface only

Hairs on the leaf of E pirottae are present on theabaxial surface only and are both unicellular andmulticellular Euphorbia granulata is similar to E pirottae in having both unicellular and multicellular hairs but herethey occur on both surfaces of the leaf (Figure 8B)

On the other hand in E scordiifolia E hirta and Ehypericifolia the hairs are multicellular and on bothsurfaces in the first two species and on the abaxial only inthe latter (Figure 7) Moreover E hypericifolia and Ehirta have long hairs when compared with the otherspecies (Figure 6)

No hairs were observed and leaves were glabrous inthe following species E serpens E arabica E peplusE inaequilatera E falcata E helioscopia E cuneata Eretusa E grossheimii E schimperiana E schimperiE chamaepeplus and the new variety of E granulata

Wax

The cuticle layer and wax in the Euphorbia species undeinvestigation vary from smooth to densely cover in waxscales This may be a thick smooth layer of wax whichsometimes has upright scales In some species only thecell surface has a smooth wax layer whereas the walbetween cells has upright scales of wax as in Eschimperiana especially around the stomata pores

(Figure 9) In contrast the anticlinal walls of somespecies have a smooth layer of wax whereas the celsurface is densely covered with upright wax flakes or withsome stellate wax flakes in other species

Stomata

Stomata were observed through LM and ESEM In thespecies under investigation stomata commonly appeaon both leaf surfaces (amphistomatic leaves) excepin E cuneata where the stomata are located only on the

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 179

983109

983105

983108983107

983106

Figure 9 Wax formation in Euphorbia species in Saudi Arabia (A+B) smooth andthick in E peplus and E schimperi (C+D) smooth on top of cell while formingupright scale flakes in between especially around stomata as in E schimperiana smooth in anticlinal walls (E) upright scale flakes in E balsamifera

abaxial surface (hypostomatic leaves) The presence andposition of subsidiary cells are variable in Euphorbia

species Subsidiary cells were found to be of unequalsize and shape whether the stoma was anisocyticparacytic actinocytic or tetracytic While some specieslack them around the stomata others have more thanone arrangement of subsidiary cells

The most common stomata type is anomocytic whilethe rare type is actinocytic recorded only in Ehelioscopia Moreover sometimes stomata of more thanone type (up to three or four types) have beenencountered on the same leaf surface as in E scordiifolia (three types on the abaxial surface) and E hirta (five

types on the abaxial and three on the adaxial) Thedistribution and type of stomata are shown in Table 2

In addition in some species such as E falcata and Ehelioscopia the stomata are deeply sunken andrepresented only by cuticular ledges surrounding theminute pores as seen in ESEM

Examples of species lacking subsidiary cells aroundstomata are E peplus E chamaepeplus E granulataE dracunculoides and E acalyphoides Therefore theyall have anomocytic stomata

On the other hand E balsamifera subsp adenensishas the mononcytic type on both leaf surfaces andanisocytic and paracytic only on the abaxial surface In

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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180 Int Res J Plant Sci

Table 2 Stomata type in Euphorbia taxa in Saudi Arabia

Ab abaxial surface + stomata presentAd adaxial surface

addition E schimperiana has anomocytic andtetracytic on the abaxial only In contrast Ecuneata has three types of stomata anomocyticparacytic and anisocytic only on the lower surfaceAs well ldquoE aff consobrinardquo ldquoE aff schimperirdquo

and E schimperi have paracytic and anisocyticonly in the abaxial on the first two and in both leafsurfaces on the latter Therefore ldquoE affconsobrinardquo and ldquoE aff schimperirdquo are the same taxa Then comparing to E schimperi results

from ansubspec

Eupsimilar and an

Subgenus

Stomata

Anomocytic Hemiparacytic Paracytic Anisocytic T

Species Ad Ab Ad Ab Ad Ab Ad Ab AT i r u c al l i

E balsamifera subsp adenensis + + +

E schimperi + + + +

ldquoEsp aff consobrinardquo + +

ldquoE sp aff schimperirdquo + +

E cuneata + + +

ErmophytonE acalyphoides + +

E pirottae + + + +

E helioscopia + + E s ul a

E grossheimii + + + + +

E retusa + + + + +

E dracunculoides +

E falcate + + E peplus +

E chamaepeplus + +

E schimperiana + C h am a e s y c e

E Arabica + + + + + + +

E serpens + + + +

E scordiifolia + + + + + + +

E granulate + + + +

E inaequilatera + + + +

E hirta + + + + + + +

E hypericifolia + + + + + +

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Table3 Venation type in Euphorbia taxa in Saudi Arabia

Subgenus Species

Venation

Number of veins atthe leaf base

Margins Sheathed

1 2 3 4 or more Open Closed Rounded T i r u c al l i

Ebalsamiferasubspadenensis

+ + +

E schimperi + + +

E aff Schimperi + + +

E cuneata + + +

ErmophytonE acalyphoides + + +

E pirottae + + E s ul a

E grossheimii + +

E retusa + + +

E dracunculoides + + +

E falcate + + +

E peplus + +

E chamaepeplus + +

E schimperiana

E helioscopia

+

+

+

+ C h am a e s y c e

E Arabica + + +

E serpens + + +

E scordiifolia + + +

E granulate + + + +

E inaequilatera + + +

E hirta + + + E hypericifolia + + +

Poinsettia E cyathophora + +

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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182 Int Res J Plant Sci

Figure 10 Sunken stomata on the abaxial surface1) Deeply sunken (A) E helioscopia (B) E falcata (C) E schimperiana 2) Slightly sunken (D) E peplus (E) E pirottae

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 183

983105983138983137983160983145983137983148

983105

983108

983108

983106

983107

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

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983105983140983137983160983145983137983148

Figure 11 Stomata types in Euphorbia taxa in Saudi Arabia (A) E balsamifera subsp adenensis (B) E schimperi (C) E cuneata and (D) E grossheimii

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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184 Int Res J Plant Sci

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983105983138983137983160983145983137983148983105983140983137983160983145983137983148

Figure 12 Stomata types in Euphorbia taxa (A) E falcata (B) E peplus (C) E chamaepeplus and (D) Eschimperiana

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 185

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Figure 13 Figure 13 Stomata types in Euphorbia taxa in Saudi Arabia (A) E arabica (B) E serpens (C)E scordiifolia and (D) E helioscopia

distribution between the leaf surfaces (see Figure 13)While E hirta has the most types E helioscopia Edracunculoides and E acalyphoides have the least

Regarding the position of the stomata on the surfacelevel of the epidermis a variable degree of sunkenesshas been recorded Using the ESEM the stomata appear

deeply sunken in some species whilst in others they areonly slightly sunken with the outer rims of the subsidiarycells surrounding the aperture The deeply and slightlysunken species are recorded in Table 2 AdditionallyFigure 10 shows the sunken stomata in different speciesunder the ESEM

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Figure 14 Stomata types in Euphorbia taxa in Saudi Arabia (A) E hirta (B) E hypericifolia (C) E pirottae and (D)Espaff schimperi

Venation

The venation patterns in the Euphorbia species underinvestigation vary from one-veined three-veined to thosewith four or more veins The predominant type is three-veined However in E balsamifera subsp Adenensis

there are seven middle parallel veins three middle thickand four thinner lateral ones (Figure 15 A and B)Whereas in E pirottae there are three main middleparallel veins and two thinner laterals (Figure 15 C) Incontrast E chamaepeplus E schimperiana and Earabica have only one thick main midrib coming from the

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 187

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Figure 15 Figure 15 Venation patterns at base of leaves invariants with more than three veins (A amp B) E balsamiferasubsp adenensis and (C) E pirottae

petiole then branched from the side into two lateral veinsat the point when the leaf starts widening (Figure 16)Each of these veins branches again into secondary veinsat the leaf margin

The rest of the species have just three veins Egrossheimii E dracunculoides E falcata E peplus Eserpens E scordiifolia E granulata E inaequilateraE helioscopia E hirta E hypericifolia and E

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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188 Int Res J Plant Sci

Figure 16 Venation patterns with one main mid vein at the leaf base in (A) E arabica and (B) E chamaepeplus

983106983105

Figure 17 Venation patterns with three-veined leaves in (A) E falcata and (B) E granulata

cyathophora All the three veins come from the petioleand start branching into secondary veins both toward themargin and the mid vein direction (Figure 17 B)

The vein endings at the leaf margin vary from closed

to open There is a relationship between the closedmargins and the sheathed veins (midrib and secondaryveins that are surrounded by parenchyma tissue) Whenthe veins are ornamented (sheathed) the margins areclosed This can be seen in subgenus Chamaesyce inspecies such as E serpens E scordiifolia E granulataE inaequilatera E hirta E hypericifolia and E arabica Additionally these species have the three-vein venation

On the other hand when the veins are not sheathedthe veins ends are open at the leaf margin as in Ebalsamifera subsp adenensis E schimperi E cuneataE acalyphoides E grossheimii E dracunculoides E

falcata E peplus E chamaepeplus E schimperiana Ehelioscopia E pirottae ldquoE aff schimperirdquo and Ecyathophora

CONCLUSIONS

Anatomical characters are seen here to be very useful inEuphorbia classification and identification The resultsobtained from anatomical studies were useful to groupsome species and identify others

Depending on the venation type the Euphorbiaspecies under investigation are divided anatomically intothree main groups see Figure 18

Group one veins are sheathed epidermal cellsdifferentiated on adaxial and abaxial surfaces Group two

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7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Figure 18 Hierarchial relation of Euphorbia taxa in Saudi Arabia based on anatomical charac

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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veins are not sheathed and epidermal cells are alike inboth surfaces And the third group includes species thatdo not belong to any of the previous groups The firstgroup can be further divided into three-veined and one-veined while the second group into three-veined and fouror more veins

In addition E grossheimii E retusa and E pirottae are related by being three-veined open at the marginswith rounded to oval tracheas and elongated cells bothon adaxial and abaxial surfaces However in E retusa the epidermal cells have straight cell walls on the adaxialsurface They are also similar in having hemiparacyticand paracytic types of stomata either on the adaxial orthe abaxial surface

On the other hand species with two differentepidermal cells on the leaf surfaces usually have three-sheathed veins with closed margins and elongatedtrachea such as E serpens E scordiifolia E granulata E inaequilatera E hirta and E hypericifolia which belongto subgenus Chamaesyce according to (Carter andSmith 1988)

In the case of E granulata which has beendescribed as having three different variants E granulata var granulata (both leaf surfaces hairy) E granulata varglabrata (adaxial glabrous and abaxial hairy) and aglabrous variant (the whole plant glabrous) Anatomicalwork has shown especially by using the ESEM that theleaves are either hairy or glabrous on both surfaces Sowhen the leaf is hairy both surfaces have differences inhair distribution Usually the lower surface is more hairythan upper surface (E granulata var glabrata ) butsometimes they are evenly distributed (Egranulata vargranulata ) In addition hairs on both surfaces are

multicellular In contrast when the leaf is glabrous bothleaf surfaces are glabrous and the whole plant isglabrous (suggested new variety) They were similar inhaving the same venation type (three-veined)

Finally ldquoE aff schimperi ldquohas some similarity with E schimperi in having the same open margin not theornamented venation type As well they both have thesame type of stomata paracytic and anisocytic but theydiffer in their distribution on abaxial and adaxial Thusthis may support the suggestion considering ldquoE affschimperi rdquo as a new subspecies of E schimperi

REFERENCES

Baranova M (1992) Principels of comparative stomatographic studiesof flowering plants The Botanical Review 58 49-99

Carlquist S (1961) Comparative Plant Anatomy Holt Rinehart ampWinston New York

Carter S Smith AR (1988) Euphorbiaceae (Part2) In Polhill ed Floraof Tropical East Africa Rotterdam Brookfield AA Balkema 409-597

Chaudhary SA (2001) Flora of The Kingdom of Saudi Arabia Vol 3Ministry of Agriculture and Water Riyadh

Dehay C (1935) Lagraveppareil Libero-ligneux foliare des EuphorbiaceaeAnnales des Sciences Naturelles Botanique 10 147-190

Dehgan B (1982) Comparative anatomy of the petiole and infragenericrelationships in Jatropha (Euphorbiaceae) American Journal oBotany 69 1283-1295

Gaucher L (1898) Ē tude anatomique du genre Euphorbia LPKlincksieck Paris

Gaucher L (1902) Recherches anatomiques sur les EuphorbiaceacuteesAnnales des Sciences Naturelles Botanique 15 161-309

Gaucher L (1902) Recherches anatomiques sur les Euphorbiaceacutees

Annales des Sciences Naturelles Botanique 15 161-309Govaerts R Fordin D Radcliffe-Smith A (2000) World Checklist and

Bibliography of Euphorbiaceae The Royal Botanic Gardens KewHickey LJ (1973) Classification of the architecture of dicotyledonous

leaves American Journal of Botany 60 17-33Hickey LJ (1979) A revised classification of the architecture o

dicotyledonous leaves In Metcalfe CR and Chalk L eds Anatomy oDicotyledons Systematic Anatomy of the leaf and stem 2nd edClarendon Press Oxford 25-39

Kakkar L Paliwal GS (1972) Epidermis in Euphorbia Indian ScienceCongress Association Proceedings 59 326-327

Kakkar L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 5 Epidermis Proceedings of the Indian National ScienceAcademy Part B Biological Sciences 40 55-67

Koumlhler E (1993) Blattvervatur-Muster der Buxaceae Dumortierr undSimmondsiaceae Van Tieghem Feddes Repertorium 104 145-167

Levin GA (1986a) Systematic foliar morphology of Phylanthoideae(Euphorbiaceae) II Phenetic analysis Annals of Missouri BotanicaGarden 73 86-98

Levin GA (1986b) Systematic foliar morphology of Phylanthoideae IICladistic analysis Systematic Botany 11 515-530

Levin GA (1986c) Systematic foliar morphology of Phyllanthoideae(Euphorbiaceae) I conspectus Annals of Missouri Botanical Garden73 29-85

Mahlberg P (1973) Histochemistry and Scanning Electron Microscopyof Starch Grains from Latex of Euphorbia-Terracina and EuphorbiaTirucalli Proceedings of the Indiana Academy of Science 82 132

Mahlberg PG (1974) Scanning Microscopic Comparison of StarchGrains from the Latex of Euphorbia -Spp Proceedings of the IndianaAcademy of Science 83 83-84

Mahlberg PG (1975) Evolution of the Laticifer in Euphorbia asInterpreted from Starch Grain Morphology American Journal oBotany 62 577-583

Mahlberg PG (1982) Comparative Morphology of Starch Grains inLatex from Varieties of Poinsettia Euphorbia-PulcherrimEuphorbiaceae Botanical Gazette 143 206-209

Mahlberg PG Davis DG Galitz DS Manners GD (1987) Laticifers andthe Classification of Euphorbia the Chemotaxonomy of EuphorbiaEsula L Botanical Journal of the Linnean Society 94 165-180

Metcalfe C Chalk L (1950) Anatomy of the Dicotyledons At theClarendon Press Oxford

Miller K Webster GL (1962) Systematic position of Cnidoscolus andJatropha Brittonia 14 174-180

Pax F (1884) Die Anatomie der Euphorbiaceen in ihrer Beziehung zumSystem derselben Botanische Jahrbuumlcher fuumlr SystematikPflanzengeschichte und Pflanzengeographie Leipzig 5 384-421

Payne WW (1970) Helicocytic and allelocytic stomata unrecognizedpatters in the Dicotyledonae Am J Bot57 140-147

Pray TR (1955a) Foliar venation of angiosperm II Histogenesis of thevenation of Liriodendron Am J Bot 42 18-27

Pray TR 1955b Foliar venation of angiosperms III Pattern andhistology of the venation of Hosta Am J Bot 42 11-18

Radford AE Dickison WC Massey JR Bell CR (1974) Vascular PlanSystematics Harper amp Row Publishers London New York

Radlkofer L (1870) Ueber Pausandra ein neues EuphorbiaceenGenus Flora 53 81-95

Raju VS Rao PN (1977) Variation in the structure and development ofoliar stomata in the Euphorbiaceae Botanical Journal of the LinneanSociety 75 69-97

Raju VS Rao PN (1987) The Taxonomic Use of the Basic StomataType in the Generic Delimitation of Chamaesyce EuphorbiaceaeFeddes Repertorium 98 137-142

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2424

Rosowski JR (1968) Laticifer Morphology in the Mature Stem and Leafof Euphorbia supina Botanical Gazette 129 113-120

Roth-Nebelsick A Uhl D Mosbrugger V Kerp H (2001) Evolution andfunction of leaf venation architecture a review Annals of Botany 87553-566

Sehgal L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 2 Venation Patterns Botanical Journal of the Linnean Society 68 173-208

Singh G (2004) Plant Systematics An Integrated Approach SciencePublishers Enfield

Solereder H (1899) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Solereder H (1908) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Aldhebiani and Jury 191

Stace CA (1965) Cuticular studies as an aid to plant taxonomy Bulletinof the British Museum Natural History Botany 4 3-78

Stace CA (1989) Plant Taxonomy and Biosystematics Hodder ampStoughton London

Tognini F (1897) Contribuzione allo studio della organogeniacomparata degli stomi Institut de Botanique University de Geneve 41-42

Van Cotthem WR (1970) A classification of stomatal types Botanica

Journal of Linnean Society 63 235-246Wagner W (1979) Reticulate veins in the systematics of modern ferns

Taxon 1 87-95Webster GL (1994a) Classification of the Euphorbiaceae Annals of the

Missouri Botanical Garden 81 3-32

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170 Int Res J Plant Sci

(1950) had established some terms to replace therepresentative lsquofamilyrsquo name proposed by Vesque(1889) Anomocytic was substituted for theranunculaceous type anisocytic replaced cruciferousdiacytic the caryophyllaceous and finally paracytic for therubiaceous The tetracytic type which can be found in

most of the monocotyledons was added by Metcalfe(1960) Later Stace (1965) proposed the term cyclocyticfor the narrow ring of four or more subsidiary cellssurrounding the stomata Metcalfe and Chalk (1950)have named and defined the actinocytic type as stomatasurrounded by a circle of radiating cells Three moretypes were introduced by Van Cotthem (1970)hexacytic epicytic and hemiparacytic And someintermediate types were added by Payne (1970) whodescribed the helicocytic and allelocytic types in relationto mesogenous forms of anisocytic paracytic and diacyticpatterns Stace (1989) lists 35 types of stomata invascular plants Closely related families are distinguishedby the presence of a specific type of stomata such asAcanthaceae and Scrophulariaceae separated by thepresence of diacytic stomata in the former as againstanomocytic in the latter Moreover some stomatal types aredistinctive of certain families for example Ranuculaceaehas the anomocytic type Brassicaceae the anisocyticCaryophyllaceae diacytic Rubiaceae paracytic and finallyPoaceae has the graminaceous type (Singh 2004)

According to Metcalfe and Chalk (1950) the maturestomata of Euphorbiaceae are anomocytic paracytic andanisocytic They are usually confined to the lower leafsurface more rarely on both surfaces of the laminaParacytic stomata were reported by Tognini (1897)which are mesogenous in development in E variegata

and Ricinus communis On the other hand according toRaju and Rao (1977) stomata in the Euphorbiaceaeshow considerable variation They found that the woodytaxa have predominantly the paracytic stomata typewhile the anisocytic stomata are characteristic of theherbaceous Phyllanthoideae Moreover they indicatedthat Chamaesyce has a high percentage of anomocyticstomata (Raju and Rao 1987) Finally a considerablediversity of stomatal types were found in Euphorbia byKakkar and Paliwal (1974) They reported that thecommon type of stomata in Euphorbia species is theanomocytic even though stomata of paracytic andanisocytic have been also observed

Venation

Some features that give the leaf its structure are forinstance leaf shape margin type and venation patternsThese together form the leaf architecture as discussed byHickey (1979) A more sophisticated knowledge of leafarchitecture has allowed a start in discriminatingphylogenetic trends from leaves (Hickey 1973 Roth-Nebelsick et al 2001) Due to its importance especiallyfor systematic classification attention has been paid

largely to the architectural properties of leaf venation(Koumlhler 1993) The careful description of venation incooperation with studies of other leaf anatomy details canprovide valuable taxonomic evidence The works of Pray(1955a 1955b) suggest that the venation can be useful ina comparative analysis Moreover Wagner (1979) has

proved that the vein patterns were significant forclassification at the level of species genus and family inferns Of course this is more noticed and observed inangiosperms especially dicotyledons where thehierarchical network patterns of the veins are much moreadvanced and complicated

Leaf venation has been neglected for a long time inthe Euphorbiaceae until the work of Levin (1986a1986b 1986c) Levin developed a new insight intorelationships within subfamily Phyllanthoideae Nodetailed studies have been provided for othesubfamilies except the work of Sehgal and Paliwa(1974) on the tribe Euphorbieae In part II of their studyon the leaf anatomy of Euphorbia Sehgal and Paliwadesignated the venation in petiolate leaves as uni- bi-and tri-veined according to the number of strandsentering the petiole or lamina base They noticed that inthe 150 species under investigation the majority bearleaves belonging to the tri-veined category Moreoverthey divided the latter into ornamented andunornamented veins When the veins are surrounded bya parenchymatous sheath it is called ornamented Andwhen this sheath is absent it is unornamented Besidessometimes trachedial nodules are encountered at theapices and in the serrations of the leaf Small veinsusually form networks in the lamina These networksmay vary in size and shape and subdivide the area o

the mesophyll The thinnest branches of the bundlesbounded by the smallest areas or regions are calledareoles These areoles usually contain blind veinendings The degree of branching of these vein endingsvaries in the leaves of different species Therefore thiscan be a useful identification tool

MATERIAL AND METHODS

Herbarium specimens were obtained from Kew (KewGarden herbarium UK) and RIY (National Herbarium inRiyadh) Since leaves are early deciduous in succulen

species most of the herbarium sheets or spirit collectionsof these species have no leaves As a result thesespecies were omitted from this anatomy study except fothe stick-like Euphorbias E balsamifera subspadenensis E schimperi ldquoE aff schimperirdquo ldquoE affconsobrinardquo and E cuneata Only mature leaves wereused for studying the epidermis stomata and venation

Epidermal peel

This method was used to observe stomata epidermal

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cells and hairs1- Leaves were rehydrated using polyoxyethylenesorbitan monolaurate (Tween20) in water and heated for30 minutes Then washed by water and conserved in70 ethanol for future work2- Both surfaces of the leaf were scraped and

peeled off under a dissection microscope with the aid offine forceps and a razor blade to remove loose cells3- Samples were bleached by Jefferyrsquos Solution orthe bleaching agent Vortex for 10-20 minutes dependingon leaf thickness Sometimes this was overnight4- Samples were washed mounted and observedunder a light microscope5- Photographs were taken using a Leitz Diaphlanpolarizing microscope or a Reichert Polyvar 2Microscope

Staining by alucin blue was tried but withoutsuccess Samples became too dense or obscure due tothe presence of tannins and other compounds in theherbarium specimens Therefore some samples were leftwithout staining

Leaf clearing

The following method was adopted from Radford et al(1974) with some modifications for use on herbariummaterials1- The leaf was placed in a Petri dish and coveredwith 5 Sodium Hydroxide (NaOH) The dish waswrapped in cling film leaving a small gap in one area forventilation2- This was microwaved for five seconds on

medium power3- The Petri dishes were left on a hot plate at 35-37degrees centigrade for a week or until the leaf wastransparent The NaOH was changed twice a day foreach sample4- When the leaf had cleared sufficiently it waswashed in water and covered in 90 bleaching agentVortex for ten minutes to one hour depending on thereaction of the sample5- The leaf was washed again in water6- Samples were dehydrated in ethanol series andstained as followinga) 30 ethanol 2 minutes

b) 50 ethanol 2 minutesc) 1 safranin in 50 ethanol 4 minutesd) Wash in water three timese) 30 ethanol 2 minutes repeated 3 timesf) 50 ethanol 2 minutes repeated twiceg) 70 ethanol 2 minutesh) 90 ethanol 2 minutesi) 100 ethanol 2 minutes

j) Absolute ethanol histoclear 11 2 minutesk) Histoclear 2 minutes

Aldhebiani and Jury 171

l) Mounted in 100 glycerol and sealed with Candabalsamm) Observed under light microscopen) Photographs were taken using a Leitz Diaphlanpolarizing microscope or Reichert Polyvar 2 Microscope

Scanning Electron Microscopy

1- Dry specimens were mounted surface up onscanning electron microscope stubs using Bostik No1adhesive2- Stubs were sputter coated for 2-3 minutes with agold palladium alloy using an Edwards sputter coater togive a coating about 15-20 nm thick3- Samples were examined in FEI Quanta FEG 600Environmental Scanning Electron Microscope (ESEM) inhigh vacuum SEM mode4- Photographs were taken using the computerizeddigital system of the microscope

RESULTS

Epidermis

Results of the epidermal characters in Euphorbia speciesunder investigation are summarized in Table 1

Cell shape

Depending upon the shape of the cell wall and

considering the results from Kakkar and Paliwal (1974)the shape of the epidermal cell varies polygonalrectangular undulate or elongated Moreover the celshape relies on the cell location on a leaf ie the middleregion the margin the apex or above the veinFurthermore in some cases both leaf surfaces have thesame cell shape but more often they are unlike Thiscombination of cell shape characters gives each speciesits unique leaf surface appearance

The normal straight cell walls in both adaxial andabaxial surfaces are shown in the following taxa Eschimperi E cuneata E helioscopia E dracunculoidesand E balsamifera subsp adenensis see Figure 1 In

addition ldquoE aff schimperirdquo and ldquoE aff consobrinardquo havealso the same straight cell wall on both surfacesMeanwhile E grossheimii E retusa and E pirottae haveelongated cells on both leaf surfaces (Figure 2 a and b)On the other hand the plicate cell walls on both adaxiaand abaxial surfaces occur in E chamaepeplus Ehypericifolia and E peplus (Figure 2 c and d) but thewalls are more folded on the lower surface In somespecies the cell wall on the adaxial surface is straightwhilst it is plicate on the abaxial such as E hirta E

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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172 Int Res J Plant Sci

Table 1 Epidermal characters of Euphorbia taxa in Saudi Arabia

Ab abaxial + PresentAd adaxial ++ the plicate folded wall is morefully developed in the lower surface than in the upper one

+ hardly observable by LM only

Subgenus

Cell shape (cell wall) Hairs

Straight Elongated Plicate Unicellular MultSpecies Ad Ab Ad Ab Ad Ab Ad Ab Ad T

i r u c al l i

E balsamifera subspadenensis + + +

E schimperi + +

ldquoEaff schimperirdquo + +

ldquoE aff consobrinardquo + +

E cuneata + +

ErmophytonE acalyphoides + + +

E pirottae + + +

E helioscopia + + E

s ul a

E grossheimii+ +

E retusa + +

E dracunculoides + +

E falcate + +

E peplus + ++

E chamaepeplus

E schimperiana +

+ ++

+ C h am a e s y c e

E Arabica + +

E serpens + +

E scordiifolia + + +

E granulate + + + + +

E inaequilatera + +

E hirta + + +

E hypericifolia + ++

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 173

983106

983108

983109

983105

983107

983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

Figure 1 Normal irregular epidermal cells in both adaxial and abaxial surfaces of the leaf (LM)(A) E balsamifera subsp adenensis (B) E schimperi (C) E cuneata (D) E falcata (E) Abaxial in E dracunculoides

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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174 Int Res J Plant Sci

983105

983106

983107

983108

983106 983106

983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

Figure 2 Error No text of specified style in document Both adaxial and abaxial surfaceshave the same shape of epidermal cells (LM) Elongated in A ampB plicate in CampD (A) Egrossheimii (B) E pirottae (C) E chamaepeplus and (D) E peplus

schimperiana E helioscopia E inaequilatera Earabica E granulata E serpens and E scordiifolia see Figure 3 and Figure 4 This last group of species belongto subgenus Chamaesyce except for E helioscopia and E schimperiana

Hairs

Hairs on the leaf epidermis can be both unicellular and

multicellular and may appear only on one surface or bothFurthermore hairs in the Euphorbia species undeinvestigation are generally simple unbranched and with awarty ornamentation on their surface Papillae appear inESEM as outgrowths whereas in the surface view theygive the impression of being rounded structures at thecentre of the cell Prominent papillae occur only in onespecies of Euphorbia in Saudi Arabia this species is Ehypericifolia (Figure 5 and 6) whereas in E arabica theywere poorly distinguishable using LM

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 175

983105983140983137983160983145983137983148

983107

983108

983105

983106

983105983138983137983160983145983137983148

Figure 3 Epidermal cells vary on leaf surfaces adaxial surface has irregular cells whilethe abaxial surface has plicate (LM)(A) E schimperiana (B) E arabica (C) E serpens (D) E scordiifolia

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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176 Int Res J Plant Sci

983105983140983137983160983145983137983148

983107

983105

983106

983105983138983137983160983145983137983148

Figure 4 Epidermal cell shape varies on the adaxial andthe abaxial surfaces straight cell walls in the formerwhile folded plicate walls occur on the latter (LM) (A) Egranulata (B) E inaequilatera and (C) E helioscopia

983112983141983149983145983152983137983154983137983139983161983156983145983139 983155983156983151983149983137983156983137

983151983150 983138983151983156983144 983155983157983154983142983137983139983141983155

Figure 5 E hypericifolia (LM)Above photo the adaxial surface with irregular epidermal cells andvery clear papillae as a circle in the middle of the cellLower photo the abaxial surface with plicate cells and papillae ahair base is obvious in the right side of the picture

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 177

983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

983120983137983152983145983148983148983137983141

983117983157983148983156983145983139983141983148983148983157983148983137983154 983144983137983145983154983155 983145983150 983137983138983137983160983145983137983148 983155983157983154983142983137983139983141983086

Figure 6 E hypericifolia (ESEM) papillae on the adaxial and abaxial surfacesmulticellular hairs are on the abaxial surface of the leaf only

Figure7 Multicellular hairs on adaxial and abaxial surfaces of E hirta

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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178 Int Res J Plant Sci

Figure 8 Hairs wart ornamentation on hair surfaces of (A) E scordiifolia and (B) E pirottae (C) Short hairs onabaxial surface of E balsamifera subsp adenensis

In E balsamifera subsp adenensis the hairs are shortand unicellular (Figure 8C) and they are located mostly atthe leaf margins and the mid-vein area on the abaxialsurface only

Hairs on the leaf of E pirottae are present on theabaxial surface only and are both unicellular andmulticellular Euphorbia granulata is similar to E pirottae in having both unicellular and multicellular hairs but herethey occur on both surfaces of the leaf (Figure 8B)

On the other hand in E scordiifolia E hirta and Ehypericifolia the hairs are multicellular and on bothsurfaces in the first two species and on the abaxial only inthe latter (Figure 7) Moreover E hypericifolia and Ehirta have long hairs when compared with the otherspecies (Figure 6)

No hairs were observed and leaves were glabrous inthe following species E serpens E arabica E peplusE inaequilatera E falcata E helioscopia E cuneata Eretusa E grossheimii E schimperiana E schimperiE chamaepeplus and the new variety of E granulata

Wax

The cuticle layer and wax in the Euphorbia species undeinvestigation vary from smooth to densely cover in waxscales This may be a thick smooth layer of wax whichsometimes has upright scales In some species only thecell surface has a smooth wax layer whereas the walbetween cells has upright scales of wax as in Eschimperiana especially around the stomata pores

(Figure 9) In contrast the anticlinal walls of somespecies have a smooth layer of wax whereas the celsurface is densely covered with upright wax flakes or withsome stellate wax flakes in other species

Stomata

Stomata were observed through LM and ESEM In thespecies under investigation stomata commonly appeaon both leaf surfaces (amphistomatic leaves) excepin E cuneata where the stomata are located only on the

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 179

983109

983105

983108983107

983106

Figure 9 Wax formation in Euphorbia species in Saudi Arabia (A+B) smooth andthick in E peplus and E schimperi (C+D) smooth on top of cell while formingupright scale flakes in between especially around stomata as in E schimperiana smooth in anticlinal walls (E) upright scale flakes in E balsamifera

abaxial surface (hypostomatic leaves) The presence andposition of subsidiary cells are variable in Euphorbia

species Subsidiary cells were found to be of unequalsize and shape whether the stoma was anisocyticparacytic actinocytic or tetracytic While some specieslack them around the stomata others have more thanone arrangement of subsidiary cells

The most common stomata type is anomocytic whilethe rare type is actinocytic recorded only in Ehelioscopia Moreover sometimes stomata of more thanone type (up to three or four types) have beenencountered on the same leaf surface as in E scordiifolia (three types on the abaxial surface) and E hirta (five

types on the abaxial and three on the adaxial) Thedistribution and type of stomata are shown in Table 2

In addition in some species such as E falcata and Ehelioscopia the stomata are deeply sunken andrepresented only by cuticular ledges surrounding theminute pores as seen in ESEM

Examples of species lacking subsidiary cells aroundstomata are E peplus E chamaepeplus E granulataE dracunculoides and E acalyphoides Therefore theyall have anomocytic stomata

On the other hand E balsamifera subsp adenensishas the mononcytic type on both leaf surfaces andanisocytic and paracytic only on the abaxial surface In

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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180 Int Res J Plant Sci

Table 2 Stomata type in Euphorbia taxa in Saudi Arabia

Ab abaxial surface + stomata presentAd adaxial surface

addition E schimperiana has anomocytic andtetracytic on the abaxial only In contrast Ecuneata has three types of stomata anomocyticparacytic and anisocytic only on the lower surfaceAs well ldquoE aff consobrinardquo ldquoE aff schimperirdquo

and E schimperi have paracytic and anisocyticonly in the abaxial on the first two and in both leafsurfaces on the latter Therefore ldquoE affconsobrinardquo and ldquoE aff schimperirdquo are the same taxa Then comparing to E schimperi results

from ansubspec

Eupsimilar and an

Subgenus

Stomata

Anomocytic Hemiparacytic Paracytic Anisocytic T

Species Ad Ab Ad Ab Ad Ab Ad Ab AT i r u c al l i

E balsamifera subsp adenensis + + +

E schimperi + + + +

ldquoEsp aff consobrinardquo + +

ldquoE sp aff schimperirdquo + +

E cuneata + + +

ErmophytonE acalyphoides + +

E pirottae + + + +

E helioscopia + + E s ul a

E grossheimii + + + + +

E retusa + + + + +

E dracunculoides +

E falcate + + E peplus +

E chamaepeplus + +

E schimperiana + C h am a e s y c e

E Arabica + + + + + + +

E serpens + + + +

E scordiifolia + + + + + + +

E granulate + + + +

E inaequilatera + + + +

E hirta + + + + + + +

E hypericifolia + + + + + +

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Table3 Venation type in Euphorbia taxa in Saudi Arabia

Subgenus Species

Venation

Number of veins atthe leaf base

Margins Sheathed

1 2 3 4 or more Open Closed Rounded T i r u c al l i

Ebalsamiferasubspadenensis

+ + +

E schimperi + + +

E aff Schimperi + + +

E cuneata + + +

ErmophytonE acalyphoides + + +

E pirottae + + E s ul a

E grossheimii + +

E retusa + + +

E dracunculoides + + +

E falcate + + +

E peplus + +

E chamaepeplus + +

E schimperiana

E helioscopia

+

+

+

+ C h am a e s y c e

E Arabica + + +

E serpens + + +

E scordiifolia + + +

E granulate + + + +

E inaequilatera + + +

E hirta + + + E hypericifolia + + +

Poinsettia E cyathophora + +

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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182 Int Res J Plant Sci

Figure 10 Sunken stomata on the abaxial surface1) Deeply sunken (A) E helioscopia (B) E falcata (C) E schimperiana 2) Slightly sunken (D) E peplus (E) E pirottae

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 183

983105983138983137983160983145983137983148

983105

983108

983108

983106

983107

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983140983137983160983145983137983148

Figure 11 Stomata types in Euphorbia taxa in Saudi Arabia (A) E balsamifera subsp adenensis (B) E schimperi (C) E cuneata and (D) E grossheimii

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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184 Int Res J Plant Sci

983105

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983105983150983151983149983151983139983161983156983145983139983105983138983137983160

983145983137983148983112983141983149983145983152983137983154983137983139983161983156983145983139

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983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983138983137983160983145983137983148983105983140983137983160983145983137983148

Figure 12 Stomata types in Euphorbia taxa (A) E falcata (B) E peplus (C) E chamaepeplus and (D) Eschimperiana

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 185

983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

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983124983141983156983154983137983139983161983156983145983139

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983105

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983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

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983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

Figure 13 Figure 13 Stomata types in Euphorbia taxa in Saudi Arabia (A) E arabica (B) E serpens (C)E scordiifolia and (D) E helioscopia

distribution between the leaf surfaces (see Figure 13)While E hirta has the most types E helioscopia Edracunculoides and E acalyphoides have the least

Regarding the position of the stomata on the surfacelevel of the epidermis a variable degree of sunkenesshas been recorded Using the ESEM the stomata appear

deeply sunken in some species whilst in others they areonly slightly sunken with the outer rims of the subsidiarycells surrounding the aperture The deeply and slightlysunken species are recorded in Table 2 AdditionallyFigure 10 shows the sunken stomata in different speciesunder the ESEM

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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186 Int Res J Plant Sci

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983112983141983149983145983152983137983154983137983139983161983156983145983139

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983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

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983105983150983145983155983151983139983161983156983145983139

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983107

983106

983105

983105983140983137983160983145983137983148983105983138983137983160983145983137983148

Figure 14 Stomata types in Euphorbia taxa in Saudi Arabia (A) E hirta (B) E hypericifolia (C) E pirottae and (D)Espaff schimperi

Venation

The venation patterns in the Euphorbia species underinvestigation vary from one-veined three-veined to thosewith four or more veins The predominant type is three-veined However in E balsamifera subsp Adenensis

there are seven middle parallel veins three middle thickand four thinner lateral ones (Figure 15 A and B)Whereas in E pirottae there are three main middleparallel veins and two thinner laterals (Figure 15 C) Incontrast E chamaepeplus E schimperiana and Earabica have only one thick main midrib coming from the

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 187

983105

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Figure 15 Figure 15 Venation patterns at base of leaves invariants with more than three veins (A amp B) E balsamiferasubsp adenensis and (C) E pirottae

petiole then branched from the side into two lateral veinsat the point when the leaf starts widening (Figure 16)Each of these veins branches again into secondary veinsat the leaf margin

The rest of the species have just three veins Egrossheimii E dracunculoides E falcata E peplus Eserpens E scordiifolia E granulata E inaequilateraE helioscopia E hirta E hypericifolia and E

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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188 Int Res J Plant Sci

Figure 16 Venation patterns with one main mid vein at the leaf base in (A) E arabica and (B) E chamaepeplus

983106983105

Figure 17 Venation patterns with three-veined leaves in (A) E falcata and (B) E granulata

cyathophora All the three veins come from the petioleand start branching into secondary veins both toward themargin and the mid vein direction (Figure 17 B)

The vein endings at the leaf margin vary from closed

to open There is a relationship between the closedmargins and the sheathed veins (midrib and secondaryveins that are surrounded by parenchyma tissue) Whenthe veins are ornamented (sheathed) the margins areclosed This can be seen in subgenus Chamaesyce inspecies such as E serpens E scordiifolia E granulataE inaequilatera E hirta E hypericifolia and E arabica Additionally these species have the three-vein venation

On the other hand when the veins are not sheathedthe veins ends are open at the leaf margin as in Ebalsamifera subsp adenensis E schimperi E cuneataE acalyphoides E grossheimii E dracunculoides E

falcata E peplus E chamaepeplus E schimperiana Ehelioscopia E pirottae ldquoE aff schimperirdquo and Ecyathophora

CONCLUSIONS

Anatomical characters are seen here to be very useful inEuphorbia classification and identification The resultsobtained from anatomical studies were useful to groupsome species and identify others

Depending on the venation type the Euphorbiaspecies under investigation are divided anatomically intothree main groups see Figure 18

Group one veins are sheathed epidermal cellsdifferentiated on adaxial and abaxial surfaces Group two

983105 983106

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Figure 18 Hierarchial relation of Euphorbia taxa in Saudi Arabia based on anatomical charac

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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190 Int Res J Plant Sci

veins are not sheathed and epidermal cells are alike inboth surfaces And the third group includes species thatdo not belong to any of the previous groups The firstgroup can be further divided into three-veined and one-veined while the second group into three-veined and fouror more veins

In addition E grossheimii E retusa and E pirottae are related by being three-veined open at the marginswith rounded to oval tracheas and elongated cells bothon adaxial and abaxial surfaces However in E retusa the epidermal cells have straight cell walls on the adaxialsurface They are also similar in having hemiparacyticand paracytic types of stomata either on the adaxial orthe abaxial surface

On the other hand species with two differentepidermal cells on the leaf surfaces usually have three-sheathed veins with closed margins and elongatedtrachea such as E serpens E scordiifolia E granulata E inaequilatera E hirta and E hypericifolia which belongto subgenus Chamaesyce according to (Carter andSmith 1988)

In the case of E granulata which has beendescribed as having three different variants E granulata var granulata (both leaf surfaces hairy) E granulata varglabrata (adaxial glabrous and abaxial hairy) and aglabrous variant (the whole plant glabrous) Anatomicalwork has shown especially by using the ESEM that theleaves are either hairy or glabrous on both surfaces Sowhen the leaf is hairy both surfaces have differences inhair distribution Usually the lower surface is more hairythan upper surface (E granulata var glabrata ) butsometimes they are evenly distributed (Egranulata vargranulata ) In addition hairs on both surfaces are

multicellular In contrast when the leaf is glabrous bothleaf surfaces are glabrous and the whole plant isglabrous (suggested new variety) They were similar inhaving the same venation type (three-veined)

Finally ldquoE aff schimperi ldquohas some similarity with E schimperi in having the same open margin not theornamented venation type As well they both have thesame type of stomata paracytic and anisocytic but theydiffer in their distribution on abaxial and adaxial Thusthis may support the suggestion considering ldquoE affschimperi rdquo as a new subspecies of E schimperi

REFERENCES

Baranova M (1992) Principels of comparative stomatographic studiesof flowering plants The Botanical Review 58 49-99

Carlquist S (1961) Comparative Plant Anatomy Holt Rinehart ampWinston New York

Carter S Smith AR (1988) Euphorbiaceae (Part2) In Polhill ed Floraof Tropical East Africa Rotterdam Brookfield AA Balkema 409-597

Chaudhary SA (2001) Flora of The Kingdom of Saudi Arabia Vol 3Ministry of Agriculture and Water Riyadh

Dehay C (1935) Lagraveppareil Libero-ligneux foliare des EuphorbiaceaeAnnales des Sciences Naturelles Botanique 10 147-190

Dehgan B (1982) Comparative anatomy of the petiole and infragenericrelationships in Jatropha (Euphorbiaceae) American Journal oBotany 69 1283-1295

Gaucher L (1898) Ē tude anatomique du genre Euphorbia LPKlincksieck Paris

Gaucher L (1902) Recherches anatomiques sur les EuphorbiaceacuteesAnnales des Sciences Naturelles Botanique 15 161-309

Gaucher L (1902) Recherches anatomiques sur les Euphorbiaceacutees

Annales des Sciences Naturelles Botanique 15 161-309Govaerts R Fordin D Radcliffe-Smith A (2000) World Checklist and

Bibliography of Euphorbiaceae The Royal Botanic Gardens KewHickey LJ (1973) Classification of the architecture of dicotyledonous

leaves American Journal of Botany 60 17-33Hickey LJ (1979) A revised classification of the architecture o

dicotyledonous leaves In Metcalfe CR and Chalk L eds Anatomy oDicotyledons Systematic Anatomy of the leaf and stem 2nd edClarendon Press Oxford 25-39

Kakkar L Paliwal GS (1972) Epidermis in Euphorbia Indian ScienceCongress Association Proceedings 59 326-327

Kakkar L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 5 Epidermis Proceedings of the Indian National ScienceAcademy Part B Biological Sciences 40 55-67

Koumlhler E (1993) Blattvervatur-Muster der Buxaceae Dumortierr undSimmondsiaceae Van Tieghem Feddes Repertorium 104 145-167

Levin GA (1986a) Systematic foliar morphology of Phylanthoideae(Euphorbiaceae) II Phenetic analysis Annals of Missouri BotanicaGarden 73 86-98

Levin GA (1986b) Systematic foliar morphology of Phylanthoideae IICladistic analysis Systematic Botany 11 515-530

Levin GA (1986c) Systematic foliar morphology of Phyllanthoideae(Euphorbiaceae) I conspectus Annals of Missouri Botanical Garden73 29-85

Mahlberg P (1973) Histochemistry and Scanning Electron Microscopyof Starch Grains from Latex of Euphorbia-Terracina and EuphorbiaTirucalli Proceedings of the Indiana Academy of Science 82 132

Mahlberg PG (1974) Scanning Microscopic Comparison of StarchGrains from the Latex of Euphorbia -Spp Proceedings of the IndianaAcademy of Science 83 83-84

Mahlberg PG (1975) Evolution of the Laticifer in Euphorbia asInterpreted from Starch Grain Morphology American Journal oBotany 62 577-583

Mahlberg PG (1982) Comparative Morphology of Starch Grains inLatex from Varieties of Poinsettia Euphorbia-PulcherrimEuphorbiaceae Botanical Gazette 143 206-209

Mahlberg PG Davis DG Galitz DS Manners GD (1987) Laticifers andthe Classification of Euphorbia the Chemotaxonomy of EuphorbiaEsula L Botanical Journal of the Linnean Society 94 165-180

Metcalfe C Chalk L (1950) Anatomy of the Dicotyledons At theClarendon Press Oxford

Miller K Webster GL (1962) Systematic position of Cnidoscolus andJatropha Brittonia 14 174-180

Pax F (1884) Die Anatomie der Euphorbiaceen in ihrer Beziehung zumSystem derselben Botanische Jahrbuumlcher fuumlr SystematikPflanzengeschichte und Pflanzengeographie Leipzig 5 384-421

Payne WW (1970) Helicocytic and allelocytic stomata unrecognizedpatters in the Dicotyledonae Am J Bot57 140-147

Pray TR (1955a) Foliar venation of angiosperm II Histogenesis of thevenation of Liriodendron Am J Bot 42 18-27

Pray TR 1955b Foliar venation of angiosperms III Pattern andhistology of the venation of Hosta Am J Bot 42 11-18

Radford AE Dickison WC Massey JR Bell CR (1974) Vascular PlanSystematics Harper amp Row Publishers London New York

Radlkofer L (1870) Ueber Pausandra ein neues EuphorbiaceenGenus Flora 53 81-95

Raju VS Rao PN (1977) Variation in the structure and development ofoliar stomata in the Euphorbiaceae Botanical Journal of the LinneanSociety 75 69-97

Raju VS Rao PN (1987) The Taxonomic Use of the Basic StomataType in the Generic Delimitation of Chamaesyce EuphorbiaceaeFeddes Repertorium 98 137-142

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2424

Rosowski JR (1968) Laticifer Morphology in the Mature Stem and Leafof Euphorbia supina Botanical Gazette 129 113-120

Roth-Nebelsick A Uhl D Mosbrugger V Kerp H (2001) Evolution andfunction of leaf venation architecture a review Annals of Botany 87553-566

Sehgal L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 2 Venation Patterns Botanical Journal of the Linnean Society 68 173-208

Singh G (2004) Plant Systematics An Integrated Approach SciencePublishers Enfield

Solereder H (1899) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Solereder H (1908) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Aldhebiani and Jury 191

Stace CA (1965) Cuticular studies as an aid to plant taxonomy Bulletinof the British Museum Natural History Botany 4 3-78

Stace CA (1989) Plant Taxonomy and Biosystematics Hodder ampStoughton London

Tognini F (1897) Contribuzione allo studio della organogeniacomparata degli stomi Institut de Botanique University de Geneve 41-42

Van Cotthem WR (1970) A classification of stomatal types Botanica

Journal of Linnean Society 63 235-246Wagner W (1979) Reticulate veins in the systematics of modern ferns

Taxon 1 87-95Webster GL (1994a) Classification of the Euphorbiaceae Annals of the

Missouri Botanical Garden 81 3-32

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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cells and hairs1- Leaves were rehydrated using polyoxyethylenesorbitan monolaurate (Tween20) in water and heated for30 minutes Then washed by water and conserved in70 ethanol for future work2- Both surfaces of the leaf were scraped and

peeled off under a dissection microscope with the aid offine forceps and a razor blade to remove loose cells3- Samples were bleached by Jefferyrsquos Solution orthe bleaching agent Vortex for 10-20 minutes dependingon leaf thickness Sometimes this was overnight4- Samples were washed mounted and observedunder a light microscope5- Photographs were taken using a Leitz Diaphlanpolarizing microscope or a Reichert Polyvar 2Microscope

Staining by alucin blue was tried but withoutsuccess Samples became too dense or obscure due tothe presence of tannins and other compounds in theherbarium specimens Therefore some samples were leftwithout staining

Leaf clearing

The following method was adopted from Radford et al(1974) with some modifications for use on herbariummaterials1- The leaf was placed in a Petri dish and coveredwith 5 Sodium Hydroxide (NaOH) The dish waswrapped in cling film leaving a small gap in one area forventilation2- This was microwaved for five seconds on

medium power3- The Petri dishes were left on a hot plate at 35-37degrees centigrade for a week or until the leaf wastransparent The NaOH was changed twice a day foreach sample4- When the leaf had cleared sufficiently it waswashed in water and covered in 90 bleaching agentVortex for ten minutes to one hour depending on thereaction of the sample5- The leaf was washed again in water6- Samples were dehydrated in ethanol series andstained as followinga) 30 ethanol 2 minutes

b) 50 ethanol 2 minutesc) 1 safranin in 50 ethanol 4 minutesd) Wash in water three timese) 30 ethanol 2 minutes repeated 3 timesf) 50 ethanol 2 minutes repeated twiceg) 70 ethanol 2 minutesh) 90 ethanol 2 minutesi) 100 ethanol 2 minutes

j) Absolute ethanol histoclear 11 2 minutesk) Histoclear 2 minutes

Aldhebiani and Jury 171

l) Mounted in 100 glycerol and sealed with Candabalsamm) Observed under light microscopen) Photographs were taken using a Leitz Diaphlanpolarizing microscope or Reichert Polyvar 2 Microscope

Scanning Electron Microscopy

1- Dry specimens were mounted surface up onscanning electron microscope stubs using Bostik No1adhesive2- Stubs were sputter coated for 2-3 minutes with agold palladium alloy using an Edwards sputter coater togive a coating about 15-20 nm thick3- Samples were examined in FEI Quanta FEG 600Environmental Scanning Electron Microscope (ESEM) inhigh vacuum SEM mode4- Photographs were taken using the computerizeddigital system of the microscope

RESULTS

Epidermis

Results of the epidermal characters in Euphorbia speciesunder investigation are summarized in Table 1

Cell shape

Depending upon the shape of the cell wall and

considering the results from Kakkar and Paliwal (1974)the shape of the epidermal cell varies polygonalrectangular undulate or elongated Moreover the celshape relies on the cell location on a leaf ie the middleregion the margin the apex or above the veinFurthermore in some cases both leaf surfaces have thesame cell shape but more often they are unlike Thiscombination of cell shape characters gives each speciesits unique leaf surface appearance

The normal straight cell walls in both adaxial andabaxial surfaces are shown in the following taxa Eschimperi E cuneata E helioscopia E dracunculoidesand E balsamifera subsp adenensis see Figure 1 In

addition ldquoE aff schimperirdquo and ldquoE aff consobrinardquo havealso the same straight cell wall on both surfacesMeanwhile E grossheimii E retusa and E pirottae haveelongated cells on both leaf surfaces (Figure 2 a and b)On the other hand the plicate cell walls on both adaxiaand abaxial surfaces occur in E chamaepeplus Ehypericifolia and E peplus (Figure 2 c and d) but thewalls are more folded on the lower surface In somespecies the cell wall on the adaxial surface is straightwhilst it is plicate on the abaxial such as E hirta E

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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172 Int Res J Plant Sci

Table 1 Epidermal characters of Euphorbia taxa in Saudi Arabia

Ab abaxial + PresentAd adaxial ++ the plicate folded wall is morefully developed in the lower surface than in the upper one

+ hardly observable by LM only

Subgenus

Cell shape (cell wall) Hairs

Straight Elongated Plicate Unicellular MultSpecies Ad Ab Ad Ab Ad Ab Ad Ab Ad T

i r u c al l i

E balsamifera subspadenensis + + +

E schimperi + +

ldquoEaff schimperirdquo + +

ldquoE aff consobrinardquo + +

E cuneata + +

ErmophytonE acalyphoides + + +

E pirottae + + +

E helioscopia + + E

s ul a

E grossheimii+ +

E retusa + +

E dracunculoides + +

E falcate + +

E peplus + ++

E chamaepeplus

E schimperiana +

+ ++

+ C h am a e s y c e

E Arabica + +

E serpens + +

E scordiifolia + + +

E granulate + + + + +

E inaequilatera + +

E hirta + + +

E hypericifolia + ++

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 173

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Figure 1 Normal irregular epidermal cells in both adaxial and abaxial surfaces of the leaf (LM)(A) E balsamifera subsp adenensis (B) E schimperi (C) E cuneata (D) E falcata (E) Abaxial in E dracunculoides

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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174 Int Res J Plant Sci

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983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

Figure 2 Error No text of specified style in document Both adaxial and abaxial surfaceshave the same shape of epidermal cells (LM) Elongated in A ampB plicate in CampD (A) Egrossheimii (B) E pirottae (C) E chamaepeplus and (D) E peplus

schimperiana E helioscopia E inaequilatera Earabica E granulata E serpens and E scordiifolia see Figure 3 and Figure 4 This last group of species belongto subgenus Chamaesyce except for E helioscopia and E schimperiana

Hairs

Hairs on the leaf epidermis can be both unicellular and

multicellular and may appear only on one surface or bothFurthermore hairs in the Euphorbia species undeinvestigation are generally simple unbranched and with awarty ornamentation on their surface Papillae appear inESEM as outgrowths whereas in the surface view theygive the impression of being rounded structures at thecentre of the cell Prominent papillae occur only in onespecies of Euphorbia in Saudi Arabia this species is Ehypericifolia (Figure 5 and 6) whereas in E arabica theywere poorly distinguishable using LM

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 175

983105983140983137983160983145983137983148

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983105983138983137983160983145983137983148

Figure 3 Epidermal cells vary on leaf surfaces adaxial surface has irregular cells whilethe abaxial surface has plicate (LM)(A) E schimperiana (B) E arabica (C) E serpens (D) E scordiifolia

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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176 Int Res J Plant Sci

983105983140983137983160983145983137983148

983107

983105

983106

983105983138983137983160983145983137983148

Figure 4 Epidermal cell shape varies on the adaxial andthe abaxial surfaces straight cell walls in the formerwhile folded plicate walls occur on the latter (LM) (A) Egranulata (B) E inaequilatera and (C) E helioscopia

983112983141983149983145983152983137983154983137983139983161983156983145983139 983155983156983151983149983137983156983137

983151983150 983138983151983156983144 983155983157983154983142983137983139983141983155

Figure 5 E hypericifolia (LM)Above photo the adaxial surface with irregular epidermal cells andvery clear papillae as a circle in the middle of the cellLower photo the abaxial surface with plicate cells and papillae ahair base is obvious in the right side of the picture

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 177

983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

983120983137983152983145983148983148983137983141

983117983157983148983156983145983139983141983148983148983157983148983137983154 983144983137983145983154983155 983145983150 983137983138983137983160983145983137983148 983155983157983154983142983137983139983141983086

Figure 6 E hypericifolia (ESEM) papillae on the adaxial and abaxial surfacesmulticellular hairs are on the abaxial surface of the leaf only

Figure7 Multicellular hairs on adaxial and abaxial surfaces of E hirta

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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178 Int Res J Plant Sci

Figure 8 Hairs wart ornamentation on hair surfaces of (A) E scordiifolia and (B) E pirottae (C) Short hairs onabaxial surface of E balsamifera subsp adenensis

In E balsamifera subsp adenensis the hairs are shortand unicellular (Figure 8C) and they are located mostly atthe leaf margins and the mid-vein area on the abaxialsurface only

Hairs on the leaf of E pirottae are present on theabaxial surface only and are both unicellular andmulticellular Euphorbia granulata is similar to E pirottae in having both unicellular and multicellular hairs but herethey occur on both surfaces of the leaf (Figure 8B)

On the other hand in E scordiifolia E hirta and Ehypericifolia the hairs are multicellular and on bothsurfaces in the first two species and on the abaxial only inthe latter (Figure 7) Moreover E hypericifolia and Ehirta have long hairs when compared with the otherspecies (Figure 6)

No hairs were observed and leaves were glabrous inthe following species E serpens E arabica E peplusE inaequilatera E falcata E helioscopia E cuneata Eretusa E grossheimii E schimperiana E schimperiE chamaepeplus and the new variety of E granulata

Wax

The cuticle layer and wax in the Euphorbia species undeinvestigation vary from smooth to densely cover in waxscales This may be a thick smooth layer of wax whichsometimes has upright scales In some species only thecell surface has a smooth wax layer whereas the walbetween cells has upright scales of wax as in Eschimperiana especially around the stomata pores

(Figure 9) In contrast the anticlinal walls of somespecies have a smooth layer of wax whereas the celsurface is densely covered with upright wax flakes or withsome stellate wax flakes in other species

Stomata

Stomata were observed through LM and ESEM In thespecies under investigation stomata commonly appeaon both leaf surfaces (amphistomatic leaves) excepin E cuneata where the stomata are located only on the

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 179

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Figure 9 Wax formation in Euphorbia species in Saudi Arabia (A+B) smooth andthick in E peplus and E schimperi (C+D) smooth on top of cell while formingupright scale flakes in between especially around stomata as in E schimperiana smooth in anticlinal walls (E) upright scale flakes in E balsamifera

abaxial surface (hypostomatic leaves) The presence andposition of subsidiary cells are variable in Euphorbia

species Subsidiary cells were found to be of unequalsize and shape whether the stoma was anisocyticparacytic actinocytic or tetracytic While some specieslack them around the stomata others have more thanone arrangement of subsidiary cells

The most common stomata type is anomocytic whilethe rare type is actinocytic recorded only in Ehelioscopia Moreover sometimes stomata of more thanone type (up to three or four types) have beenencountered on the same leaf surface as in E scordiifolia (three types on the abaxial surface) and E hirta (five

types on the abaxial and three on the adaxial) Thedistribution and type of stomata are shown in Table 2

In addition in some species such as E falcata and Ehelioscopia the stomata are deeply sunken andrepresented only by cuticular ledges surrounding theminute pores as seen in ESEM

Examples of species lacking subsidiary cells aroundstomata are E peplus E chamaepeplus E granulataE dracunculoides and E acalyphoides Therefore theyall have anomocytic stomata

On the other hand E balsamifera subsp adenensishas the mononcytic type on both leaf surfaces andanisocytic and paracytic only on the abaxial surface In

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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180 Int Res J Plant Sci

Table 2 Stomata type in Euphorbia taxa in Saudi Arabia

Ab abaxial surface + stomata presentAd adaxial surface

addition E schimperiana has anomocytic andtetracytic on the abaxial only In contrast Ecuneata has three types of stomata anomocyticparacytic and anisocytic only on the lower surfaceAs well ldquoE aff consobrinardquo ldquoE aff schimperirdquo

and E schimperi have paracytic and anisocyticonly in the abaxial on the first two and in both leafsurfaces on the latter Therefore ldquoE affconsobrinardquo and ldquoE aff schimperirdquo are the same taxa Then comparing to E schimperi results

from ansubspec

Eupsimilar and an

Subgenus

Stomata

Anomocytic Hemiparacytic Paracytic Anisocytic T

Species Ad Ab Ad Ab Ad Ab Ad Ab AT i r u c al l i

E balsamifera subsp adenensis + + +

E schimperi + + + +

ldquoEsp aff consobrinardquo + +

ldquoE sp aff schimperirdquo + +

E cuneata + + +

ErmophytonE acalyphoides + +

E pirottae + + + +

E helioscopia + + E s ul a

E grossheimii + + + + +

E retusa + + + + +

E dracunculoides +

E falcate + + E peplus +

E chamaepeplus + +

E schimperiana + C h am a e s y c e

E Arabica + + + + + + +

E serpens + + + +

E scordiifolia + + + + + + +

E granulate + + + +

E inaequilatera + + + +

E hirta + + + + + + +

E hypericifolia + + + + + +

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Table3 Venation type in Euphorbia taxa in Saudi Arabia

Subgenus Species

Venation

Number of veins atthe leaf base

Margins Sheathed

1 2 3 4 or more Open Closed Rounded T i r u c al l i

Ebalsamiferasubspadenensis

+ + +

E schimperi + + +

E aff Schimperi + + +

E cuneata + + +

ErmophytonE acalyphoides + + +

E pirottae + + E s ul a

E grossheimii + +

E retusa + + +

E dracunculoides + + +

E falcate + + +

E peplus + +

E chamaepeplus + +

E schimperiana

E helioscopia

+

+

+

+ C h am a e s y c e

E Arabica + + +

E serpens + + +

E scordiifolia + + +

E granulate + + + +

E inaequilatera + + +

E hirta + + + E hypericifolia + + +

Poinsettia E cyathophora + +

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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182 Int Res J Plant Sci

Figure 10 Sunken stomata on the abaxial surface1) Deeply sunken (A) E helioscopia (B) E falcata (C) E schimperiana 2) Slightly sunken (D) E peplus (E) E pirottae

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 183

983105983138983137983160983145983137983148

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983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

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983112983141983149983145983152983137983154983137983139983161983156983145983139983120983137983154983137983139983161983156983145983139

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983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983140983137983160983145983137983148

Figure 11 Stomata types in Euphorbia taxa in Saudi Arabia (A) E balsamifera subsp adenensis (B) E schimperi (C) E cuneata and (D) E grossheimii

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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184 Int Res J Plant Sci

983105

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983105983150983151983149983151983139983161983156983145983139983105983138983137983160

983145983137983148983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

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983112983141983149983145983152983137983154983137983139983161983156983145983139

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983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983138983137983160983145983137983148983105983140983137983160983145983137983148

Figure 12 Stomata types in Euphorbia taxa (A) E falcata (B) E peplus (C) E chamaepeplus and (D) Eschimperiana

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 185

983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

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983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105

983108

983106

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983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

Figure 13 Figure 13 Stomata types in Euphorbia taxa in Saudi Arabia (A) E arabica (B) E serpens (C)E scordiifolia and (D) E helioscopia

distribution between the leaf surfaces (see Figure 13)While E hirta has the most types E helioscopia Edracunculoides and E acalyphoides have the least

Regarding the position of the stomata on the surfacelevel of the epidermis a variable degree of sunkenesshas been recorded Using the ESEM the stomata appear

deeply sunken in some species whilst in others they areonly slightly sunken with the outer rims of the subsidiarycells surrounding the aperture The deeply and slightlysunken species are recorded in Table 2 AdditionallyFigure 10 shows the sunken stomata in different speciesunder the ESEM

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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186 Int Res J Plant Sci

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

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983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

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983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983107

983106

983105

983105983140983137983160983145983137983148983105983138983137983160983145983137983148

Figure 14 Stomata types in Euphorbia taxa in Saudi Arabia (A) E hirta (B) E hypericifolia (C) E pirottae and (D)Espaff schimperi

Venation

The venation patterns in the Euphorbia species underinvestigation vary from one-veined three-veined to thosewith four or more veins The predominant type is three-veined However in E balsamifera subsp Adenensis

there are seven middle parallel veins three middle thickand four thinner lateral ones (Figure 15 A and B)Whereas in E pirottae there are three main middleparallel veins and two thinner laterals (Figure 15 C) Incontrast E chamaepeplus E schimperiana and Earabica have only one thick main midrib coming from the

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 187

983105

983106

983107

Figure 15 Figure 15 Venation patterns at base of leaves invariants with more than three veins (A amp B) E balsamiferasubsp adenensis and (C) E pirottae

petiole then branched from the side into two lateral veinsat the point when the leaf starts widening (Figure 16)Each of these veins branches again into secondary veinsat the leaf margin

The rest of the species have just three veins Egrossheimii E dracunculoides E falcata E peplus Eserpens E scordiifolia E granulata E inaequilateraE helioscopia E hirta E hypericifolia and E

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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188 Int Res J Plant Sci

Figure 16 Venation patterns with one main mid vein at the leaf base in (A) E arabica and (B) E chamaepeplus

983106983105

Figure 17 Venation patterns with three-veined leaves in (A) E falcata and (B) E granulata

cyathophora All the three veins come from the petioleand start branching into secondary veins both toward themargin and the mid vein direction (Figure 17 B)

The vein endings at the leaf margin vary from closed

to open There is a relationship between the closedmargins and the sheathed veins (midrib and secondaryveins that are surrounded by parenchyma tissue) Whenthe veins are ornamented (sheathed) the margins areclosed This can be seen in subgenus Chamaesyce inspecies such as E serpens E scordiifolia E granulataE inaequilatera E hirta E hypericifolia and E arabica Additionally these species have the three-vein venation

On the other hand when the veins are not sheathedthe veins ends are open at the leaf margin as in Ebalsamifera subsp adenensis E schimperi E cuneataE acalyphoides E grossheimii E dracunculoides E

falcata E peplus E chamaepeplus E schimperiana Ehelioscopia E pirottae ldquoE aff schimperirdquo and Ecyathophora

CONCLUSIONS

Anatomical characters are seen here to be very useful inEuphorbia classification and identification The resultsobtained from anatomical studies were useful to groupsome species and identify others

Depending on the venation type the Euphorbiaspecies under investigation are divided anatomically intothree main groups see Figure 18

Group one veins are sheathed epidermal cellsdifferentiated on adaxial and abaxial surfaces Group two

983105 983106

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Figure 18 Hierarchial relation of Euphorbia taxa in Saudi Arabia based on anatomical charac

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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190 Int Res J Plant Sci

veins are not sheathed and epidermal cells are alike inboth surfaces And the third group includes species thatdo not belong to any of the previous groups The firstgroup can be further divided into three-veined and one-veined while the second group into three-veined and fouror more veins

In addition E grossheimii E retusa and E pirottae are related by being three-veined open at the marginswith rounded to oval tracheas and elongated cells bothon adaxial and abaxial surfaces However in E retusa the epidermal cells have straight cell walls on the adaxialsurface They are also similar in having hemiparacyticand paracytic types of stomata either on the adaxial orthe abaxial surface

On the other hand species with two differentepidermal cells on the leaf surfaces usually have three-sheathed veins with closed margins and elongatedtrachea such as E serpens E scordiifolia E granulata E inaequilatera E hirta and E hypericifolia which belongto subgenus Chamaesyce according to (Carter andSmith 1988)

In the case of E granulata which has beendescribed as having three different variants E granulata var granulata (both leaf surfaces hairy) E granulata varglabrata (adaxial glabrous and abaxial hairy) and aglabrous variant (the whole plant glabrous) Anatomicalwork has shown especially by using the ESEM that theleaves are either hairy or glabrous on both surfaces Sowhen the leaf is hairy both surfaces have differences inhair distribution Usually the lower surface is more hairythan upper surface (E granulata var glabrata ) butsometimes they are evenly distributed (Egranulata vargranulata ) In addition hairs on both surfaces are

multicellular In contrast when the leaf is glabrous bothleaf surfaces are glabrous and the whole plant isglabrous (suggested new variety) They were similar inhaving the same venation type (three-veined)

Finally ldquoE aff schimperi ldquohas some similarity with E schimperi in having the same open margin not theornamented venation type As well they both have thesame type of stomata paracytic and anisocytic but theydiffer in their distribution on abaxial and adaxial Thusthis may support the suggestion considering ldquoE affschimperi rdquo as a new subspecies of E schimperi

REFERENCES

Baranova M (1992) Principels of comparative stomatographic studiesof flowering plants The Botanical Review 58 49-99

Carlquist S (1961) Comparative Plant Anatomy Holt Rinehart ampWinston New York

Carter S Smith AR (1988) Euphorbiaceae (Part2) In Polhill ed Floraof Tropical East Africa Rotterdam Brookfield AA Balkema 409-597

Chaudhary SA (2001) Flora of The Kingdom of Saudi Arabia Vol 3Ministry of Agriculture and Water Riyadh

Dehay C (1935) Lagraveppareil Libero-ligneux foliare des EuphorbiaceaeAnnales des Sciences Naturelles Botanique 10 147-190

Dehgan B (1982) Comparative anatomy of the petiole and infragenericrelationships in Jatropha (Euphorbiaceae) American Journal oBotany 69 1283-1295

Gaucher L (1898) Ē tude anatomique du genre Euphorbia LPKlincksieck Paris

Gaucher L (1902) Recherches anatomiques sur les EuphorbiaceacuteesAnnales des Sciences Naturelles Botanique 15 161-309

Gaucher L (1902) Recherches anatomiques sur les Euphorbiaceacutees

Annales des Sciences Naturelles Botanique 15 161-309Govaerts R Fordin D Radcliffe-Smith A (2000) World Checklist and

Bibliography of Euphorbiaceae The Royal Botanic Gardens KewHickey LJ (1973) Classification of the architecture of dicotyledonous

leaves American Journal of Botany 60 17-33Hickey LJ (1979) A revised classification of the architecture o

dicotyledonous leaves In Metcalfe CR and Chalk L eds Anatomy oDicotyledons Systematic Anatomy of the leaf and stem 2nd edClarendon Press Oxford 25-39

Kakkar L Paliwal GS (1972) Epidermis in Euphorbia Indian ScienceCongress Association Proceedings 59 326-327

Kakkar L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 5 Epidermis Proceedings of the Indian National ScienceAcademy Part B Biological Sciences 40 55-67

Koumlhler E (1993) Blattvervatur-Muster der Buxaceae Dumortierr undSimmondsiaceae Van Tieghem Feddes Repertorium 104 145-167

Levin GA (1986a) Systematic foliar morphology of Phylanthoideae(Euphorbiaceae) II Phenetic analysis Annals of Missouri BotanicaGarden 73 86-98

Levin GA (1986b) Systematic foliar morphology of Phylanthoideae IICladistic analysis Systematic Botany 11 515-530

Levin GA (1986c) Systematic foliar morphology of Phyllanthoideae(Euphorbiaceae) I conspectus Annals of Missouri Botanical Garden73 29-85

Mahlberg P (1973) Histochemistry and Scanning Electron Microscopyof Starch Grains from Latex of Euphorbia-Terracina and EuphorbiaTirucalli Proceedings of the Indiana Academy of Science 82 132

Mahlberg PG (1974) Scanning Microscopic Comparison of StarchGrains from the Latex of Euphorbia -Spp Proceedings of the IndianaAcademy of Science 83 83-84

Mahlberg PG (1975) Evolution of the Laticifer in Euphorbia asInterpreted from Starch Grain Morphology American Journal oBotany 62 577-583

Mahlberg PG (1982) Comparative Morphology of Starch Grains inLatex from Varieties of Poinsettia Euphorbia-PulcherrimEuphorbiaceae Botanical Gazette 143 206-209

Mahlberg PG Davis DG Galitz DS Manners GD (1987) Laticifers andthe Classification of Euphorbia the Chemotaxonomy of EuphorbiaEsula L Botanical Journal of the Linnean Society 94 165-180

Metcalfe C Chalk L (1950) Anatomy of the Dicotyledons At theClarendon Press Oxford

Miller K Webster GL (1962) Systematic position of Cnidoscolus andJatropha Brittonia 14 174-180

Pax F (1884) Die Anatomie der Euphorbiaceen in ihrer Beziehung zumSystem derselben Botanische Jahrbuumlcher fuumlr SystematikPflanzengeschichte und Pflanzengeographie Leipzig 5 384-421

Payne WW (1970) Helicocytic and allelocytic stomata unrecognizedpatters in the Dicotyledonae Am J Bot57 140-147

Pray TR (1955a) Foliar venation of angiosperm II Histogenesis of thevenation of Liriodendron Am J Bot 42 18-27

Pray TR 1955b Foliar venation of angiosperms III Pattern andhistology of the venation of Hosta Am J Bot 42 11-18

Radford AE Dickison WC Massey JR Bell CR (1974) Vascular PlanSystematics Harper amp Row Publishers London New York

Radlkofer L (1870) Ueber Pausandra ein neues EuphorbiaceenGenus Flora 53 81-95

Raju VS Rao PN (1977) Variation in the structure and development ofoliar stomata in the Euphorbiaceae Botanical Journal of the LinneanSociety 75 69-97

Raju VS Rao PN (1987) The Taxonomic Use of the Basic StomataType in the Generic Delimitation of Chamaesyce EuphorbiaceaeFeddes Repertorium 98 137-142

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2424

Rosowski JR (1968) Laticifer Morphology in the Mature Stem and Leafof Euphorbia supina Botanical Gazette 129 113-120

Roth-Nebelsick A Uhl D Mosbrugger V Kerp H (2001) Evolution andfunction of leaf venation architecture a review Annals of Botany 87553-566

Sehgal L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 2 Venation Patterns Botanical Journal of the Linnean Society 68 173-208

Singh G (2004) Plant Systematics An Integrated Approach SciencePublishers Enfield

Solereder H (1899) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Solereder H (1908) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Aldhebiani and Jury 191

Stace CA (1965) Cuticular studies as an aid to plant taxonomy Bulletinof the British Museum Natural History Botany 4 3-78

Stace CA (1989) Plant Taxonomy and Biosystematics Hodder ampStoughton London

Tognini F (1897) Contribuzione allo studio della organogeniacomparata degli stomi Institut de Botanique University de Geneve 41-42

Van Cotthem WR (1970) A classification of stomatal types Botanica

Journal of Linnean Society 63 235-246Wagner W (1979) Reticulate veins in the systematics of modern ferns

Taxon 1 87-95Webster GL (1994a) Classification of the Euphorbiaceae Annals of the

Missouri Botanical Garden 81 3-32

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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172 Int Res J Plant Sci

Table 1 Epidermal characters of Euphorbia taxa in Saudi Arabia

Ab abaxial + PresentAd adaxial ++ the plicate folded wall is morefully developed in the lower surface than in the upper one

+ hardly observable by LM only

Subgenus

Cell shape (cell wall) Hairs

Straight Elongated Plicate Unicellular MultSpecies Ad Ab Ad Ab Ad Ab Ad Ab Ad T

i r u c al l i

E balsamifera subspadenensis + + +

E schimperi + +

ldquoEaff schimperirdquo + +

ldquoE aff consobrinardquo + +

E cuneata + +

ErmophytonE acalyphoides + + +

E pirottae + + +

E helioscopia + + E

s ul a

E grossheimii+ +

E retusa + +

E dracunculoides + +

E falcate + +

E peplus + ++

E chamaepeplus

E schimperiana +

+ ++

+ C h am a e s y c e

E Arabica + +

E serpens + +

E scordiifolia + + +

E granulate + + + + +

E inaequilatera + +

E hirta + + +

E hypericifolia + ++

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 173

983106

983108

983109

983105

983107

983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

Figure 1 Normal irregular epidermal cells in both adaxial and abaxial surfaces of the leaf (LM)(A) E balsamifera subsp adenensis (B) E schimperi (C) E cuneata (D) E falcata (E) Abaxial in E dracunculoides

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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174 Int Res J Plant Sci

983105

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983106 983106

983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

Figure 2 Error No text of specified style in document Both adaxial and abaxial surfaceshave the same shape of epidermal cells (LM) Elongated in A ampB plicate in CampD (A) Egrossheimii (B) E pirottae (C) E chamaepeplus and (D) E peplus

schimperiana E helioscopia E inaequilatera Earabica E granulata E serpens and E scordiifolia see Figure 3 and Figure 4 This last group of species belongto subgenus Chamaesyce except for E helioscopia and E schimperiana

Hairs

Hairs on the leaf epidermis can be both unicellular and

multicellular and may appear only on one surface or bothFurthermore hairs in the Euphorbia species undeinvestigation are generally simple unbranched and with awarty ornamentation on their surface Papillae appear inESEM as outgrowths whereas in the surface view theygive the impression of being rounded structures at thecentre of the cell Prominent papillae occur only in onespecies of Euphorbia in Saudi Arabia this species is Ehypericifolia (Figure 5 and 6) whereas in E arabica theywere poorly distinguishable using LM

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 175

983105983140983137983160983145983137983148

983107

983108

983105

983106

983105983138983137983160983145983137983148

Figure 3 Epidermal cells vary on leaf surfaces adaxial surface has irregular cells whilethe abaxial surface has plicate (LM)(A) E schimperiana (B) E arabica (C) E serpens (D) E scordiifolia

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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176 Int Res J Plant Sci

983105983140983137983160983145983137983148

983107

983105

983106

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Figure 4 Epidermal cell shape varies on the adaxial andthe abaxial surfaces straight cell walls in the formerwhile folded plicate walls occur on the latter (LM) (A) Egranulata (B) E inaequilatera and (C) E helioscopia

983112983141983149983145983152983137983154983137983139983161983156983145983139 983155983156983151983149983137983156983137

983151983150 983138983151983156983144 983155983157983154983142983137983139983141983155

Figure 5 E hypericifolia (LM)Above photo the adaxial surface with irregular epidermal cells andvery clear papillae as a circle in the middle of the cellLower photo the abaxial surface with plicate cells and papillae ahair base is obvious in the right side of the picture

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 177

983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

983120983137983152983145983148983148983137983141

983117983157983148983156983145983139983141983148983148983157983148983137983154 983144983137983145983154983155 983145983150 983137983138983137983160983145983137983148 983155983157983154983142983137983139983141983086

Figure 6 E hypericifolia (ESEM) papillae on the adaxial and abaxial surfacesmulticellular hairs are on the abaxial surface of the leaf only

Figure7 Multicellular hairs on adaxial and abaxial surfaces of E hirta

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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178 Int Res J Plant Sci

Figure 8 Hairs wart ornamentation on hair surfaces of (A) E scordiifolia and (B) E pirottae (C) Short hairs onabaxial surface of E balsamifera subsp adenensis

In E balsamifera subsp adenensis the hairs are shortand unicellular (Figure 8C) and they are located mostly atthe leaf margins and the mid-vein area on the abaxialsurface only

Hairs on the leaf of E pirottae are present on theabaxial surface only and are both unicellular andmulticellular Euphorbia granulata is similar to E pirottae in having both unicellular and multicellular hairs but herethey occur on both surfaces of the leaf (Figure 8B)

On the other hand in E scordiifolia E hirta and Ehypericifolia the hairs are multicellular and on bothsurfaces in the first two species and on the abaxial only inthe latter (Figure 7) Moreover E hypericifolia and Ehirta have long hairs when compared with the otherspecies (Figure 6)

No hairs were observed and leaves were glabrous inthe following species E serpens E arabica E peplusE inaequilatera E falcata E helioscopia E cuneata Eretusa E grossheimii E schimperiana E schimperiE chamaepeplus and the new variety of E granulata

Wax

The cuticle layer and wax in the Euphorbia species undeinvestigation vary from smooth to densely cover in waxscales This may be a thick smooth layer of wax whichsometimes has upright scales In some species only thecell surface has a smooth wax layer whereas the walbetween cells has upright scales of wax as in Eschimperiana especially around the stomata pores

(Figure 9) In contrast the anticlinal walls of somespecies have a smooth layer of wax whereas the celsurface is densely covered with upright wax flakes or withsome stellate wax flakes in other species

Stomata

Stomata were observed through LM and ESEM In thespecies under investigation stomata commonly appeaon both leaf surfaces (amphistomatic leaves) excepin E cuneata where the stomata are located only on the

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 179

983109

983105

983108983107

983106

Figure 9 Wax formation in Euphorbia species in Saudi Arabia (A+B) smooth andthick in E peplus and E schimperi (C+D) smooth on top of cell while formingupright scale flakes in between especially around stomata as in E schimperiana smooth in anticlinal walls (E) upright scale flakes in E balsamifera

abaxial surface (hypostomatic leaves) The presence andposition of subsidiary cells are variable in Euphorbia

species Subsidiary cells were found to be of unequalsize and shape whether the stoma was anisocyticparacytic actinocytic or tetracytic While some specieslack them around the stomata others have more thanone arrangement of subsidiary cells

The most common stomata type is anomocytic whilethe rare type is actinocytic recorded only in Ehelioscopia Moreover sometimes stomata of more thanone type (up to three or four types) have beenencountered on the same leaf surface as in E scordiifolia (three types on the abaxial surface) and E hirta (five

types on the abaxial and three on the adaxial) Thedistribution and type of stomata are shown in Table 2

In addition in some species such as E falcata and Ehelioscopia the stomata are deeply sunken andrepresented only by cuticular ledges surrounding theminute pores as seen in ESEM

Examples of species lacking subsidiary cells aroundstomata are E peplus E chamaepeplus E granulataE dracunculoides and E acalyphoides Therefore theyall have anomocytic stomata

On the other hand E balsamifera subsp adenensishas the mononcytic type on both leaf surfaces andanisocytic and paracytic only on the abaxial surface In

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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180 Int Res J Plant Sci

Table 2 Stomata type in Euphorbia taxa in Saudi Arabia

Ab abaxial surface + stomata presentAd adaxial surface

addition E schimperiana has anomocytic andtetracytic on the abaxial only In contrast Ecuneata has three types of stomata anomocyticparacytic and anisocytic only on the lower surfaceAs well ldquoE aff consobrinardquo ldquoE aff schimperirdquo

and E schimperi have paracytic and anisocyticonly in the abaxial on the first two and in both leafsurfaces on the latter Therefore ldquoE affconsobrinardquo and ldquoE aff schimperirdquo are the same taxa Then comparing to E schimperi results

from ansubspec

Eupsimilar and an

Subgenus

Stomata

Anomocytic Hemiparacytic Paracytic Anisocytic T

Species Ad Ab Ad Ab Ad Ab Ad Ab AT i r u c al l i

E balsamifera subsp adenensis + + +

E schimperi + + + +

ldquoEsp aff consobrinardquo + +

ldquoE sp aff schimperirdquo + +

E cuneata + + +

ErmophytonE acalyphoides + +

E pirottae + + + +

E helioscopia + + E s ul a

E grossheimii + + + + +

E retusa + + + + +

E dracunculoides +

E falcate + + E peplus +

E chamaepeplus + +

E schimperiana + C h am a e s y c e

E Arabica + + + + + + +

E serpens + + + +

E scordiifolia + + + + + + +

E granulate + + + +

E inaequilatera + + + +

E hirta + + + + + + +

E hypericifolia + + + + + +

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Table3 Venation type in Euphorbia taxa in Saudi Arabia

Subgenus Species

Venation

Number of veins atthe leaf base

Margins Sheathed

1 2 3 4 or more Open Closed Rounded T i r u c al l i

Ebalsamiferasubspadenensis

+ + +

E schimperi + + +

E aff Schimperi + + +

E cuneata + + +

ErmophytonE acalyphoides + + +

E pirottae + + E s ul a

E grossheimii + +

E retusa + + +

E dracunculoides + + +

E falcate + + +

E peplus + +

E chamaepeplus + +

E schimperiana

E helioscopia

+

+

+

+ C h am a e s y c e

E Arabica + + +

E serpens + + +

E scordiifolia + + +

E granulate + + + +

E inaequilatera + + +

E hirta + + + E hypericifolia + + +

Poinsettia E cyathophora + +

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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182 Int Res J Plant Sci

Figure 10 Sunken stomata on the abaxial surface1) Deeply sunken (A) E helioscopia (B) E falcata (C) E schimperiana 2) Slightly sunken (D) E peplus (E) E pirottae

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 183

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983112983141983149983145983152983137983154983137983139983161983156983145983139

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983105983139983156983145983150983151983139983161983156983145983139

983105983140983137983160983145983137983148

Figure 11 Stomata types in Euphorbia taxa in Saudi Arabia (A) E balsamifera subsp adenensis (B) E schimperi (C) E cuneata and (D) E grossheimii

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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184 Int Res J Plant Sci

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983124983141983156983154983137983139983161983156983145983139

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983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983138983137983160983145983137983148983105983140983137983160983145983137983148

Figure 12 Stomata types in Euphorbia taxa (A) E falcata (B) E peplus (C) E chamaepeplus and (D) Eschimperiana

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 185

983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

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983112983141983149983145983152983137983154983137983139983161983156983145983139

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983124983141983156983154983137983139983161983156983145983139

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983105983150983151983149983151983139983161983156983145983139

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983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

Figure 13 Figure 13 Stomata types in Euphorbia taxa in Saudi Arabia (A) E arabica (B) E serpens (C)E scordiifolia and (D) E helioscopia

distribution between the leaf surfaces (see Figure 13)While E hirta has the most types E helioscopia Edracunculoides and E acalyphoides have the least

Regarding the position of the stomata on the surfacelevel of the epidermis a variable degree of sunkenesshas been recorded Using the ESEM the stomata appear

deeply sunken in some species whilst in others they areonly slightly sunken with the outer rims of the subsidiarycells surrounding the aperture The deeply and slightlysunken species are recorded in Table 2 AdditionallyFigure 10 shows the sunken stomata in different speciesunder the ESEM

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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186 Int Res J Plant Sci

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983120983137983154983137983139983161983156983145983139

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983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

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983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

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983124983141983156983154983137983139983161983156983145983139

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983107

983106

983105

983105983140983137983160983145983137983148983105983138983137983160983145983137983148

Figure 14 Stomata types in Euphorbia taxa in Saudi Arabia (A) E hirta (B) E hypericifolia (C) E pirottae and (D)Espaff schimperi

Venation

The venation patterns in the Euphorbia species underinvestigation vary from one-veined three-veined to thosewith four or more veins The predominant type is three-veined However in E balsamifera subsp Adenensis

there are seven middle parallel veins three middle thickand four thinner lateral ones (Figure 15 A and B)Whereas in E pirottae there are three main middleparallel veins and two thinner laterals (Figure 15 C) Incontrast E chamaepeplus E schimperiana and Earabica have only one thick main midrib coming from the

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 187

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Figure 15 Figure 15 Venation patterns at base of leaves invariants with more than three veins (A amp B) E balsamiferasubsp adenensis and (C) E pirottae

petiole then branched from the side into two lateral veinsat the point when the leaf starts widening (Figure 16)Each of these veins branches again into secondary veinsat the leaf margin

The rest of the species have just three veins Egrossheimii E dracunculoides E falcata E peplus Eserpens E scordiifolia E granulata E inaequilateraE helioscopia E hirta E hypericifolia and E

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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188 Int Res J Plant Sci

Figure 16 Venation patterns with one main mid vein at the leaf base in (A) E arabica and (B) E chamaepeplus

983106983105

Figure 17 Venation patterns with three-veined leaves in (A) E falcata and (B) E granulata

cyathophora All the three veins come from the petioleand start branching into secondary veins both toward themargin and the mid vein direction (Figure 17 B)

The vein endings at the leaf margin vary from closed

to open There is a relationship between the closedmargins and the sheathed veins (midrib and secondaryveins that are surrounded by parenchyma tissue) Whenthe veins are ornamented (sheathed) the margins areclosed This can be seen in subgenus Chamaesyce inspecies such as E serpens E scordiifolia E granulataE inaequilatera E hirta E hypericifolia and E arabica Additionally these species have the three-vein venation

On the other hand when the veins are not sheathedthe veins ends are open at the leaf margin as in Ebalsamifera subsp adenensis E schimperi E cuneataE acalyphoides E grossheimii E dracunculoides E

falcata E peplus E chamaepeplus E schimperiana Ehelioscopia E pirottae ldquoE aff schimperirdquo and Ecyathophora

CONCLUSIONS

Anatomical characters are seen here to be very useful inEuphorbia classification and identification The resultsobtained from anatomical studies were useful to groupsome species and identify others

Depending on the venation type the Euphorbiaspecies under investigation are divided anatomically intothree main groups see Figure 18

Group one veins are sheathed epidermal cellsdifferentiated on adaxial and abaxial surfaces Group two

983105 983106

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Figure 18 Hierarchial relation of Euphorbia taxa in Saudi Arabia based on anatomical charac

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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190 Int Res J Plant Sci

veins are not sheathed and epidermal cells are alike inboth surfaces And the third group includes species thatdo not belong to any of the previous groups The firstgroup can be further divided into three-veined and one-veined while the second group into three-veined and fouror more veins

In addition E grossheimii E retusa and E pirottae are related by being three-veined open at the marginswith rounded to oval tracheas and elongated cells bothon adaxial and abaxial surfaces However in E retusa the epidermal cells have straight cell walls on the adaxialsurface They are also similar in having hemiparacyticand paracytic types of stomata either on the adaxial orthe abaxial surface

On the other hand species with two differentepidermal cells on the leaf surfaces usually have three-sheathed veins with closed margins and elongatedtrachea such as E serpens E scordiifolia E granulata E inaequilatera E hirta and E hypericifolia which belongto subgenus Chamaesyce according to (Carter andSmith 1988)

In the case of E granulata which has beendescribed as having three different variants E granulata var granulata (both leaf surfaces hairy) E granulata varglabrata (adaxial glabrous and abaxial hairy) and aglabrous variant (the whole plant glabrous) Anatomicalwork has shown especially by using the ESEM that theleaves are either hairy or glabrous on both surfaces Sowhen the leaf is hairy both surfaces have differences inhair distribution Usually the lower surface is more hairythan upper surface (E granulata var glabrata ) butsometimes they are evenly distributed (Egranulata vargranulata ) In addition hairs on both surfaces are

multicellular In contrast when the leaf is glabrous bothleaf surfaces are glabrous and the whole plant isglabrous (suggested new variety) They were similar inhaving the same venation type (three-veined)

Finally ldquoE aff schimperi ldquohas some similarity with E schimperi in having the same open margin not theornamented venation type As well they both have thesame type of stomata paracytic and anisocytic but theydiffer in their distribution on abaxial and adaxial Thusthis may support the suggestion considering ldquoE affschimperi rdquo as a new subspecies of E schimperi

REFERENCES

Baranova M (1992) Principels of comparative stomatographic studiesof flowering plants The Botanical Review 58 49-99

Carlquist S (1961) Comparative Plant Anatomy Holt Rinehart ampWinston New York

Carter S Smith AR (1988) Euphorbiaceae (Part2) In Polhill ed Floraof Tropical East Africa Rotterdam Brookfield AA Balkema 409-597

Chaudhary SA (2001) Flora of The Kingdom of Saudi Arabia Vol 3Ministry of Agriculture and Water Riyadh

Dehay C (1935) Lagraveppareil Libero-ligneux foliare des EuphorbiaceaeAnnales des Sciences Naturelles Botanique 10 147-190

Dehgan B (1982) Comparative anatomy of the petiole and infragenericrelationships in Jatropha (Euphorbiaceae) American Journal oBotany 69 1283-1295

Gaucher L (1898) Ē tude anatomique du genre Euphorbia LPKlincksieck Paris

Gaucher L (1902) Recherches anatomiques sur les EuphorbiaceacuteesAnnales des Sciences Naturelles Botanique 15 161-309

Gaucher L (1902) Recherches anatomiques sur les Euphorbiaceacutees

Annales des Sciences Naturelles Botanique 15 161-309Govaerts R Fordin D Radcliffe-Smith A (2000) World Checklist and

Bibliography of Euphorbiaceae The Royal Botanic Gardens KewHickey LJ (1973) Classification of the architecture of dicotyledonous

leaves American Journal of Botany 60 17-33Hickey LJ (1979) A revised classification of the architecture o

dicotyledonous leaves In Metcalfe CR and Chalk L eds Anatomy oDicotyledons Systematic Anatomy of the leaf and stem 2nd edClarendon Press Oxford 25-39

Kakkar L Paliwal GS (1972) Epidermis in Euphorbia Indian ScienceCongress Association Proceedings 59 326-327

Kakkar L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 5 Epidermis Proceedings of the Indian National ScienceAcademy Part B Biological Sciences 40 55-67

Koumlhler E (1993) Blattvervatur-Muster der Buxaceae Dumortierr undSimmondsiaceae Van Tieghem Feddes Repertorium 104 145-167

Levin GA (1986a) Systematic foliar morphology of Phylanthoideae(Euphorbiaceae) II Phenetic analysis Annals of Missouri BotanicaGarden 73 86-98

Levin GA (1986b) Systematic foliar morphology of Phylanthoideae IICladistic analysis Systematic Botany 11 515-530

Levin GA (1986c) Systematic foliar morphology of Phyllanthoideae(Euphorbiaceae) I conspectus Annals of Missouri Botanical Garden73 29-85

Mahlberg P (1973) Histochemistry and Scanning Electron Microscopyof Starch Grains from Latex of Euphorbia-Terracina and EuphorbiaTirucalli Proceedings of the Indiana Academy of Science 82 132

Mahlberg PG (1974) Scanning Microscopic Comparison of StarchGrains from the Latex of Euphorbia -Spp Proceedings of the IndianaAcademy of Science 83 83-84

Mahlberg PG (1975) Evolution of the Laticifer in Euphorbia asInterpreted from Starch Grain Morphology American Journal oBotany 62 577-583

Mahlberg PG (1982) Comparative Morphology of Starch Grains inLatex from Varieties of Poinsettia Euphorbia-PulcherrimEuphorbiaceae Botanical Gazette 143 206-209

Mahlberg PG Davis DG Galitz DS Manners GD (1987) Laticifers andthe Classification of Euphorbia the Chemotaxonomy of EuphorbiaEsula L Botanical Journal of the Linnean Society 94 165-180

Metcalfe C Chalk L (1950) Anatomy of the Dicotyledons At theClarendon Press Oxford

Miller K Webster GL (1962) Systematic position of Cnidoscolus andJatropha Brittonia 14 174-180

Pax F (1884) Die Anatomie der Euphorbiaceen in ihrer Beziehung zumSystem derselben Botanische Jahrbuumlcher fuumlr SystematikPflanzengeschichte und Pflanzengeographie Leipzig 5 384-421

Payne WW (1970) Helicocytic and allelocytic stomata unrecognizedpatters in the Dicotyledonae Am J Bot57 140-147

Pray TR (1955a) Foliar venation of angiosperm II Histogenesis of thevenation of Liriodendron Am J Bot 42 18-27

Pray TR 1955b Foliar venation of angiosperms III Pattern andhistology of the venation of Hosta Am J Bot 42 11-18

Radford AE Dickison WC Massey JR Bell CR (1974) Vascular PlanSystematics Harper amp Row Publishers London New York

Radlkofer L (1870) Ueber Pausandra ein neues EuphorbiaceenGenus Flora 53 81-95

Raju VS Rao PN (1977) Variation in the structure and development ofoliar stomata in the Euphorbiaceae Botanical Journal of the LinneanSociety 75 69-97

Raju VS Rao PN (1987) The Taxonomic Use of the Basic StomataType in the Generic Delimitation of Chamaesyce EuphorbiaceaeFeddes Repertorium 98 137-142

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2424

Rosowski JR (1968) Laticifer Morphology in the Mature Stem and Leafof Euphorbia supina Botanical Gazette 129 113-120

Roth-Nebelsick A Uhl D Mosbrugger V Kerp H (2001) Evolution andfunction of leaf venation architecture a review Annals of Botany 87553-566

Sehgal L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 2 Venation Patterns Botanical Journal of the Linnean Society 68 173-208

Singh G (2004) Plant Systematics An Integrated Approach SciencePublishers Enfield

Solereder H (1899) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Solereder H (1908) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Aldhebiani and Jury 191

Stace CA (1965) Cuticular studies as an aid to plant taxonomy Bulletinof the British Museum Natural History Botany 4 3-78

Stace CA (1989) Plant Taxonomy and Biosystematics Hodder ampStoughton London

Tognini F (1897) Contribuzione allo studio della organogeniacomparata degli stomi Institut de Botanique University de Geneve 41-42

Van Cotthem WR (1970) A classification of stomatal types Botanica

Journal of Linnean Society 63 235-246Wagner W (1979) Reticulate veins in the systematics of modern ferns

Taxon 1 87-95Webster GL (1994a) Classification of the Euphorbiaceae Annals of the

Missouri Botanical Garden 81 3-32

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 173

983106

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983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

Figure 1 Normal irregular epidermal cells in both adaxial and abaxial surfaces of the leaf (LM)(A) E balsamifera subsp adenensis (B) E schimperi (C) E cuneata (D) E falcata (E) Abaxial in E dracunculoides

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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174 Int Res J Plant Sci

983105

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983108

983106 983106

983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

Figure 2 Error No text of specified style in document Both adaxial and abaxial surfaceshave the same shape of epidermal cells (LM) Elongated in A ampB plicate in CampD (A) Egrossheimii (B) E pirottae (C) E chamaepeplus and (D) E peplus

schimperiana E helioscopia E inaequilatera Earabica E granulata E serpens and E scordiifolia see Figure 3 and Figure 4 This last group of species belongto subgenus Chamaesyce except for E helioscopia and E schimperiana

Hairs

Hairs on the leaf epidermis can be both unicellular and

multicellular and may appear only on one surface or bothFurthermore hairs in the Euphorbia species undeinvestigation are generally simple unbranched and with awarty ornamentation on their surface Papillae appear inESEM as outgrowths whereas in the surface view theygive the impression of being rounded structures at thecentre of the cell Prominent papillae occur only in onespecies of Euphorbia in Saudi Arabia this species is Ehypericifolia (Figure 5 and 6) whereas in E arabica theywere poorly distinguishable using LM

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 175

983105983140983137983160983145983137983148

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983105983138983137983160983145983137983148

Figure 3 Epidermal cells vary on leaf surfaces adaxial surface has irregular cells whilethe abaxial surface has plicate (LM)(A) E schimperiana (B) E arabica (C) E serpens (D) E scordiifolia

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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176 Int Res J Plant Sci

983105983140983137983160983145983137983148

983107

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Figure 4 Epidermal cell shape varies on the adaxial andthe abaxial surfaces straight cell walls in the formerwhile folded plicate walls occur on the latter (LM) (A) Egranulata (B) E inaequilatera and (C) E helioscopia

983112983141983149983145983152983137983154983137983139983161983156983145983139 983155983156983151983149983137983156983137

983151983150 983138983151983156983144 983155983157983154983142983137983139983141983155

Figure 5 E hypericifolia (LM)Above photo the adaxial surface with irregular epidermal cells andvery clear papillae as a circle in the middle of the cellLower photo the abaxial surface with plicate cells and papillae ahair base is obvious in the right side of the picture

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 177

983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

983120983137983152983145983148983148983137983141

983117983157983148983156983145983139983141983148983148983157983148983137983154 983144983137983145983154983155 983145983150 983137983138983137983160983145983137983148 983155983157983154983142983137983139983141983086

Figure 6 E hypericifolia (ESEM) papillae on the adaxial and abaxial surfacesmulticellular hairs are on the abaxial surface of the leaf only

Figure7 Multicellular hairs on adaxial and abaxial surfaces of E hirta

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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178 Int Res J Plant Sci

Figure 8 Hairs wart ornamentation on hair surfaces of (A) E scordiifolia and (B) E pirottae (C) Short hairs onabaxial surface of E balsamifera subsp adenensis

In E balsamifera subsp adenensis the hairs are shortand unicellular (Figure 8C) and they are located mostly atthe leaf margins and the mid-vein area on the abaxialsurface only

Hairs on the leaf of E pirottae are present on theabaxial surface only and are both unicellular andmulticellular Euphorbia granulata is similar to E pirottae in having both unicellular and multicellular hairs but herethey occur on both surfaces of the leaf (Figure 8B)

On the other hand in E scordiifolia E hirta and Ehypericifolia the hairs are multicellular and on bothsurfaces in the first two species and on the abaxial only inthe latter (Figure 7) Moreover E hypericifolia and Ehirta have long hairs when compared with the otherspecies (Figure 6)

No hairs were observed and leaves were glabrous inthe following species E serpens E arabica E peplusE inaequilatera E falcata E helioscopia E cuneata Eretusa E grossheimii E schimperiana E schimperiE chamaepeplus and the new variety of E granulata

Wax

The cuticle layer and wax in the Euphorbia species undeinvestigation vary from smooth to densely cover in waxscales This may be a thick smooth layer of wax whichsometimes has upright scales In some species only thecell surface has a smooth wax layer whereas the walbetween cells has upright scales of wax as in Eschimperiana especially around the stomata pores

(Figure 9) In contrast the anticlinal walls of somespecies have a smooth layer of wax whereas the celsurface is densely covered with upright wax flakes or withsome stellate wax flakes in other species

Stomata

Stomata were observed through LM and ESEM In thespecies under investigation stomata commonly appeaon both leaf surfaces (amphistomatic leaves) excepin E cuneata where the stomata are located only on the

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 179

983109

983105

983108983107

983106

Figure 9 Wax formation in Euphorbia species in Saudi Arabia (A+B) smooth andthick in E peplus and E schimperi (C+D) smooth on top of cell while formingupright scale flakes in between especially around stomata as in E schimperiana smooth in anticlinal walls (E) upright scale flakes in E balsamifera

abaxial surface (hypostomatic leaves) The presence andposition of subsidiary cells are variable in Euphorbia

species Subsidiary cells were found to be of unequalsize and shape whether the stoma was anisocyticparacytic actinocytic or tetracytic While some specieslack them around the stomata others have more thanone arrangement of subsidiary cells

The most common stomata type is anomocytic whilethe rare type is actinocytic recorded only in Ehelioscopia Moreover sometimes stomata of more thanone type (up to three or four types) have beenencountered on the same leaf surface as in E scordiifolia (three types on the abaxial surface) and E hirta (five

types on the abaxial and three on the adaxial) Thedistribution and type of stomata are shown in Table 2

In addition in some species such as E falcata and Ehelioscopia the stomata are deeply sunken andrepresented only by cuticular ledges surrounding theminute pores as seen in ESEM

Examples of species lacking subsidiary cells aroundstomata are E peplus E chamaepeplus E granulataE dracunculoides and E acalyphoides Therefore theyall have anomocytic stomata

On the other hand E balsamifera subsp adenensishas the mononcytic type on both leaf surfaces andanisocytic and paracytic only on the abaxial surface In

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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180 Int Res J Plant Sci

Table 2 Stomata type in Euphorbia taxa in Saudi Arabia

Ab abaxial surface + stomata presentAd adaxial surface

addition E schimperiana has anomocytic andtetracytic on the abaxial only In contrast Ecuneata has three types of stomata anomocyticparacytic and anisocytic only on the lower surfaceAs well ldquoE aff consobrinardquo ldquoE aff schimperirdquo

and E schimperi have paracytic and anisocyticonly in the abaxial on the first two and in both leafsurfaces on the latter Therefore ldquoE affconsobrinardquo and ldquoE aff schimperirdquo are the same taxa Then comparing to E schimperi results

from ansubspec

Eupsimilar and an

Subgenus

Stomata

Anomocytic Hemiparacytic Paracytic Anisocytic T

Species Ad Ab Ad Ab Ad Ab Ad Ab AT i r u c al l i

E balsamifera subsp adenensis + + +

E schimperi + + + +

ldquoEsp aff consobrinardquo + +

ldquoE sp aff schimperirdquo + +

E cuneata + + +

ErmophytonE acalyphoides + +

E pirottae + + + +

E helioscopia + + E s ul a

E grossheimii + + + + +

E retusa + + + + +

E dracunculoides +

E falcate + + E peplus +

E chamaepeplus + +

E schimperiana + C h am a e s y c e

E Arabica + + + + + + +

E serpens + + + +

E scordiifolia + + + + + + +

E granulate + + + +

E inaequilatera + + + +

E hirta + + + + + + +

E hypericifolia + + + + + +

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Table3 Venation type in Euphorbia taxa in Saudi Arabia

Subgenus Species

Venation

Number of veins atthe leaf base

Margins Sheathed

1 2 3 4 or more Open Closed Rounded T i r u c al l i

Ebalsamiferasubspadenensis

+ + +

E schimperi + + +

E aff Schimperi + + +

E cuneata + + +

ErmophytonE acalyphoides + + +

E pirottae + + E s ul a

E grossheimii + +

E retusa + + +

E dracunculoides + + +

E falcate + + +

E peplus + +

E chamaepeplus + +

E schimperiana

E helioscopia

+

+

+

+ C h am a e s y c e

E Arabica + + +

E serpens + + +

E scordiifolia + + +

E granulate + + + +

E inaequilatera + + +

E hirta + + + E hypericifolia + + +

Poinsettia E cyathophora + +

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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182 Int Res J Plant Sci

Figure 10 Sunken stomata on the abaxial surface1) Deeply sunken (A) E helioscopia (B) E falcata (C) E schimperiana 2) Slightly sunken (D) E peplus (E) E pirottae

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 183

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983105983140983137983160983145983137983148

Figure 11 Stomata types in Euphorbia taxa in Saudi Arabia (A) E balsamifera subsp adenensis (B) E schimperi (C) E cuneata and (D) E grossheimii

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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184 Int Res J Plant Sci

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Figure 12 Stomata types in Euphorbia taxa (A) E falcata (B) E peplus (C) E chamaepeplus and (D) Eschimperiana

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 185

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Figure 13 Figure 13 Stomata types in Euphorbia taxa in Saudi Arabia (A) E arabica (B) E serpens (C)E scordiifolia and (D) E helioscopia

distribution between the leaf surfaces (see Figure 13)While E hirta has the most types E helioscopia Edracunculoides and E acalyphoides have the least

Regarding the position of the stomata on the surfacelevel of the epidermis a variable degree of sunkenesshas been recorded Using the ESEM the stomata appear

deeply sunken in some species whilst in others they areonly slightly sunken with the outer rims of the subsidiarycells surrounding the aperture The deeply and slightlysunken species are recorded in Table 2 AdditionallyFigure 10 shows the sunken stomata in different speciesunder the ESEM

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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186 Int Res J Plant Sci

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Figure 14 Stomata types in Euphorbia taxa in Saudi Arabia (A) E hirta (B) E hypericifolia (C) E pirottae and (D)Espaff schimperi

Venation

The venation patterns in the Euphorbia species underinvestigation vary from one-veined three-veined to thosewith four or more veins The predominant type is three-veined However in E balsamifera subsp Adenensis

there are seven middle parallel veins three middle thickand four thinner lateral ones (Figure 15 A and B)Whereas in E pirottae there are three main middleparallel veins and two thinner laterals (Figure 15 C) Incontrast E chamaepeplus E schimperiana and Earabica have only one thick main midrib coming from the

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 187

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Figure 15 Figure 15 Venation patterns at base of leaves invariants with more than three veins (A amp B) E balsamiferasubsp adenensis and (C) E pirottae

petiole then branched from the side into two lateral veinsat the point when the leaf starts widening (Figure 16)Each of these veins branches again into secondary veinsat the leaf margin

The rest of the species have just three veins Egrossheimii E dracunculoides E falcata E peplus Eserpens E scordiifolia E granulata E inaequilateraE helioscopia E hirta E hypericifolia and E

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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188 Int Res J Plant Sci

Figure 16 Venation patterns with one main mid vein at the leaf base in (A) E arabica and (B) E chamaepeplus

983106983105

Figure 17 Venation patterns with three-veined leaves in (A) E falcata and (B) E granulata

cyathophora All the three veins come from the petioleand start branching into secondary veins both toward themargin and the mid vein direction (Figure 17 B)

The vein endings at the leaf margin vary from closed

to open There is a relationship between the closedmargins and the sheathed veins (midrib and secondaryveins that are surrounded by parenchyma tissue) Whenthe veins are ornamented (sheathed) the margins areclosed This can be seen in subgenus Chamaesyce inspecies such as E serpens E scordiifolia E granulataE inaequilatera E hirta E hypericifolia and E arabica Additionally these species have the three-vein venation

On the other hand when the veins are not sheathedthe veins ends are open at the leaf margin as in Ebalsamifera subsp adenensis E schimperi E cuneataE acalyphoides E grossheimii E dracunculoides E

falcata E peplus E chamaepeplus E schimperiana Ehelioscopia E pirottae ldquoE aff schimperirdquo and Ecyathophora

CONCLUSIONS

Anatomical characters are seen here to be very useful inEuphorbia classification and identification The resultsobtained from anatomical studies were useful to groupsome species and identify others

Depending on the venation type the Euphorbiaspecies under investigation are divided anatomically intothree main groups see Figure 18

Group one veins are sheathed epidermal cellsdifferentiated on adaxial and abaxial surfaces Group two

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7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Figure 18 Hierarchial relation of Euphorbia taxa in Saudi Arabia based on anatomical charac

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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190 Int Res J Plant Sci

veins are not sheathed and epidermal cells are alike inboth surfaces And the third group includes species thatdo not belong to any of the previous groups The firstgroup can be further divided into three-veined and one-veined while the second group into three-veined and fouror more veins

In addition E grossheimii E retusa and E pirottae are related by being three-veined open at the marginswith rounded to oval tracheas and elongated cells bothon adaxial and abaxial surfaces However in E retusa the epidermal cells have straight cell walls on the adaxialsurface They are also similar in having hemiparacyticand paracytic types of stomata either on the adaxial orthe abaxial surface

On the other hand species with two differentepidermal cells on the leaf surfaces usually have three-sheathed veins with closed margins and elongatedtrachea such as E serpens E scordiifolia E granulata E inaequilatera E hirta and E hypericifolia which belongto subgenus Chamaesyce according to (Carter andSmith 1988)

In the case of E granulata which has beendescribed as having three different variants E granulata var granulata (both leaf surfaces hairy) E granulata varglabrata (adaxial glabrous and abaxial hairy) and aglabrous variant (the whole plant glabrous) Anatomicalwork has shown especially by using the ESEM that theleaves are either hairy or glabrous on both surfaces Sowhen the leaf is hairy both surfaces have differences inhair distribution Usually the lower surface is more hairythan upper surface (E granulata var glabrata ) butsometimes they are evenly distributed (Egranulata vargranulata ) In addition hairs on both surfaces are

multicellular In contrast when the leaf is glabrous bothleaf surfaces are glabrous and the whole plant isglabrous (suggested new variety) They were similar inhaving the same venation type (three-veined)

Finally ldquoE aff schimperi ldquohas some similarity with E schimperi in having the same open margin not theornamented venation type As well they both have thesame type of stomata paracytic and anisocytic but theydiffer in their distribution on abaxial and adaxial Thusthis may support the suggestion considering ldquoE affschimperi rdquo as a new subspecies of E schimperi

REFERENCES

Baranova M (1992) Principels of comparative stomatographic studiesof flowering plants The Botanical Review 58 49-99

Carlquist S (1961) Comparative Plant Anatomy Holt Rinehart ampWinston New York

Carter S Smith AR (1988) Euphorbiaceae (Part2) In Polhill ed Floraof Tropical East Africa Rotterdam Brookfield AA Balkema 409-597

Chaudhary SA (2001) Flora of The Kingdom of Saudi Arabia Vol 3Ministry of Agriculture and Water Riyadh

Dehay C (1935) Lagraveppareil Libero-ligneux foliare des EuphorbiaceaeAnnales des Sciences Naturelles Botanique 10 147-190

Dehgan B (1982) Comparative anatomy of the petiole and infragenericrelationships in Jatropha (Euphorbiaceae) American Journal oBotany 69 1283-1295

Gaucher L (1898) Ē tude anatomique du genre Euphorbia LPKlincksieck Paris

Gaucher L (1902) Recherches anatomiques sur les EuphorbiaceacuteesAnnales des Sciences Naturelles Botanique 15 161-309

Gaucher L (1902) Recherches anatomiques sur les Euphorbiaceacutees

Annales des Sciences Naturelles Botanique 15 161-309Govaerts R Fordin D Radcliffe-Smith A (2000) World Checklist and

Bibliography of Euphorbiaceae The Royal Botanic Gardens KewHickey LJ (1973) Classification of the architecture of dicotyledonous

leaves American Journal of Botany 60 17-33Hickey LJ (1979) A revised classification of the architecture o

dicotyledonous leaves In Metcalfe CR and Chalk L eds Anatomy oDicotyledons Systematic Anatomy of the leaf and stem 2nd edClarendon Press Oxford 25-39

Kakkar L Paliwal GS (1972) Epidermis in Euphorbia Indian ScienceCongress Association Proceedings 59 326-327

Kakkar L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 5 Epidermis Proceedings of the Indian National ScienceAcademy Part B Biological Sciences 40 55-67

Koumlhler E (1993) Blattvervatur-Muster der Buxaceae Dumortierr undSimmondsiaceae Van Tieghem Feddes Repertorium 104 145-167

Levin GA (1986a) Systematic foliar morphology of Phylanthoideae(Euphorbiaceae) II Phenetic analysis Annals of Missouri BotanicaGarden 73 86-98

Levin GA (1986b) Systematic foliar morphology of Phylanthoideae IICladistic analysis Systematic Botany 11 515-530

Levin GA (1986c) Systematic foliar morphology of Phyllanthoideae(Euphorbiaceae) I conspectus Annals of Missouri Botanical Garden73 29-85

Mahlberg P (1973) Histochemistry and Scanning Electron Microscopyof Starch Grains from Latex of Euphorbia-Terracina and EuphorbiaTirucalli Proceedings of the Indiana Academy of Science 82 132

Mahlberg PG (1974) Scanning Microscopic Comparison of StarchGrains from the Latex of Euphorbia -Spp Proceedings of the IndianaAcademy of Science 83 83-84

Mahlberg PG (1975) Evolution of the Laticifer in Euphorbia asInterpreted from Starch Grain Morphology American Journal oBotany 62 577-583

Mahlberg PG (1982) Comparative Morphology of Starch Grains inLatex from Varieties of Poinsettia Euphorbia-PulcherrimEuphorbiaceae Botanical Gazette 143 206-209

Mahlberg PG Davis DG Galitz DS Manners GD (1987) Laticifers andthe Classification of Euphorbia the Chemotaxonomy of EuphorbiaEsula L Botanical Journal of the Linnean Society 94 165-180

Metcalfe C Chalk L (1950) Anatomy of the Dicotyledons At theClarendon Press Oxford

Miller K Webster GL (1962) Systematic position of Cnidoscolus andJatropha Brittonia 14 174-180

Pax F (1884) Die Anatomie der Euphorbiaceen in ihrer Beziehung zumSystem derselben Botanische Jahrbuumlcher fuumlr SystematikPflanzengeschichte und Pflanzengeographie Leipzig 5 384-421

Payne WW (1970) Helicocytic and allelocytic stomata unrecognizedpatters in the Dicotyledonae Am J Bot57 140-147

Pray TR (1955a) Foliar venation of angiosperm II Histogenesis of thevenation of Liriodendron Am J Bot 42 18-27

Pray TR 1955b Foliar venation of angiosperms III Pattern andhistology of the venation of Hosta Am J Bot 42 11-18

Radford AE Dickison WC Massey JR Bell CR (1974) Vascular PlanSystematics Harper amp Row Publishers London New York

Radlkofer L (1870) Ueber Pausandra ein neues EuphorbiaceenGenus Flora 53 81-95

Raju VS Rao PN (1977) Variation in the structure and development ofoliar stomata in the Euphorbiaceae Botanical Journal of the LinneanSociety 75 69-97

Raju VS Rao PN (1987) The Taxonomic Use of the Basic StomataType in the Generic Delimitation of Chamaesyce EuphorbiaceaeFeddes Repertorium 98 137-142

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2424

Rosowski JR (1968) Laticifer Morphology in the Mature Stem and Leafof Euphorbia supina Botanical Gazette 129 113-120

Roth-Nebelsick A Uhl D Mosbrugger V Kerp H (2001) Evolution andfunction of leaf venation architecture a review Annals of Botany 87553-566

Sehgal L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 2 Venation Patterns Botanical Journal of the Linnean Society 68 173-208

Singh G (2004) Plant Systematics An Integrated Approach SciencePublishers Enfield

Solereder H (1899) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Solereder H (1908) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Aldhebiani and Jury 191

Stace CA (1965) Cuticular studies as an aid to plant taxonomy Bulletinof the British Museum Natural History Botany 4 3-78

Stace CA (1989) Plant Taxonomy and Biosystematics Hodder ampStoughton London

Tognini F (1897) Contribuzione allo studio della organogeniacomparata degli stomi Institut de Botanique University de Geneve 41-42

Van Cotthem WR (1970) A classification of stomatal types Botanica

Journal of Linnean Society 63 235-246Wagner W (1979) Reticulate veins in the systematics of modern ferns

Taxon 1 87-95Webster GL (1994a) Classification of the Euphorbiaceae Annals of the

Missouri Botanical Garden 81 3-32

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Figure 2 Error No text of specified style in document Both adaxial and abaxial surfaceshave the same shape of epidermal cells (LM) Elongated in A ampB plicate in CampD (A) Egrossheimii (B) E pirottae (C) E chamaepeplus and (D) E peplus

schimperiana E helioscopia E inaequilatera Earabica E granulata E serpens and E scordiifolia see Figure 3 and Figure 4 This last group of species belongto subgenus Chamaesyce except for E helioscopia and E schimperiana

Hairs

Hairs on the leaf epidermis can be both unicellular and

multicellular and may appear only on one surface or bothFurthermore hairs in the Euphorbia species undeinvestigation are generally simple unbranched and with awarty ornamentation on their surface Papillae appear inESEM as outgrowths whereas in the surface view theygive the impression of being rounded structures at thecentre of the cell Prominent papillae occur only in onespecies of Euphorbia in Saudi Arabia this species is Ehypericifolia (Figure 5 and 6) whereas in E arabica theywere poorly distinguishable using LM

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Figure 3 Epidermal cells vary on leaf surfaces adaxial surface has irregular cells whilethe abaxial surface has plicate (LM)(A) E schimperiana (B) E arabica (C) E serpens (D) E scordiifolia

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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176 Int Res J Plant Sci

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Figure 4 Epidermal cell shape varies on the adaxial andthe abaxial surfaces straight cell walls in the formerwhile folded plicate walls occur on the latter (LM) (A) Egranulata (B) E inaequilatera and (C) E helioscopia

983112983141983149983145983152983137983154983137983139983161983156983145983139 983155983156983151983149983137983156983137

983151983150 983138983151983156983144 983155983157983154983142983137983139983141983155

Figure 5 E hypericifolia (LM)Above photo the adaxial surface with irregular epidermal cells andvery clear papillae as a circle in the middle of the cellLower photo the abaxial surface with plicate cells and papillae ahair base is obvious in the right side of the picture

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 177

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Figure 6 E hypericifolia (ESEM) papillae on the adaxial and abaxial surfacesmulticellular hairs are on the abaxial surface of the leaf only

Figure7 Multicellular hairs on adaxial and abaxial surfaces of E hirta

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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178 Int Res J Plant Sci

Figure 8 Hairs wart ornamentation on hair surfaces of (A) E scordiifolia and (B) E pirottae (C) Short hairs onabaxial surface of E balsamifera subsp adenensis

In E balsamifera subsp adenensis the hairs are shortand unicellular (Figure 8C) and they are located mostly atthe leaf margins and the mid-vein area on the abaxialsurface only

Hairs on the leaf of E pirottae are present on theabaxial surface only and are both unicellular andmulticellular Euphorbia granulata is similar to E pirottae in having both unicellular and multicellular hairs but herethey occur on both surfaces of the leaf (Figure 8B)

On the other hand in E scordiifolia E hirta and Ehypericifolia the hairs are multicellular and on bothsurfaces in the first two species and on the abaxial only inthe latter (Figure 7) Moreover E hypericifolia and Ehirta have long hairs when compared with the otherspecies (Figure 6)

No hairs were observed and leaves were glabrous inthe following species E serpens E arabica E peplusE inaequilatera E falcata E helioscopia E cuneata Eretusa E grossheimii E schimperiana E schimperiE chamaepeplus and the new variety of E granulata

Wax

The cuticle layer and wax in the Euphorbia species undeinvestigation vary from smooth to densely cover in waxscales This may be a thick smooth layer of wax whichsometimes has upright scales In some species only thecell surface has a smooth wax layer whereas the walbetween cells has upright scales of wax as in Eschimperiana especially around the stomata pores

(Figure 9) In contrast the anticlinal walls of somespecies have a smooth layer of wax whereas the celsurface is densely covered with upright wax flakes or withsome stellate wax flakes in other species

Stomata

Stomata were observed through LM and ESEM In thespecies under investigation stomata commonly appeaon both leaf surfaces (amphistomatic leaves) excepin E cuneata where the stomata are located only on the

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 179

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Figure 9 Wax formation in Euphorbia species in Saudi Arabia (A+B) smooth andthick in E peplus and E schimperi (C+D) smooth on top of cell while formingupright scale flakes in between especially around stomata as in E schimperiana smooth in anticlinal walls (E) upright scale flakes in E balsamifera

abaxial surface (hypostomatic leaves) The presence andposition of subsidiary cells are variable in Euphorbia

species Subsidiary cells were found to be of unequalsize and shape whether the stoma was anisocyticparacytic actinocytic or tetracytic While some specieslack them around the stomata others have more thanone arrangement of subsidiary cells

The most common stomata type is anomocytic whilethe rare type is actinocytic recorded only in Ehelioscopia Moreover sometimes stomata of more thanone type (up to three or four types) have beenencountered on the same leaf surface as in E scordiifolia (three types on the abaxial surface) and E hirta (five

types on the abaxial and three on the adaxial) Thedistribution and type of stomata are shown in Table 2

In addition in some species such as E falcata and Ehelioscopia the stomata are deeply sunken andrepresented only by cuticular ledges surrounding theminute pores as seen in ESEM

Examples of species lacking subsidiary cells aroundstomata are E peplus E chamaepeplus E granulataE dracunculoides and E acalyphoides Therefore theyall have anomocytic stomata

On the other hand E balsamifera subsp adenensishas the mononcytic type on both leaf surfaces andanisocytic and paracytic only on the abaxial surface In

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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180 Int Res J Plant Sci

Table 2 Stomata type in Euphorbia taxa in Saudi Arabia

Ab abaxial surface + stomata presentAd adaxial surface

addition E schimperiana has anomocytic andtetracytic on the abaxial only In contrast Ecuneata has three types of stomata anomocyticparacytic and anisocytic only on the lower surfaceAs well ldquoE aff consobrinardquo ldquoE aff schimperirdquo

and E schimperi have paracytic and anisocyticonly in the abaxial on the first two and in both leafsurfaces on the latter Therefore ldquoE affconsobrinardquo and ldquoE aff schimperirdquo are the same taxa Then comparing to E schimperi results

from ansubspec

Eupsimilar and an

Subgenus

Stomata

Anomocytic Hemiparacytic Paracytic Anisocytic T

Species Ad Ab Ad Ab Ad Ab Ad Ab AT i r u c al l i

E balsamifera subsp adenensis + + +

E schimperi + + + +

ldquoEsp aff consobrinardquo + +

ldquoE sp aff schimperirdquo + +

E cuneata + + +

ErmophytonE acalyphoides + +

E pirottae + + + +

E helioscopia + + E s ul a

E grossheimii + + + + +

E retusa + + + + +

E dracunculoides +

E falcate + + E peplus +

E chamaepeplus + +

E schimperiana + C h am a e s y c e

E Arabica + + + + + + +

E serpens + + + +

E scordiifolia + + + + + + +

E granulate + + + +

E inaequilatera + + + +

E hirta + + + + + + +

E hypericifolia + + + + + +

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Table3 Venation type in Euphorbia taxa in Saudi Arabia

Subgenus Species

Venation

Number of veins atthe leaf base

Margins Sheathed

1 2 3 4 or more Open Closed Rounded T i r u c al l i

Ebalsamiferasubspadenensis

+ + +

E schimperi + + +

E aff Schimperi + + +

E cuneata + + +

ErmophytonE acalyphoides + + +

E pirottae + + E s ul a

E grossheimii + +

E retusa + + +

E dracunculoides + + +

E falcate + + +

E peplus + +

E chamaepeplus + +

E schimperiana

E helioscopia

+

+

+

+ C h am a e s y c e

E Arabica + + +

E serpens + + +

E scordiifolia + + +

E granulate + + + +

E inaequilatera + + +

E hirta + + + E hypericifolia + + +

Poinsettia E cyathophora + +

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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182 Int Res J Plant Sci

Figure 10 Sunken stomata on the abaxial surface1) Deeply sunken (A) E helioscopia (B) E falcata (C) E schimperiana 2) Slightly sunken (D) E peplus (E) E pirottae

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 183

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983105983139983156983145983150983151983139983161983156983145983139

983105983140983137983160983145983137983148

Figure 11 Stomata types in Euphorbia taxa in Saudi Arabia (A) E balsamifera subsp adenensis (B) E schimperi (C) E cuneata and (D) E grossheimii

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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184 Int Res J Plant Sci

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983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139983105983150983145983155983151983139983161983156983145983139

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983105983139983156983145983150983151983139983161983156983145983139

983105983138983137983160983145983137983148983105983140983137983160983145983137983148

Figure 12 Stomata types in Euphorbia taxa (A) E falcata (B) E peplus (C) E chamaepeplus and (D) Eschimperiana

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 185

983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

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983105983139983156983145983150983151983139983161983156983145983139

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983112983141983149983145983152983137983154983137983139983161983156983145983139

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983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

Figure 13 Figure 13 Stomata types in Euphorbia taxa in Saudi Arabia (A) E arabica (B) E serpens (C)E scordiifolia and (D) E helioscopia

distribution between the leaf surfaces (see Figure 13)While E hirta has the most types E helioscopia Edracunculoides and E acalyphoides have the least

Regarding the position of the stomata on the surfacelevel of the epidermis a variable degree of sunkenesshas been recorded Using the ESEM the stomata appear

deeply sunken in some species whilst in others they areonly slightly sunken with the outer rims of the subsidiarycells surrounding the aperture The deeply and slightlysunken species are recorded in Table 2 AdditionallyFigure 10 shows the sunken stomata in different speciesunder the ESEM

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983105983150983151983149983151983139983161983156983145983139

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983112983141983149983145983152983137983154983137983139983161983156983145983139

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983124983141983156983154983137983139983161983156983145983139

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983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983107

983106

983105

983105983140983137983160983145983137983148983105983138983137983160983145983137983148

Figure 14 Stomata types in Euphorbia taxa in Saudi Arabia (A) E hirta (B) E hypericifolia (C) E pirottae and (D)Espaff schimperi

Venation

The venation patterns in the Euphorbia species underinvestigation vary from one-veined three-veined to thosewith four or more veins The predominant type is three-veined However in E balsamifera subsp Adenensis

there are seven middle parallel veins three middle thickand four thinner lateral ones (Figure 15 A and B)Whereas in E pirottae there are three main middleparallel veins and two thinner laterals (Figure 15 C) Incontrast E chamaepeplus E schimperiana and Earabica have only one thick main midrib coming from the

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 187

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983106

983107

Figure 15 Figure 15 Venation patterns at base of leaves invariants with more than three veins (A amp B) E balsamiferasubsp adenensis and (C) E pirottae

petiole then branched from the side into two lateral veinsat the point when the leaf starts widening (Figure 16)Each of these veins branches again into secondary veinsat the leaf margin

The rest of the species have just three veins Egrossheimii E dracunculoides E falcata E peplus Eserpens E scordiifolia E granulata E inaequilateraE helioscopia E hirta E hypericifolia and E

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188 Int Res J Plant Sci

Figure 16 Venation patterns with one main mid vein at the leaf base in (A) E arabica and (B) E chamaepeplus

983106983105

Figure 17 Venation patterns with three-veined leaves in (A) E falcata and (B) E granulata

cyathophora All the three veins come from the petioleand start branching into secondary veins both toward themargin and the mid vein direction (Figure 17 B)

The vein endings at the leaf margin vary from closed

to open There is a relationship between the closedmargins and the sheathed veins (midrib and secondaryveins that are surrounded by parenchyma tissue) Whenthe veins are ornamented (sheathed) the margins areclosed This can be seen in subgenus Chamaesyce inspecies such as E serpens E scordiifolia E granulataE inaequilatera E hirta E hypericifolia and E arabica Additionally these species have the three-vein venation

On the other hand when the veins are not sheathedthe veins ends are open at the leaf margin as in Ebalsamifera subsp adenensis E schimperi E cuneataE acalyphoides E grossheimii E dracunculoides E

falcata E peplus E chamaepeplus E schimperiana Ehelioscopia E pirottae ldquoE aff schimperirdquo and Ecyathophora

CONCLUSIONS

Anatomical characters are seen here to be very useful inEuphorbia classification and identification The resultsobtained from anatomical studies were useful to groupsome species and identify others

Depending on the venation type the Euphorbiaspecies under investigation are divided anatomically intothree main groups see Figure 18

Group one veins are sheathed epidermal cellsdifferentiated on adaxial and abaxial surfaces Group two

983105 983106

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Figure 18 Hierarchial relation of Euphorbia taxa in Saudi Arabia based on anatomical charac

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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190 Int Res J Plant Sci

veins are not sheathed and epidermal cells are alike inboth surfaces And the third group includes species thatdo not belong to any of the previous groups The firstgroup can be further divided into three-veined and one-veined while the second group into three-veined and fouror more veins

In addition E grossheimii E retusa and E pirottae are related by being three-veined open at the marginswith rounded to oval tracheas and elongated cells bothon adaxial and abaxial surfaces However in E retusa the epidermal cells have straight cell walls on the adaxialsurface They are also similar in having hemiparacyticand paracytic types of stomata either on the adaxial orthe abaxial surface

On the other hand species with two differentepidermal cells on the leaf surfaces usually have three-sheathed veins with closed margins and elongatedtrachea such as E serpens E scordiifolia E granulata E inaequilatera E hirta and E hypericifolia which belongto subgenus Chamaesyce according to (Carter andSmith 1988)

In the case of E granulata which has beendescribed as having three different variants E granulata var granulata (both leaf surfaces hairy) E granulata varglabrata (adaxial glabrous and abaxial hairy) and aglabrous variant (the whole plant glabrous) Anatomicalwork has shown especially by using the ESEM that theleaves are either hairy or glabrous on both surfaces Sowhen the leaf is hairy both surfaces have differences inhair distribution Usually the lower surface is more hairythan upper surface (E granulata var glabrata ) butsometimes they are evenly distributed (Egranulata vargranulata ) In addition hairs on both surfaces are

multicellular In contrast when the leaf is glabrous bothleaf surfaces are glabrous and the whole plant isglabrous (suggested new variety) They were similar inhaving the same venation type (three-veined)

Finally ldquoE aff schimperi ldquohas some similarity with E schimperi in having the same open margin not theornamented venation type As well they both have thesame type of stomata paracytic and anisocytic but theydiffer in their distribution on abaxial and adaxial Thusthis may support the suggestion considering ldquoE affschimperi rdquo as a new subspecies of E schimperi

REFERENCES

Baranova M (1992) Principels of comparative stomatographic studiesof flowering plants The Botanical Review 58 49-99

Carlquist S (1961) Comparative Plant Anatomy Holt Rinehart ampWinston New York

Carter S Smith AR (1988) Euphorbiaceae (Part2) In Polhill ed Floraof Tropical East Africa Rotterdam Brookfield AA Balkema 409-597

Chaudhary SA (2001) Flora of The Kingdom of Saudi Arabia Vol 3Ministry of Agriculture and Water Riyadh

Dehay C (1935) Lagraveppareil Libero-ligneux foliare des EuphorbiaceaeAnnales des Sciences Naturelles Botanique 10 147-190

Dehgan B (1982) Comparative anatomy of the petiole and infragenericrelationships in Jatropha (Euphorbiaceae) American Journal oBotany 69 1283-1295

Gaucher L (1898) Ē tude anatomique du genre Euphorbia LPKlincksieck Paris

Gaucher L (1902) Recherches anatomiques sur les EuphorbiaceacuteesAnnales des Sciences Naturelles Botanique 15 161-309

Gaucher L (1902) Recherches anatomiques sur les Euphorbiaceacutees

Annales des Sciences Naturelles Botanique 15 161-309Govaerts R Fordin D Radcliffe-Smith A (2000) World Checklist and

Bibliography of Euphorbiaceae The Royal Botanic Gardens KewHickey LJ (1973) Classification of the architecture of dicotyledonous

leaves American Journal of Botany 60 17-33Hickey LJ (1979) A revised classification of the architecture o

dicotyledonous leaves In Metcalfe CR and Chalk L eds Anatomy oDicotyledons Systematic Anatomy of the leaf and stem 2nd edClarendon Press Oxford 25-39

Kakkar L Paliwal GS (1972) Epidermis in Euphorbia Indian ScienceCongress Association Proceedings 59 326-327

Kakkar L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 5 Epidermis Proceedings of the Indian National ScienceAcademy Part B Biological Sciences 40 55-67

Koumlhler E (1993) Blattvervatur-Muster der Buxaceae Dumortierr undSimmondsiaceae Van Tieghem Feddes Repertorium 104 145-167

Levin GA (1986a) Systematic foliar morphology of Phylanthoideae(Euphorbiaceae) II Phenetic analysis Annals of Missouri BotanicaGarden 73 86-98

Levin GA (1986b) Systematic foliar morphology of Phylanthoideae IICladistic analysis Systematic Botany 11 515-530

Levin GA (1986c) Systematic foliar morphology of Phyllanthoideae(Euphorbiaceae) I conspectus Annals of Missouri Botanical Garden73 29-85

Mahlberg P (1973) Histochemistry and Scanning Electron Microscopyof Starch Grains from Latex of Euphorbia-Terracina and EuphorbiaTirucalli Proceedings of the Indiana Academy of Science 82 132

Mahlberg PG (1974) Scanning Microscopic Comparison of StarchGrains from the Latex of Euphorbia -Spp Proceedings of the IndianaAcademy of Science 83 83-84

Mahlberg PG (1975) Evolution of the Laticifer in Euphorbia asInterpreted from Starch Grain Morphology American Journal oBotany 62 577-583

Mahlberg PG (1982) Comparative Morphology of Starch Grains inLatex from Varieties of Poinsettia Euphorbia-PulcherrimEuphorbiaceae Botanical Gazette 143 206-209

Mahlberg PG Davis DG Galitz DS Manners GD (1987) Laticifers andthe Classification of Euphorbia the Chemotaxonomy of EuphorbiaEsula L Botanical Journal of the Linnean Society 94 165-180

Metcalfe C Chalk L (1950) Anatomy of the Dicotyledons At theClarendon Press Oxford

Miller K Webster GL (1962) Systematic position of Cnidoscolus andJatropha Brittonia 14 174-180

Pax F (1884) Die Anatomie der Euphorbiaceen in ihrer Beziehung zumSystem derselben Botanische Jahrbuumlcher fuumlr SystematikPflanzengeschichte und Pflanzengeographie Leipzig 5 384-421

Payne WW (1970) Helicocytic and allelocytic stomata unrecognizedpatters in the Dicotyledonae Am J Bot57 140-147

Pray TR (1955a) Foliar venation of angiosperm II Histogenesis of thevenation of Liriodendron Am J Bot 42 18-27

Pray TR 1955b Foliar venation of angiosperms III Pattern andhistology of the venation of Hosta Am J Bot 42 11-18

Radford AE Dickison WC Massey JR Bell CR (1974) Vascular PlanSystematics Harper amp Row Publishers London New York

Radlkofer L (1870) Ueber Pausandra ein neues EuphorbiaceenGenus Flora 53 81-95

Raju VS Rao PN (1977) Variation in the structure and development ofoliar stomata in the Euphorbiaceae Botanical Journal of the LinneanSociety 75 69-97

Raju VS Rao PN (1987) The Taxonomic Use of the Basic StomataType in the Generic Delimitation of Chamaesyce EuphorbiaceaeFeddes Repertorium 98 137-142

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2424

Rosowski JR (1968) Laticifer Morphology in the Mature Stem and Leafof Euphorbia supina Botanical Gazette 129 113-120

Roth-Nebelsick A Uhl D Mosbrugger V Kerp H (2001) Evolution andfunction of leaf venation architecture a review Annals of Botany 87553-566

Sehgal L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 2 Venation Patterns Botanical Journal of the Linnean Society 68 173-208

Singh G (2004) Plant Systematics An Integrated Approach SciencePublishers Enfield

Solereder H (1899) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Solereder H (1908) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Aldhebiani and Jury 191

Stace CA (1965) Cuticular studies as an aid to plant taxonomy Bulletinof the British Museum Natural History Botany 4 3-78

Stace CA (1989) Plant Taxonomy and Biosystematics Hodder ampStoughton London

Tognini F (1897) Contribuzione allo studio della organogeniacomparata degli stomi Institut de Botanique University de Geneve 41-42

Van Cotthem WR (1970) A classification of stomatal types Botanica

Journal of Linnean Society 63 235-246Wagner W (1979) Reticulate veins in the systematics of modern ferns

Taxon 1 87-95Webster GL (1994a) Classification of the Euphorbiaceae Annals of the

Missouri Botanical Garden 81 3-32

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 824

Aldhebiani and Jury 175

983105983140983137983160983145983137983148

983107

983108

983105

983106

983105983138983137983160983145983137983148

Figure 3 Epidermal cells vary on leaf surfaces adaxial surface has irregular cells whilethe abaxial surface has plicate (LM)(A) E schimperiana (B) E arabica (C) E serpens (D) E scordiifolia

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 924

176 Int Res J Plant Sci

983105983140983137983160983145983137983148

983107

983105

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Figure 4 Epidermal cell shape varies on the adaxial andthe abaxial surfaces straight cell walls in the formerwhile folded plicate walls occur on the latter (LM) (A) Egranulata (B) E inaequilatera and (C) E helioscopia

983112983141983149983145983152983137983154983137983139983161983156983145983139 983155983156983151983149983137983156983137

983151983150 983138983151983156983144 983155983157983154983142983137983139983141983155

Figure 5 E hypericifolia (LM)Above photo the adaxial surface with irregular epidermal cells andvery clear papillae as a circle in the middle of the cellLower photo the abaxial surface with plicate cells and papillae ahair base is obvious in the right side of the picture

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 177

983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

983120983137983152983145983148983148983137983141

983117983157983148983156983145983139983141983148983148983157983148983137983154 983144983137983145983154983155 983145983150 983137983138983137983160983145983137983148 983155983157983154983142983137983139983141983086

Figure 6 E hypericifolia (ESEM) papillae on the adaxial and abaxial surfacesmulticellular hairs are on the abaxial surface of the leaf only

Figure7 Multicellular hairs on adaxial and abaxial surfaces of E hirta

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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178 Int Res J Plant Sci

Figure 8 Hairs wart ornamentation on hair surfaces of (A) E scordiifolia and (B) E pirottae (C) Short hairs onabaxial surface of E balsamifera subsp adenensis

In E balsamifera subsp adenensis the hairs are shortand unicellular (Figure 8C) and they are located mostly atthe leaf margins and the mid-vein area on the abaxialsurface only

Hairs on the leaf of E pirottae are present on theabaxial surface only and are both unicellular andmulticellular Euphorbia granulata is similar to E pirottae in having both unicellular and multicellular hairs but herethey occur on both surfaces of the leaf (Figure 8B)

On the other hand in E scordiifolia E hirta and Ehypericifolia the hairs are multicellular and on bothsurfaces in the first two species and on the abaxial only inthe latter (Figure 7) Moreover E hypericifolia and Ehirta have long hairs when compared with the otherspecies (Figure 6)

No hairs were observed and leaves were glabrous inthe following species E serpens E arabica E peplusE inaequilatera E falcata E helioscopia E cuneata Eretusa E grossheimii E schimperiana E schimperiE chamaepeplus and the new variety of E granulata

Wax

The cuticle layer and wax in the Euphorbia species undeinvestigation vary from smooth to densely cover in waxscales This may be a thick smooth layer of wax whichsometimes has upright scales In some species only thecell surface has a smooth wax layer whereas the walbetween cells has upright scales of wax as in Eschimperiana especially around the stomata pores

(Figure 9) In contrast the anticlinal walls of somespecies have a smooth layer of wax whereas the celsurface is densely covered with upright wax flakes or withsome stellate wax flakes in other species

Stomata

Stomata were observed through LM and ESEM In thespecies under investigation stomata commonly appeaon both leaf surfaces (amphistomatic leaves) excepin E cuneata where the stomata are located only on the

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 1224

Aldhebiani and Jury 179

983109

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983108983107

983106

Figure 9 Wax formation in Euphorbia species in Saudi Arabia (A+B) smooth andthick in E peplus and E schimperi (C+D) smooth on top of cell while formingupright scale flakes in between especially around stomata as in E schimperiana smooth in anticlinal walls (E) upright scale flakes in E balsamifera

abaxial surface (hypostomatic leaves) The presence andposition of subsidiary cells are variable in Euphorbia

species Subsidiary cells were found to be of unequalsize and shape whether the stoma was anisocyticparacytic actinocytic or tetracytic While some specieslack them around the stomata others have more thanone arrangement of subsidiary cells

The most common stomata type is anomocytic whilethe rare type is actinocytic recorded only in Ehelioscopia Moreover sometimes stomata of more thanone type (up to three or four types) have beenencountered on the same leaf surface as in E scordiifolia (three types on the abaxial surface) and E hirta (five

types on the abaxial and three on the adaxial) Thedistribution and type of stomata are shown in Table 2

In addition in some species such as E falcata and Ehelioscopia the stomata are deeply sunken andrepresented only by cuticular ledges surrounding theminute pores as seen in ESEM

Examples of species lacking subsidiary cells aroundstomata are E peplus E chamaepeplus E granulataE dracunculoides and E acalyphoides Therefore theyall have anomocytic stomata

On the other hand E balsamifera subsp adenensishas the mononcytic type on both leaf surfaces andanisocytic and paracytic only on the abaxial surface In

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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180 Int Res J Plant Sci

Table 2 Stomata type in Euphorbia taxa in Saudi Arabia

Ab abaxial surface + stomata presentAd adaxial surface

addition E schimperiana has anomocytic andtetracytic on the abaxial only In contrast Ecuneata has three types of stomata anomocyticparacytic and anisocytic only on the lower surfaceAs well ldquoE aff consobrinardquo ldquoE aff schimperirdquo

and E schimperi have paracytic and anisocyticonly in the abaxial on the first two and in both leafsurfaces on the latter Therefore ldquoE affconsobrinardquo and ldquoE aff schimperirdquo are the same taxa Then comparing to E schimperi results

from ansubspec

Eupsimilar and an

Subgenus

Stomata

Anomocytic Hemiparacytic Paracytic Anisocytic T

Species Ad Ab Ad Ab Ad Ab Ad Ab AT i r u c al l i

E balsamifera subsp adenensis + + +

E schimperi + + + +

ldquoEsp aff consobrinardquo + +

ldquoE sp aff schimperirdquo + +

E cuneata + + +

ErmophytonE acalyphoides + +

E pirottae + + + +

E helioscopia + + E s ul a

E grossheimii + + + + +

E retusa + + + + +

E dracunculoides +

E falcate + + E peplus +

E chamaepeplus + +

E schimperiana + C h am a e s y c e

E Arabica + + + + + + +

E serpens + + + +

E scordiifolia + + + + + + +

E granulate + + + +

E inaequilatera + + + +

E hirta + + + + + + +

E hypericifolia + + + + + +

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Table3 Venation type in Euphorbia taxa in Saudi Arabia

Subgenus Species

Venation

Number of veins atthe leaf base

Margins Sheathed

1 2 3 4 or more Open Closed Rounded T i r u c al l i

Ebalsamiferasubspadenensis

+ + +

E schimperi + + +

E aff Schimperi + + +

E cuneata + + +

ErmophytonE acalyphoides + + +

E pirottae + + E s ul a

E grossheimii + +

E retusa + + +

E dracunculoides + + +

E falcate + + +

E peplus + +

E chamaepeplus + +

E schimperiana

E helioscopia

+

+

+

+ C h am a e s y c e

E Arabica + + +

E serpens + + +

E scordiifolia + + +

E granulate + + + +

E inaequilatera + + +

E hirta + + + E hypericifolia + + +

Poinsettia E cyathophora + +

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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182 Int Res J Plant Sci

Figure 10 Sunken stomata on the abaxial surface1) Deeply sunken (A) E helioscopia (B) E falcata (C) E schimperiana 2) Slightly sunken (D) E peplus (E) E pirottae

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 1624

Aldhebiani and Jury 183

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983105983150983145983155983151983139983161983156983145983139983124983141983156983154983137983139983161983156983145983139

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983105983139983156983145983150983151983139983161983156983145983139

983105983140983137983160983145983137983148

Figure 11 Stomata types in Euphorbia taxa in Saudi Arabia (A) E balsamifera subsp adenensis (B) E schimperi (C) E cuneata and (D) E grossheimii

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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184 Int Res J Plant Sci

983105

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983124983141983156983154983137983139983161983156983145983139

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983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

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983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983138983137983160983145983137983148983105983140983137983160983145983137983148

Figure 12 Stomata types in Euphorbia taxa (A) E falcata (B) E peplus (C) E chamaepeplus and (D) Eschimperiana

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 185

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983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

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983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

Figure 13 Figure 13 Stomata types in Euphorbia taxa in Saudi Arabia (A) E arabica (B) E serpens (C)E scordiifolia and (D) E helioscopia

distribution between the leaf surfaces (see Figure 13)While E hirta has the most types E helioscopia Edracunculoides and E acalyphoides have the least

Regarding the position of the stomata on the surfacelevel of the epidermis a variable degree of sunkenesshas been recorded Using the ESEM the stomata appear

deeply sunken in some species whilst in others they areonly slightly sunken with the outer rims of the subsidiarycells surrounding the aperture The deeply and slightlysunken species are recorded in Table 2 AdditionallyFigure 10 shows the sunken stomata in different speciesunder the ESEM

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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186 Int Res J Plant Sci

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983112983141983149983145983152983137983154983137983139983161983156983145983139

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983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

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983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

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983107

983106

983105

983105983140983137983160983145983137983148983105983138983137983160983145983137983148

Figure 14 Stomata types in Euphorbia taxa in Saudi Arabia (A) E hirta (B) E hypericifolia (C) E pirottae and (D)Espaff schimperi

Venation

The venation patterns in the Euphorbia species underinvestigation vary from one-veined three-veined to thosewith four or more veins The predominant type is three-veined However in E balsamifera subsp Adenensis

there are seven middle parallel veins three middle thickand four thinner lateral ones (Figure 15 A and B)Whereas in E pirottae there are three main middleparallel veins and two thinner laterals (Figure 15 C) Incontrast E chamaepeplus E schimperiana and Earabica have only one thick main midrib coming from the

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 187

983105

983106

983107

Figure 15 Figure 15 Venation patterns at base of leaves invariants with more than three veins (A amp B) E balsamiferasubsp adenensis and (C) E pirottae

petiole then branched from the side into two lateral veinsat the point when the leaf starts widening (Figure 16)Each of these veins branches again into secondary veinsat the leaf margin

The rest of the species have just three veins Egrossheimii E dracunculoides E falcata E peplus Eserpens E scordiifolia E granulata E inaequilateraE helioscopia E hirta E hypericifolia and E

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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188 Int Res J Plant Sci

Figure 16 Venation patterns with one main mid vein at the leaf base in (A) E arabica and (B) E chamaepeplus

983106983105

Figure 17 Venation patterns with three-veined leaves in (A) E falcata and (B) E granulata

cyathophora All the three veins come from the petioleand start branching into secondary veins both toward themargin and the mid vein direction (Figure 17 B)

The vein endings at the leaf margin vary from closed

to open There is a relationship between the closedmargins and the sheathed veins (midrib and secondaryveins that are surrounded by parenchyma tissue) Whenthe veins are ornamented (sheathed) the margins areclosed This can be seen in subgenus Chamaesyce inspecies such as E serpens E scordiifolia E granulataE inaequilatera E hirta E hypericifolia and E arabica Additionally these species have the three-vein venation

On the other hand when the veins are not sheathedthe veins ends are open at the leaf margin as in Ebalsamifera subsp adenensis E schimperi E cuneataE acalyphoides E grossheimii E dracunculoides E

falcata E peplus E chamaepeplus E schimperiana Ehelioscopia E pirottae ldquoE aff schimperirdquo and Ecyathophora

CONCLUSIONS

Anatomical characters are seen here to be very useful inEuphorbia classification and identification The resultsobtained from anatomical studies were useful to groupsome species and identify others

Depending on the venation type the Euphorbiaspecies under investigation are divided anatomically intothree main groups see Figure 18

Group one veins are sheathed epidermal cellsdifferentiated on adaxial and abaxial surfaces Group two

983105 983106

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Figure 18 Hierarchial relation of Euphorbia taxa in Saudi Arabia based on anatomical charac

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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190 Int Res J Plant Sci

veins are not sheathed and epidermal cells are alike inboth surfaces And the third group includes species thatdo not belong to any of the previous groups The firstgroup can be further divided into three-veined and one-veined while the second group into three-veined and fouror more veins

In addition E grossheimii E retusa and E pirottae are related by being three-veined open at the marginswith rounded to oval tracheas and elongated cells bothon adaxial and abaxial surfaces However in E retusa the epidermal cells have straight cell walls on the adaxialsurface They are also similar in having hemiparacyticand paracytic types of stomata either on the adaxial orthe abaxial surface

On the other hand species with two differentepidermal cells on the leaf surfaces usually have three-sheathed veins with closed margins and elongatedtrachea such as E serpens E scordiifolia E granulata E inaequilatera E hirta and E hypericifolia which belongto subgenus Chamaesyce according to (Carter andSmith 1988)

In the case of E granulata which has beendescribed as having three different variants E granulata var granulata (both leaf surfaces hairy) E granulata varglabrata (adaxial glabrous and abaxial hairy) and aglabrous variant (the whole plant glabrous) Anatomicalwork has shown especially by using the ESEM that theleaves are either hairy or glabrous on both surfaces Sowhen the leaf is hairy both surfaces have differences inhair distribution Usually the lower surface is more hairythan upper surface (E granulata var glabrata ) butsometimes they are evenly distributed (Egranulata vargranulata ) In addition hairs on both surfaces are

multicellular In contrast when the leaf is glabrous bothleaf surfaces are glabrous and the whole plant isglabrous (suggested new variety) They were similar inhaving the same venation type (three-veined)

Finally ldquoE aff schimperi ldquohas some similarity with E schimperi in having the same open margin not theornamented venation type As well they both have thesame type of stomata paracytic and anisocytic but theydiffer in their distribution on abaxial and adaxial Thusthis may support the suggestion considering ldquoE affschimperi rdquo as a new subspecies of E schimperi

REFERENCES

Baranova M (1992) Principels of comparative stomatographic studiesof flowering plants The Botanical Review 58 49-99

Carlquist S (1961) Comparative Plant Anatomy Holt Rinehart ampWinston New York

Carter S Smith AR (1988) Euphorbiaceae (Part2) In Polhill ed Floraof Tropical East Africa Rotterdam Brookfield AA Balkema 409-597

Chaudhary SA (2001) Flora of The Kingdom of Saudi Arabia Vol 3Ministry of Agriculture and Water Riyadh

Dehay C (1935) Lagraveppareil Libero-ligneux foliare des EuphorbiaceaeAnnales des Sciences Naturelles Botanique 10 147-190

Dehgan B (1982) Comparative anatomy of the petiole and infragenericrelationships in Jatropha (Euphorbiaceae) American Journal oBotany 69 1283-1295

Gaucher L (1898) Ē tude anatomique du genre Euphorbia LPKlincksieck Paris

Gaucher L (1902) Recherches anatomiques sur les EuphorbiaceacuteesAnnales des Sciences Naturelles Botanique 15 161-309

Gaucher L (1902) Recherches anatomiques sur les Euphorbiaceacutees

Annales des Sciences Naturelles Botanique 15 161-309Govaerts R Fordin D Radcliffe-Smith A (2000) World Checklist and

Bibliography of Euphorbiaceae The Royal Botanic Gardens KewHickey LJ (1973) Classification of the architecture of dicotyledonous

leaves American Journal of Botany 60 17-33Hickey LJ (1979) A revised classification of the architecture o

dicotyledonous leaves In Metcalfe CR and Chalk L eds Anatomy oDicotyledons Systematic Anatomy of the leaf and stem 2nd edClarendon Press Oxford 25-39

Kakkar L Paliwal GS (1972) Epidermis in Euphorbia Indian ScienceCongress Association Proceedings 59 326-327

Kakkar L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 5 Epidermis Proceedings of the Indian National ScienceAcademy Part B Biological Sciences 40 55-67

Koumlhler E (1993) Blattvervatur-Muster der Buxaceae Dumortierr undSimmondsiaceae Van Tieghem Feddes Repertorium 104 145-167

Levin GA (1986a) Systematic foliar morphology of Phylanthoideae(Euphorbiaceae) II Phenetic analysis Annals of Missouri BotanicaGarden 73 86-98

Levin GA (1986b) Systematic foliar morphology of Phylanthoideae IICladistic analysis Systematic Botany 11 515-530

Levin GA (1986c) Systematic foliar morphology of Phyllanthoideae(Euphorbiaceae) I conspectus Annals of Missouri Botanical Garden73 29-85

Mahlberg P (1973) Histochemistry and Scanning Electron Microscopyof Starch Grains from Latex of Euphorbia-Terracina and EuphorbiaTirucalli Proceedings of the Indiana Academy of Science 82 132

Mahlberg PG (1974) Scanning Microscopic Comparison of StarchGrains from the Latex of Euphorbia -Spp Proceedings of the IndianaAcademy of Science 83 83-84

Mahlberg PG (1975) Evolution of the Laticifer in Euphorbia asInterpreted from Starch Grain Morphology American Journal oBotany 62 577-583

Mahlberg PG (1982) Comparative Morphology of Starch Grains inLatex from Varieties of Poinsettia Euphorbia-PulcherrimEuphorbiaceae Botanical Gazette 143 206-209

Mahlberg PG Davis DG Galitz DS Manners GD (1987) Laticifers andthe Classification of Euphorbia the Chemotaxonomy of EuphorbiaEsula L Botanical Journal of the Linnean Society 94 165-180

Metcalfe C Chalk L (1950) Anatomy of the Dicotyledons At theClarendon Press Oxford

Miller K Webster GL (1962) Systematic position of Cnidoscolus andJatropha Brittonia 14 174-180

Pax F (1884) Die Anatomie der Euphorbiaceen in ihrer Beziehung zumSystem derselben Botanische Jahrbuumlcher fuumlr SystematikPflanzengeschichte und Pflanzengeographie Leipzig 5 384-421

Payne WW (1970) Helicocytic and allelocytic stomata unrecognizedpatters in the Dicotyledonae Am J Bot57 140-147

Pray TR (1955a) Foliar venation of angiosperm II Histogenesis of thevenation of Liriodendron Am J Bot 42 18-27

Pray TR 1955b Foliar venation of angiosperms III Pattern andhistology of the venation of Hosta Am J Bot 42 11-18

Radford AE Dickison WC Massey JR Bell CR (1974) Vascular PlanSystematics Harper amp Row Publishers London New York

Radlkofer L (1870) Ueber Pausandra ein neues EuphorbiaceenGenus Flora 53 81-95

Raju VS Rao PN (1977) Variation in the structure and development ofoliar stomata in the Euphorbiaceae Botanical Journal of the LinneanSociety 75 69-97

Raju VS Rao PN (1987) The Taxonomic Use of the Basic StomataType in the Generic Delimitation of Chamaesyce EuphorbiaceaeFeddes Repertorium 98 137-142

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2424

Rosowski JR (1968) Laticifer Morphology in the Mature Stem and Leafof Euphorbia supina Botanical Gazette 129 113-120

Roth-Nebelsick A Uhl D Mosbrugger V Kerp H (2001) Evolution andfunction of leaf venation architecture a review Annals of Botany 87553-566

Sehgal L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 2 Venation Patterns Botanical Journal of the Linnean Society 68 173-208

Singh G (2004) Plant Systematics An Integrated Approach SciencePublishers Enfield

Solereder H (1899) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Solereder H (1908) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Aldhebiani and Jury 191

Stace CA (1965) Cuticular studies as an aid to plant taxonomy Bulletinof the British Museum Natural History Botany 4 3-78

Stace CA (1989) Plant Taxonomy and Biosystematics Hodder ampStoughton London

Tognini F (1897) Contribuzione allo studio della organogeniacomparata degli stomi Institut de Botanique University de Geneve 41-42

Van Cotthem WR (1970) A classification of stomatal types Botanica

Journal of Linnean Society 63 235-246Wagner W (1979) Reticulate veins in the systematics of modern ferns

Taxon 1 87-95Webster GL (1994a) Classification of the Euphorbiaceae Annals of the

Missouri Botanical Garden 81 3-32

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 924

176 Int Res J Plant Sci

983105983140983137983160983145983137983148

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Figure 4 Epidermal cell shape varies on the adaxial andthe abaxial surfaces straight cell walls in the formerwhile folded plicate walls occur on the latter (LM) (A) Egranulata (B) E inaequilatera and (C) E helioscopia

983112983141983149983145983152983137983154983137983139983161983156983145983139 983155983156983151983149983137983156983137

983151983150 983138983151983156983144 983155983157983154983142983137983139983141983155

Figure 5 E hypericifolia (LM)Above photo the adaxial surface with irregular epidermal cells andvery clear papillae as a circle in the middle of the cellLower photo the abaxial surface with plicate cells and papillae ahair base is obvious in the right side of the picture

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 177

983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

983120983137983152983145983148983148983137983141

983117983157983148983156983145983139983141983148983148983157983148983137983154 983144983137983145983154983155 983145983150 983137983138983137983160983145983137983148 983155983157983154983142983137983139983141983086

Figure 6 E hypericifolia (ESEM) papillae on the adaxial and abaxial surfacesmulticellular hairs are on the abaxial surface of the leaf only

Figure7 Multicellular hairs on adaxial and abaxial surfaces of E hirta

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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178 Int Res J Plant Sci

Figure 8 Hairs wart ornamentation on hair surfaces of (A) E scordiifolia and (B) E pirottae (C) Short hairs onabaxial surface of E balsamifera subsp adenensis

In E balsamifera subsp adenensis the hairs are shortand unicellular (Figure 8C) and they are located mostly atthe leaf margins and the mid-vein area on the abaxialsurface only

Hairs on the leaf of E pirottae are present on theabaxial surface only and are both unicellular andmulticellular Euphorbia granulata is similar to E pirottae in having both unicellular and multicellular hairs but herethey occur on both surfaces of the leaf (Figure 8B)

On the other hand in E scordiifolia E hirta and Ehypericifolia the hairs are multicellular and on bothsurfaces in the first two species and on the abaxial only inthe latter (Figure 7) Moreover E hypericifolia and Ehirta have long hairs when compared with the otherspecies (Figure 6)

No hairs were observed and leaves were glabrous inthe following species E serpens E arabica E peplusE inaequilatera E falcata E helioscopia E cuneata Eretusa E grossheimii E schimperiana E schimperiE chamaepeplus and the new variety of E granulata

Wax

The cuticle layer and wax in the Euphorbia species undeinvestigation vary from smooth to densely cover in waxscales This may be a thick smooth layer of wax whichsometimes has upright scales In some species only thecell surface has a smooth wax layer whereas the walbetween cells has upright scales of wax as in Eschimperiana especially around the stomata pores

(Figure 9) In contrast the anticlinal walls of somespecies have a smooth layer of wax whereas the celsurface is densely covered with upright wax flakes or withsome stellate wax flakes in other species

Stomata

Stomata were observed through LM and ESEM In thespecies under investigation stomata commonly appeaon both leaf surfaces (amphistomatic leaves) excepin E cuneata where the stomata are located only on the

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 179

983109

983105

983108983107

983106

Figure 9 Wax formation in Euphorbia species in Saudi Arabia (A+B) smooth andthick in E peplus and E schimperi (C+D) smooth on top of cell while formingupright scale flakes in between especially around stomata as in E schimperiana smooth in anticlinal walls (E) upright scale flakes in E balsamifera

abaxial surface (hypostomatic leaves) The presence andposition of subsidiary cells are variable in Euphorbia

species Subsidiary cells were found to be of unequalsize and shape whether the stoma was anisocyticparacytic actinocytic or tetracytic While some specieslack them around the stomata others have more thanone arrangement of subsidiary cells

The most common stomata type is anomocytic whilethe rare type is actinocytic recorded only in Ehelioscopia Moreover sometimes stomata of more thanone type (up to three or four types) have beenencountered on the same leaf surface as in E scordiifolia (three types on the abaxial surface) and E hirta (five

types on the abaxial and three on the adaxial) Thedistribution and type of stomata are shown in Table 2

In addition in some species such as E falcata and Ehelioscopia the stomata are deeply sunken andrepresented only by cuticular ledges surrounding theminute pores as seen in ESEM

Examples of species lacking subsidiary cells aroundstomata are E peplus E chamaepeplus E granulataE dracunculoides and E acalyphoides Therefore theyall have anomocytic stomata

On the other hand E balsamifera subsp adenensishas the mononcytic type on both leaf surfaces andanisocytic and paracytic only on the abaxial surface In

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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180 Int Res J Plant Sci

Table 2 Stomata type in Euphorbia taxa in Saudi Arabia

Ab abaxial surface + stomata presentAd adaxial surface

addition E schimperiana has anomocytic andtetracytic on the abaxial only In contrast Ecuneata has three types of stomata anomocyticparacytic and anisocytic only on the lower surfaceAs well ldquoE aff consobrinardquo ldquoE aff schimperirdquo

and E schimperi have paracytic and anisocyticonly in the abaxial on the first two and in both leafsurfaces on the latter Therefore ldquoE affconsobrinardquo and ldquoE aff schimperirdquo are the same taxa Then comparing to E schimperi results

from ansubspec

Eupsimilar and an

Subgenus

Stomata

Anomocytic Hemiparacytic Paracytic Anisocytic T

Species Ad Ab Ad Ab Ad Ab Ad Ab AT i r u c al l i

E balsamifera subsp adenensis + + +

E schimperi + + + +

ldquoEsp aff consobrinardquo + +

ldquoE sp aff schimperirdquo + +

E cuneata + + +

ErmophytonE acalyphoides + +

E pirottae + + + +

E helioscopia + + E s ul a

E grossheimii + + + + +

E retusa + + + + +

E dracunculoides +

E falcate + + E peplus +

E chamaepeplus + +

E schimperiana + C h am a e s y c e

E Arabica + + + + + + +

E serpens + + + +

E scordiifolia + + + + + + +

E granulate + + + +

E inaequilatera + + + +

E hirta + + + + + + +

E hypericifolia + + + + + +

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Table3 Venation type in Euphorbia taxa in Saudi Arabia

Subgenus Species

Venation

Number of veins atthe leaf base

Margins Sheathed

1 2 3 4 or more Open Closed Rounded T i r u c al l i

Ebalsamiferasubspadenensis

+ + +

E schimperi + + +

E aff Schimperi + + +

E cuneata + + +

ErmophytonE acalyphoides + + +

E pirottae + + E s ul a

E grossheimii + +

E retusa + + +

E dracunculoides + + +

E falcate + + +

E peplus + +

E chamaepeplus + +

E schimperiana

E helioscopia

+

+

+

+ C h am a e s y c e

E Arabica + + +

E serpens + + +

E scordiifolia + + +

E granulate + + + +

E inaequilatera + + +

E hirta + + + E hypericifolia + + +

Poinsettia E cyathophora + +

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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182 Int Res J Plant Sci

Figure 10 Sunken stomata on the abaxial surface1) Deeply sunken (A) E helioscopia (B) E falcata (C) E schimperiana 2) Slightly sunken (D) E peplus (E) E pirottae

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 183

983105983138983137983160983145983137983148

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983108

983108

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983112983141983149983145983152983137983154983137983139983161983156983145983139

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983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

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983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

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983105983140983137983160983145983137983148

Figure 11 Stomata types in Euphorbia taxa in Saudi Arabia (A) E balsamifera subsp adenensis (B) E schimperi (C) E cuneata and (D) E grossheimii

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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184 Int Res J Plant Sci

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983105983138983137983160983145983137983148983105983140983137983160983145983137983148

Figure 12 Stomata types in Euphorbia taxa (A) E falcata (B) E peplus (C) E chamaepeplus and (D) Eschimperiana

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 185

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Figure 13 Figure 13 Stomata types in Euphorbia taxa in Saudi Arabia (A) E arabica (B) E serpens (C)E scordiifolia and (D) E helioscopia

distribution between the leaf surfaces (see Figure 13)While E hirta has the most types E helioscopia Edracunculoides and E acalyphoides have the least

Regarding the position of the stomata on the surfacelevel of the epidermis a variable degree of sunkenesshas been recorded Using the ESEM the stomata appear

deeply sunken in some species whilst in others they areonly slightly sunken with the outer rims of the subsidiarycells surrounding the aperture The deeply and slightlysunken species are recorded in Table 2 AdditionallyFigure 10 shows the sunken stomata in different speciesunder the ESEM

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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186 Int Res J Plant Sci

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Figure 14 Stomata types in Euphorbia taxa in Saudi Arabia (A) E hirta (B) E hypericifolia (C) E pirottae and (D)Espaff schimperi

Venation

The venation patterns in the Euphorbia species underinvestigation vary from one-veined three-veined to thosewith four or more veins The predominant type is three-veined However in E balsamifera subsp Adenensis

there are seven middle parallel veins three middle thickand four thinner lateral ones (Figure 15 A and B)Whereas in E pirottae there are three main middleparallel veins and two thinner laterals (Figure 15 C) Incontrast E chamaepeplus E schimperiana and Earabica have only one thick main midrib coming from the

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 187

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Figure 15 Figure 15 Venation patterns at base of leaves invariants with more than three veins (A amp B) E balsamiferasubsp adenensis and (C) E pirottae

petiole then branched from the side into two lateral veinsat the point when the leaf starts widening (Figure 16)Each of these veins branches again into secondary veinsat the leaf margin

The rest of the species have just three veins Egrossheimii E dracunculoides E falcata E peplus Eserpens E scordiifolia E granulata E inaequilateraE helioscopia E hirta E hypericifolia and E

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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188 Int Res J Plant Sci

Figure 16 Venation patterns with one main mid vein at the leaf base in (A) E arabica and (B) E chamaepeplus

983106983105

Figure 17 Venation patterns with three-veined leaves in (A) E falcata and (B) E granulata

cyathophora All the three veins come from the petioleand start branching into secondary veins both toward themargin and the mid vein direction (Figure 17 B)

The vein endings at the leaf margin vary from closed

to open There is a relationship between the closedmargins and the sheathed veins (midrib and secondaryveins that are surrounded by parenchyma tissue) Whenthe veins are ornamented (sheathed) the margins areclosed This can be seen in subgenus Chamaesyce inspecies such as E serpens E scordiifolia E granulataE inaequilatera E hirta E hypericifolia and E arabica Additionally these species have the three-vein venation

On the other hand when the veins are not sheathedthe veins ends are open at the leaf margin as in Ebalsamifera subsp adenensis E schimperi E cuneataE acalyphoides E grossheimii E dracunculoides E

falcata E peplus E chamaepeplus E schimperiana Ehelioscopia E pirottae ldquoE aff schimperirdquo and Ecyathophora

CONCLUSIONS

Anatomical characters are seen here to be very useful inEuphorbia classification and identification The resultsobtained from anatomical studies were useful to groupsome species and identify others

Depending on the venation type the Euphorbiaspecies under investigation are divided anatomically intothree main groups see Figure 18

Group one veins are sheathed epidermal cellsdifferentiated on adaxial and abaxial surfaces Group two

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7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Figure 18 Hierarchial relation of Euphorbia taxa in Saudi Arabia based on anatomical charac

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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190 Int Res J Plant Sci

veins are not sheathed and epidermal cells are alike inboth surfaces And the third group includes species thatdo not belong to any of the previous groups The firstgroup can be further divided into three-veined and one-veined while the second group into three-veined and fouror more veins

In addition E grossheimii E retusa and E pirottae are related by being three-veined open at the marginswith rounded to oval tracheas and elongated cells bothon adaxial and abaxial surfaces However in E retusa the epidermal cells have straight cell walls on the adaxialsurface They are also similar in having hemiparacyticand paracytic types of stomata either on the adaxial orthe abaxial surface

On the other hand species with two differentepidermal cells on the leaf surfaces usually have three-sheathed veins with closed margins and elongatedtrachea such as E serpens E scordiifolia E granulata E inaequilatera E hirta and E hypericifolia which belongto subgenus Chamaesyce according to (Carter andSmith 1988)

In the case of E granulata which has beendescribed as having three different variants E granulata var granulata (both leaf surfaces hairy) E granulata varglabrata (adaxial glabrous and abaxial hairy) and aglabrous variant (the whole plant glabrous) Anatomicalwork has shown especially by using the ESEM that theleaves are either hairy or glabrous on both surfaces Sowhen the leaf is hairy both surfaces have differences inhair distribution Usually the lower surface is more hairythan upper surface (E granulata var glabrata ) butsometimes they are evenly distributed (Egranulata vargranulata ) In addition hairs on both surfaces are

multicellular In contrast when the leaf is glabrous bothleaf surfaces are glabrous and the whole plant isglabrous (suggested new variety) They were similar inhaving the same venation type (three-veined)

Finally ldquoE aff schimperi ldquohas some similarity with E schimperi in having the same open margin not theornamented venation type As well they both have thesame type of stomata paracytic and anisocytic but theydiffer in their distribution on abaxial and adaxial Thusthis may support the suggestion considering ldquoE affschimperi rdquo as a new subspecies of E schimperi

REFERENCES

Baranova M (1992) Principels of comparative stomatographic studiesof flowering plants The Botanical Review 58 49-99

Carlquist S (1961) Comparative Plant Anatomy Holt Rinehart ampWinston New York

Carter S Smith AR (1988) Euphorbiaceae (Part2) In Polhill ed Floraof Tropical East Africa Rotterdam Brookfield AA Balkema 409-597

Chaudhary SA (2001) Flora of The Kingdom of Saudi Arabia Vol 3Ministry of Agriculture and Water Riyadh

Dehay C (1935) Lagraveppareil Libero-ligneux foliare des EuphorbiaceaeAnnales des Sciences Naturelles Botanique 10 147-190

Dehgan B (1982) Comparative anatomy of the petiole and infragenericrelationships in Jatropha (Euphorbiaceae) American Journal oBotany 69 1283-1295

Gaucher L (1898) Ē tude anatomique du genre Euphorbia LPKlincksieck Paris

Gaucher L (1902) Recherches anatomiques sur les EuphorbiaceacuteesAnnales des Sciences Naturelles Botanique 15 161-309

Gaucher L (1902) Recherches anatomiques sur les Euphorbiaceacutees

Annales des Sciences Naturelles Botanique 15 161-309Govaerts R Fordin D Radcliffe-Smith A (2000) World Checklist and

Bibliography of Euphorbiaceae The Royal Botanic Gardens KewHickey LJ (1973) Classification of the architecture of dicotyledonous

leaves American Journal of Botany 60 17-33Hickey LJ (1979) A revised classification of the architecture o

dicotyledonous leaves In Metcalfe CR and Chalk L eds Anatomy oDicotyledons Systematic Anatomy of the leaf and stem 2nd edClarendon Press Oxford 25-39

Kakkar L Paliwal GS (1972) Epidermis in Euphorbia Indian ScienceCongress Association Proceedings 59 326-327

Kakkar L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 5 Epidermis Proceedings of the Indian National ScienceAcademy Part B Biological Sciences 40 55-67

Koumlhler E (1993) Blattvervatur-Muster der Buxaceae Dumortierr undSimmondsiaceae Van Tieghem Feddes Repertorium 104 145-167

Levin GA (1986a) Systematic foliar morphology of Phylanthoideae(Euphorbiaceae) II Phenetic analysis Annals of Missouri BotanicaGarden 73 86-98

Levin GA (1986b) Systematic foliar morphology of Phylanthoideae IICladistic analysis Systematic Botany 11 515-530

Levin GA (1986c) Systematic foliar morphology of Phyllanthoideae(Euphorbiaceae) I conspectus Annals of Missouri Botanical Garden73 29-85

Mahlberg P (1973) Histochemistry and Scanning Electron Microscopyof Starch Grains from Latex of Euphorbia-Terracina and EuphorbiaTirucalli Proceedings of the Indiana Academy of Science 82 132

Mahlberg PG (1974) Scanning Microscopic Comparison of StarchGrains from the Latex of Euphorbia -Spp Proceedings of the IndianaAcademy of Science 83 83-84

Mahlberg PG (1975) Evolution of the Laticifer in Euphorbia asInterpreted from Starch Grain Morphology American Journal oBotany 62 577-583

Mahlberg PG (1982) Comparative Morphology of Starch Grains inLatex from Varieties of Poinsettia Euphorbia-PulcherrimEuphorbiaceae Botanical Gazette 143 206-209

Mahlberg PG Davis DG Galitz DS Manners GD (1987) Laticifers andthe Classification of Euphorbia the Chemotaxonomy of EuphorbiaEsula L Botanical Journal of the Linnean Society 94 165-180

Metcalfe C Chalk L (1950) Anatomy of the Dicotyledons At theClarendon Press Oxford

Miller K Webster GL (1962) Systematic position of Cnidoscolus andJatropha Brittonia 14 174-180

Pax F (1884) Die Anatomie der Euphorbiaceen in ihrer Beziehung zumSystem derselben Botanische Jahrbuumlcher fuumlr SystematikPflanzengeschichte und Pflanzengeographie Leipzig 5 384-421

Payne WW (1970) Helicocytic and allelocytic stomata unrecognizedpatters in the Dicotyledonae Am J Bot57 140-147

Pray TR (1955a) Foliar venation of angiosperm II Histogenesis of thevenation of Liriodendron Am J Bot 42 18-27

Pray TR 1955b Foliar venation of angiosperms III Pattern andhistology of the venation of Hosta Am J Bot 42 11-18

Radford AE Dickison WC Massey JR Bell CR (1974) Vascular PlanSystematics Harper amp Row Publishers London New York

Radlkofer L (1870) Ueber Pausandra ein neues EuphorbiaceenGenus Flora 53 81-95

Raju VS Rao PN (1977) Variation in the structure and development ofoliar stomata in the Euphorbiaceae Botanical Journal of the LinneanSociety 75 69-97

Raju VS Rao PN (1987) The Taxonomic Use of the Basic StomataType in the Generic Delimitation of Chamaesyce EuphorbiaceaeFeddes Repertorium 98 137-142

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2424

Rosowski JR (1968) Laticifer Morphology in the Mature Stem and Leafof Euphorbia supina Botanical Gazette 129 113-120

Roth-Nebelsick A Uhl D Mosbrugger V Kerp H (2001) Evolution andfunction of leaf venation architecture a review Annals of Botany 87553-566

Sehgal L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 2 Venation Patterns Botanical Journal of the Linnean Society 68 173-208

Singh G (2004) Plant Systematics An Integrated Approach SciencePublishers Enfield

Solereder H (1899) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Solereder H (1908) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Aldhebiani and Jury 191

Stace CA (1965) Cuticular studies as an aid to plant taxonomy Bulletinof the British Museum Natural History Botany 4 3-78

Stace CA (1989) Plant Taxonomy and Biosystematics Hodder ampStoughton London

Tognini F (1897) Contribuzione allo studio della organogeniacomparata degli stomi Institut de Botanique University de Geneve 41-42

Van Cotthem WR (1970) A classification of stomatal types Botanica

Journal of Linnean Society 63 235-246Wagner W (1979) Reticulate veins in the systematics of modern ferns

Taxon 1 87-95Webster GL (1994a) Classification of the Euphorbiaceae Annals of the

Missouri Botanical Garden 81 3-32

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 177

983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

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Figure 6 E hypericifolia (ESEM) papillae on the adaxial and abaxial surfacesmulticellular hairs are on the abaxial surface of the leaf only

Figure7 Multicellular hairs on adaxial and abaxial surfaces of E hirta

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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178 Int Res J Plant Sci

Figure 8 Hairs wart ornamentation on hair surfaces of (A) E scordiifolia and (B) E pirottae (C) Short hairs onabaxial surface of E balsamifera subsp adenensis

In E balsamifera subsp adenensis the hairs are shortand unicellular (Figure 8C) and they are located mostly atthe leaf margins and the mid-vein area on the abaxialsurface only

Hairs on the leaf of E pirottae are present on theabaxial surface only and are both unicellular andmulticellular Euphorbia granulata is similar to E pirottae in having both unicellular and multicellular hairs but herethey occur on both surfaces of the leaf (Figure 8B)

On the other hand in E scordiifolia E hirta and Ehypericifolia the hairs are multicellular and on bothsurfaces in the first two species and on the abaxial only inthe latter (Figure 7) Moreover E hypericifolia and Ehirta have long hairs when compared with the otherspecies (Figure 6)

No hairs were observed and leaves were glabrous inthe following species E serpens E arabica E peplusE inaequilatera E falcata E helioscopia E cuneata Eretusa E grossheimii E schimperiana E schimperiE chamaepeplus and the new variety of E granulata

Wax

The cuticle layer and wax in the Euphorbia species undeinvestigation vary from smooth to densely cover in waxscales This may be a thick smooth layer of wax whichsometimes has upright scales In some species only thecell surface has a smooth wax layer whereas the walbetween cells has upright scales of wax as in Eschimperiana especially around the stomata pores

(Figure 9) In contrast the anticlinal walls of somespecies have a smooth layer of wax whereas the celsurface is densely covered with upright wax flakes or withsome stellate wax flakes in other species

Stomata

Stomata were observed through LM and ESEM In thespecies under investigation stomata commonly appeaon both leaf surfaces (amphistomatic leaves) excepin E cuneata where the stomata are located only on the

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 179

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Figure 9 Wax formation in Euphorbia species in Saudi Arabia (A+B) smooth andthick in E peplus and E schimperi (C+D) smooth on top of cell while formingupright scale flakes in between especially around stomata as in E schimperiana smooth in anticlinal walls (E) upright scale flakes in E balsamifera

abaxial surface (hypostomatic leaves) The presence andposition of subsidiary cells are variable in Euphorbia

species Subsidiary cells were found to be of unequalsize and shape whether the stoma was anisocyticparacytic actinocytic or tetracytic While some specieslack them around the stomata others have more thanone arrangement of subsidiary cells

The most common stomata type is anomocytic whilethe rare type is actinocytic recorded only in Ehelioscopia Moreover sometimes stomata of more thanone type (up to three or four types) have beenencountered on the same leaf surface as in E scordiifolia (three types on the abaxial surface) and E hirta (five

types on the abaxial and three on the adaxial) Thedistribution and type of stomata are shown in Table 2

In addition in some species such as E falcata and Ehelioscopia the stomata are deeply sunken andrepresented only by cuticular ledges surrounding theminute pores as seen in ESEM

Examples of species lacking subsidiary cells aroundstomata are E peplus E chamaepeplus E granulataE dracunculoides and E acalyphoides Therefore theyall have anomocytic stomata

On the other hand E balsamifera subsp adenensishas the mononcytic type on both leaf surfaces andanisocytic and paracytic only on the abaxial surface In

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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180 Int Res J Plant Sci

Table 2 Stomata type in Euphorbia taxa in Saudi Arabia

Ab abaxial surface + stomata presentAd adaxial surface

addition E schimperiana has anomocytic andtetracytic on the abaxial only In contrast Ecuneata has three types of stomata anomocyticparacytic and anisocytic only on the lower surfaceAs well ldquoE aff consobrinardquo ldquoE aff schimperirdquo

and E schimperi have paracytic and anisocyticonly in the abaxial on the first two and in both leafsurfaces on the latter Therefore ldquoE affconsobrinardquo and ldquoE aff schimperirdquo are the same taxa Then comparing to E schimperi results

from ansubspec

Eupsimilar and an

Subgenus

Stomata

Anomocytic Hemiparacytic Paracytic Anisocytic T

Species Ad Ab Ad Ab Ad Ab Ad Ab AT i r u c al l i

E balsamifera subsp adenensis + + +

E schimperi + + + +

ldquoEsp aff consobrinardquo + +

ldquoE sp aff schimperirdquo + +

E cuneata + + +

ErmophytonE acalyphoides + +

E pirottae + + + +

E helioscopia + + E s ul a

E grossheimii + + + + +

E retusa + + + + +

E dracunculoides +

E falcate + + E peplus +

E chamaepeplus + +

E schimperiana + C h am a e s y c e

E Arabica + + + + + + +

E serpens + + + +

E scordiifolia + + + + + + +

E granulate + + + +

E inaequilatera + + + +

E hirta + + + + + + +

E hypericifolia + + + + + +

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Table3 Venation type in Euphorbia taxa in Saudi Arabia

Subgenus Species

Venation

Number of veins atthe leaf base

Margins Sheathed

1 2 3 4 or more Open Closed Rounded T i r u c al l i

Ebalsamiferasubspadenensis

+ + +

E schimperi + + +

E aff Schimperi + + +

E cuneata + + +

ErmophytonE acalyphoides + + +

E pirottae + + E s ul a

E grossheimii + +

E retusa + + +

E dracunculoides + + +

E falcate + + +

E peplus + +

E chamaepeplus + +

E schimperiana

E helioscopia

+

+

+

+ C h am a e s y c e

E Arabica + + +

E serpens + + +

E scordiifolia + + +

E granulate + + + +

E inaequilatera + + +

E hirta + + + E hypericifolia + + +

Poinsettia E cyathophora + +

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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182 Int Res J Plant Sci

Figure 10 Sunken stomata on the abaxial surface1) Deeply sunken (A) E helioscopia (B) E falcata (C) E schimperiana 2) Slightly sunken (D) E peplus (E) E pirottae

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 183

983105983138983137983160983145983137983148

983105

983108

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983106

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983112983141983149983145983152983137983154983137983139983161983156983145983139

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983112983141983149983145983152983137983154983137983139983161983156983145983139

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983112983141983149983145983152983137983154983137983139983161983156983145983139983120983137983154983137983139983161983156983145983139

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983105983139983156983145983150983151983139983161983156983145983139

983105983140983137983160983145983137983148

Figure 11 Stomata types in Euphorbia taxa in Saudi Arabia (A) E balsamifera subsp adenensis (B) E schimperi (C) E cuneata and (D) E grossheimii

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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184 Int Res J Plant Sci

983105

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983105983150983151983149983151983139983161983156983145983139983105983138983137983160

983145983137983148983112983141983149983145983152983137983154983137983139983161983156983145983139

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983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

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983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

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983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983138983137983160983145983137983148983105983140983137983160983145983137983148

Figure 12 Stomata types in Euphorbia taxa (A) E falcata (B) E peplus (C) E chamaepeplus and (D) Eschimperiana

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 185

983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

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983112983141983149983145983152983137983154983137983139983161983156983145983139

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983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105

983108

983106

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983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

Figure 13 Figure 13 Stomata types in Euphorbia taxa in Saudi Arabia (A) E arabica (B) E serpens (C)E scordiifolia and (D) E helioscopia

distribution between the leaf surfaces (see Figure 13)While E hirta has the most types E helioscopia Edracunculoides and E acalyphoides have the least

Regarding the position of the stomata on the surfacelevel of the epidermis a variable degree of sunkenesshas been recorded Using the ESEM the stomata appear

deeply sunken in some species whilst in others they areonly slightly sunken with the outer rims of the subsidiarycells surrounding the aperture The deeply and slightlysunken species are recorded in Table 2 AdditionallyFigure 10 shows the sunken stomata in different speciesunder the ESEM

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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186 Int Res J Plant Sci

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

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983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

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983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983107

983106

983105

983105983140983137983160983145983137983148983105983138983137983160983145983137983148

Figure 14 Stomata types in Euphorbia taxa in Saudi Arabia (A) E hirta (B) E hypericifolia (C) E pirottae and (D)Espaff schimperi

Venation

The venation patterns in the Euphorbia species underinvestigation vary from one-veined three-veined to thosewith four or more veins The predominant type is three-veined However in E balsamifera subsp Adenensis

there are seven middle parallel veins three middle thickand four thinner lateral ones (Figure 15 A and B)Whereas in E pirottae there are three main middleparallel veins and two thinner laterals (Figure 15 C) Incontrast E chamaepeplus E schimperiana and Earabica have only one thick main midrib coming from the

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 187

983105

983106

983107

Figure 15 Figure 15 Venation patterns at base of leaves invariants with more than three veins (A amp B) E balsamiferasubsp adenensis and (C) E pirottae

petiole then branched from the side into two lateral veinsat the point when the leaf starts widening (Figure 16)Each of these veins branches again into secondary veinsat the leaf margin

The rest of the species have just three veins Egrossheimii E dracunculoides E falcata E peplus Eserpens E scordiifolia E granulata E inaequilateraE helioscopia E hirta E hypericifolia and E

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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188 Int Res J Plant Sci

Figure 16 Venation patterns with one main mid vein at the leaf base in (A) E arabica and (B) E chamaepeplus

983106983105

Figure 17 Venation patterns with three-veined leaves in (A) E falcata and (B) E granulata

cyathophora All the three veins come from the petioleand start branching into secondary veins both toward themargin and the mid vein direction (Figure 17 B)

The vein endings at the leaf margin vary from closed

to open There is a relationship between the closedmargins and the sheathed veins (midrib and secondaryveins that are surrounded by parenchyma tissue) Whenthe veins are ornamented (sheathed) the margins areclosed This can be seen in subgenus Chamaesyce inspecies such as E serpens E scordiifolia E granulataE inaequilatera E hirta E hypericifolia and E arabica Additionally these species have the three-vein venation

On the other hand when the veins are not sheathedthe veins ends are open at the leaf margin as in Ebalsamifera subsp adenensis E schimperi E cuneataE acalyphoides E grossheimii E dracunculoides E

falcata E peplus E chamaepeplus E schimperiana Ehelioscopia E pirottae ldquoE aff schimperirdquo and Ecyathophora

CONCLUSIONS

Anatomical characters are seen here to be very useful inEuphorbia classification and identification The resultsobtained from anatomical studies were useful to groupsome species and identify others

Depending on the venation type the Euphorbiaspecies under investigation are divided anatomically intothree main groups see Figure 18

Group one veins are sheathed epidermal cellsdifferentiated on adaxial and abaxial surfaces Group two

983105 983106

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Figure 18 Hierarchial relation of Euphorbia taxa in Saudi Arabia based on anatomical charac

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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190 Int Res J Plant Sci

veins are not sheathed and epidermal cells are alike inboth surfaces And the third group includes species thatdo not belong to any of the previous groups The firstgroup can be further divided into three-veined and one-veined while the second group into three-veined and fouror more veins

In addition E grossheimii E retusa and E pirottae are related by being three-veined open at the marginswith rounded to oval tracheas and elongated cells bothon adaxial and abaxial surfaces However in E retusa the epidermal cells have straight cell walls on the adaxialsurface They are also similar in having hemiparacyticand paracytic types of stomata either on the adaxial orthe abaxial surface

On the other hand species with two differentepidermal cells on the leaf surfaces usually have three-sheathed veins with closed margins and elongatedtrachea such as E serpens E scordiifolia E granulata E inaequilatera E hirta and E hypericifolia which belongto subgenus Chamaesyce according to (Carter andSmith 1988)

In the case of E granulata which has beendescribed as having three different variants E granulata var granulata (both leaf surfaces hairy) E granulata varglabrata (adaxial glabrous and abaxial hairy) and aglabrous variant (the whole plant glabrous) Anatomicalwork has shown especially by using the ESEM that theleaves are either hairy or glabrous on both surfaces Sowhen the leaf is hairy both surfaces have differences inhair distribution Usually the lower surface is more hairythan upper surface (E granulata var glabrata ) butsometimes they are evenly distributed (Egranulata vargranulata ) In addition hairs on both surfaces are

multicellular In contrast when the leaf is glabrous bothleaf surfaces are glabrous and the whole plant isglabrous (suggested new variety) They were similar inhaving the same venation type (three-veined)

Finally ldquoE aff schimperi ldquohas some similarity with E schimperi in having the same open margin not theornamented venation type As well they both have thesame type of stomata paracytic and anisocytic but theydiffer in their distribution on abaxial and adaxial Thusthis may support the suggestion considering ldquoE affschimperi rdquo as a new subspecies of E schimperi

REFERENCES

Baranova M (1992) Principels of comparative stomatographic studiesof flowering plants The Botanical Review 58 49-99

Carlquist S (1961) Comparative Plant Anatomy Holt Rinehart ampWinston New York

Carter S Smith AR (1988) Euphorbiaceae (Part2) In Polhill ed Floraof Tropical East Africa Rotterdam Brookfield AA Balkema 409-597

Chaudhary SA (2001) Flora of The Kingdom of Saudi Arabia Vol 3Ministry of Agriculture and Water Riyadh

Dehay C (1935) Lagraveppareil Libero-ligneux foliare des EuphorbiaceaeAnnales des Sciences Naturelles Botanique 10 147-190

Dehgan B (1982) Comparative anatomy of the petiole and infragenericrelationships in Jatropha (Euphorbiaceae) American Journal oBotany 69 1283-1295

Gaucher L (1898) Ē tude anatomique du genre Euphorbia LPKlincksieck Paris

Gaucher L (1902) Recherches anatomiques sur les EuphorbiaceacuteesAnnales des Sciences Naturelles Botanique 15 161-309

Gaucher L (1902) Recherches anatomiques sur les Euphorbiaceacutees

Annales des Sciences Naturelles Botanique 15 161-309Govaerts R Fordin D Radcliffe-Smith A (2000) World Checklist and

Bibliography of Euphorbiaceae The Royal Botanic Gardens KewHickey LJ (1973) Classification of the architecture of dicotyledonous

leaves American Journal of Botany 60 17-33Hickey LJ (1979) A revised classification of the architecture o

dicotyledonous leaves In Metcalfe CR and Chalk L eds Anatomy oDicotyledons Systematic Anatomy of the leaf and stem 2nd edClarendon Press Oxford 25-39

Kakkar L Paliwal GS (1972) Epidermis in Euphorbia Indian ScienceCongress Association Proceedings 59 326-327

Kakkar L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 5 Epidermis Proceedings of the Indian National ScienceAcademy Part B Biological Sciences 40 55-67

Koumlhler E (1993) Blattvervatur-Muster der Buxaceae Dumortierr undSimmondsiaceae Van Tieghem Feddes Repertorium 104 145-167

Levin GA (1986a) Systematic foliar morphology of Phylanthoideae(Euphorbiaceae) II Phenetic analysis Annals of Missouri BotanicaGarden 73 86-98

Levin GA (1986b) Systematic foliar morphology of Phylanthoideae IICladistic analysis Systematic Botany 11 515-530

Levin GA (1986c) Systematic foliar morphology of Phyllanthoideae(Euphorbiaceae) I conspectus Annals of Missouri Botanical Garden73 29-85

Mahlberg P (1973) Histochemistry and Scanning Electron Microscopyof Starch Grains from Latex of Euphorbia-Terracina and EuphorbiaTirucalli Proceedings of the Indiana Academy of Science 82 132

Mahlberg PG (1974) Scanning Microscopic Comparison of StarchGrains from the Latex of Euphorbia -Spp Proceedings of the IndianaAcademy of Science 83 83-84

Mahlberg PG (1975) Evolution of the Laticifer in Euphorbia asInterpreted from Starch Grain Morphology American Journal oBotany 62 577-583

Mahlberg PG (1982) Comparative Morphology of Starch Grains inLatex from Varieties of Poinsettia Euphorbia-PulcherrimEuphorbiaceae Botanical Gazette 143 206-209

Mahlberg PG Davis DG Galitz DS Manners GD (1987) Laticifers andthe Classification of Euphorbia the Chemotaxonomy of EuphorbiaEsula L Botanical Journal of the Linnean Society 94 165-180

Metcalfe C Chalk L (1950) Anatomy of the Dicotyledons At theClarendon Press Oxford

Miller K Webster GL (1962) Systematic position of Cnidoscolus andJatropha Brittonia 14 174-180

Pax F (1884) Die Anatomie der Euphorbiaceen in ihrer Beziehung zumSystem derselben Botanische Jahrbuumlcher fuumlr SystematikPflanzengeschichte und Pflanzengeographie Leipzig 5 384-421

Payne WW (1970) Helicocytic and allelocytic stomata unrecognizedpatters in the Dicotyledonae Am J Bot57 140-147

Pray TR (1955a) Foliar venation of angiosperm II Histogenesis of thevenation of Liriodendron Am J Bot 42 18-27

Pray TR 1955b Foliar venation of angiosperms III Pattern andhistology of the venation of Hosta Am J Bot 42 11-18

Radford AE Dickison WC Massey JR Bell CR (1974) Vascular PlanSystematics Harper amp Row Publishers London New York

Radlkofer L (1870) Ueber Pausandra ein neues EuphorbiaceenGenus Flora 53 81-95

Raju VS Rao PN (1977) Variation in the structure and development ofoliar stomata in the Euphorbiaceae Botanical Journal of the LinneanSociety 75 69-97

Raju VS Rao PN (1987) The Taxonomic Use of the Basic StomataType in the Generic Delimitation of Chamaesyce EuphorbiaceaeFeddes Repertorium 98 137-142

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2424

Rosowski JR (1968) Laticifer Morphology in the Mature Stem and Leafof Euphorbia supina Botanical Gazette 129 113-120

Roth-Nebelsick A Uhl D Mosbrugger V Kerp H (2001) Evolution andfunction of leaf venation architecture a review Annals of Botany 87553-566

Sehgal L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 2 Venation Patterns Botanical Journal of the Linnean Society 68 173-208

Singh G (2004) Plant Systematics An Integrated Approach SciencePublishers Enfield

Solereder H (1899) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Solereder H (1908) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Aldhebiani and Jury 191

Stace CA (1965) Cuticular studies as an aid to plant taxonomy Bulletinof the British Museum Natural History Botany 4 3-78

Stace CA (1989) Plant Taxonomy and Biosystematics Hodder ampStoughton London

Tognini F (1897) Contribuzione allo studio della organogeniacomparata degli stomi Institut de Botanique University de Geneve 41-42

Van Cotthem WR (1970) A classification of stomatal types Botanica

Journal of Linnean Society 63 235-246Wagner W (1979) Reticulate veins in the systematics of modern ferns

Taxon 1 87-95Webster GL (1994a) Classification of the Euphorbiaceae Annals of the

Missouri Botanical Garden 81 3-32

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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178 Int Res J Plant Sci

Figure 8 Hairs wart ornamentation on hair surfaces of (A) E scordiifolia and (B) E pirottae (C) Short hairs onabaxial surface of E balsamifera subsp adenensis

In E balsamifera subsp adenensis the hairs are shortand unicellular (Figure 8C) and they are located mostly atthe leaf margins and the mid-vein area on the abaxialsurface only

Hairs on the leaf of E pirottae are present on theabaxial surface only and are both unicellular andmulticellular Euphorbia granulata is similar to E pirottae in having both unicellular and multicellular hairs but herethey occur on both surfaces of the leaf (Figure 8B)

On the other hand in E scordiifolia E hirta and Ehypericifolia the hairs are multicellular and on bothsurfaces in the first two species and on the abaxial only inthe latter (Figure 7) Moreover E hypericifolia and Ehirta have long hairs when compared with the otherspecies (Figure 6)

No hairs were observed and leaves were glabrous inthe following species E serpens E arabica E peplusE inaequilatera E falcata E helioscopia E cuneata Eretusa E grossheimii E schimperiana E schimperiE chamaepeplus and the new variety of E granulata

Wax

The cuticle layer and wax in the Euphorbia species undeinvestigation vary from smooth to densely cover in waxscales This may be a thick smooth layer of wax whichsometimes has upright scales In some species only thecell surface has a smooth wax layer whereas the walbetween cells has upright scales of wax as in Eschimperiana especially around the stomata pores

(Figure 9) In contrast the anticlinal walls of somespecies have a smooth layer of wax whereas the celsurface is densely covered with upright wax flakes or withsome stellate wax flakes in other species

Stomata

Stomata were observed through LM and ESEM In thespecies under investigation stomata commonly appeaon both leaf surfaces (amphistomatic leaves) excepin E cuneata where the stomata are located only on the

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 179

983109

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Figure 9 Wax formation in Euphorbia species in Saudi Arabia (A+B) smooth andthick in E peplus and E schimperi (C+D) smooth on top of cell while formingupright scale flakes in between especially around stomata as in E schimperiana smooth in anticlinal walls (E) upright scale flakes in E balsamifera

abaxial surface (hypostomatic leaves) The presence andposition of subsidiary cells are variable in Euphorbia

species Subsidiary cells were found to be of unequalsize and shape whether the stoma was anisocyticparacytic actinocytic or tetracytic While some specieslack them around the stomata others have more thanone arrangement of subsidiary cells

The most common stomata type is anomocytic whilethe rare type is actinocytic recorded only in Ehelioscopia Moreover sometimes stomata of more thanone type (up to three or four types) have beenencountered on the same leaf surface as in E scordiifolia (three types on the abaxial surface) and E hirta (five

types on the abaxial and three on the adaxial) Thedistribution and type of stomata are shown in Table 2

In addition in some species such as E falcata and Ehelioscopia the stomata are deeply sunken andrepresented only by cuticular ledges surrounding theminute pores as seen in ESEM

Examples of species lacking subsidiary cells aroundstomata are E peplus E chamaepeplus E granulataE dracunculoides and E acalyphoides Therefore theyall have anomocytic stomata

On the other hand E balsamifera subsp adenensishas the mononcytic type on both leaf surfaces andanisocytic and paracytic only on the abaxial surface In

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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180 Int Res J Plant Sci

Table 2 Stomata type in Euphorbia taxa in Saudi Arabia

Ab abaxial surface + stomata presentAd adaxial surface

addition E schimperiana has anomocytic andtetracytic on the abaxial only In contrast Ecuneata has three types of stomata anomocyticparacytic and anisocytic only on the lower surfaceAs well ldquoE aff consobrinardquo ldquoE aff schimperirdquo

and E schimperi have paracytic and anisocyticonly in the abaxial on the first two and in both leafsurfaces on the latter Therefore ldquoE affconsobrinardquo and ldquoE aff schimperirdquo are the same taxa Then comparing to E schimperi results

from ansubspec

Eupsimilar and an

Subgenus

Stomata

Anomocytic Hemiparacytic Paracytic Anisocytic T

Species Ad Ab Ad Ab Ad Ab Ad Ab AT i r u c al l i

E balsamifera subsp adenensis + + +

E schimperi + + + +

ldquoEsp aff consobrinardquo + +

ldquoE sp aff schimperirdquo + +

E cuneata + + +

ErmophytonE acalyphoides + +

E pirottae + + + +

E helioscopia + + E s ul a

E grossheimii + + + + +

E retusa + + + + +

E dracunculoides +

E falcate + + E peplus +

E chamaepeplus + +

E schimperiana + C h am a e s y c e

E Arabica + + + + + + +

E serpens + + + +

E scordiifolia + + + + + + +

E granulate + + + +

E inaequilatera + + + +

E hirta + + + + + + +

E hypericifolia + + + + + +

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Table3 Venation type in Euphorbia taxa in Saudi Arabia

Subgenus Species

Venation

Number of veins atthe leaf base

Margins Sheathed

1 2 3 4 or more Open Closed Rounded T i r u c al l i

Ebalsamiferasubspadenensis

+ + +

E schimperi + + +

E aff Schimperi + + +

E cuneata + + +

ErmophytonE acalyphoides + + +

E pirottae + + E s ul a

E grossheimii + +

E retusa + + +

E dracunculoides + + +

E falcate + + +

E peplus + +

E chamaepeplus + +

E schimperiana

E helioscopia

+

+

+

+ C h am a e s y c e

E Arabica + + +

E serpens + + +

E scordiifolia + + +

E granulate + + + +

E inaequilatera + + +

E hirta + + + E hypericifolia + + +

Poinsettia E cyathophora + +

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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182 Int Res J Plant Sci

Figure 10 Sunken stomata on the abaxial surface1) Deeply sunken (A) E helioscopia (B) E falcata (C) E schimperiana 2) Slightly sunken (D) E peplus (E) E pirottae

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 183

983105983138983137983160983145983137983148

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983112983141983149983145983152983137983154983137983139983161983156983145983139

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983112983141983149983145983152983137983154983137983139983161983156983145983139983120983137983154983137983139983161983156983145983139

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983105983139983156983145983150983151983139983161983156983145983139

983105983140983137983160983145983137983148

Figure 11 Stomata types in Euphorbia taxa in Saudi Arabia (A) E balsamifera subsp adenensis (B) E schimperi (C) E cuneata and (D) E grossheimii

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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184 Int Res J Plant Sci

983105

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983105983150983151983149983151983139983161983156983145983139983105983138983137983160

983145983137983148983112983141983149983145983152983137983154983137983139983161983156983145983139

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983112983141983149983145983152983137983154983137983139983161983156983145983139

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983124983141983156983154983137983139983161983156983145983139

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983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983138983137983160983145983137983148983105983140983137983160983145983137983148

Figure 12 Stomata types in Euphorbia taxa (A) E falcata (B) E peplus (C) E chamaepeplus and (D) Eschimperiana

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 185

983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

983105983150983151983149983151983139983161983156983145983139

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983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

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983106

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983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

Figure 13 Figure 13 Stomata types in Euphorbia taxa in Saudi Arabia (A) E arabica (B) E serpens (C)E scordiifolia and (D) E helioscopia

distribution between the leaf surfaces (see Figure 13)While E hirta has the most types E helioscopia Edracunculoides and E acalyphoides have the least

Regarding the position of the stomata on the surfacelevel of the epidermis a variable degree of sunkenesshas been recorded Using the ESEM the stomata appear

deeply sunken in some species whilst in others they areonly slightly sunken with the outer rims of the subsidiarycells surrounding the aperture The deeply and slightlysunken species are recorded in Table 2 AdditionallyFigure 10 shows the sunken stomata in different speciesunder the ESEM

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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186 Int Res J Plant Sci

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

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983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

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983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983107

983106

983105

983105983140983137983160983145983137983148983105983138983137983160983145983137983148

Figure 14 Stomata types in Euphorbia taxa in Saudi Arabia (A) E hirta (B) E hypericifolia (C) E pirottae and (D)Espaff schimperi

Venation

The venation patterns in the Euphorbia species underinvestigation vary from one-veined three-veined to thosewith four or more veins The predominant type is three-veined However in E balsamifera subsp Adenensis

there are seven middle parallel veins three middle thickand four thinner lateral ones (Figure 15 A and B)Whereas in E pirottae there are three main middleparallel veins and two thinner laterals (Figure 15 C) Incontrast E chamaepeplus E schimperiana and Earabica have only one thick main midrib coming from the

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 187

983105

983106

983107

Figure 15 Figure 15 Venation patterns at base of leaves invariants with more than three veins (A amp B) E balsamiferasubsp adenensis and (C) E pirottae

petiole then branched from the side into two lateral veinsat the point when the leaf starts widening (Figure 16)Each of these veins branches again into secondary veinsat the leaf margin

The rest of the species have just three veins Egrossheimii E dracunculoides E falcata E peplus Eserpens E scordiifolia E granulata E inaequilateraE helioscopia E hirta E hypericifolia and E

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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188 Int Res J Plant Sci

Figure 16 Venation patterns with one main mid vein at the leaf base in (A) E arabica and (B) E chamaepeplus

983106983105

Figure 17 Venation patterns with three-veined leaves in (A) E falcata and (B) E granulata

cyathophora All the three veins come from the petioleand start branching into secondary veins both toward themargin and the mid vein direction (Figure 17 B)

The vein endings at the leaf margin vary from closed

to open There is a relationship between the closedmargins and the sheathed veins (midrib and secondaryveins that are surrounded by parenchyma tissue) Whenthe veins are ornamented (sheathed) the margins areclosed This can be seen in subgenus Chamaesyce inspecies such as E serpens E scordiifolia E granulataE inaequilatera E hirta E hypericifolia and E arabica Additionally these species have the three-vein venation

On the other hand when the veins are not sheathedthe veins ends are open at the leaf margin as in Ebalsamifera subsp adenensis E schimperi E cuneataE acalyphoides E grossheimii E dracunculoides E

falcata E peplus E chamaepeplus E schimperiana Ehelioscopia E pirottae ldquoE aff schimperirdquo and Ecyathophora

CONCLUSIONS

Anatomical characters are seen here to be very useful inEuphorbia classification and identification The resultsobtained from anatomical studies were useful to groupsome species and identify others

Depending on the venation type the Euphorbiaspecies under investigation are divided anatomically intothree main groups see Figure 18

Group one veins are sheathed epidermal cellsdifferentiated on adaxial and abaxial surfaces Group two

983105 983106

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Figure 18 Hierarchial relation of Euphorbia taxa in Saudi Arabia based on anatomical charac

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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190 Int Res J Plant Sci

veins are not sheathed and epidermal cells are alike inboth surfaces And the third group includes species thatdo not belong to any of the previous groups The firstgroup can be further divided into three-veined and one-veined while the second group into three-veined and fouror more veins

In addition E grossheimii E retusa and E pirottae are related by being three-veined open at the marginswith rounded to oval tracheas and elongated cells bothon adaxial and abaxial surfaces However in E retusa the epidermal cells have straight cell walls on the adaxialsurface They are also similar in having hemiparacyticand paracytic types of stomata either on the adaxial orthe abaxial surface

On the other hand species with two differentepidermal cells on the leaf surfaces usually have three-sheathed veins with closed margins and elongatedtrachea such as E serpens E scordiifolia E granulata E inaequilatera E hirta and E hypericifolia which belongto subgenus Chamaesyce according to (Carter andSmith 1988)

In the case of E granulata which has beendescribed as having three different variants E granulata var granulata (both leaf surfaces hairy) E granulata varglabrata (adaxial glabrous and abaxial hairy) and aglabrous variant (the whole plant glabrous) Anatomicalwork has shown especially by using the ESEM that theleaves are either hairy or glabrous on both surfaces Sowhen the leaf is hairy both surfaces have differences inhair distribution Usually the lower surface is more hairythan upper surface (E granulata var glabrata ) butsometimes they are evenly distributed (Egranulata vargranulata ) In addition hairs on both surfaces are

multicellular In contrast when the leaf is glabrous bothleaf surfaces are glabrous and the whole plant isglabrous (suggested new variety) They were similar inhaving the same venation type (three-veined)

Finally ldquoE aff schimperi ldquohas some similarity with E schimperi in having the same open margin not theornamented venation type As well they both have thesame type of stomata paracytic and anisocytic but theydiffer in their distribution on abaxial and adaxial Thusthis may support the suggestion considering ldquoE affschimperi rdquo as a new subspecies of E schimperi

REFERENCES

Baranova M (1992) Principels of comparative stomatographic studiesof flowering plants The Botanical Review 58 49-99

Carlquist S (1961) Comparative Plant Anatomy Holt Rinehart ampWinston New York

Carter S Smith AR (1988) Euphorbiaceae (Part2) In Polhill ed Floraof Tropical East Africa Rotterdam Brookfield AA Balkema 409-597

Chaudhary SA (2001) Flora of The Kingdom of Saudi Arabia Vol 3Ministry of Agriculture and Water Riyadh

Dehay C (1935) Lagraveppareil Libero-ligneux foliare des EuphorbiaceaeAnnales des Sciences Naturelles Botanique 10 147-190

Dehgan B (1982) Comparative anatomy of the petiole and infragenericrelationships in Jatropha (Euphorbiaceae) American Journal oBotany 69 1283-1295

Gaucher L (1898) Ē tude anatomique du genre Euphorbia LPKlincksieck Paris

Gaucher L (1902) Recherches anatomiques sur les EuphorbiaceacuteesAnnales des Sciences Naturelles Botanique 15 161-309

Gaucher L (1902) Recherches anatomiques sur les Euphorbiaceacutees

Annales des Sciences Naturelles Botanique 15 161-309Govaerts R Fordin D Radcliffe-Smith A (2000) World Checklist and

Bibliography of Euphorbiaceae The Royal Botanic Gardens KewHickey LJ (1973) Classification of the architecture of dicotyledonous

leaves American Journal of Botany 60 17-33Hickey LJ (1979) A revised classification of the architecture o

dicotyledonous leaves In Metcalfe CR and Chalk L eds Anatomy oDicotyledons Systematic Anatomy of the leaf and stem 2nd edClarendon Press Oxford 25-39

Kakkar L Paliwal GS (1972) Epidermis in Euphorbia Indian ScienceCongress Association Proceedings 59 326-327

Kakkar L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 5 Epidermis Proceedings of the Indian National ScienceAcademy Part B Biological Sciences 40 55-67

Koumlhler E (1993) Blattvervatur-Muster der Buxaceae Dumortierr undSimmondsiaceae Van Tieghem Feddes Repertorium 104 145-167

Levin GA (1986a) Systematic foliar morphology of Phylanthoideae(Euphorbiaceae) II Phenetic analysis Annals of Missouri BotanicaGarden 73 86-98

Levin GA (1986b) Systematic foliar morphology of Phylanthoideae IICladistic analysis Systematic Botany 11 515-530

Levin GA (1986c) Systematic foliar morphology of Phyllanthoideae(Euphorbiaceae) I conspectus Annals of Missouri Botanical Garden73 29-85

Mahlberg P (1973) Histochemistry and Scanning Electron Microscopyof Starch Grains from Latex of Euphorbia-Terracina and EuphorbiaTirucalli Proceedings of the Indiana Academy of Science 82 132

Mahlberg PG (1974) Scanning Microscopic Comparison of StarchGrains from the Latex of Euphorbia -Spp Proceedings of the IndianaAcademy of Science 83 83-84

Mahlberg PG (1975) Evolution of the Laticifer in Euphorbia asInterpreted from Starch Grain Morphology American Journal oBotany 62 577-583

Mahlberg PG (1982) Comparative Morphology of Starch Grains inLatex from Varieties of Poinsettia Euphorbia-PulcherrimEuphorbiaceae Botanical Gazette 143 206-209

Mahlberg PG Davis DG Galitz DS Manners GD (1987) Laticifers andthe Classification of Euphorbia the Chemotaxonomy of EuphorbiaEsula L Botanical Journal of the Linnean Society 94 165-180

Metcalfe C Chalk L (1950) Anatomy of the Dicotyledons At theClarendon Press Oxford

Miller K Webster GL (1962) Systematic position of Cnidoscolus andJatropha Brittonia 14 174-180

Pax F (1884) Die Anatomie der Euphorbiaceen in ihrer Beziehung zumSystem derselben Botanische Jahrbuumlcher fuumlr SystematikPflanzengeschichte und Pflanzengeographie Leipzig 5 384-421

Payne WW (1970) Helicocytic and allelocytic stomata unrecognizedpatters in the Dicotyledonae Am J Bot57 140-147

Pray TR (1955a) Foliar venation of angiosperm II Histogenesis of thevenation of Liriodendron Am J Bot 42 18-27

Pray TR 1955b Foliar venation of angiosperms III Pattern andhistology of the venation of Hosta Am J Bot 42 11-18

Radford AE Dickison WC Massey JR Bell CR (1974) Vascular PlanSystematics Harper amp Row Publishers London New York

Radlkofer L (1870) Ueber Pausandra ein neues EuphorbiaceenGenus Flora 53 81-95

Raju VS Rao PN (1977) Variation in the structure and development ofoliar stomata in the Euphorbiaceae Botanical Journal of the LinneanSociety 75 69-97

Raju VS Rao PN (1987) The Taxonomic Use of the Basic StomataType in the Generic Delimitation of Chamaesyce EuphorbiaceaeFeddes Repertorium 98 137-142

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2424

Rosowski JR (1968) Laticifer Morphology in the Mature Stem and Leafof Euphorbia supina Botanical Gazette 129 113-120

Roth-Nebelsick A Uhl D Mosbrugger V Kerp H (2001) Evolution andfunction of leaf venation architecture a review Annals of Botany 87553-566

Sehgal L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 2 Venation Patterns Botanical Journal of the Linnean Society 68 173-208

Singh G (2004) Plant Systematics An Integrated Approach SciencePublishers Enfield

Solereder H (1899) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Solereder H (1908) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Aldhebiani and Jury 191

Stace CA (1965) Cuticular studies as an aid to plant taxonomy Bulletinof the British Museum Natural History Botany 4 3-78

Stace CA (1989) Plant Taxonomy and Biosystematics Hodder ampStoughton London

Tognini F (1897) Contribuzione allo studio della organogeniacomparata degli stomi Institut de Botanique University de Geneve 41-42

Van Cotthem WR (1970) A classification of stomatal types Botanica

Journal of Linnean Society 63 235-246Wagner W (1979) Reticulate veins in the systematics of modern ferns

Taxon 1 87-95Webster GL (1994a) Classification of the Euphorbiaceae Annals of the

Missouri Botanical Garden 81 3-32

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 179

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Figure 9 Wax formation in Euphorbia species in Saudi Arabia (A+B) smooth andthick in E peplus and E schimperi (C+D) smooth on top of cell while formingupright scale flakes in between especially around stomata as in E schimperiana smooth in anticlinal walls (E) upright scale flakes in E balsamifera

abaxial surface (hypostomatic leaves) The presence andposition of subsidiary cells are variable in Euphorbia

species Subsidiary cells were found to be of unequalsize and shape whether the stoma was anisocyticparacytic actinocytic or tetracytic While some specieslack them around the stomata others have more thanone arrangement of subsidiary cells

The most common stomata type is anomocytic whilethe rare type is actinocytic recorded only in Ehelioscopia Moreover sometimes stomata of more thanone type (up to three or four types) have beenencountered on the same leaf surface as in E scordiifolia (three types on the abaxial surface) and E hirta (five

types on the abaxial and three on the adaxial) Thedistribution and type of stomata are shown in Table 2

In addition in some species such as E falcata and Ehelioscopia the stomata are deeply sunken andrepresented only by cuticular ledges surrounding theminute pores as seen in ESEM

Examples of species lacking subsidiary cells aroundstomata are E peplus E chamaepeplus E granulataE dracunculoides and E acalyphoides Therefore theyall have anomocytic stomata

On the other hand E balsamifera subsp adenensishas the mononcytic type on both leaf surfaces andanisocytic and paracytic only on the abaxial surface In

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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180 Int Res J Plant Sci

Table 2 Stomata type in Euphorbia taxa in Saudi Arabia

Ab abaxial surface + stomata presentAd adaxial surface

addition E schimperiana has anomocytic andtetracytic on the abaxial only In contrast Ecuneata has three types of stomata anomocyticparacytic and anisocytic only on the lower surfaceAs well ldquoE aff consobrinardquo ldquoE aff schimperirdquo

and E schimperi have paracytic and anisocyticonly in the abaxial on the first two and in both leafsurfaces on the latter Therefore ldquoE affconsobrinardquo and ldquoE aff schimperirdquo are the same taxa Then comparing to E schimperi results

from ansubspec

Eupsimilar and an

Subgenus

Stomata

Anomocytic Hemiparacytic Paracytic Anisocytic T

Species Ad Ab Ad Ab Ad Ab Ad Ab AT i r u c al l i

E balsamifera subsp adenensis + + +

E schimperi + + + +

ldquoEsp aff consobrinardquo + +

ldquoE sp aff schimperirdquo + +

E cuneata + + +

ErmophytonE acalyphoides + +

E pirottae + + + +

E helioscopia + + E s ul a

E grossheimii + + + + +

E retusa + + + + +

E dracunculoides +

E falcate + + E peplus +

E chamaepeplus + +

E schimperiana + C h am a e s y c e

E Arabica + + + + + + +

E serpens + + + +

E scordiifolia + + + + + + +

E granulate + + + +

E inaequilatera + + + +

E hirta + + + + + + +

E hypericifolia + + + + + +

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Table3 Venation type in Euphorbia taxa in Saudi Arabia

Subgenus Species

Venation

Number of veins atthe leaf base

Margins Sheathed

1 2 3 4 or more Open Closed Rounded T i r u c al l i

Ebalsamiferasubspadenensis

+ + +

E schimperi + + +

E aff Schimperi + + +

E cuneata + + +

ErmophytonE acalyphoides + + +

E pirottae + + E s ul a

E grossheimii + +

E retusa + + +

E dracunculoides + + +

E falcate + + +

E peplus + +

E chamaepeplus + +

E schimperiana

E helioscopia

+

+

+

+ C h am a e s y c e

E Arabica + + +

E serpens + + +

E scordiifolia + + +

E granulate + + + +

E inaequilatera + + +

E hirta + + + E hypericifolia + + +

Poinsettia E cyathophora + +

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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182 Int Res J Plant Sci

Figure 10 Sunken stomata on the abaxial surface1) Deeply sunken (A) E helioscopia (B) E falcata (C) E schimperiana 2) Slightly sunken (D) E peplus (E) E pirottae

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 183

983105983138983137983160983145983137983148

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983105983139983156983145983150983151983139983161983156983145983139

983105983140983137983160983145983137983148

Figure 11 Stomata types in Euphorbia taxa in Saudi Arabia (A) E balsamifera subsp adenensis (B) E schimperi (C) E cuneata and (D) E grossheimii

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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184 Int Res J Plant Sci

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983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139983105983150983145983155983151983139983161983156983145983139

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983105983139983156983145983150983151983139983161983156983145983139

983105983138983137983160983145983137983148983105983140983137983160983145983137983148

Figure 12 Stomata types in Euphorbia taxa (A) E falcata (B) E peplus (C) E chamaepeplus and (D) Eschimperiana

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 185

983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

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983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

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983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

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983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

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983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

Figure 13 Figure 13 Stomata types in Euphorbia taxa in Saudi Arabia (A) E arabica (B) E serpens (C)E scordiifolia and (D) E helioscopia

distribution between the leaf surfaces (see Figure 13)While E hirta has the most types E helioscopia Edracunculoides and E acalyphoides have the least

Regarding the position of the stomata on the surfacelevel of the epidermis a variable degree of sunkenesshas been recorded Using the ESEM the stomata appear

deeply sunken in some species whilst in others they areonly slightly sunken with the outer rims of the subsidiarycells surrounding the aperture The deeply and slightlysunken species are recorded in Table 2 AdditionallyFigure 10 shows the sunken stomata in different speciesunder the ESEM

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186 Int Res J Plant Sci

983105983150983151983149983151983139983161983156983145983139

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983124983141983156983154983137983139983161983156983145983139

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983112983141983149983145983152983137983154983137983139983161983156983145983139

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983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983107

983106

983105

983105983140983137983160983145983137983148983105983138983137983160983145983137983148

Figure 14 Stomata types in Euphorbia taxa in Saudi Arabia (A) E hirta (B) E hypericifolia (C) E pirottae and (D)Espaff schimperi

Venation

The venation patterns in the Euphorbia species underinvestigation vary from one-veined three-veined to thosewith four or more veins The predominant type is three-veined However in E balsamifera subsp Adenensis

there are seven middle parallel veins three middle thickand four thinner lateral ones (Figure 15 A and B)Whereas in E pirottae there are three main middleparallel veins and two thinner laterals (Figure 15 C) Incontrast E chamaepeplus E schimperiana and Earabica have only one thick main midrib coming from the

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 187

983105

983106

983107

Figure 15 Figure 15 Venation patterns at base of leaves invariants with more than three veins (A amp B) E balsamiferasubsp adenensis and (C) E pirottae

petiole then branched from the side into two lateral veinsat the point when the leaf starts widening (Figure 16)Each of these veins branches again into secondary veinsat the leaf margin

The rest of the species have just three veins Egrossheimii E dracunculoides E falcata E peplus Eserpens E scordiifolia E granulata E inaequilateraE helioscopia E hirta E hypericifolia and E

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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188 Int Res J Plant Sci

Figure 16 Venation patterns with one main mid vein at the leaf base in (A) E arabica and (B) E chamaepeplus

983106983105

Figure 17 Venation patterns with three-veined leaves in (A) E falcata and (B) E granulata

cyathophora All the three veins come from the petioleand start branching into secondary veins both toward themargin and the mid vein direction (Figure 17 B)

The vein endings at the leaf margin vary from closed

to open There is a relationship between the closedmargins and the sheathed veins (midrib and secondaryveins that are surrounded by parenchyma tissue) Whenthe veins are ornamented (sheathed) the margins areclosed This can be seen in subgenus Chamaesyce inspecies such as E serpens E scordiifolia E granulataE inaequilatera E hirta E hypericifolia and E arabica Additionally these species have the three-vein venation

On the other hand when the veins are not sheathedthe veins ends are open at the leaf margin as in Ebalsamifera subsp adenensis E schimperi E cuneataE acalyphoides E grossheimii E dracunculoides E

falcata E peplus E chamaepeplus E schimperiana Ehelioscopia E pirottae ldquoE aff schimperirdquo and Ecyathophora

CONCLUSIONS

Anatomical characters are seen here to be very useful inEuphorbia classification and identification The resultsobtained from anatomical studies were useful to groupsome species and identify others

Depending on the venation type the Euphorbiaspecies under investigation are divided anatomically intothree main groups see Figure 18

Group one veins are sheathed epidermal cellsdifferentiated on adaxial and abaxial surfaces Group two

983105 983106

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Figure 18 Hierarchial relation of Euphorbia taxa in Saudi Arabia based on anatomical charac

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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190 Int Res J Plant Sci

veins are not sheathed and epidermal cells are alike inboth surfaces And the third group includes species thatdo not belong to any of the previous groups The firstgroup can be further divided into three-veined and one-veined while the second group into three-veined and fouror more veins

In addition E grossheimii E retusa and E pirottae are related by being three-veined open at the marginswith rounded to oval tracheas and elongated cells bothon adaxial and abaxial surfaces However in E retusa the epidermal cells have straight cell walls on the adaxialsurface They are also similar in having hemiparacyticand paracytic types of stomata either on the adaxial orthe abaxial surface

On the other hand species with two differentepidermal cells on the leaf surfaces usually have three-sheathed veins with closed margins and elongatedtrachea such as E serpens E scordiifolia E granulata E inaequilatera E hirta and E hypericifolia which belongto subgenus Chamaesyce according to (Carter andSmith 1988)

In the case of E granulata which has beendescribed as having three different variants E granulata var granulata (both leaf surfaces hairy) E granulata varglabrata (adaxial glabrous and abaxial hairy) and aglabrous variant (the whole plant glabrous) Anatomicalwork has shown especially by using the ESEM that theleaves are either hairy or glabrous on both surfaces Sowhen the leaf is hairy both surfaces have differences inhair distribution Usually the lower surface is more hairythan upper surface (E granulata var glabrata ) butsometimes they are evenly distributed (Egranulata vargranulata ) In addition hairs on both surfaces are

multicellular In contrast when the leaf is glabrous bothleaf surfaces are glabrous and the whole plant isglabrous (suggested new variety) They were similar inhaving the same venation type (three-veined)

Finally ldquoE aff schimperi ldquohas some similarity with E schimperi in having the same open margin not theornamented venation type As well they both have thesame type of stomata paracytic and anisocytic but theydiffer in their distribution on abaxial and adaxial Thusthis may support the suggestion considering ldquoE affschimperi rdquo as a new subspecies of E schimperi

REFERENCES

Baranova M (1992) Principels of comparative stomatographic studiesof flowering plants The Botanical Review 58 49-99

Carlquist S (1961) Comparative Plant Anatomy Holt Rinehart ampWinston New York

Carter S Smith AR (1988) Euphorbiaceae (Part2) In Polhill ed Floraof Tropical East Africa Rotterdam Brookfield AA Balkema 409-597

Chaudhary SA (2001) Flora of The Kingdom of Saudi Arabia Vol 3Ministry of Agriculture and Water Riyadh

Dehay C (1935) Lagraveppareil Libero-ligneux foliare des EuphorbiaceaeAnnales des Sciences Naturelles Botanique 10 147-190

Dehgan B (1982) Comparative anatomy of the petiole and infragenericrelationships in Jatropha (Euphorbiaceae) American Journal oBotany 69 1283-1295

Gaucher L (1898) Ē tude anatomique du genre Euphorbia LPKlincksieck Paris

Gaucher L (1902) Recherches anatomiques sur les EuphorbiaceacuteesAnnales des Sciences Naturelles Botanique 15 161-309

Gaucher L (1902) Recherches anatomiques sur les Euphorbiaceacutees

Annales des Sciences Naturelles Botanique 15 161-309Govaerts R Fordin D Radcliffe-Smith A (2000) World Checklist and

Bibliography of Euphorbiaceae The Royal Botanic Gardens KewHickey LJ (1973) Classification of the architecture of dicotyledonous

leaves American Journal of Botany 60 17-33Hickey LJ (1979) A revised classification of the architecture o

dicotyledonous leaves In Metcalfe CR and Chalk L eds Anatomy oDicotyledons Systematic Anatomy of the leaf and stem 2nd edClarendon Press Oxford 25-39

Kakkar L Paliwal GS (1972) Epidermis in Euphorbia Indian ScienceCongress Association Proceedings 59 326-327

Kakkar L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 5 Epidermis Proceedings of the Indian National ScienceAcademy Part B Biological Sciences 40 55-67

Koumlhler E (1993) Blattvervatur-Muster der Buxaceae Dumortierr undSimmondsiaceae Van Tieghem Feddes Repertorium 104 145-167

Levin GA (1986a) Systematic foliar morphology of Phylanthoideae(Euphorbiaceae) II Phenetic analysis Annals of Missouri BotanicaGarden 73 86-98

Levin GA (1986b) Systematic foliar morphology of Phylanthoideae IICladistic analysis Systematic Botany 11 515-530

Levin GA (1986c) Systematic foliar morphology of Phyllanthoideae(Euphorbiaceae) I conspectus Annals of Missouri Botanical Garden73 29-85

Mahlberg P (1973) Histochemistry and Scanning Electron Microscopyof Starch Grains from Latex of Euphorbia-Terracina and EuphorbiaTirucalli Proceedings of the Indiana Academy of Science 82 132

Mahlberg PG (1974) Scanning Microscopic Comparison of StarchGrains from the Latex of Euphorbia -Spp Proceedings of the IndianaAcademy of Science 83 83-84

Mahlberg PG (1975) Evolution of the Laticifer in Euphorbia asInterpreted from Starch Grain Morphology American Journal oBotany 62 577-583

Mahlberg PG (1982) Comparative Morphology of Starch Grains inLatex from Varieties of Poinsettia Euphorbia-PulcherrimEuphorbiaceae Botanical Gazette 143 206-209

Mahlberg PG Davis DG Galitz DS Manners GD (1987) Laticifers andthe Classification of Euphorbia the Chemotaxonomy of EuphorbiaEsula L Botanical Journal of the Linnean Society 94 165-180

Metcalfe C Chalk L (1950) Anatomy of the Dicotyledons At theClarendon Press Oxford

Miller K Webster GL (1962) Systematic position of Cnidoscolus andJatropha Brittonia 14 174-180

Pax F (1884) Die Anatomie der Euphorbiaceen in ihrer Beziehung zumSystem derselben Botanische Jahrbuumlcher fuumlr SystematikPflanzengeschichte und Pflanzengeographie Leipzig 5 384-421

Payne WW (1970) Helicocytic and allelocytic stomata unrecognizedpatters in the Dicotyledonae Am J Bot57 140-147

Pray TR (1955a) Foliar venation of angiosperm II Histogenesis of thevenation of Liriodendron Am J Bot 42 18-27

Pray TR 1955b Foliar venation of angiosperms III Pattern andhistology of the venation of Hosta Am J Bot 42 11-18

Radford AE Dickison WC Massey JR Bell CR (1974) Vascular PlanSystematics Harper amp Row Publishers London New York

Radlkofer L (1870) Ueber Pausandra ein neues EuphorbiaceenGenus Flora 53 81-95

Raju VS Rao PN (1977) Variation in the structure and development ofoliar stomata in the Euphorbiaceae Botanical Journal of the LinneanSociety 75 69-97

Raju VS Rao PN (1987) The Taxonomic Use of the Basic StomataType in the Generic Delimitation of Chamaesyce EuphorbiaceaeFeddes Repertorium 98 137-142

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2424

Rosowski JR (1968) Laticifer Morphology in the Mature Stem and Leafof Euphorbia supina Botanical Gazette 129 113-120

Roth-Nebelsick A Uhl D Mosbrugger V Kerp H (2001) Evolution andfunction of leaf venation architecture a review Annals of Botany 87553-566

Sehgal L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 2 Venation Patterns Botanical Journal of the Linnean Society 68 173-208

Singh G (2004) Plant Systematics An Integrated Approach SciencePublishers Enfield

Solereder H (1899) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Solereder H (1908) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Aldhebiani and Jury 191

Stace CA (1965) Cuticular studies as an aid to plant taxonomy Bulletinof the British Museum Natural History Botany 4 3-78

Stace CA (1989) Plant Taxonomy and Biosystematics Hodder ampStoughton London

Tognini F (1897) Contribuzione allo studio della organogeniacomparata degli stomi Institut de Botanique University de Geneve 41-42

Van Cotthem WR (1970) A classification of stomatal types Botanica

Journal of Linnean Society 63 235-246Wagner W (1979) Reticulate veins in the systematics of modern ferns

Taxon 1 87-95Webster GL (1994a) Classification of the Euphorbiaceae Annals of the

Missouri Botanical Garden 81 3-32

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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180 Int Res J Plant Sci

Table 2 Stomata type in Euphorbia taxa in Saudi Arabia

Ab abaxial surface + stomata presentAd adaxial surface

addition E schimperiana has anomocytic andtetracytic on the abaxial only In contrast Ecuneata has three types of stomata anomocyticparacytic and anisocytic only on the lower surfaceAs well ldquoE aff consobrinardquo ldquoE aff schimperirdquo

and E schimperi have paracytic and anisocyticonly in the abaxial on the first two and in both leafsurfaces on the latter Therefore ldquoE affconsobrinardquo and ldquoE aff schimperirdquo are the same taxa Then comparing to E schimperi results

from ansubspec

Eupsimilar and an

Subgenus

Stomata

Anomocytic Hemiparacytic Paracytic Anisocytic T

Species Ad Ab Ad Ab Ad Ab Ad Ab AT i r u c al l i

E balsamifera subsp adenensis + + +

E schimperi + + + +

ldquoEsp aff consobrinardquo + +

ldquoE sp aff schimperirdquo + +

E cuneata + + +

ErmophytonE acalyphoides + +

E pirottae + + + +

E helioscopia + + E s ul a

E grossheimii + + + + +

E retusa + + + + +

E dracunculoides +

E falcate + + E peplus +

E chamaepeplus + +

E schimperiana + C h am a e s y c e

E Arabica + + + + + + +

E serpens + + + +

E scordiifolia + + + + + + +

E granulate + + + +

E inaequilatera + + + +

E hirta + + + + + + +

E hypericifolia + + + + + +

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Table3 Venation type in Euphorbia taxa in Saudi Arabia

Subgenus Species

Venation

Number of veins atthe leaf base

Margins Sheathed

1 2 3 4 or more Open Closed Rounded T i r u c al l i

Ebalsamiferasubspadenensis

+ + +

E schimperi + + +

E aff Schimperi + + +

E cuneata + + +

ErmophytonE acalyphoides + + +

E pirottae + + E s ul a

E grossheimii + +

E retusa + + +

E dracunculoides + + +

E falcate + + +

E peplus + +

E chamaepeplus + +

E schimperiana

E helioscopia

+

+

+

+ C h am a e s y c e

E Arabica + + +

E serpens + + +

E scordiifolia + + +

E granulate + + + +

E inaequilatera + + +

E hirta + + + E hypericifolia + + +

Poinsettia E cyathophora + +

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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182 Int Res J Plant Sci

Figure 10 Sunken stomata on the abaxial surface1) Deeply sunken (A) E helioscopia (B) E falcata (C) E schimperiana 2) Slightly sunken (D) E peplus (E) E pirottae

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 183

983105983138983137983160983145983137983148

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983105983139983156983145983150983151983139983161983156983145983139

983105983140983137983160983145983137983148

Figure 11 Stomata types in Euphorbia taxa in Saudi Arabia (A) E balsamifera subsp adenensis (B) E schimperi (C) E cuneata and (D) E grossheimii

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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184 Int Res J Plant Sci

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983112983141983149983145983152983137983154983137983139983161983156983145983139

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983112983141983149983145983152983137983154983137983139983161983156983145983139

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983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

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983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983138983137983160983145983137983148983105983140983137983160983145983137983148

Figure 12 Stomata types in Euphorbia taxa (A) E falcata (B) E peplus (C) E chamaepeplus and (D) Eschimperiana

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 185

983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

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983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105

983108

983106

983107

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

Figure 13 Figure 13 Stomata types in Euphorbia taxa in Saudi Arabia (A) E arabica (B) E serpens (C)E scordiifolia and (D) E helioscopia

distribution between the leaf surfaces (see Figure 13)While E hirta has the most types E helioscopia Edracunculoides and E acalyphoides have the least

Regarding the position of the stomata on the surfacelevel of the epidermis a variable degree of sunkenesshas been recorded Using the ESEM the stomata appear

deeply sunken in some species whilst in others they areonly slightly sunken with the outer rims of the subsidiarycells surrounding the aperture The deeply and slightlysunken species are recorded in Table 2 AdditionallyFigure 10 shows the sunken stomata in different speciesunder the ESEM

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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186 Int Res J Plant Sci

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

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983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

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983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

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983124983141983156983154983137983139983161983156983145983139

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983112983141983149983145983152983137983154983137983139983161983156983145983139

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983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983107

983106

983105

983105983140983137983160983145983137983148983105983138983137983160983145983137983148

Figure 14 Stomata types in Euphorbia taxa in Saudi Arabia (A) E hirta (B) E hypericifolia (C) E pirottae and (D)Espaff schimperi

Venation

The venation patterns in the Euphorbia species underinvestigation vary from one-veined three-veined to thosewith four or more veins The predominant type is three-veined However in E balsamifera subsp Adenensis

there are seven middle parallel veins three middle thickand four thinner lateral ones (Figure 15 A and B)Whereas in E pirottae there are three main middleparallel veins and two thinner laterals (Figure 15 C) Incontrast E chamaepeplus E schimperiana and Earabica have only one thick main midrib coming from the

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 187

983105

983106

983107

Figure 15 Figure 15 Venation patterns at base of leaves invariants with more than three veins (A amp B) E balsamiferasubsp adenensis and (C) E pirottae

petiole then branched from the side into two lateral veinsat the point when the leaf starts widening (Figure 16)Each of these veins branches again into secondary veinsat the leaf margin

The rest of the species have just three veins Egrossheimii E dracunculoides E falcata E peplus Eserpens E scordiifolia E granulata E inaequilateraE helioscopia E hirta E hypericifolia and E

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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188 Int Res J Plant Sci

Figure 16 Venation patterns with one main mid vein at the leaf base in (A) E arabica and (B) E chamaepeplus

983106983105

Figure 17 Venation patterns with three-veined leaves in (A) E falcata and (B) E granulata

cyathophora All the three veins come from the petioleand start branching into secondary veins both toward themargin and the mid vein direction (Figure 17 B)

The vein endings at the leaf margin vary from closed

to open There is a relationship between the closedmargins and the sheathed veins (midrib and secondaryveins that are surrounded by parenchyma tissue) Whenthe veins are ornamented (sheathed) the margins areclosed This can be seen in subgenus Chamaesyce inspecies such as E serpens E scordiifolia E granulataE inaequilatera E hirta E hypericifolia and E arabica Additionally these species have the three-vein venation

On the other hand when the veins are not sheathedthe veins ends are open at the leaf margin as in Ebalsamifera subsp adenensis E schimperi E cuneataE acalyphoides E grossheimii E dracunculoides E

falcata E peplus E chamaepeplus E schimperiana Ehelioscopia E pirottae ldquoE aff schimperirdquo and Ecyathophora

CONCLUSIONS

Anatomical characters are seen here to be very useful inEuphorbia classification and identification The resultsobtained from anatomical studies were useful to groupsome species and identify others

Depending on the venation type the Euphorbiaspecies under investigation are divided anatomically intothree main groups see Figure 18

Group one veins are sheathed epidermal cellsdifferentiated on adaxial and abaxial surfaces Group two

983105 983106

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Figure 18 Hierarchial relation of Euphorbia taxa in Saudi Arabia based on anatomical charac

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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190 Int Res J Plant Sci

veins are not sheathed and epidermal cells are alike inboth surfaces And the third group includes species thatdo not belong to any of the previous groups The firstgroup can be further divided into three-veined and one-veined while the second group into three-veined and fouror more veins

In addition E grossheimii E retusa and E pirottae are related by being three-veined open at the marginswith rounded to oval tracheas and elongated cells bothon adaxial and abaxial surfaces However in E retusa the epidermal cells have straight cell walls on the adaxialsurface They are also similar in having hemiparacyticand paracytic types of stomata either on the adaxial orthe abaxial surface

On the other hand species with two differentepidermal cells on the leaf surfaces usually have three-sheathed veins with closed margins and elongatedtrachea such as E serpens E scordiifolia E granulata E inaequilatera E hirta and E hypericifolia which belongto subgenus Chamaesyce according to (Carter andSmith 1988)

In the case of E granulata which has beendescribed as having three different variants E granulata var granulata (both leaf surfaces hairy) E granulata varglabrata (adaxial glabrous and abaxial hairy) and aglabrous variant (the whole plant glabrous) Anatomicalwork has shown especially by using the ESEM that theleaves are either hairy or glabrous on both surfaces Sowhen the leaf is hairy both surfaces have differences inhair distribution Usually the lower surface is more hairythan upper surface (E granulata var glabrata ) butsometimes they are evenly distributed (Egranulata vargranulata ) In addition hairs on both surfaces are

multicellular In contrast when the leaf is glabrous bothleaf surfaces are glabrous and the whole plant isglabrous (suggested new variety) They were similar inhaving the same venation type (three-veined)

Finally ldquoE aff schimperi ldquohas some similarity with E schimperi in having the same open margin not theornamented venation type As well they both have thesame type of stomata paracytic and anisocytic but theydiffer in their distribution on abaxial and adaxial Thusthis may support the suggestion considering ldquoE affschimperi rdquo as a new subspecies of E schimperi

REFERENCES

Baranova M (1992) Principels of comparative stomatographic studiesof flowering plants The Botanical Review 58 49-99

Carlquist S (1961) Comparative Plant Anatomy Holt Rinehart ampWinston New York

Carter S Smith AR (1988) Euphorbiaceae (Part2) In Polhill ed Floraof Tropical East Africa Rotterdam Brookfield AA Balkema 409-597

Chaudhary SA (2001) Flora of The Kingdom of Saudi Arabia Vol 3Ministry of Agriculture and Water Riyadh

Dehay C (1935) Lagraveppareil Libero-ligneux foliare des EuphorbiaceaeAnnales des Sciences Naturelles Botanique 10 147-190

Dehgan B (1982) Comparative anatomy of the petiole and infragenericrelationships in Jatropha (Euphorbiaceae) American Journal oBotany 69 1283-1295

Gaucher L (1898) Ē tude anatomique du genre Euphorbia LPKlincksieck Paris

Gaucher L (1902) Recherches anatomiques sur les EuphorbiaceacuteesAnnales des Sciences Naturelles Botanique 15 161-309

Gaucher L (1902) Recherches anatomiques sur les Euphorbiaceacutees

Annales des Sciences Naturelles Botanique 15 161-309Govaerts R Fordin D Radcliffe-Smith A (2000) World Checklist and

Bibliography of Euphorbiaceae The Royal Botanic Gardens KewHickey LJ (1973) Classification of the architecture of dicotyledonous

leaves American Journal of Botany 60 17-33Hickey LJ (1979) A revised classification of the architecture o

dicotyledonous leaves In Metcalfe CR and Chalk L eds Anatomy oDicotyledons Systematic Anatomy of the leaf and stem 2nd edClarendon Press Oxford 25-39

Kakkar L Paliwal GS (1972) Epidermis in Euphorbia Indian ScienceCongress Association Proceedings 59 326-327

Kakkar L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 5 Epidermis Proceedings of the Indian National ScienceAcademy Part B Biological Sciences 40 55-67

Koumlhler E (1993) Blattvervatur-Muster der Buxaceae Dumortierr undSimmondsiaceae Van Tieghem Feddes Repertorium 104 145-167

Levin GA (1986a) Systematic foliar morphology of Phylanthoideae(Euphorbiaceae) II Phenetic analysis Annals of Missouri BotanicaGarden 73 86-98

Levin GA (1986b) Systematic foliar morphology of Phylanthoideae IICladistic analysis Systematic Botany 11 515-530

Levin GA (1986c) Systematic foliar morphology of Phyllanthoideae(Euphorbiaceae) I conspectus Annals of Missouri Botanical Garden73 29-85

Mahlberg P (1973) Histochemistry and Scanning Electron Microscopyof Starch Grains from Latex of Euphorbia-Terracina and EuphorbiaTirucalli Proceedings of the Indiana Academy of Science 82 132

Mahlberg PG (1974) Scanning Microscopic Comparison of StarchGrains from the Latex of Euphorbia -Spp Proceedings of the IndianaAcademy of Science 83 83-84

Mahlberg PG (1975) Evolution of the Laticifer in Euphorbia asInterpreted from Starch Grain Morphology American Journal oBotany 62 577-583

Mahlberg PG (1982) Comparative Morphology of Starch Grains inLatex from Varieties of Poinsettia Euphorbia-PulcherrimEuphorbiaceae Botanical Gazette 143 206-209

Mahlberg PG Davis DG Galitz DS Manners GD (1987) Laticifers andthe Classification of Euphorbia the Chemotaxonomy of EuphorbiaEsula L Botanical Journal of the Linnean Society 94 165-180

Metcalfe C Chalk L (1950) Anatomy of the Dicotyledons At theClarendon Press Oxford

Miller K Webster GL (1962) Systematic position of Cnidoscolus andJatropha Brittonia 14 174-180

Pax F (1884) Die Anatomie der Euphorbiaceen in ihrer Beziehung zumSystem derselben Botanische Jahrbuumlcher fuumlr SystematikPflanzengeschichte und Pflanzengeographie Leipzig 5 384-421

Payne WW (1970) Helicocytic and allelocytic stomata unrecognizedpatters in the Dicotyledonae Am J Bot57 140-147

Pray TR (1955a) Foliar venation of angiosperm II Histogenesis of thevenation of Liriodendron Am J Bot 42 18-27

Pray TR 1955b Foliar venation of angiosperms III Pattern andhistology of the venation of Hosta Am J Bot 42 11-18

Radford AE Dickison WC Massey JR Bell CR (1974) Vascular PlanSystematics Harper amp Row Publishers London New York

Radlkofer L (1870) Ueber Pausandra ein neues EuphorbiaceenGenus Flora 53 81-95

Raju VS Rao PN (1977) Variation in the structure and development ofoliar stomata in the Euphorbiaceae Botanical Journal of the LinneanSociety 75 69-97

Raju VS Rao PN (1987) The Taxonomic Use of the Basic StomataType in the Generic Delimitation of Chamaesyce EuphorbiaceaeFeddes Repertorium 98 137-142

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2424

Rosowski JR (1968) Laticifer Morphology in the Mature Stem and Leafof Euphorbia supina Botanical Gazette 129 113-120

Roth-Nebelsick A Uhl D Mosbrugger V Kerp H (2001) Evolution andfunction of leaf venation architecture a review Annals of Botany 87553-566

Sehgal L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 2 Venation Patterns Botanical Journal of the Linnean Society 68 173-208

Singh G (2004) Plant Systematics An Integrated Approach SciencePublishers Enfield

Solereder H (1899) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Solereder H (1908) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Aldhebiani and Jury 191

Stace CA (1965) Cuticular studies as an aid to plant taxonomy Bulletinof the British Museum Natural History Botany 4 3-78

Stace CA (1989) Plant Taxonomy and Biosystematics Hodder ampStoughton London

Tognini F (1897) Contribuzione allo studio della organogeniacomparata degli stomi Institut de Botanique University de Geneve 41-42

Van Cotthem WR (1970) A classification of stomatal types Botanica

Journal of Linnean Society 63 235-246Wagner W (1979) Reticulate veins in the systematics of modern ferns

Taxon 1 87-95Webster GL (1994a) Classification of the Euphorbiaceae Annals of the

Missouri Botanical Garden 81 3-32

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Table3 Venation type in Euphorbia taxa in Saudi Arabia

Subgenus Species

Venation

Number of veins atthe leaf base

Margins Sheathed

1 2 3 4 or more Open Closed Rounded T i r u c al l i

Ebalsamiferasubspadenensis

+ + +

E schimperi + + +

E aff Schimperi + + +

E cuneata + + +

ErmophytonE acalyphoides + + +

E pirottae + + E s ul a

E grossheimii + +

E retusa + + +

E dracunculoides + + +

E falcate + + +

E peplus + +

E chamaepeplus + +

E schimperiana

E helioscopia

+

+

+

+ C h am a e s y c e

E Arabica + + +

E serpens + + +

E scordiifolia + + +

E granulate + + + +

E inaequilatera + + +

E hirta + + + E hypericifolia + + +

Poinsettia E cyathophora + +

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182 Int Res J Plant Sci

Figure 10 Sunken stomata on the abaxial surface1) Deeply sunken (A) E helioscopia (B) E falcata (C) E schimperiana 2) Slightly sunken (D) E peplus (E) E pirottae

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 183

983105983138983137983160983145983137983148

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983105983140983137983160983145983137983148

Figure 11 Stomata types in Euphorbia taxa in Saudi Arabia (A) E balsamifera subsp adenensis (B) E schimperi (C) E cuneata and (D) E grossheimii

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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184 Int Res J Plant Sci

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983105983138983137983160983145983137983148983105983140983137983160983145983137983148

Figure 12 Stomata types in Euphorbia taxa (A) E falcata (B) E peplus (C) E chamaepeplus and (D) Eschimperiana

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 185

983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

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Figure 13 Figure 13 Stomata types in Euphorbia taxa in Saudi Arabia (A) E arabica (B) E serpens (C)E scordiifolia and (D) E helioscopia

distribution between the leaf surfaces (see Figure 13)While E hirta has the most types E helioscopia Edracunculoides and E acalyphoides have the least

Regarding the position of the stomata on the surfacelevel of the epidermis a variable degree of sunkenesshas been recorded Using the ESEM the stomata appear

deeply sunken in some species whilst in others they areonly slightly sunken with the outer rims of the subsidiarycells surrounding the aperture The deeply and slightlysunken species are recorded in Table 2 AdditionallyFigure 10 shows the sunken stomata in different speciesunder the ESEM

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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186 Int Res J Plant Sci

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983107

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983105983140983137983160983145983137983148983105983138983137983160983145983137983148

Figure 14 Stomata types in Euphorbia taxa in Saudi Arabia (A) E hirta (B) E hypericifolia (C) E pirottae and (D)Espaff schimperi

Venation

The venation patterns in the Euphorbia species underinvestigation vary from one-veined three-veined to thosewith four or more veins The predominant type is three-veined However in E balsamifera subsp Adenensis

there are seven middle parallel veins three middle thickand four thinner lateral ones (Figure 15 A and B)Whereas in E pirottae there are three main middleparallel veins and two thinner laterals (Figure 15 C) Incontrast E chamaepeplus E schimperiana and Earabica have only one thick main midrib coming from the

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 187

983105

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Figure 15 Figure 15 Venation patterns at base of leaves invariants with more than three veins (A amp B) E balsamiferasubsp adenensis and (C) E pirottae

petiole then branched from the side into two lateral veinsat the point when the leaf starts widening (Figure 16)Each of these veins branches again into secondary veinsat the leaf margin

The rest of the species have just three veins Egrossheimii E dracunculoides E falcata E peplus Eserpens E scordiifolia E granulata E inaequilateraE helioscopia E hirta E hypericifolia and E

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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188 Int Res J Plant Sci

Figure 16 Venation patterns with one main mid vein at the leaf base in (A) E arabica and (B) E chamaepeplus

983106983105

Figure 17 Venation patterns with three-veined leaves in (A) E falcata and (B) E granulata

cyathophora All the three veins come from the petioleand start branching into secondary veins both toward themargin and the mid vein direction (Figure 17 B)

The vein endings at the leaf margin vary from closed

to open There is a relationship between the closedmargins and the sheathed veins (midrib and secondaryveins that are surrounded by parenchyma tissue) Whenthe veins are ornamented (sheathed) the margins areclosed This can be seen in subgenus Chamaesyce inspecies such as E serpens E scordiifolia E granulataE inaequilatera E hirta E hypericifolia and E arabica Additionally these species have the three-vein venation

On the other hand when the veins are not sheathedthe veins ends are open at the leaf margin as in Ebalsamifera subsp adenensis E schimperi E cuneataE acalyphoides E grossheimii E dracunculoides E

falcata E peplus E chamaepeplus E schimperiana Ehelioscopia E pirottae ldquoE aff schimperirdquo and Ecyathophora

CONCLUSIONS

Anatomical characters are seen here to be very useful inEuphorbia classification and identification The resultsobtained from anatomical studies were useful to groupsome species and identify others

Depending on the venation type the Euphorbiaspecies under investigation are divided anatomically intothree main groups see Figure 18

Group one veins are sheathed epidermal cellsdifferentiated on adaxial and abaxial surfaces Group two

983105 983106

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Figure 18 Hierarchial relation of Euphorbia taxa in Saudi Arabia based on anatomical charac

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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190 Int Res J Plant Sci

veins are not sheathed and epidermal cells are alike inboth surfaces And the third group includes species thatdo not belong to any of the previous groups The firstgroup can be further divided into three-veined and one-veined while the second group into three-veined and fouror more veins

In addition E grossheimii E retusa and E pirottae are related by being three-veined open at the marginswith rounded to oval tracheas and elongated cells bothon adaxial and abaxial surfaces However in E retusa the epidermal cells have straight cell walls on the adaxialsurface They are also similar in having hemiparacyticand paracytic types of stomata either on the adaxial orthe abaxial surface

On the other hand species with two differentepidermal cells on the leaf surfaces usually have three-sheathed veins with closed margins and elongatedtrachea such as E serpens E scordiifolia E granulata E inaequilatera E hirta and E hypericifolia which belongto subgenus Chamaesyce according to (Carter andSmith 1988)

In the case of E granulata which has beendescribed as having three different variants E granulata var granulata (both leaf surfaces hairy) E granulata varglabrata (adaxial glabrous and abaxial hairy) and aglabrous variant (the whole plant glabrous) Anatomicalwork has shown especially by using the ESEM that theleaves are either hairy or glabrous on both surfaces Sowhen the leaf is hairy both surfaces have differences inhair distribution Usually the lower surface is more hairythan upper surface (E granulata var glabrata ) butsometimes they are evenly distributed (Egranulata vargranulata ) In addition hairs on both surfaces are

multicellular In contrast when the leaf is glabrous bothleaf surfaces are glabrous and the whole plant isglabrous (suggested new variety) They were similar inhaving the same venation type (three-veined)

Finally ldquoE aff schimperi ldquohas some similarity with E schimperi in having the same open margin not theornamented venation type As well they both have thesame type of stomata paracytic and anisocytic but theydiffer in their distribution on abaxial and adaxial Thusthis may support the suggestion considering ldquoE affschimperi rdquo as a new subspecies of E schimperi

REFERENCES

Baranova M (1992) Principels of comparative stomatographic studiesof flowering plants The Botanical Review 58 49-99

Carlquist S (1961) Comparative Plant Anatomy Holt Rinehart ampWinston New York

Carter S Smith AR (1988) Euphorbiaceae (Part2) In Polhill ed Floraof Tropical East Africa Rotterdam Brookfield AA Balkema 409-597

Chaudhary SA (2001) Flora of The Kingdom of Saudi Arabia Vol 3Ministry of Agriculture and Water Riyadh

Dehay C (1935) Lagraveppareil Libero-ligneux foliare des EuphorbiaceaeAnnales des Sciences Naturelles Botanique 10 147-190

Dehgan B (1982) Comparative anatomy of the petiole and infragenericrelationships in Jatropha (Euphorbiaceae) American Journal oBotany 69 1283-1295

Gaucher L (1898) Ē tude anatomique du genre Euphorbia LPKlincksieck Paris

Gaucher L (1902) Recherches anatomiques sur les EuphorbiaceacuteesAnnales des Sciences Naturelles Botanique 15 161-309

Gaucher L (1902) Recherches anatomiques sur les Euphorbiaceacutees

Annales des Sciences Naturelles Botanique 15 161-309Govaerts R Fordin D Radcliffe-Smith A (2000) World Checklist and

Bibliography of Euphorbiaceae The Royal Botanic Gardens KewHickey LJ (1973) Classification of the architecture of dicotyledonous

leaves American Journal of Botany 60 17-33Hickey LJ (1979) A revised classification of the architecture o

dicotyledonous leaves In Metcalfe CR and Chalk L eds Anatomy oDicotyledons Systematic Anatomy of the leaf and stem 2nd edClarendon Press Oxford 25-39

Kakkar L Paliwal GS (1972) Epidermis in Euphorbia Indian ScienceCongress Association Proceedings 59 326-327

Kakkar L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 5 Epidermis Proceedings of the Indian National ScienceAcademy Part B Biological Sciences 40 55-67

Koumlhler E (1993) Blattvervatur-Muster der Buxaceae Dumortierr undSimmondsiaceae Van Tieghem Feddes Repertorium 104 145-167

Levin GA (1986a) Systematic foliar morphology of Phylanthoideae(Euphorbiaceae) II Phenetic analysis Annals of Missouri BotanicaGarden 73 86-98

Levin GA (1986b) Systematic foliar morphology of Phylanthoideae IICladistic analysis Systematic Botany 11 515-530

Levin GA (1986c) Systematic foliar morphology of Phyllanthoideae(Euphorbiaceae) I conspectus Annals of Missouri Botanical Garden73 29-85

Mahlberg P (1973) Histochemistry and Scanning Electron Microscopyof Starch Grains from Latex of Euphorbia-Terracina and EuphorbiaTirucalli Proceedings of the Indiana Academy of Science 82 132

Mahlberg PG (1974) Scanning Microscopic Comparison of StarchGrains from the Latex of Euphorbia -Spp Proceedings of the IndianaAcademy of Science 83 83-84

Mahlberg PG (1975) Evolution of the Laticifer in Euphorbia asInterpreted from Starch Grain Morphology American Journal oBotany 62 577-583

Mahlberg PG (1982) Comparative Morphology of Starch Grains inLatex from Varieties of Poinsettia Euphorbia-PulcherrimEuphorbiaceae Botanical Gazette 143 206-209

Mahlberg PG Davis DG Galitz DS Manners GD (1987) Laticifers andthe Classification of Euphorbia the Chemotaxonomy of EuphorbiaEsula L Botanical Journal of the Linnean Society 94 165-180

Metcalfe C Chalk L (1950) Anatomy of the Dicotyledons At theClarendon Press Oxford

Miller K Webster GL (1962) Systematic position of Cnidoscolus andJatropha Brittonia 14 174-180

Pax F (1884) Die Anatomie der Euphorbiaceen in ihrer Beziehung zumSystem derselben Botanische Jahrbuumlcher fuumlr SystematikPflanzengeschichte und Pflanzengeographie Leipzig 5 384-421

Payne WW (1970) Helicocytic and allelocytic stomata unrecognizedpatters in the Dicotyledonae Am J Bot57 140-147

Pray TR (1955a) Foliar venation of angiosperm II Histogenesis of thevenation of Liriodendron Am J Bot 42 18-27

Pray TR 1955b Foliar venation of angiosperms III Pattern andhistology of the venation of Hosta Am J Bot 42 11-18

Radford AE Dickison WC Massey JR Bell CR (1974) Vascular PlanSystematics Harper amp Row Publishers London New York

Radlkofer L (1870) Ueber Pausandra ein neues EuphorbiaceenGenus Flora 53 81-95

Raju VS Rao PN (1977) Variation in the structure and development ofoliar stomata in the Euphorbiaceae Botanical Journal of the LinneanSociety 75 69-97

Raju VS Rao PN (1987) The Taxonomic Use of the Basic StomataType in the Generic Delimitation of Chamaesyce EuphorbiaceaeFeddes Repertorium 98 137-142

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2424

Rosowski JR (1968) Laticifer Morphology in the Mature Stem and Leafof Euphorbia supina Botanical Gazette 129 113-120

Roth-Nebelsick A Uhl D Mosbrugger V Kerp H (2001) Evolution andfunction of leaf venation architecture a review Annals of Botany 87553-566

Sehgal L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 2 Venation Patterns Botanical Journal of the Linnean Society 68 173-208

Singh G (2004) Plant Systematics An Integrated Approach SciencePublishers Enfield

Solereder H (1899) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Solereder H (1908) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Aldhebiani and Jury 191

Stace CA (1965) Cuticular studies as an aid to plant taxonomy Bulletinof the British Museum Natural History Botany 4 3-78

Stace CA (1989) Plant Taxonomy and Biosystematics Hodder ampStoughton London

Tognini F (1897) Contribuzione allo studio della organogeniacomparata degli stomi Institut de Botanique University de Geneve 41-42

Van Cotthem WR (1970) A classification of stomatal types Botanica

Journal of Linnean Society 63 235-246Wagner W (1979) Reticulate veins in the systematics of modern ferns

Taxon 1 87-95Webster GL (1994a) Classification of the Euphorbiaceae Annals of the

Missouri Botanical Garden 81 3-32

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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182 Int Res J Plant Sci

Figure 10 Sunken stomata on the abaxial surface1) Deeply sunken (A) E helioscopia (B) E falcata (C) E schimperiana 2) Slightly sunken (D) E peplus (E) E pirottae

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 1624

Aldhebiani and Jury 183

983105983138983137983160983145983137983148

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983105983140983137983160983145983137983148

Figure 11 Stomata types in Euphorbia taxa in Saudi Arabia (A) E balsamifera subsp adenensis (B) E schimperi (C) E cuneata and (D) E grossheimii

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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184 Int Res J Plant Sci

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983105983138983137983160983145983137983148983105983140983137983160983145983137983148

Figure 12 Stomata types in Euphorbia taxa (A) E falcata (B) E peplus (C) E chamaepeplus and (D) Eschimperiana

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 185

983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

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Figure 13 Figure 13 Stomata types in Euphorbia taxa in Saudi Arabia (A) E arabica (B) E serpens (C)E scordiifolia and (D) E helioscopia

distribution between the leaf surfaces (see Figure 13)While E hirta has the most types E helioscopia Edracunculoides and E acalyphoides have the least

Regarding the position of the stomata on the surfacelevel of the epidermis a variable degree of sunkenesshas been recorded Using the ESEM the stomata appear

deeply sunken in some species whilst in others they areonly slightly sunken with the outer rims of the subsidiarycells surrounding the aperture The deeply and slightlysunken species are recorded in Table 2 AdditionallyFigure 10 shows the sunken stomata in different speciesunder the ESEM

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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186 Int Res J Plant Sci

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983105983140983137983160983145983137983148983105983138983137983160983145983137983148

Figure 14 Stomata types in Euphorbia taxa in Saudi Arabia (A) E hirta (B) E hypericifolia (C) E pirottae and (D)Espaff schimperi

Venation

The venation patterns in the Euphorbia species underinvestigation vary from one-veined three-veined to thosewith four or more veins The predominant type is three-veined However in E balsamifera subsp Adenensis

there are seven middle parallel veins three middle thickand four thinner lateral ones (Figure 15 A and B)Whereas in E pirottae there are three main middleparallel veins and two thinner laterals (Figure 15 C) Incontrast E chamaepeplus E schimperiana and Earabica have only one thick main midrib coming from the

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 187

983105

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Figure 15 Figure 15 Venation patterns at base of leaves invariants with more than three veins (A amp B) E balsamiferasubsp adenensis and (C) E pirottae

petiole then branched from the side into two lateral veinsat the point when the leaf starts widening (Figure 16)Each of these veins branches again into secondary veinsat the leaf margin

The rest of the species have just three veins Egrossheimii E dracunculoides E falcata E peplus Eserpens E scordiifolia E granulata E inaequilateraE helioscopia E hirta E hypericifolia and E

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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188 Int Res J Plant Sci

Figure 16 Venation patterns with one main mid vein at the leaf base in (A) E arabica and (B) E chamaepeplus

983106983105

Figure 17 Venation patterns with three-veined leaves in (A) E falcata and (B) E granulata

cyathophora All the three veins come from the petioleand start branching into secondary veins both toward themargin and the mid vein direction (Figure 17 B)

The vein endings at the leaf margin vary from closed

to open There is a relationship between the closedmargins and the sheathed veins (midrib and secondaryveins that are surrounded by parenchyma tissue) Whenthe veins are ornamented (sheathed) the margins areclosed This can be seen in subgenus Chamaesyce inspecies such as E serpens E scordiifolia E granulataE inaequilatera E hirta E hypericifolia and E arabica Additionally these species have the three-vein venation

On the other hand when the veins are not sheathedthe veins ends are open at the leaf margin as in Ebalsamifera subsp adenensis E schimperi E cuneataE acalyphoides E grossheimii E dracunculoides E

falcata E peplus E chamaepeplus E schimperiana Ehelioscopia E pirottae ldquoE aff schimperirdquo and Ecyathophora

CONCLUSIONS

Anatomical characters are seen here to be very useful inEuphorbia classification and identification The resultsobtained from anatomical studies were useful to groupsome species and identify others

Depending on the venation type the Euphorbiaspecies under investigation are divided anatomically intothree main groups see Figure 18

Group one veins are sheathed epidermal cellsdifferentiated on adaxial and abaxial surfaces Group two

983105 983106

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Figure 18 Hierarchial relation of Euphorbia taxa in Saudi Arabia based on anatomical charac

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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190 Int Res J Plant Sci

veins are not sheathed and epidermal cells are alike inboth surfaces And the third group includes species thatdo not belong to any of the previous groups The firstgroup can be further divided into three-veined and one-veined while the second group into three-veined and fouror more veins

In addition E grossheimii E retusa and E pirottae are related by being three-veined open at the marginswith rounded to oval tracheas and elongated cells bothon adaxial and abaxial surfaces However in E retusa the epidermal cells have straight cell walls on the adaxialsurface They are also similar in having hemiparacyticand paracytic types of stomata either on the adaxial orthe abaxial surface

On the other hand species with two differentepidermal cells on the leaf surfaces usually have three-sheathed veins with closed margins and elongatedtrachea such as E serpens E scordiifolia E granulata E inaequilatera E hirta and E hypericifolia which belongto subgenus Chamaesyce according to (Carter andSmith 1988)

In the case of E granulata which has beendescribed as having three different variants E granulata var granulata (both leaf surfaces hairy) E granulata varglabrata (adaxial glabrous and abaxial hairy) and aglabrous variant (the whole plant glabrous) Anatomicalwork has shown especially by using the ESEM that theleaves are either hairy or glabrous on both surfaces Sowhen the leaf is hairy both surfaces have differences inhair distribution Usually the lower surface is more hairythan upper surface (E granulata var glabrata ) butsometimes they are evenly distributed (Egranulata vargranulata ) In addition hairs on both surfaces are

multicellular In contrast when the leaf is glabrous bothleaf surfaces are glabrous and the whole plant isglabrous (suggested new variety) They were similar inhaving the same venation type (three-veined)

Finally ldquoE aff schimperi ldquohas some similarity with E schimperi in having the same open margin not theornamented venation type As well they both have thesame type of stomata paracytic and anisocytic but theydiffer in their distribution on abaxial and adaxial Thusthis may support the suggestion considering ldquoE affschimperi rdquo as a new subspecies of E schimperi

REFERENCES

Baranova M (1992) Principels of comparative stomatographic studiesof flowering plants The Botanical Review 58 49-99

Carlquist S (1961) Comparative Plant Anatomy Holt Rinehart ampWinston New York

Carter S Smith AR (1988) Euphorbiaceae (Part2) In Polhill ed Floraof Tropical East Africa Rotterdam Brookfield AA Balkema 409-597

Chaudhary SA (2001) Flora of The Kingdom of Saudi Arabia Vol 3Ministry of Agriculture and Water Riyadh

Dehay C (1935) Lagraveppareil Libero-ligneux foliare des EuphorbiaceaeAnnales des Sciences Naturelles Botanique 10 147-190

Dehgan B (1982) Comparative anatomy of the petiole and infragenericrelationships in Jatropha (Euphorbiaceae) American Journal oBotany 69 1283-1295

Gaucher L (1898) Ē tude anatomique du genre Euphorbia LPKlincksieck Paris

Gaucher L (1902) Recherches anatomiques sur les EuphorbiaceacuteesAnnales des Sciences Naturelles Botanique 15 161-309

Gaucher L (1902) Recherches anatomiques sur les Euphorbiaceacutees

Annales des Sciences Naturelles Botanique 15 161-309Govaerts R Fordin D Radcliffe-Smith A (2000) World Checklist and

Bibliography of Euphorbiaceae The Royal Botanic Gardens KewHickey LJ (1973) Classification of the architecture of dicotyledonous

leaves American Journal of Botany 60 17-33Hickey LJ (1979) A revised classification of the architecture o

dicotyledonous leaves In Metcalfe CR and Chalk L eds Anatomy oDicotyledons Systematic Anatomy of the leaf and stem 2nd edClarendon Press Oxford 25-39

Kakkar L Paliwal GS (1972) Epidermis in Euphorbia Indian ScienceCongress Association Proceedings 59 326-327

Kakkar L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 5 Epidermis Proceedings of the Indian National ScienceAcademy Part B Biological Sciences 40 55-67

Koumlhler E (1993) Blattvervatur-Muster der Buxaceae Dumortierr undSimmondsiaceae Van Tieghem Feddes Repertorium 104 145-167

Levin GA (1986a) Systematic foliar morphology of Phylanthoideae(Euphorbiaceae) II Phenetic analysis Annals of Missouri BotanicaGarden 73 86-98

Levin GA (1986b) Systematic foliar morphology of Phylanthoideae IICladistic analysis Systematic Botany 11 515-530

Levin GA (1986c) Systematic foliar morphology of Phyllanthoideae(Euphorbiaceae) I conspectus Annals of Missouri Botanical Garden73 29-85

Mahlberg P (1973) Histochemistry and Scanning Electron Microscopyof Starch Grains from Latex of Euphorbia-Terracina and EuphorbiaTirucalli Proceedings of the Indiana Academy of Science 82 132

Mahlberg PG (1974) Scanning Microscopic Comparison of StarchGrains from the Latex of Euphorbia -Spp Proceedings of the IndianaAcademy of Science 83 83-84

Mahlberg PG (1975) Evolution of the Laticifer in Euphorbia asInterpreted from Starch Grain Morphology American Journal oBotany 62 577-583

Mahlberg PG (1982) Comparative Morphology of Starch Grains inLatex from Varieties of Poinsettia Euphorbia-PulcherrimEuphorbiaceae Botanical Gazette 143 206-209

Mahlberg PG Davis DG Galitz DS Manners GD (1987) Laticifers andthe Classification of Euphorbia the Chemotaxonomy of EuphorbiaEsula L Botanical Journal of the Linnean Society 94 165-180

Metcalfe C Chalk L (1950) Anatomy of the Dicotyledons At theClarendon Press Oxford

Miller K Webster GL (1962) Systematic position of Cnidoscolus andJatropha Brittonia 14 174-180

Pax F (1884) Die Anatomie der Euphorbiaceen in ihrer Beziehung zumSystem derselben Botanische Jahrbuumlcher fuumlr SystematikPflanzengeschichte und Pflanzengeographie Leipzig 5 384-421

Payne WW (1970) Helicocytic and allelocytic stomata unrecognizedpatters in the Dicotyledonae Am J Bot57 140-147

Pray TR (1955a) Foliar venation of angiosperm II Histogenesis of thevenation of Liriodendron Am J Bot 42 18-27

Pray TR 1955b Foliar venation of angiosperms III Pattern andhistology of the venation of Hosta Am J Bot 42 11-18

Radford AE Dickison WC Massey JR Bell CR (1974) Vascular PlanSystematics Harper amp Row Publishers London New York

Radlkofer L (1870) Ueber Pausandra ein neues EuphorbiaceenGenus Flora 53 81-95

Raju VS Rao PN (1977) Variation in the structure and development ofoliar stomata in the Euphorbiaceae Botanical Journal of the LinneanSociety 75 69-97

Raju VS Rao PN (1987) The Taxonomic Use of the Basic StomataType in the Generic Delimitation of Chamaesyce EuphorbiaceaeFeddes Repertorium 98 137-142

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2424

Rosowski JR (1968) Laticifer Morphology in the Mature Stem and Leafof Euphorbia supina Botanical Gazette 129 113-120

Roth-Nebelsick A Uhl D Mosbrugger V Kerp H (2001) Evolution andfunction of leaf venation architecture a review Annals of Botany 87553-566

Sehgal L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 2 Venation Patterns Botanical Journal of the Linnean Society 68 173-208

Singh G (2004) Plant Systematics An Integrated Approach SciencePublishers Enfield

Solereder H (1899) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Solereder H (1908) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Aldhebiani and Jury 191

Stace CA (1965) Cuticular studies as an aid to plant taxonomy Bulletinof the British Museum Natural History Botany 4 3-78

Stace CA (1989) Plant Taxonomy and Biosystematics Hodder ampStoughton London

Tognini F (1897) Contribuzione allo studio della organogeniacomparata degli stomi Institut de Botanique University de Geneve 41-42

Van Cotthem WR (1970) A classification of stomatal types Botanica

Journal of Linnean Society 63 235-246Wagner W (1979) Reticulate veins in the systematics of modern ferns

Taxon 1 87-95Webster GL (1994a) Classification of the Euphorbiaceae Annals of the

Missouri Botanical Garden 81 3-32

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Figure 11 Stomata types in Euphorbia taxa in Saudi Arabia (A) E balsamifera subsp adenensis (B) E schimperi (C) E cuneata and (D) E grossheimii

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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184 Int Res J Plant Sci

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Figure 12 Stomata types in Euphorbia taxa (A) E falcata (B) E peplus (C) E chamaepeplus and (D) Eschimperiana

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 185

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Figure 13 Figure 13 Stomata types in Euphorbia taxa in Saudi Arabia (A) E arabica (B) E serpens (C)E scordiifolia and (D) E helioscopia

distribution between the leaf surfaces (see Figure 13)While E hirta has the most types E helioscopia Edracunculoides and E acalyphoides have the least

Regarding the position of the stomata on the surfacelevel of the epidermis a variable degree of sunkenesshas been recorded Using the ESEM the stomata appear

deeply sunken in some species whilst in others they areonly slightly sunken with the outer rims of the subsidiarycells surrounding the aperture The deeply and slightlysunken species are recorded in Table 2 AdditionallyFigure 10 shows the sunken stomata in different speciesunder the ESEM

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Figure 14 Stomata types in Euphorbia taxa in Saudi Arabia (A) E hirta (B) E hypericifolia (C) E pirottae and (D)Espaff schimperi

Venation

The venation patterns in the Euphorbia species underinvestigation vary from one-veined three-veined to thosewith four or more veins The predominant type is three-veined However in E balsamifera subsp Adenensis

there are seven middle parallel veins three middle thickand four thinner lateral ones (Figure 15 A and B)Whereas in E pirottae there are three main middleparallel veins and two thinner laterals (Figure 15 C) Incontrast E chamaepeplus E schimperiana and Earabica have only one thick main midrib coming from the

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 187

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Figure 15 Figure 15 Venation patterns at base of leaves invariants with more than three veins (A amp B) E balsamiferasubsp adenensis and (C) E pirottae

petiole then branched from the side into two lateral veinsat the point when the leaf starts widening (Figure 16)Each of these veins branches again into secondary veinsat the leaf margin

The rest of the species have just three veins Egrossheimii E dracunculoides E falcata E peplus Eserpens E scordiifolia E granulata E inaequilateraE helioscopia E hirta E hypericifolia and E

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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188 Int Res J Plant Sci

Figure 16 Venation patterns with one main mid vein at the leaf base in (A) E arabica and (B) E chamaepeplus

983106983105

Figure 17 Venation patterns with three-veined leaves in (A) E falcata and (B) E granulata

cyathophora All the three veins come from the petioleand start branching into secondary veins both toward themargin and the mid vein direction (Figure 17 B)

The vein endings at the leaf margin vary from closed

to open There is a relationship between the closedmargins and the sheathed veins (midrib and secondaryveins that are surrounded by parenchyma tissue) Whenthe veins are ornamented (sheathed) the margins areclosed This can be seen in subgenus Chamaesyce inspecies such as E serpens E scordiifolia E granulataE inaequilatera E hirta E hypericifolia and E arabica Additionally these species have the three-vein venation

On the other hand when the veins are not sheathedthe veins ends are open at the leaf margin as in Ebalsamifera subsp adenensis E schimperi E cuneataE acalyphoides E grossheimii E dracunculoides E

falcata E peplus E chamaepeplus E schimperiana Ehelioscopia E pirottae ldquoE aff schimperirdquo and Ecyathophora

CONCLUSIONS

Anatomical characters are seen here to be very useful inEuphorbia classification and identification The resultsobtained from anatomical studies were useful to groupsome species and identify others

Depending on the venation type the Euphorbiaspecies under investigation are divided anatomically intothree main groups see Figure 18

Group one veins are sheathed epidermal cellsdifferentiated on adaxial and abaxial surfaces Group two

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7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Figure 18 Hierarchial relation of Euphorbia taxa in Saudi Arabia based on anatomical charac

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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190 Int Res J Plant Sci

veins are not sheathed and epidermal cells are alike inboth surfaces And the third group includes species thatdo not belong to any of the previous groups The firstgroup can be further divided into three-veined and one-veined while the second group into three-veined and fouror more veins

In addition E grossheimii E retusa and E pirottae are related by being three-veined open at the marginswith rounded to oval tracheas and elongated cells bothon adaxial and abaxial surfaces However in E retusa the epidermal cells have straight cell walls on the adaxialsurface They are also similar in having hemiparacyticand paracytic types of stomata either on the adaxial orthe abaxial surface

On the other hand species with two differentepidermal cells on the leaf surfaces usually have three-sheathed veins with closed margins and elongatedtrachea such as E serpens E scordiifolia E granulata E inaequilatera E hirta and E hypericifolia which belongto subgenus Chamaesyce according to (Carter andSmith 1988)

In the case of E granulata which has beendescribed as having three different variants E granulata var granulata (both leaf surfaces hairy) E granulata varglabrata (adaxial glabrous and abaxial hairy) and aglabrous variant (the whole plant glabrous) Anatomicalwork has shown especially by using the ESEM that theleaves are either hairy or glabrous on both surfaces Sowhen the leaf is hairy both surfaces have differences inhair distribution Usually the lower surface is more hairythan upper surface (E granulata var glabrata ) butsometimes they are evenly distributed (Egranulata vargranulata ) In addition hairs on both surfaces are

multicellular In contrast when the leaf is glabrous bothleaf surfaces are glabrous and the whole plant isglabrous (suggested new variety) They were similar inhaving the same venation type (three-veined)

Finally ldquoE aff schimperi ldquohas some similarity with E schimperi in having the same open margin not theornamented venation type As well they both have thesame type of stomata paracytic and anisocytic but theydiffer in their distribution on abaxial and adaxial Thusthis may support the suggestion considering ldquoE affschimperi rdquo as a new subspecies of E schimperi

REFERENCES

Baranova M (1992) Principels of comparative stomatographic studiesof flowering plants The Botanical Review 58 49-99

Carlquist S (1961) Comparative Plant Anatomy Holt Rinehart ampWinston New York

Carter S Smith AR (1988) Euphorbiaceae (Part2) In Polhill ed Floraof Tropical East Africa Rotterdam Brookfield AA Balkema 409-597

Chaudhary SA (2001) Flora of The Kingdom of Saudi Arabia Vol 3Ministry of Agriculture and Water Riyadh

Dehay C (1935) Lagraveppareil Libero-ligneux foliare des EuphorbiaceaeAnnales des Sciences Naturelles Botanique 10 147-190

Dehgan B (1982) Comparative anatomy of the petiole and infragenericrelationships in Jatropha (Euphorbiaceae) American Journal oBotany 69 1283-1295

Gaucher L (1898) Ē tude anatomique du genre Euphorbia LPKlincksieck Paris

Gaucher L (1902) Recherches anatomiques sur les EuphorbiaceacuteesAnnales des Sciences Naturelles Botanique 15 161-309

Gaucher L (1902) Recherches anatomiques sur les Euphorbiaceacutees

Annales des Sciences Naturelles Botanique 15 161-309Govaerts R Fordin D Radcliffe-Smith A (2000) World Checklist and

Bibliography of Euphorbiaceae The Royal Botanic Gardens KewHickey LJ (1973) Classification of the architecture of dicotyledonous

leaves American Journal of Botany 60 17-33Hickey LJ (1979) A revised classification of the architecture o

dicotyledonous leaves In Metcalfe CR and Chalk L eds Anatomy oDicotyledons Systematic Anatomy of the leaf and stem 2nd edClarendon Press Oxford 25-39

Kakkar L Paliwal GS (1972) Epidermis in Euphorbia Indian ScienceCongress Association Proceedings 59 326-327

Kakkar L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 5 Epidermis Proceedings of the Indian National ScienceAcademy Part B Biological Sciences 40 55-67

Koumlhler E (1993) Blattvervatur-Muster der Buxaceae Dumortierr undSimmondsiaceae Van Tieghem Feddes Repertorium 104 145-167

Levin GA (1986a) Systematic foliar morphology of Phylanthoideae(Euphorbiaceae) II Phenetic analysis Annals of Missouri BotanicaGarden 73 86-98

Levin GA (1986b) Systematic foliar morphology of Phylanthoideae IICladistic analysis Systematic Botany 11 515-530

Levin GA (1986c) Systematic foliar morphology of Phyllanthoideae(Euphorbiaceae) I conspectus Annals of Missouri Botanical Garden73 29-85

Mahlberg P (1973) Histochemistry and Scanning Electron Microscopyof Starch Grains from Latex of Euphorbia-Terracina and EuphorbiaTirucalli Proceedings of the Indiana Academy of Science 82 132

Mahlberg PG (1974) Scanning Microscopic Comparison of StarchGrains from the Latex of Euphorbia -Spp Proceedings of the IndianaAcademy of Science 83 83-84

Mahlberg PG (1975) Evolution of the Laticifer in Euphorbia asInterpreted from Starch Grain Morphology American Journal oBotany 62 577-583

Mahlberg PG (1982) Comparative Morphology of Starch Grains inLatex from Varieties of Poinsettia Euphorbia-PulcherrimEuphorbiaceae Botanical Gazette 143 206-209

Mahlberg PG Davis DG Galitz DS Manners GD (1987) Laticifers andthe Classification of Euphorbia the Chemotaxonomy of EuphorbiaEsula L Botanical Journal of the Linnean Society 94 165-180

Metcalfe C Chalk L (1950) Anatomy of the Dicotyledons At theClarendon Press Oxford

Miller K Webster GL (1962) Systematic position of Cnidoscolus andJatropha Brittonia 14 174-180

Pax F (1884) Die Anatomie der Euphorbiaceen in ihrer Beziehung zumSystem derselben Botanische Jahrbuumlcher fuumlr SystematikPflanzengeschichte und Pflanzengeographie Leipzig 5 384-421

Payne WW (1970) Helicocytic and allelocytic stomata unrecognizedpatters in the Dicotyledonae Am J Bot57 140-147

Pray TR (1955a) Foliar venation of angiosperm II Histogenesis of thevenation of Liriodendron Am J Bot 42 18-27

Pray TR 1955b Foliar venation of angiosperms III Pattern andhistology of the venation of Hosta Am J Bot 42 11-18

Radford AE Dickison WC Massey JR Bell CR (1974) Vascular PlanSystematics Harper amp Row Publishers London New York

Radlkofer L (1870) Ueber Pausandra ein neues EuphorbiaceenGenus Flora 53 81-95

Raju VS Rao PN (1977) Variation in the structure and development ofoliar stomata in the Euphorbiaceae Botanical Journal of the LinneanSociety 75 69-97

Raju VS Rao PN (1987) The Taxonomic Use of the Basic StomataType in the Generic Delimitation of Chamaesyce EuphorbiaceaeFeddes Repertorium 98 137-142

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2424

Rosowski JR (1968) Laticifer Morphology in the Mature Stem and Leafof Euphorbia supina Botanical Gazette 129 113-120

Roth-Nebelsick A Uhl D Mosbrugger V Kerp H (2001) Evolution andfunction of leaf venation architecture a review Annals of Botany 87553-566

Sehgal L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 2 Venation Patterns Botanical Journal of the Linnean Society 68 173-208

Singh G (2004) Plant Systematics An Integrated Approach SciencePublishers Enfield

Solereder H (1899) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Solereder H (1908) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Aldhebiani and Jury 191

Stace CA (1965) Cuticular studies as an aid to plant taxonomy Bulletinof the British Museum Natural History Botany 4 3-78

Stace CA (1989) Plant Taxonomy and Biosystematics Hodder ampStoughton London

Tognini F (1897) Contribuzione allo studio della organogeniacomparata degli stomi Institut de Botanique University de Geneve 41-42

Van Cotthem WR (1970) A classification of stomatal types Botanica

Journal of Linnean Society 63 235-246Wagner W (1979) Reticulate veins in the systematics of modern ferns

Taxon 1 87-95Webster GL (1994a) Classification of the Euphorbiaceae Annals of the

Missouri Botanical Garden 81 3-32

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 1724

184 Int Res J Plant Sci

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983112983141983149983145983152983137983154983137983139983161983156983145983139

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983112983141983149983145983152983137983154983137983139983161983156983145983139

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983105983138983137983160983145983137983148983105983140983137983160983145983137983148

Figure 12 Stomata types in Euphorbia taxa (A) E falcata (B) E peplus (C) E chamaepeplus and (D) Eschimperiana

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 1824

Aldhebiani and Jury 185

983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

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Figure 13 Figure 13 Stomata types in Euphorbia taxa in Saudi Arabia (A) E arabica (B) E serpens (C)E scordiifolia and (D) E helioscopia

distribution between the leaf surfaces (see Figure 13)While E hirta has the most types E helioscopia Edracunculoides and E acalyphoides have the least

Regarding the position of the stomata on the surfacelevel of the epidermis a variable degree of sunkenesshas been recorded Using the ESEM the stomata appear

deeply sunken in some species whilst in others they areonly slightly sunken with the outer rims of the subsidiarycells surrounding the aperture The deeply and slightlysunken species are recorded in Table 2 AdditionallyFigure 10 shows the sunken stomata in different speciesunder the ESEM

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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186 Int Res J Plant Sci

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Figure 14 Stomata types in Euphorbia taxa in Saudi Arabia (A) E hirta (B) E hypericifolia (C) E pirottae and (D)Espaff schimperi

Venation

The venation patterns in the Euphorbia species underinvestigation vary from one-veined three-veined to thosewith four or more veins The predominant type is three-veined However in E balsamifera subsp Adenensis

there are seven middle parallel veins three middle thickand four thinner lateral ones (Figure 15 A and B)Whereas in E pirottae there are three main middleparallel veins and two thinner laterals (Figure 15 C) Incontrast E chamaepeplus E schimperiana and Earabica have only one thick main midrib coming from the

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Aldhebiani and Jury 187

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Figure 15 Figure 15 Venation patterns at base of leaves invariants with more than three veins (A amp B) E balsamiferasubsp adenensis and (C) E pirottae

petiole then branched from the side into two lateral veinsat the point when the leaf starts widening (Figure 16)Each of these veins branches again into secondary veinsat the leaf margin

The rest of the species have just three veins Egrossheimii E dracunculoides E falcata E peplus Eserpens E scordiifolia E granulata E inaequilateraE helioscopia E hirta E hypericifolia and E

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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188 Int Res J Plant Sci

Figure 16 Venation patterns with one main mid vein at the leaf base in (A) E arabica and (B) E chamaepeplus

983106983105

Figure 17 Venation patterns with three-veined leaves in (A) E falcata and (B) E granulata

cyathophora All the three veins come from the petioleand start branching into secondary veins both toward themargin and the mid vein direction (Figure 17 B)

The vein endings at the leaf margin vary from closed

to open There is a relationship between the closedmargins and the sheathed veins (midrib and secondaryveins that are surrounded by parenchyma tissue) Whenthe veins are ornamented (sheathed) the margins areclosed This can be seen in subgenus Chamaesyce inspecies such as E serpens E scordiifolia E granulataE inaequilatera E hirta E hypericifolia and E arabica Additionally these species have the three-vein venation

On the other hand when the veins are not sheathedthe veins ends are open at the leaf margin as in Ebalsamifera subsp adenensis E schimperi E cuneataE acalyphoides E grossheimii E dracunculoides E

falcata E peplus E chamaepeplus E schimperiana Ehelioscopia E pirottae ldquoE aff schimperirdquo and Ecyathophora

CONCLUSIONS

Anatomical characters are seen here to be very useful inEuphorbia classification and identification The resultsobtained from anatomical studies were useful to groupsome species and identify others

Depending on the venation type the Euphorbiaspecies under investigation are divided anatomically intothree main groups see Figure 18

Group one veins are sheathed epidermal cellsdifferentiated on adaxial and abaxial surfaces Group two

983105 983106

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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Figure 18 Hierarchial relation of Euphorbia taxa in Saudi Arabia based on anatomical charac

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

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190 Int Res J Plant Sci

veins are not sheathed and epidermal cells are alike inboth surfaces And the third group includes species thatdo not belong to any of the previous groups The firstgroup can be further divided into three-veined and one-veined while the second group into three-veined and fouror more veins

In addition E grossheimii E retusa and E pirottae are related by being three-veined open at the marginswith rounded to oval tracheas and elongated cells bothon adaxial and abaxial surfaces However in E retusa the epidermal cells have straight cell walls on the adaxialsurface They are also similar in having hemiparacyticand paracytic types of stomata either on the adaxial orthe abaxial surface

On the other hand species with two differentepidermal cells on the leaf surfaces usually have three-sheathed veins with closed margins and elongatedtrachea such as E serpens E scordiifolia E granulata E inaequilatera E hirta and E hypericifolia which belongto subgenus Chamaesyce according to (Carter andSmith 1988)

In the case of E granulata which has beendescribed as having three different variants E granulata var granulata (both leaf surfaces hairy) E granulata varglabrata (adaxial glabrous and abaxial hairy) and aglabrous variant (the whole plant glabrous) Anatomicalwork has shown especially by using the ESEM that theleaves are either hairy or glabrous on both surfaces Sowhen the leaf is hairy both surfaces have differences inhair distribution Usually the lower surface is more hairythan upper surface (E granulata var glabrata ) butsometimes they are evenly distributed (Egranulata vargranulata ) In addition hairs on both surfaces are

multicellular In contrast when the leaf is glabrous bothleaf surfaces are glabrous and the whole plant isglabrous (suggested new variety) They were similar inhaving the same venation type (three-veined)

Finally ldquoE aff schimperi ldquohas some similarity with E schimperi in having the same open margin not theornamented venation type As well they both have thesame type of stomata paracytic and anisocytic but theydiffer in their distribution on abaxial and adaxial Thusthis may support the suggestion considering ldquoE affschimperi rdquo as a new subspecies of E schimperi

REFERENCES

Baranova M (1992) Principels of comparative stomatographic studiesof flowering plants The Botanical Review 58 49-99

Carlquist S (1961) Comparative Plant Anatomy Holt Rinehart ampWinston New York

Carter S Smith AR (1988) Euphorbiaceae (Part2) In Polhill ed Floraof Tropical East Africa Rotterdam Brookfield AA Balkema 409-597

Chaudhary SA (2001) Flora of The Kingdom of Saudi Arabia Vol 3Ministry of Agriculture and Water Riyadh

Dehay C (1935) Lagraveppareil Libero-ligneux foliare des EuphorbiaceaeAnnales des Sciences Naturelles Botanique 10 147-190

Dehgan B (1982) Comparative anatomy of the petiole and infragenericrelationships in Jatropha (Euphorbiaceae) American Journal oBotany 69 1283-1295

Gaucher L (1898) Ē tude anatomique du genre Euphorbia LPKlincksieck Paris

Gaucher L (1902) Recherches anatomiques sur les EuphorbiaceacuteesAnnales des Sciences Naturelles Botanique 15 161-309

Gaucher L (1902) Recherches anatomiques sur les Euphorbiaceacutees

Annales des Sciences Naturelles Botanique 15 161-309Govaerts R Fordin D Radcliffe-Smith A (2000) World Checklist and

Bibliography of Euphorbiaceae The Royal Botanic Gardens KewHickey LJ (1973) Classification of the architecture of dicotyledonous

leaves American Journal of Botany 60 17-33Hickey LJ (1979) A revised classification of the architecture o

dicotyledonous leaves In Metcalfe CR and Chalk L eds Anatomy oDicotyledons Systematic Anatomy of the leaf and stem 2nd edClarendon Press Oxford 25-39

Kakkar L Paliwal GS (1972) Epidermis in Euphorbia Indian ScienceCongress Association Proceedings 59 326-327

Kakkar L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 5 Epidermis Proceedings of the Indian National ScienceAcademy Part B Biological Sciences 40 55-67

Koumlhler E (1993) Blattvervatur-Muster der Buxaceae Dumortierr undSimmondsiaceae Van Tieghem Feddes Repertorium 104 145-167

Levin GA (1986a) Systematic foliar morphology of Phylanthoideae(Euphorbiaceae) II Phenetic analysis Annals of Missouri BotanicaGarden 73 86-98

Levin GA (1986b) Systematic foliar morphology of Phylanthoideae IICladistic analysis Systematic Botany 11 515-530

Levin GA (1986c) Systematic foliar morphology of Phyllanthoideae(Euphorbiaceae) I conspectus Annals of Missouri Botanical Garden73 29-85

Mahlberg P (1973) Histochemistry and Scanning Electron Microscopyof Starch Grains from Latex of Euphorbia-Terracina and EuphorbiaTirucalli Proceedings of the Indiana Academy of Science 82 132

Mahlberg PG (1974) Scanning Microscopic Comparison of StarchGrains from the Latex of Euphorbia -Spp Proceedings of the IndianaAcademy of Science 83 83-84

Mahlberg PG (1975) Evolution of the Laticifer in Euphorbia asInterpreted from Starch Grain Morphology American Journal oBotany 62 577-583

Mahlberg PG (1982) Comparative Morphology of Starch Grains inLatex from Varieties of Poinsettia Euphorbia-PulcherrimEuphorbiaceae Botanical Gazette 143 206-209

Mahlberg PG Davis DG Galitz DS Manners GD (1987) Laticifers andthe Classification of Euphorbia the Chemotaxonomy of EuphorbiaEsula L Botanical Journal of the Linnean Society 94 165-180

Metcalfe C Chalk L (1950) Anatomy of the Dicotyledons At theClarendon Press Oxford

Miller K Webster GL (1962) Systematic position of Cnidoscolus andJatropha Brittonia 14 174-180

Pax F (1884) Die Anatomie der Euphorbiaceen in ihrer Beziehung zumSystem derselben Botanische Jahrbuumlcher fuumlr SystematikPflanzengeschichte und Pflanzengeographie Leipzig 5 384-421

Payne WW (1970) Helicocytic and allelocytic stomata unrecognizedpatters in the Dicotyledonae Am J Bot57 140-147

Pray TR (1955a) Foliar venation of angiosperm II Histogenesis of thevenation of Liriodendron Am J Bot 42 18-27

Pray TR 1955b Foliar venation of angiosperms III Pattern andhistology of the venation of Hosta Am J Bot 42 11-18

Radford AE Dickison WC Massey JR Bell CR (1974) Vascular PlanSystematics Harper amp Row Publishers London New York

Radlkofer L (1870) Ueber Pausandra ein neues EuphorbiaceenGenus Flora 53 81-95

Raju VS Rao PN (1977) Variation in the structure and development ofoliar stomata in the Euphorbiaceae Botanical Journal of the LinneanSociety 75 69-97

Raju VS Rao PN (1987) The Taxonomic Use of the Basic StomataType in the Generic Delimitation of Chamaesyce EuphorbiaceaeFeddes Repertorium 98 137-142

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2424

Rosowski JR (1968) Laticifer Morphology in the Mature Stem and Leafof Euphorbia supina Botanical Gazette 129 113-120

Roth-Nebelsick A Uhl D Mosbrugger V Kerp H (2001) Evolution andfunction of leaf venation architecture a review Annals of Botany 87553-566

Sehgal L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 2 Venation Patterns Botanical Journal of the Linnean Society 68 173-208

Singh G (2004) Plant Systematics An Integrated Approach SciencePublishers Enfield

Solereder H (1899) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Solereder H (1908) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Aldhebiani and Jury 191

Stace CA (1965) Cuticular studies as an aid to plant taxonomy Bulletinof the British Museum Natural History Botany 4 3-78

Stace CA (1989) Plant Taxonomy and Biosystematics Hodder ampStoughton London

Tognini F (1897) Contribuzione allo studio della organogeniacomparata degli stomi Institut de Botanique University de Geneve 41-42

Van Cotthem WR (1970) A classification of stomatal types Botanica

Journal of Linnean Society 63 235-246Wagner W (1979) Reticulate veins in the systematics of modern ferns

Taxon 1 87-95Webster GL (1994a) Classification of the Euphorbiaceae Annals of the

Missouri Botanical Garden 81 3-32

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 1824

Aldhebiani and Jury 185

983105983140983137983160983145983137983148 983105983138983137983160983145983137983148

983105983150983151983149983151983139983161983156983145983139

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983112983141983149983145983152983137983154983137983139983161983156983145983139

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Figure 13 Figure 13 Stomata types in Euphorbia taxa in Saudi Arabia (A) E arabica (B) E serpens (C)E scordiifolia and (D) E helioscopia

distribution between the leaf surfaces (see Figure 13)While E hirta has the most types E helioscopia Edracunculoides and E acalyphoides have the least

Regarding the position of the stomata on the surfacelevel of the epidermis a variable degree of sunkenesshas been recorded Using the ESEM the stomata appear

deeply sunken in some species whilst in others they areonly slightly sunken with the outer rims of the subsidiarycells surrounding the aperture The deeply and slightlysunken species are recorded in Table 2 AdditionallyFigure 10 shows the sunken stomata in different speciesunder the ESEM

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 1924

186 Int Res J Plant Sci

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983107

983106

983105

983105983140983137983160983145983137983148983105983138983137983160983145983137983148

Figure 14 Stomata types in Euphorbia taxa in Saudi Arabia (A) E hirta (B) E hypericifolia (C) E pirottae and (D)Espaff schimperi

Venation

The venation patterns in the Euphorbia species underinvestigation vary from one-veined three-veined to thosewith four or more veins The predominant type is three-veined However in E balsamifera subsp Adenensis

there are seven middle parallel veins three middle thickand four thinner lateral ones (Figure 15 A and B)Whereas in E pirottae there are three main middleparallel veins and two thinner laterals (Figure 15 C) Incontrast E chamaepeplus E schimperiana and Earabica have only one thick main midrib coming from the

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2024

Aldhebiani and Jury 187

983105

983106

983107

Figure 15 Figure 15 Venation patterns at base of leaves invariants with more than three veins (A amp B) E balsamiferasubsp adenensis and (C) E pirottae

petiole then branched from the side into two lateral veinsat the point when the leaf starts widening (Figure 16)Each of these veins branches again into secondary veinsat the leaf margin

The rest of the species have just three veins Egrossheimii E dracunculoides E falcata E peplus Eserpens E scordiifolia E granulata E inaequilateraE helioscopia E hirta E hypericifolia and E

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2124

188 Int Res J Plant Sci

Figure 16 Venation patterns with one main mid vein at the leaf base in (A) E arabica and (B) E chamaepeplus

983106983105

Figure 17 Venation patterns with three-veined leaves in (A) E falcata and (B) E granulata

cyathophora All the three veins come from the petioleand start branching into secondary veins both toward themargin and the mid vein direction (Figure 17 B)

The vein endings at the leaf margin vary from closed

to open There is a relationship between the closedmargins and the sheathed veins (midrib and secondaryveins that are surrounded by parenchyma tissue) Whenthe veins are ornamented (sheathed) the margins areclosed This can be seen in subgenus Chamaesyce inspecies such as E serpens E scordiifolia E granulataE inaequilatera E hirta E hypericifolia and E arabica Additionally these species have the three-vein venation

On the other hand when the veins are not sheathedthe veins ends are open at the leaf margin as in Ebalsamifera subsp adenensis E schimperi E cuneataE acalyphoides E grossheimii E dracunculoides E

falcata E peplus E chamaepeplus E schimperiana Ehelioscopia E pirottae ldquoE aff schimperirdquo and Ecyathophora

CONCLUSIONS

Anatomical characters are seen here to be very useful inEuphorbia classification and identification The resultsobtained from anatomical studies were useful to groupsome species and identify others

Depending on the venation type the Euphorbiaspecies under investigation are divided anatomically intothree main groups see Figure 18

Group one veins are sheathed epidermal cellsdifferentiated on adaxial and abaxial surfaces Group two

983105 983106

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2224

Figure 18 Hierarchial relation of Euphorbia taxa in Saudi Arabia based on anatomical charac

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2324

190 Int Res J Plant Sci

veins are not sheathed and epidermal cells are alike inboth surfaces And the third group includes species thatdo not belong to any of the previous groups The firstgroup can be further divided into three-veined and one-veined while the second group into three-veined and fouror more veins

In addition E grossheimii E retusa and E pirottae are related by being three-veined open at the marginswith rounded to oval tracheas and elongated cells bothon adaxial and abaxial surfaces However in E retusa the epidermal cells have straight cell walls on the adaxialsurface They are also similar in having hemiparacyticand paracytic types of stomata either on the adaxial orthe abaxial surface

On the other hand species with two differentepidermal cells on the leaf surfaces usually have three-sheathed veins with closed margins and elongatedtrachea such as E serpens E scordiifolia E granulata E inaequilatera E hirta and E hypericifolia which belongto subgenus Chamaesyce according to (Carter andSmith 1988)

In the case of E granulata which has beendescribed as having three different variants E granulata var granulata (both leaf surfaces hairy) E granulata varglabrata (adaxial glabrous and abaxial hairy) and aglabrous variant (the whole plant glabrous) Anatomicalwork has shown especially by using the ESEM that theleaves are either hairy or glabrous on both surfaces Sowhen the leaf is hairy both surfaces have differences inhair distribution Usually the lower surface is more hairythan upper surface (E granulata var glabrata ) butsometimes they are evenly distributed (Egranulata vargranulata ) In addition hairs on both surfaces are

multicellular In contrast when the leaf is glabrous bothleaf surfaces are glabrous and the whole plant isglabrous (suggested new variety) They were similar inhaving the same venation type (three-veined)

Finally ldquoE aff schimperi ldquohas some similarity with E schimperi in having the same open margin not theornamented venation type As well they both have thesame type of stomata paracytic and anisocytic but theydiffer in their distribution on abaxial and adaxial Thusthis may support the suggestion considering ldquoE affschimperi rdquo as a new subspecies of E schimperi

REFERENCES

Baranova M (1992) Principels of comparative stomatographic studiesof flowering plants The Botanical Review 58 49-99

Carlquist S (1961) Comparative Plant Anatomy Holt Rinehart ampWinston New York

Carter S Smith AR (1988) Euphorbiaceae (Part2) In Polhill ed Floraof Tropical East Africa Rotterdam Brookfield AA Balkema 409-597

Chaudhary SA (2001) Flora of The Kingdom of Saudi Arabia Vol 3Ministry of Agriculture and Water Riyadh

Dehay C (1935) Lagraveppareil Libero-ligneux foliare des EuphorbiaceaeAnnales des Sciences Naturelles Botanique 10 147-190

Dehgan B (1982) Comparative anatomy of the petiole and infragenericrelationships in Jatropha (Euphorbiaceae) American Journal oBotany 69 1283-1295

Gaucher L (1898) Ē tude anatomique du genre Euphorbia LPKlincksieck Paris

Gaucher L (1902) Recherches anatomiques sur les EuphorbiaceacuteesAnnales des Sciences Naturelles Botanique 15 161-309

Gaucher L (1902) Recherches anatomiques sur les Euphorbiaceacutees

Annales des Sciences Naturelles Botanique 15 161-309Govaerts R Fordin D Radcliffe-Smith A (2000) World Checklist and

Bibliography of Euphorbiaceae The Royal Botanic Gardens KewHickey LJ (1973) Classification of the architecture of dicotyledonous

leaves American Journal of Botany 60 17-33Hickey LJ (1979) A revised classification of the architecture o

dicotyledonous leaves In Metcalfe CR and Chalk L eds Anatomy oDicotyledons Systematic Anatomy of the leaf and stem 2nd edClarendon Press Oxford 25-39

Kakkar L Paliwal GS (1972) Epidermis in Euphorbia Indian ScienceCongress Association Proceedings 59 326-327

Kakkar L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 5 Epidermis Proceedings of the Indian National ScienceAcademy Part B Biological Sciences 40 55-67

Koumlhler E (1993) Blattvervatur-Muster der Buxaceae Dumortierr undSimmondsiaceae Van Tieghem Feddes Repertorium 104 145-167

Levin GA (1986a) Systematic foliar morphology of Phylanthoideae(Euphorbiaceae) II Phenetic analysis Annals of Missouri BotanicaGarden 73 86-98

Levin GA (1986b) Systematic foliar morphology of Phylanthoideae IICladistic analysis Systematic Botany 11 515-530

Levin GA (1986c) Systematic foliar morphology of Phyllanthoideae(Euphorbiaceae) I conspectus Annals of Missouri Botanical Garden73 29-85

Mahlberg P (1973) Histochemistry and Scanning Electron Microscopyof Starch Grains from Latex of Euphorbia-Terracina and EuphorbiaTirucalli Proceedings of the Indiana Academy of Science 82 132

Mahlberg PG (1974) Scanning Microscopic Comparison of StarchGrains from the Latex of Euphorbia -Spp Proceedings of the IndianaAcademy of Science 83 83-84

Mahlberg PG (1975) Evolution of the Laticifer in Euphorbia asInterpreted from Starch Grain Morphology American Journal oBotany 62 577-583

Mahlberg PG (1982) Comparative Morphology of Starch Grains inLatex from Varieties of Poinsettia Euphorbia-PulcherrimEuphorbiaceae Botanical Gazette 143 206-209

Mahlberg PG Davis DG Galitz DS Manners GD (1987) Laticifers andthe Classification of Euphorbia the Chemotaxonomy of EuphorbiaEsula L Botanical Journal of the Linnean Society 94 165-180

Metcalfe C Chalk L (1950) Anatomy of the Dicotyledons At theClarendon Press Oxford

Miller K Webster GL (1962) Systematic position of Cnidoscolus andJatropha Brittonia 14 174-180

Pax F (1884) Die Anatomie der Euphorbiaceen in ihrer Beziehung zumSystem derselben Botanische Jahrbuumlcher fuumlr SystematikPflanzengeschichte und Pflanzengeographie Leipzig 5 384-421

Payne WW (1970) Helicocytic and allelocytic stomata unrecognizedpatters in the Dicotyledonae Am J Bot57 140-147

Pray TR (1955a) Foliar venation of angiosperm II Histogenesis of thevenation of Liriodendron Am J Bot 42 18-27

Pray TR 1955b Foliar venation of angiosperms III Pattern andhistology of the venation of Hosta Am J Bot 42 11-18

Radford AE Dickison WC Massey JR Bell CR (1974) Vascular PlanSystematics Harper amp Row Publishers London New York

Radlkofer L (1870) Ueber Pausandra ein neues EuphorbiaceenGenus Flora 53 81-95

Raju VS Rao PN (1977) Variation in the structure and development ofoliar stomata in the Euphorbiaceae Botanical Journal of the LinneanSociety 75 69-97

Raju VS Rao PN (1987) The Taxonomic Use of the Basic StomataType in the Generic Delimitation of Chamaesyce EuphorbiaceaeFeddes Repertorium 98 137-142

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2424

Rosowski JR (1968) Laticifer Morphology in the Mature Stem and Leafof Euphorbia supina Botanical Gazette 129 113-120

Roth-Nebelsick A Uhl D Mosbrugger V Kerp H (2001) Evolution andfunction of leaf venation architecture a review Annals of Botany 87553-566

Sehgal L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 2 Venation Patterns Botanical Journal of the Linnean Society 68 173-208

Singh G (2004) Plant Systematics An Integrated Approach SciencePublishers Enfield

Solereder H (1899) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Solereder H (1908) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Aldhebiani and Jury 191

Stace CA (1965) Cuticular studies as an aid to plant taxonomy Bulletinof the British Museum Natural History Botany 4 3-78

Stace CA (1989) Plant Taxonomy and Biosystematics Hodder ampStoughton London

Tognini F (1897) Contribuzione allo studio della organogeniacomparata degli stomi Institut de Botanique University de Geneve 41-42

Van Cotthem WR (1970) A classification of stomatal types Botanica

Journal of Linnean Society 63 235-246Wagner W (1979) Reticulate veins in the systematics of modern ferns

Taxon 1 87-95Webster GL (1994a) Classification of the Euphorbiaceae Annals of the

Missouri Botanical Garden 81 3-32

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 1924

186 Int Res J Plant Sci

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983105983150983151983149983151983139983161983156983145983139

983112983141983149983145983152983137983154983137983139983161983156983145983139

983120983137983154983137983139983161983156983145983139

983105983150983145983155983151983139983161983156983145983139

983124983141983156983154983137983139983161983156983145983139

983105983139983156983145983150983151983139983161983156983145983139

983107

983106

983105

983105983140983137983160983145983137983148983105983138983137983160983145983137983148

Figure 14 Stomata types in Euphorbia taxa in Saudi Arabia (A) E hirta (B) E hypericifolia (C) E pirottae and (D)Espaff schimperi

Venation

The venation patterns in the Euphorbia species underinvestigation vary from one-veined three-veined to thosewith four or more veins The predominant type is three-veined However in E balsamifera subsp Adenensis

there are seven middle parallel veins three middle thickand four thinner lateral ones (Figure 15 A and B)Whereas in E pirottae there are three main middleparallel veins and two thinner laterals (Figure 15 C) Incontrast E chamaepeplus E schimperiana and Earabica have only one thick main midrib coming from the

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2024

Aldhebiani and Jury 187

983105

983106

983107

Figure 15 Figure 15 Venation patterns at base of leaves invariants with more than three veins (A amp B) E balsamiferasubsp adenensis and (C) E pirottae

petiole then branched from the side into two lateral veinsat the point when the leaf starts widening (Figure 16)Each of these veins branches again into secondary veinsat the leaf margin

The rest of the species have just three veins Egrossheimii E dracunculoides E falcata E peplus Eserpens E scordiifolia E granulata E inaequilateraE helioscopia E hirta E hypericifolia and E

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2124

188 Int Res J Plant Sci

Figure 16 Venation patterns with one main mid vein at the leaf base in (A) E arabica and (B) E chamaepeplus

983106983105

Figure 17 Venation patterns with three-veined leaves in (A) E falcata and (B) E granulata

cyathophora All the three veins come from the petioleand start branching into secondary veins both toward themargin and the mid vein direction (Figure 17 B)

The vein endings at the leaf margin vary from closed

to open There is a relationship between the closedmargins and the sheathed veins (midrib and secondaryveins that are surrounded by parenchyma tissue) Whenthe veins are ornamented (sheathed) the margins areclosed This can be seen in subgenus Chamaesyce inspecies such as E serpens E scordiifolia E granulataE inaequilatera E hirta E hypericifolia and E arabica Additionally these species have the three-vein venation

On the other hand when the veins are not sheathedthe veins ends are open at the leaf margin as in Ebalsamifera subsp adenensis E schimperi E cuneataE acalyphoides E grossheimii E dracunculoides E

falcata E peplus E chamaepeplus E schimperiana Ehelioscopia E pirottae ldquoE aff schimperirdquo and Ecyathophora

CONCLUSIONS

Anatomical characters are seen here to be very useful inEuphorbia classification and identification The resultsobtained from anatomical studies were useful to groupsome species and identify others

Depending on the venation type the Euphorbiaspecies under investigation are divided anatomically intothree main groups see Figure 18

Group one veins are sheathed epidermal cellsdifferentiated on adaxial and abaxial surfaces Group two

983105 983106

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2224

Figure 18 Hierarchial relation of Euphorbia taxa in Saudi Arabia based on anatomical charac

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2324

190 Int Res J Plant Sci

veins are not sheathed and epidermal cells are alike inboth surfaces And the third group includes species thatdo not belong to any of the previous groups The firstgroup can be further divided into three-veined and one-veined while the second group into three-veined and fouror more veins

In addition E grossheimii E retusa and E pirottae are related by being three-veined open at the marginswith rounded to oval tracheas and elongated cells bothon adaxial and abaxial surfaces However in E retusa the epidermal cells have straight cell walls on the adaxialsurface They are also similar in having hemiparacyticand paracytic types of stomata either on the adaxial orthe abaxial surface

On the other hand species with two differentepidermal cells on the leaf surfaces usually have three-sheathed veins with closed margins and elongatedtrachea such as E serpens E scordiifolia E granulata E inaequilatera E hirta and E hypericifolia which belongto subgenus Chamaesyce according to (Carter andSmith 1988)

In the case of E granulata which has beendescribed as having three different variants E granulata var granulata (both leaf surfaces hairy) E granulata varglabrata (adaxial glabrous and abaxial hairy) and aglabrous variant (the whole plant glabrous) Anatomicalwork has shown especially by using the ESEM that theleaves are either hairy or glabrous on both surfaces Sowhen the leaf is hairy both surfaces have differences inhair distribution Usually the lower surface is more hairythan upper surface (E granulata var glabrata ) butsometimes they are evenly distributed (Egranulata vargranulata ) In addition hairs on both surfaces are

multicellular In contrast when the leaf is glabrous bothleaf surfaces are glabrous and the whole plant isglabrous (suggested new variety) They were similar inhaving the same venation type (three-veined)

Finally ldquoE aff schimperi ldquohas some similarity with E schimperi in having the same open margin not theornamented venation type As well they both have thesame type of stomata paracytic and anisocytic but theydiffer in their distribution on abaxial and adaxial Thusthis may support the suggestion considering ldquoE affschimperi rdquo as a new subspecies of E schimperi

REFERENCES

Baranova M (1992) Principels of comparative stomatographic studiesof flowering plants The Botanical Review 58 49-99

Carlquist S (1961) Comparative Plant Anatomy Holt Rinehart ampWinston New York

Carter S Smith AR (1988) Euphorbiaceae (Part2) In Polhill ed Floraof Tropical East Africa Rotterdam Brookfield AA Balkema 409-597

Chaudhary SA (2001) Flora of The Kingdom of Saudi Arabia Vol 3Ministry of Agriculture and Water Riyadh

Dehay C (1935) Lagraveppareil Libero-ligneux foliare des EuphorbiaceaeAnnales des Sciences Naturelles Botanique 10 147-190

Dehgan B (1982) Comparative anatomy of the petiole and infragenericrelationships in Jatropha (Euphorbiaceae) American Journal oBotany 69 1283-1295

Gaucher L (1898) Ē tude anatomique du genre Euphorbia LPKlincksieck Paris

Gaucher L (1902) Recherches anatomiques sur les EuphorbiaceacuteesAnnales des Sciences Naturelles Botanique 15 161-309

Gaucher L (1902) Recherches anatomiques sur les Euphorbiaceacutees

Annales des Sciences Naturelles Botanique 15 161-309Govaerts R Fordin D Radcliffe-Smith A (2000) World Checklist and

Bibliography of Euphorbiaceae The Royal Botanic Gardens KewHickey LJ (1973) Classification of the architecture of dicotyledonous

leaves American Journal of Botany 60 17-33Hickey LJ (1979) A revised classification of the architecture o

dicotyledonous leaves In Metcalfe CR and Chalk L eds Anatomy oDicotyledons Systematic Anatomy of the leaf and stem 2nd edClarendon Press Oxford 25-39

Kakkar L Paliwal GS (1972) Epidermis in Euphorbia Indian ScienceCongress Association Proceedings 59 326-327

Kakkar L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 5 Epidermis Proceedings of the Indian National ScienceAcademy Part B Biological Sciences 40 55-67

Koumlhler E (1993) Blattvervatur-Muster der Buxaceae Dumortierr undSimmondsiaceae Van Tieghem Feddes Repertorium 104 145-167

Levin GA (1986a) Systematic foliar morphology of Phylanthoideae(Euphorbiaceae) II Phenetic analysis Annals of Missouri BotanicaGarden 73 86-98

Levin GA (1986b) Systematic foliar morphology of Phylanthoideae IICladistic analysis Systematic Botany 11 515-530

Levin GA (1986c) Systematic foliar morphology of Phyllanthoideae(Euphorbiaceae) I conspectus Annals of Missouri Botanical Garden73 29-85

Mahlberg P (1973) Histochemistry and Scanning Electron Microscopyof Starch Grains from Latex of Euphorbia-Terracina and EuphorbiaTirucalli Proceedings of the Indiana Academy of Science 82 132

Mahlberg PG (1974) Scanning Microscopic Comparison of StarchGrains from the Latex of Euphorbia -Spp Proceedings of the IndianaAcademy of Science 83 83-84

Mahlberg PG (1975) Evolution of the Laticifer in Euphorbia asInterpreted from Starch Grain Morphology American Journal oBotany 62 577-583

Mahlberg PG (1982) Comparative Morphology of Starch Grains inLatex from Varieties of Poinsettia Euphorbia-PulcherrimEuphorbiaceae Botanical Gazette 143 206-209

Mahlberg PG Davis DG Galitz DS Manners GD (1987) Laticifers andthe Classification of Euphorbia the Chemotaxonomy of EuphorbiaEsula L Botanical Journal of the Linnean Society 94 165-180

Metcalfe C Chalk L (1950) Anatomy of the Dicotyledons At theClarendon Press Oxford

Miller K Webster GL (1962) Systematic position of Cnidoscolus andJatropha Brittonia 14 174-180

Pax F (1884) Die Anatomie der Euphorbiaceen in ihrer Beziehung zumSystem derselben Botanische Jahrbuumlcher fuumlr SystematikPflanzengeschichte und Pflanzengeographie Leipzig 5 384-421

Payne WW (1970) Helicocytic and allelocytic stomata unrecognizedpatters in the Dicotyledonae Am J Bot57 140-147

Pray TR (1955a) Foliar venation of angiosperm II Histogenesis of thevenation of Liriodendron Am J Bot 42 18-27

Pray TR 1955b Foliar venation of angiosperms III Pattern andhistology of the venation of Hosta Am J Bot 42 11-18

Radford AE Dickison WC Massey JR Bell CR (1974) Vascular PlanSystematics Harper amp Row Publishers London New York

Radlkofer L (1870) Ueber Pausandra ein neues EuphorbiaceenGenus Flora 53 81-95

Raju VS Rao PN (1977) Variation in the structure and development ofoliar stomata in the Euphorbiaceae Botanical Journal of the LinneanSociety 75 69-97

Raju VS Rao PN (1987) The Taxonomic Use of the Basic StomataType in the Generic Delimitation of Chamaesyce EuphorbiaceaeFeddes Repertorium 98 137-142

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2424

Rosowski JR (1968) Laticifer Morphology in the Mature Stem and Leafof Euphorbia supina Botanical Gazette 129 113-120

Roth-Nebelsick A Uhl D Mosbrugger V Kerp H (2001) Evolution andfunction of leaf venation architecture a review Annals of Botany 87553-566

Sehgal L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 2 Venation Patterns Botanical Journal of the Linnean Society 68 173-208

Singh G (2004) Plant Systematics An Integrated Approach SciencePublishers Enfield

Solereder H (1899) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Solereder H (1908) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Aldhebiani and Jury 191

Stace CA (1965) Cuticular studies as an aid to plant taxonomy Bulletinof the British Museum Natural History Botany 4 3-78

Stace CA (1989) Plant Taxonomy and Biosystematics Hodder ampStoughton London

Tognini F (1897) Contribuzione allo studio della organogeniacomparata degli stomi Institut de Botanique University de Geneve 41-42

Van Cotthem WR (1970) A classification of stomatal types Botanica

Journal of Linnean Society 63 235-246Wagner W (1979) Reticulate veins in the systematics of modern ferns

Taxon 1 87-95Webster GL (1994a) Classification of the Euphorbiaceae Annals of the

Missouri Botanical Garden 81 3-32

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2024

Aldhebiani and Jury 187

983105

983106

983107

Figure 15 Figure 15 Venation patterns at base of leaves invariants with more than three veins (A amp B) E balsamiferasubsp adenensis and (C) E pirottae

petiole then branched from the side into two lateral veinsat the point when the leaf starts widening (Figure 16)Each of these veins branches again into secondary veinsat the leaf margin

The rest of the species have just three veins Egrossheimii E dracunculoides E falcata E peplus Eserpens E scordiifolia E granulata E inaequilateraE helioscopia E hirta E hypericifolia and E

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2124

188 Int Res J Plant Sci

Figure 16 Venation patterns with one main mid vein at the leaf base in (A) E arabica and (B) E chamaepeplus

983106983105

Figure 17 Venation patterns with three-veined leaves in (A) E falcata and (B) E granulata

cyathophora All the three veins come from the petioleand start branching into secondary veins both toward themargin and the mid vein direction (Figure 17 B)

The vein endings at the leaf margin vary from closed

to open There is a relationship between the closedmargins and the sheathed veins (midrib and secondaryveins that are surrounded by parenchyma tissue) Whenthe veins are ornamented (sheathed) the margins areclosed This can be seen in subgenus Chamaesyce inspecies such as E serpens E scordiifolia E granulataE inaequilatera E hirta E hypericifolia and E arabica Additionally these species have the three-vein venation

On the other hand when the veins are not sheathedthe veins ends are open at the leaf margin as in Ebalsamifera subsp adenensis E schimperi E cuneataE acalyphoides E grossheimii E dracunculoides E

falcata E peplus E chamaepeplus E schimperiana Ehelioscopia E pirottae ldquoE aff schimperirdquo and Ecyathophora

CONCLUSIONS

Anatomical characters are seen here to be very useful inEuphorbia classification and identification The resultsobtained from anatomical studies were useful to groupsome species and identify others

Depending on the venation type the Euphorbiaspecies under investigation are divided anatomically intothree main groups see Figure 18

Group one veins are sheathed epidermal cellsdifferentiated on adaxial and abaxial surfaces Group two

983105 983106

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2224

Figure 18 Hierarchial relation of Euphorbia taxa in Saudi Arabia based on anatomical charac

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2324

190 Int Res J Plant Sci

veins are not sheathed and epidermal cells are alike inboth surfaces And the third group includes species thatdo not belong to any of the previous groups The firstgroup can be further divided into three-veined and one-veined while the second group into three-veined and fouror more veins

In addition E grossheimii E retusa and E pirottae are related by being three-veined open at the marginswith rounded to oval tracheas and elongated cells bothon adaxial and abaxial surfaces However in E retusa the epidermal cells have straight cell walls on the adaxialsurface They are also similar in having hemiparacyticand paracytic types of stomata either on the adaxial orthe abaxial surface

On the other hand species with two differentepidermal cells on the leaf surfaces usually have three-sheathed veins with closed margins and elongatedtrachea such as E serpens E scordiifolia E granulata E inaequilatera E hirta and E hypericifolia which belongto subgenus Chamaesyce according to (Carter andSmith 1988)

In the case of E granulata which has beendescribed as having three different variants E granulata var granulata (both leaf surfaces hairy) E granulata varglabrata (adaxial glabrous and abaxial hairy) and aglabrous variant (the whole plant glabrous) Anatomicalwork has shown especially by using the ESEM that theleaves are either hairy or glabrous on both surfaces Sowhen the leaf is hairy both surfaces have differences inhair distribution Usually the lower surface is more hairythan upper surface (E granulata var glabrata ) butsometimes they are evenly distributed (Egranulata vargranulata ) In addition hairs on both surfaces are

multicellular In contrast when the leaf is glabrous bothleaf surfaces are glabrous and the whole plant isglabrous (suggested new variety) They were similar inhaving the same venation type (three-veined)

Finally ldquoE aff schimperi ldquohas some similarity with E schimperi in having the same open margin not theornamented venation type As well they both have thesame type of stomata paracytic and anisocytic but theydiffer in their distribution on abaxial and adaxial Thusthis may support the suggestion considering ldquoE affschimperi rdquo as a new subspecies of E schimperi

REFERENCES

Baranova M (1992) Principels of comparative stomatographic studiesof flowering plants The Botanical Review 58 49-99

Carlquist S (1961) Comparative Plant Anatomy Holt Rinehart ampWinston New York

Carter S Smith AR (1988) Euphorbiaceae (Part2) In Polhill ed Floraof Tropical East Africa Rotterdam Brookfield AA Balkema 409-597

Chaudhary SA (2001) Flora of The Kingdom of Saudi Arabia Vol 3Ministry of Agriculture and Water Riyadh

Dehay C (1935) Lagraveppareil Libero-ligneux foliare des EuphorbiaceaeAnnales des Sciences Naturelles Botanique 10 147-190

Dehgan B (1982) Comparative anatomy of the petiole and infragenericrelationships in Jatropha (Euphorbiaceae) American Journal oBotany 69 1283-1295

Gaucher L (1898) Ē tude anatomique du genre Euphorbia LPKlincksieck Paris

Gaucher L (1902) Recherches anatomiques sur les EuphorbiaceacuteesAnnales des Sciences Naturelles Botanique 15 161-309

Gaucher L (1902) Recherches anatomiques sur les Euphorbiaceacutees

Annales des Sciences Naturelles Botanique 15 161-309Govaerts R Fordin D Radcliffe-Smith A (2000) World Checklist and

Bibliography of Euphorbiaceae The Royal Botanic Gardens KewHickey LJ (1973) Classification of the architecture of dicotyledonous

leaves American Journal of Botany 60 17-33Hickey LJ (1979) A revised classification of the architecture o

dicotyledonous leaves In Metcalfe CR and Chalk L eds Anatomy oDicotyledons Systematic Anatomy of the leaf and stem 2nd edClarendon Press Oxford 25-39

Kakkar L Paliwal GS (1972) Epidermis in Euphorbia Indian ScienceCongress Association Proceedings 59 326-327

Kakkar L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 5 Epidermis Proceedings of the Indian National ScienceAcademy Part B Biological Sciences 40 55-67

Koumlhler E (1993) Blattvervatur-Muster der Buxaceae Dumortierr undSimmondsiaceae Van Tieghem Feddes Repertorium 104 145-167

Levin GA (1986a) Systematic foliar morphology of Phylanthoideae(Euphorbiaceae) II Phenetic analysis Annals of Missouri BotanicaGarden 73 86-98

Levin GA (1986b) Systematic foliar morphology of Phylanthoideae IICladistic analysis Systematic Botany 11 515-530

Levin GA (1986c) Systematic foliar morphology of Phyllanthoideae(Euphorbiaceae) I conspectus Annals of Missouri Botanical Garden73 29-85

Mahlberg P (1973) Histochemistry and Scanning Electron Microscopyof Starch Grains from Latex of Euphorbia-Terracina and EuphorbiaTirucalli Proceedings of the Indiana Academy of Science 82 132

Mahlberg PG (1974) Scanning Microscopic Comparison of StarchGrains from the Latex of Euphorbia -Spp Proceedings of the IndianaAcademy of Science 83 83-84

Mahlberg PG (1975) Evolution of the Laticifer in Euphorbia asInterpreted from Starch Grain Morphology American Journal oBotany 62 577-583

Mahlberg PG (1982) Comparative Morphology of Starch Grains inLatex from Varieties of Poinsettia Euphorbia-PulcherrimEuphorbiaceae Botanical Gazette 143 206-209

Mahlberg PG Davis DG Galitz DS Manners GD (1987) Laticifers andthe Classification of Euphorbia the Chemotaxonomy of EuphorbiaEsula L Botanical Journal of the Linnean Society 94 165-180

Metcalfe C Chalk L (1950) Anatomy of the Dicotyledons At theClarendon Press Oxford

Miller K Webster GL (1962) Systematic position of Cnidoscolus andJatropha Brittonia 14 174-180

Pax F (1884) Die Anatomie der Euphorbiaceen in ihrer Beziehung zumSystem derselben Botanische Jahrbuumlcher fuumlr SystematikPflanzengeschichte und Pflanzengeographie Leipzig 5 384-421

Payne WW (1970) Helicocytic and allelocytic stomata unrecognizedpatters in the Dicotyledonae Am J Bot57 140-147

Pray TR (1955a) Foliar venation of angiosperm II Histogenesis of thevenation of Liriodendron Am J Bot 42 18-27

Pray TR 1955b Foliar venation of angiosperms III Pattern andhistology of the venation of Hosta Am J Bot 42 11-18

Radford AE Dickison WC Massey JR Bell CR (1974) Vascular PlanSystematics Harper amp Row Publishers London New York

Radlkofer L (1870) Ueber Pausandra ein neues EuphorbiaceenGenus Flora 53 81-95

Raju VS Rao PN (1977) Variation in the structure and development ofoliar stomata in the Euphorbiaceae Botanical Journal of the LinneanSociety 75 69-97

Raju VS Rao PN (1987) The Taxonomic Use of the Basic StomataType in the Generic Delimitation of Chamaesyce EuphorbiaceaeFeddes Repertorium 98 137-142

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2424

Rosowski JR (1968) Laticifer Morphology in the Mature Stem and Leafof Euphorbia supina Botanical Gazette 129 113-120

Roth-Nebelsick A Uhl D Mosbrugger V Kerp H (2001) Evolution andfunction of leaf venation architecture a review Annals of Botany 87553-566

Sehgal L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 2 Venation Patterns Botanical Journal of the Linnean Society 68 173-208

Singh G (2004) Plant Systematics An Integrated Approach SciencePublishers Enfield

Solereder H (1899) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Solereder H (1908) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Aldhebiani and Jury 191

Stace CA (1965) Cuticular studies as an aid to plant taxonomy Bulletinof the British Museum Natural History Botany 4 3-78

Stace CA (1989) Plant Taxonomy and Biosystematics Hodder ampStoughton London

Tognini F (1897) Contribuzione allo studio della organogeniacomparata degli stomi Institut de Botanique University de Geneve 41-42

Van Cotthem WR (1970) A classification of stomatal types Botanica

Journal of Linnean Society 63 235-246Wagner W (1979) Reticulate veins in the systematics of modern ferns

Taxon 1 87-95Webster GL (1994a) Classification of the Euphorbiaceae Annals of the

Missouri Botanical Garden 81 3-32

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2124

188 Int Res J Plant Sci

Figure 16 Venation patterns with one main mid vein at the leaf base in (A) E arabica and (B) E chamaepeplus

983106983105

Figure 17 Venation patterns with three-veined leaves in (A) E falcata and (B) E granulata

cyathophora All the three veins come from the petioleand start branching into secondary veins both toward themargin and the mid vein direction (Figure 17 B)

The vein endings at the leaf margin vary from closed

to open There is a relationship between the closedmargins and the sheathed veins (midrib and secondaryveins that are surrounded by parenchyma tissue) Whenthe veins are ornamented (sheathed) the margins areclosed This can be seen in subgenus Chamaesyce inspecies such as E serpens E scordiifolia E granulataE inaequilatera E hirta E hypericifolia and E arabica Additionally these species have the three-vein venation

On the other hand when the veins are not sheathedthe veins ends are open at the leaf margin as in Ebalsamifera subsp adenensis E schimperi E cuneataE acalyphoides E grossheimii E dracunculoides E

falcata E peplus E chamaepeplus E schimperiana Ehelioscopia E pirottae ldquoE aff schimperirdquo and Ecyathophora

CONCLUSIONS

Anatomical characters are seen here to be very useful inEuphorbia classification and identification The resultsobtained from anatomical studies were useful to groupsome species and identify others

Depending on the venation type the Euphorbiaspecies under investigation are divided anatomically intothree main groups see Figure 18

Group one veins are sheathed epidermal cellsdifferentiated on adaxial and abaxial surfaces Group two

983105 983106

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2224

Figure 18 Hierarchial relation of Euphorbia taxa in Saudi Arabia based on anatomical charac

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2324

190 Int Res J Plant Sci

veins are not sheathed and epidermal cells are alike inboth surfaces And the third group includes species thatdo not belong to any of the previous groups The firstgroup can be further divided into three-veined and one-veined while the second group into three-veined and fouror more veins

In addition E grossheimii E retusa and E pirottae are related by being three-veined open at the marginswith rounded to oval tracheas and elongated cells bothon adaxial and abaxial surfaces However in E retusa the epidermal cells have straight cell walls on the adaxialsurface They are also similar in having hemiparacyticand paracytic types of stomata either on the adaxial orthe abaxial surface

On the other hand species with two differentepidermal cells on the leaf surfaces usually have three-sheathed veins with closed margins and elongatedtrachea such as E serpens E scordiifolia E granulata E inaequilatera E hirta and E hypericifolia which belongto subgenus Chamaesyce according to (Carter andSmith 1988)

In the case of E granulata which has beendescribed as having three different variants E granulata var granulata (both leaf surfaces hairy) E granulata varglabrata (adaxial glabrous and abaxial hairy) and aglabrous variant (the whole plant glabrous) Anatomicalwork has shown especially by using the ESEM that theleaves are either hairy or glabrous on both surfaces Sowhen the leaf is hairy both surfaces have differences inhair distribution Usually the lower surface is more hairythan upper surface (E granulata var glabrata ) butsometimes they are evenly distributed (Egranulata vargranulata ) In addition hairs on both surfaces are

multicellular In contrast when the leaf is glabrous bothleaf surfaces are glabrous and the whole plant isglabrous (suggested new variety) They were similar inhaving the same venation type (three-veined)

Finally ldquoE aff schimperi ldquohas some similarity with E schimperi in having the same open margin not theornamented venation type As well they both have thesame type of stomata paracytic and anisocytic but theydiffer in their distribution on abaxial and adaxial Thusthis may support the suggestion considering ldquoE affschimperi rdquo as a new subspecies of E schimperi

REFERENCES

Baranova M (1992) Principels of comparative stomatographic studiesof flowering plants The Botanical Review 58 49-99

Carlquist S (1961) Comparative Plant Anatomy Holt Rinehart ampWinston New York

Carter S Smith AR (1988) Euphorbiaceae (Part2) In Polhill ed Floraof Tropical East Africa Rotterdam Brookfield AA Balkema 409-597

Chaudhary SA (2001) Flora of The Kingdom of Saudi Arabia Vol 3Ministry of Agriculture and Water Riyadh

Dehay C (1935) Lagraveppareil Libero-ligneux foliare des EuphorbiaceaeAnnales des Sciences Naturelles Botanique 10 147-190

Dehgan B (1982) Comparative anatomy of the petiole and infragenericrelationships in Jatropha (Euphorbiaceae) American Journal oBotany 69 1283-1295

Gaucher L (1898) Ē tude anatomique du genre Euphorbia LPKlincksieck Paris

Gaucher L (1902) Recherches anatomiques sur les EuphorbiaceacuteesAnnales des Sciences Naturelles Botanique 15 161-309

Gaucher L (1902) Recherches anatomiques sur les Euphorbiaceacutees

Annales des Sciences Naturelles Botanique 15 161-309Govaerts R Fordin D Radcliffe-Smith A (2000) World Checklist and

Bibliography of Euphorbiaceae The Royal Botanic Gardens KewHickey LJ (1973) Classification of the architecture of dicotyledonous

leaves American Journal of Botany 60 17-33Hickey LJ (1979) A revised classification of the architecture o

dicotyledonous leaves In Metcalfe CR and Chalk L eds Anatomy oDicotyledons Systematic Anatomy of the leaf and stem 2nd edClarendon Press Oxford 25-39

Kakkar L Paliwal GS (1972) Epidermis in Euphorbia Indian ScienceCongress Association Proceedings 59 326-327

Kakkar L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 5 Epidermis Proceedings of the Indian National ScienceAcademy Part B Biological Sciences 40 55-67

Koumlhler E (1993) Blattvervatur-Muster der Buxaceae Dumortierr undSimmondsiaceae Van Tieghem Feddes Repertorium 104 145-167

Levin GA (1986a) Systematic foliar morphology of Phylanthoideae(Euphorbiaceae) II Phenetic analysis Annals of Missouri BotanicaGarden 73 86-98

Levin GA (1986b) Systematic foliar morphology of Phylanthoideae IICladistic analysis Systematic Botany 11 515-530

Levin GA (1986c) Systematic foliar morphology of Phyllanthoideae(Euphorbiaceae) I conspectus Annals of Missouri Botanical Garden73 29-85

Mahlberg P (1973) Histochemistry and Scanning Electron Microscopyof Starch Grains from Latex of Euphorbia-Terracina and EuphorbiaTirucalli Proceedings of the Indiana Academy of Science 82 132

Mahlberg PG (1974) Scanning Microscopic Comparison of StarchGrains from the Latex of Euphorbia -Spp Proceedings of the IndianaAcademy of Science 83 83-84

Mahlberg PG (1975) Evolution of the Laticifer in Euphorbia asInterpreted from Starch Grain Morphology American Journal oBotany 62 577-583

Mahlberg PG (1982) Comparative Morphology of Starch Grains inLatex from Varieties of Poinsettia Euphorbia-PulcherrimEuphorbiaceae Botanical Gazette 143 206-209

Mahlberg PG Davis DG Galitz DS Manners GD (1987) Laticifers andthe Classification of Euphorbia the Chemotaxonomy of EuphorbiaEsula L Botanical Journal of the Linnean Society 94 165-180

Metcalfe C Chalk L (1950) Anatomy of the Dicotyledons At theClarendon Press Oxford

Miller K Webster GL (1962) Systematic position of Cnidoscolus andJatropha Brittonia 14 174-180

Pax F (1884) Die Anatomie der Euphorbiaceen in ihrer Beziehung zumSystem derselben Botanische Jahrbuumlcher fuumlr SystematikPflanzengeschichte und Pflanzengeographie Leipzig 5 384-421

Payne WW (1970) Helicocytic and allelocytic stomata unrecognizedpatters in the Dicotyledonae Am J Bot57 140-147

Pray TR (1955a) Foliar venation of angiosperm II Histogenesis of thevenation of Liriodendron Am J Bot 42 18-27

Pray TR 1955b Foliar venation of angiosperms III Pattern andhistology of the venation of Hosta Am J Bot 42 11-18

Radford AE Dickison WC Massey JR Bell CR (1974) Vascular PlanSystematics Harper amp Row Publishers London New York

Radlkofer L (1870) Ueber Pausandra ein neues EuphorbiaceenGenus Flora 53 81-95

Raju VS Rao PN (1977) Variation in the structure and development ofoliar stomata in the Euphorbiaceae Botanical Journal of the LinneanSociety 75 69-97

Raju VS Rao PN (1987) The Taxonomic Use of the Basic StomataType in the Generic Delimitation of Chamaesyce EuphorbiaceaeFeddes Repertorium 98 137-142

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2424

Rosowski JR (1968) Laticifer Morphology in the Mature Stem and Leafof Euphorbia supina Botanical Gazette 129 113-120

Roth-Nebelsick A Uhl D Mosbrugger V Kerp H (2001) Evolution andfunction of leaf venation architecture a review Annals of Botany 87553-566

Sehgal L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 2 Venation Patterns Botanical Journal of the Linnean Society 68 173-208

Singh G (2004) Plant Systematics An Integrated Approach SciencePublishers Enfield

Solereder H (1899) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Solereder H (1908) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Aldhebiani and Jury 191

Stace CA (1965) Cuticular studies as an aid to plant taxonomy Bulletinof the British Museum Natural History Botany 4 3-78

Stace CA (1989) Plant Taxonomy and Biosystematics Hodder ampStoughton London

Tognini F (1897) Contribuzione allo studio della organogeniacomparata degli stomi Institut de Botanique University de Geneve 41-42

Van Cotthem WR (1970) A classification of stomatal types Botanica

Journal of Linnean Society 63 235-246Wagner W (1979) Reticulate veins in the systematics of modern ferns

Taxon 1 87-95Webster GL (1994a) Classification of the Euphorbiaceae Annals of the

Missouri Botanical Garden 81 3-32

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2224

Figure 18 Hierarchial relation of Euphorbia taxa in Saudi Arabia based on anatomical charac

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2324

190 Int Res J Plant Sci

veins are not sheathed and epidermal cells are alike inboth surfaces And the third group includes species thatdo not belong to any of the previous groups The firstgroup can be further divided into three-veined and one-veined while the second group into three-veined and fouror more veins

In addition E grossheimii E retusa and E pirottae are related by being three-veined open at the marginswith rounded to oval tracheas and elongated cells bothon adaxial and abaxial surfaces However in E retusa the epidermal cells have straight cell walls on the adaxialsurface They are also similar in having hemiparacyticand paracytic types of stomata either on the adaxial orthe abaxial surface

On the other hand species with two differentepidermal cells on the leaf surfaces usually have three-sheathed veins with closed margins and elongatedtrachea such as E serpens E scordiifolia E granulata E inaequilatera E hirta and E hypericifolia which belongto subgenus Chamaesyce according to (Carter andSmith 1988)

In the case of E granulata which has beendescribed as having three different variants E granulata var granulata (both leaf surfaces hairy) E granulata varglabrata (adaxial glabrous and abaxial hairy) and aglabrous variant (the whole plant glabrous) Anatomicalwork has shown especially by using the ESEM that theleaves are either hairy or glabrous on both surfaces Sowhen the leaf is hairy both surfaces have differences inhair distribution Usually the lower surface is more hairythan upper surface (E granulata var glabrata ) butsometimes they are evenly distributed (Egranulata vargranulata ) In addition hairs on both surfaces are

multicellular In contrast when the leaf is glabrous bothleaf surfaces are glabrous and the whole plant isglabrous (suggested new variety) They were similar inhaving the same venation type (three-veined)

Finally ldquoE aff schimperi ldquohas some similarity with E schimperi in having the same open margin not theornamented venation type As well they both have thesame type of stomata paracytic and anisocytic but theydiffer in their distribution on abaxial and adaxial Thusthis may support the suggestion considering ldquoE affschimperi rdquo as a new subspecies of E schimperi

REFERENCES

Baranova M (1992) Principels of comparative stomatographic studiesof flowering plants The Botanical Review 58 49-99

Carlquist S (1961) Comparative Plant Anatomy Holt Rinehart ampWinston New York

Carter S Smith AR (1988) Euphorbiaceae (Part2) In Polhill ed Floraof Tropical East Africa Rotterdam Brookfield AA Balkema 409-597

Chaudhary SA (2001) Flora of The Kingdom of Saudi Arabia Vol 3Ministry of Agriculture and Water Riyadh

Dehay C (1935) Lagraveppareil Libero-ligneux foliare des EuphorbiaceaeAnnales des Sciences Naturelles Botanique 10 147-190

Dehgan B (1982) Comparative anatomy of the petiole and infragenericrelationships in Jatropha (Euphorbiaceae) American Journal oBotany 69 1283-1295

Gaucher L (1898) Ē tude anatomique du genre Euphorbia LPKlincksieck Paris

Gaucher L (1902) Recherches anatomiques sur les EuphorbiaceacuteesAnnales des Sciences Naturelles Botanique 15 161-309

Gaucher L (1902) Recherches anatomiques sur les Euphorbiaceacutees

Annales des Sciences Naturelles Botanique 15 161-309Govaerts R Fordin D Radcliffe-Smith A (2000) World Checklist and

Bibliography of Euphorbiaceae The Royal Botanic Gardens KewHickey LJ (1973) Classification of the architecture of dicotyledonous

leaves American Journal of Botany 60 17-33Hickey LJ (1979) A revised classification of the architecture o

dicotyledonous leaves In Metcalfe CR and Chalk L eds Anatomy oDicotyledons Systematic Anatomy of the leaf and stem 2nd edClarendon Press Oxford 25-39

Kakkar L Paliwal GS (1972) Epidermis in Euphorbia Indian ScienceCongress Association Proceedings 59 326-327

Kakkar L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 5 Epidermis Proceedings of the Indian National ScienceAcademy Part B Biological Sciences 40 55-67

Koumlhler E (1993) Blattvervatur-Muster der Buxaceae Dumortierr undSimmondsiaceae Van Tieghem Feddes Repertorium 104 145-167

Levin GA (1986a) Systematic foliar morphology of Phylanthoideae(Euphorbiaceae) II Phenetic analysis Annals of Missouri BotanicaGarden 73 86-98

Levin GA (1986b) Systematic foliar morphology of Phylanthoideae IICladistic analysis Systematic Botany 11 515-530

Levin GA (1986c) Systematic foliar morphology of Phyllanthoideae(Euphorbiaceae) I conspectus Annals of Missouri Botanical Garden73 29-85

Mahlberg P (1973) Histochemistry and Scanning Electron Microscopyof Starch Grains from Latex of Euphorbia-Terracina and EuphorbiaTirucalli Proceedings of the Indiana Academy of Science 82 132

Mahlberg PG (1974) Scanning Microscopic Comparison of StarchGrains from the Latex of Euphorbia -Spp Proceedings of the IndianaAcademy of Science 83 83-84

Mahlberg PG (1975) Evolution of the Laticifer in Euphorbia asInterpreted from Starch Grain Morphology American Journal oBotany 62 577-583

Mahlberg PG (1982) Comparative Morphology of Starch Grains inLatex from Varieties of Poinsettia Euphorbia-PulcherrimEuphorbiaceae Botanical Gazette 143 206-209

Mahlberg PG Davis DG Galitz DS Manners GD (1987) Laticifers andthe Classification of Euphorbia the Chemotaxonomy of EuphorbiaEsula L Botanical Journal of the Linnean Society 94 165-180

Metcalfe C Chalk L (1950) Anatomy of the Dicotyledons At theClarendon Press Oxford

Miller K Webster GL (1962) Systematic position of Cnidoscolus andJatropha Brittonia 14 174-180

Pax F (1884) Die Anatomie der Euphorbiaceen in ihrer Beziehung zumSystem derselben Botanische Jahrbuumlcher fuumlr SystematikPflanzengeschichte und Pflanzengeographie Leipzig 5 384-421

Payne WW (1970) Helicocytic and allelocytic stomata unrecognizedpatters in the Dicotyledonae Am J Bot57 140-147

Pray TR (1955a) Foliar venation of angiosperm II Histogenesis of thevenation of Liriodendron Am J Bot 42 18-27

Pray TR 1955b Foliar venation of angiosperms III Pattern andhistology of the venation of Hosta Am J Bot 42 11-18

Radford AE Dickison WC Massey JR Bell CR (1974) Vascular PlanSystematics Harper amp Row Publishers London New York

Radlkofer L (1870) Ueber Pausandra ein neues EuphorbiaceenGenus Flora 53 81-95

Raju VS Rao PN (1977) Variation in the structure and development ofoliar stomata in the Euphorbiaceae Botanical Journal of the LinneanSociety 75 69-97

Raju VS Rao PN (1987) The Taxonomic Use of the Basic StomataType in the Generic Delimitation of Chamaesyce EuphorbiaceaeFeddes Repertorium 98 137-142

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2424

Rosowski JR (1968) Laticifer Morphology in the Mature Stem and Leafof Euphorbia supina Botanical Gazette 129 113-120

Roth-Nebelsick A Uhl D Mosbrugger V Kerp H (2001) Evolution andfunction of leaf venation architecture a review Annals of Botany 87553-566

Sehgal L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 2 Venation Patterns Botanical Journal of the Linnean Society 68 173-208

Singh G (2004) Plant Systematics An Integrated Approach SciencePublishers Enfield

Solereder H (1899) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Solereder H (1908) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Aldhebiani and Jury 191

Stace CA (1965) Cuticular studies as an aid to plant taxonomy Bulletinof the British Museum Natural History Botany 4 3-78

Stace CA (1989) Plant Taxonomy and Biosystematics Hodder ampStoughton London

Tognini F (1897) Contribuzione allo studio della organogeniacomparata degli stomi Institut de Botanique University de Geneve 41-42

Van Cotthem WR (1970) A classification of stomatal types Botanica

Journal of Linnean Society 63 235-246Wagner W (1979) Reticulate veins in the systematics of modern ferns

Taxon 1 87-95Webster GL (1994a) Classification of the Euphorbiaceae Annals of the

Missouri Botanical Garden 81 3-32

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2324

190 Int Res J Plant Sci

veins are not sheathed and epidermal cells are alike inboth surfaces And the third group includes species thatdo not belong to any of the previous groups The firstgroup can be further divided into three-veined and one-veined while the second group into three-veined and fouror more veins

In addition E grossheimii E retusa and E pirottae are related by being three-veined open at the marginswith rounded to oval tracheas and elongated cells bothon adaxial and abaxial surfaces However in E retusa the epidermal cells have straight cell walls on the adaxialsurface They are also similar in having hemiparacyticand paracytic types of stomata either on the adaxial orthe abaxial surface

On the other hand species with two differentepidermal cells on the leaf surfaces usually have three-sheathed veins with closed margins and elongatedtrachea such as E serpens E scordiifolia E granulata E inaequilatera E hirta and E hypericifolia which belongto subgenus Chamaesyce according to (Carter andSmith 1988)

In the case of E granulata which has beendescribed as having three different variants E granulata var granulata (both leaf surfaces hairy) E granulata varglabrata (adaxial glabrous and abaxial hairy) and aglabrous variant (the whole plant glabrous) Anatomicalwork has shown especially by using the ESEM that theleaves are either hairy or glabrous on both surfaces Sowhen the leaf is hairy both surfaces have differences inhair distribution Usually the lower surface is more hairythan upper surface (E granulata var glabrata ) butsometimes they are evenly distributed (Egranulata vargranulata ) In addition hairs on both surfaces are

multicellular In contrast when the leaf is glabrous bothleaf surfaces are glabrous and the whole plant isglabrous (suggested new variety) They were similar inhaving the same venation type (three-veined)

Finally ldquoE aff schimperi ldquohas some similarity with E schimperi in having the same open margin not theornamented venation type As well they both have thesame type of stomata paracytic and anisocytic but theydiffer in their distribution on abaxial and adaxial Thusthis may support the suggestion considering ldquoE affschimperi rdquo as a new subspecies of E schimperi

REFERENCES

Baranova M (1992) Principels of comparative stomatographic studiesof flowering plants The Botanical Review 58 49-99

Carlquist S (1961) Comparative Plant Anatomy Holt Rinehart ampWinston New York

Carter S Smith AR (1988) Euphorbiaceae (Part2) In Polhill ed Floraof Tropical East Africa Rotterdam Brookfield AA Balkema 409-597

Chaudhary SA (2001) Flora of The Kingdom of Saudi Arabia Vol 3Ministry of Agriculture and Water Riyadh

Dehay C (1935) Lagraveppareil Libero-ligneux foliare des EuphorbiaceaeAnnales des Sciences Naturelles Botanique 10 147-190

Dehgan B (1982) Comparative anatomy of the petiole and infragenericrelationships in Jatropha (Euphorbiaceae) American Journal oBotany 69 1283-1295

Gaucher L (1898) Ē tude anatomique du genre Euphorbia LPKlincksieck Paris

Gaucher L (1902) Recherches anatomiques sur les EuphorbiaceacuteesAnnales des Sciences Naturelles Botanique 15 161-309

Gaucher L (1902) Recherches anatomiques sur les Euphorbiaceacutees

Annales des Sciences Naturelles Botanique 15 161-309Govaerts R Fordin D Radcliffe-Smith A (2000) World Checklist and

Bibliography of Euphorbiaceae The Royal Botanic Gardens KewHickey LJ (1973) Classification of the architecture of dicotyledonous

leaves American Journal of Botany 60 17-33Hickey LJ (1979) A revised classification of the architecture o

dicotyledonous leaves In Metcalfe CR and Chalk L eds Anatomy oDicotyledons Systematic Anatomy of the leaf and stem 2nd edClarendon Press Oxford 25-39

Kakkar L Paliwal GS (1972) Epidermis in Euphorbia Indian ScienceCongress Association Proceedings 59 326-327

Kakkar L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 5 Epidermis Proceedings of the Indian National ScienceAcademy Part B Biological Sciences 40 55-67

Koumlhler E (1993) Blattvervatur-Muster der Buxaceae Dumortierr undSimmondsiaceae Van Tieghem Feddes Repertorium 104 145-167

Levin GA (1986a) Systematic foliar morphology of Phylanthoideae(Euphorbiaceae) II Phenetic analysis Annals of Missouri BotanicaGarden 73 86-98

Levin GA (1986b) Systematic foliar morphology of Phylanthoideae IICladistic analysis Systematic Botany 11 515-530

Levin GA (1986c) Systematic foliar morphology of Phyllanthoideae(Euphorbiaceae) I conspectus Annals of Missouri Botanical Garden73 29-85

Mahlberg P (1973) Histochemistry and Scanning Electron Microscopyof Starch Grains from Latex of Euphorbia-Terracina and EuphorbiaTirucalli Proceedings of the Indiana Academy of Science 82 132

Mahlberg PG (1974) Scanning Microscopic Comparison of StarchGrains from the Latex of Euphorbia -Spp Proceedings of the IndianaAcademy of Science 83 83-84

Mahlberg PG (1975) Evolution of the Laticifer in Euphorbia asInterpreted from Starch Grain Morphology American Journal oBotany 62 577-583

Mahlberg PG (1982) Comparative Morphology of Starch Grains inLatex from Varieties of Poinsettia Euphorbia-PulcherrimEuphorbiaceae Botanical Gazette 143 206-209

Mahlberg PG Davis DG Galitz DS Manners GD (1987) Laticifers andthe Classification of Euphorbia the Chemotaxonomy of EuphorbiaEsula L Botanical Journal of the Linnean Society 94 165-180

Metcalfe C Chalk L (1950) Anatomy of the Dicotyledons At theClarendon Press Oxford

Miller K Webster GL (1962) Systematic position of Cnidoscolus andJatropha Brittonia 14 174-180

Pax F (1884) Die Anatomie der Euphorbiaceen in ihrer Beziehung zumSystem derselben Botanische Jahrbuumlcher fuumlr SystematikPflanzengeschichte und Pflanzengeographie Leipzig 5 384-421

Payne WW (1970) Helicocytic and allelocytic stomata unrecognizedpatters in the Dicotyledonae Am J Bot57 140-147

Pray TR (1955a) Foliar venation of angiosperm II Histogenesis of thevenation of Liriodendron Am J Bot 42 18-27

Pray TR 1955b Foliar venation of angiosperms III Pattern andhistology of the venation of Hosta Am J Bot 42 11-18

Radford AE Dickison WC Massey JR Bell CR (1974) Vascular PlanSystematics Harper amp Row Publishers London New York

Radlkofer L (1870) Ueber Pausandra ein neues EuphorbiaceenGenus Flora 53 81-95

Raju VS Rao PN (1977) Variation in the structure and development ofoliar stomata in the Euphorbiaceae Botanical Journal of the LinneanSociety 75 69-97

Raju VS Rao PN (1987) The Taxonomic Use of the Basic StomataType in the Generic Delimitation of Chamaesyce EuphorbiaceaeFeddes Repertorium 98 137-142

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2424

Rosowski JR (1968) Laticifer Morphology in the Mature Stem and Leafof Euphorbia supina Botanical Gazette 129 113-120

Roth-Nebelsick A Uhl D Mosbrugger V Kerp H (2001) Evolution andfunction of leaf venation architecture a review Annals of Botany 87553-566

Sehgal L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 2 Venation Patterns Botanical Journal of the Linnean Society 68 173-208

Singh G (2004) Plant Systematics An Integrated Approach SciencePublishers Enfield

Solereder H (1899) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Solereder H (1908) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Aldhebiani and Jury 191

Stace CA (1965) Cuticular studies as an aid to plant taxonomy Bulletinof the British Museum Natural History Botany 4 3-78

Stace CA (1989) Plant Taxonomy and Biosystematics Hodder ampStoughton London

Tognini F (1897) Contribuzione allo studio della organogeniacomparata degli stomi Institut de Botanique University de Geneve 41-42

Van Cotthem WR (1970) A classification of stomatal types Botanica

Journal of Linnean Society 63 235-246Wagner W (1979) Reticulate veins in the systematics of modern ferns

Taxon 1 87-95Webster GL (1994a) Classification of the Euphorbiaceae Annals of the

Missouri Botanical Garden 81 3-32

7232019 Anatomical Studies on the Genus Euphorbia l Saudi Arabia Subgenera Triucalli Ermophyton Esula and Chamaesyce

httpslidepdfcomreaderfullanatomical-studies-on-the-genus-euphorbia-l-saudi-arabia-subgenera-triucalli 2424

Rosowski JR (1968) Laticifer Morphology in the Mature Stem and Leafof Euphorbia supina Botanical Gazette 129 113-120

Roth-Nebelsick A Uhl D Mosbrugger V Kerp H (2001) Evolution andfunction of leaf venation architecture a review Annals of Botany 87553-566

Sehgal L Paliwal GS (1974) Studies on the Leaf Anatomy of EuphorbiaPart 2 Venation Patterns Botanical Journal of the Linnean Society 68 173-208

Singh G (2004) Plant Systematics An Integrated Approach SciencePublishers Enfield

Solereder H (1899) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Solereder H (1908) Systematische Anatomie der Dicotyledonen Ergaumlnzungsband Enke Stuttgart

Aldhebiani and Jury 191

Stace CA (1965) Cuticular studies as an aid to plant taxonomy Bulletinof the British Museum Natural History Botany 4 3-78

Stace CA (1989) Plant Taxonomy and Biosystematics Hodder ampStoughton London

Tognini F (1897) Contribuzione allo studio della organogeniacomparata degli stomi Institut de Botanique University de Geneve 41-42

Van Cotthem WR (1970) A classification of stomatal types Botanica

Journal of Linnean Society 63 235-246Wagner W (1979) Reticulate veins in the systematics of modern ferns

Taxon 1 87-95Webster GL (1994a) Classification of the Euphorbiaceae Annals of the

Missouri Botanical Garden 81 3-32