Survey
* Your assessment is very important for improving the workof artificial intelligence, which forms the content of this project
* Your assessment is very important for improving the workof artificial intelligence, which forms the content of this project
115-173 27(1981) BLUMEA Comparative leaf anatomy of the Asiatic J. Koster Rijksherbarium, Myristicaceae & P. Leiden, Baas The Netherlands Contents Summary 116 Introduction 116 Materials A survey 117 and methods of the leaf anatomical characters in the Asiatic Myristicaceae 118 The indumentum 120 The cuticle The 118 unspecialized epidermal Some miscellaneous 121 cells epidermal features 122 122 The stomatal The hypodermis 123 The mesophyll 124 The midrib 124 The veins 125 The petiole 126 complex 126 Crystals Secretory 127 structures 128 Sclereids Generic 128 descriptions Explanatory 128 notes Gymnacranthera 129 Horsfieldia 131 Knema 136 Hair 140 types of,Knema 141 Myristica Value of the characters Taxonomic Introductory Below 146 151 implications the remarks 151 level 151 genus Gymnacranthera 151 Horsfieldia 152 Knema. 153 A tentative phylogeny of the hair types in Knema 155 156 Myristica On the On the Remarks genus level family on 158 level 159 161 ecology 162 Keys Synoptical key to the Malesian Synoptical key to the groups species of Knema and varieties species of Gymnacranthera 162 163 Acknowledgements 164 References 164 VOL. BLUMEA 116 27, I, No. 1981 Summary The hairs, of leaf anatomy Knema Horsfieldia, the cells of which characters important proper on Winteraceae. Guard and Knema of an abaxial free The cells and Myristicaceae bundles of the Many relation in the family (e.g. within can oil the mesic markedly cells. The subsidiary alveolar material hitherto layer was a of the overlying complex in The vascular system of the midrib is Myristica. latter is absent from into only reported for stomatal paracytic are arms be classified further can cutinaceous, such length by typically arranged bordering cells, leaving a (the and be separated on Leaf paper). anatomy and Canellaceae, cells, order other many anatomical variation easily star- composed and there Gymnacranthera). are varying be can leaf anatomical used for characters is species grouping leaf anatomical characters (see lends close little the although leaf paracytic stomata) to support shows family anatomy a points table and of typically isolated fairly a and affinity several to V, Magnoliales. of the leaf anatomical characters to the in value of these of the leaf Annonaceae and with ring of layer of Knema uniseriate characteristic, centre. of this end a bundle the the four genera at a have cell and the relative angiosperms overarched forming Much Magnolialean characters position and and taxonomic identification, and have often embedded in the are is sunken discussed. synoptical keys Myristicaceae adaxial collateral phloem diagnostic analysed of Myristicaceae (Gymnacranthera, Myristicaceae The hairs genera. the abaxial leaf surface. In Knema and and the Asiatic of the genera detail. The number of arms per arms. to separate Myristica shaped opening in always have in is described Myristica) Many species different types. the cuticle of the four Asiatic species 60 c. and ecology of Myristicaceae of the highly xeromorphic. are This is discussed in family. Introduction In 1897 his published Warburg of the monograph another recognized 15 attention the remarkable hairs, present in this revision to of (Sinclair, In genera. the four Asiatic 1958a and b, 1968 his work died in and has 1961, started the as species. Complementary explore the leaf anatomical can often add affinity, in Solereder publication recorded fragrans. (1895) strands on Asiatic genera, with an hairs species the studied. Of one species a of the new has been in studied the and it per genus De and recognized five (1950). patterns some to of The leaf can be Warburg's Worsdell petiole 83 since leaf specimens. foliar sclereids in Armstrong (1975) two been Wilde worthwhile Myristicaceae, Chalk midrib of (1908) Myristica species of the included leaf anatomy in Horsfieldia respectively. c. 60 species, belonging Knema. In order species, and seemed mentioned before. the of Knema and on material has A short characterization Metcalfe and in When he Asiatic genera for Flora the Asiatic scarce. species 6 years, a Malesiana Horsfieldia. 1979), work Flora of taxonomic boundaries and is and he paid special recognized several Wilde, revision leaf anatomy of emphasis last (De within phloem (1975c) anatomical variation within were and in the identification of sterile 1908) (1966) and Wilson study comprises Wilde's Myristicaceae the their studies of several This 1975) revision genus knowledge medullary Rao and Chin family. Siddiqi new diversity help (1899, on of De to our and be of anatomical literature found to a first for and about 50 in the available 1895) Myristicaceae Myristica he Sinclair worked towards family. remained unfinished. Since much Malesiana, with Knema natural the in which Myristicaceae, (Warburg, 1974 and 1968, become (Rijksherbarium, Leiden) anatomy of genera 37 in Knema, 72 in Gymnacranthera, collected publication to specimens or ten gain of some most specimens to the four impression species were or of leaf varieties studied. J. Koster & P. Baas: Leaf MATERIALS The material reduce the chance of in the or material from of Knema the Rijksherbarium, from the Kew Herbarium. To came leaves but selected, were the with flowers, specimens material identified De Wilde by , Fully developed sometimes case in specimens present misidentification, only identified by Sinclair, 117 Myristicaceae METHODS the material of Knema korthalsii Leiden; only taken. AND from dried taken was of Asiatic anatomy of the study younger material, when the hairs had been shed from the older required mounted in Euparal, leaves. Permanent slides, were made of transverse microtome sections of the central part of the lamina and both distal and basal end of the and of free hand made and For were ether and leaf electron hour in an warm Measurementsand at their widest linear eye measurements were were measured, veins were of distributed over counted; made at (Warb.) Sinclair; Borneo: Rossum63. van -- Jacobs 5259; G. var. van (Houtt.) Warb.; 14970; Ashton & Java: — Subramanian Warb. var. Borneo: a between the veins areas least 7 at on areas (0.063 2 mm margin ), was made. were 18969; H. 1407. — curtisii; Wood SAN var. K. S var. eugeniifolia; Borneo: H. — Sanusi b. Tahir S cult. BW2796; Philippines: Merrill4031. — Robinson Koster 235; Warb.; Borneo: BW 1123; ex. 3944; Rosli Warb. bb 23779. Theunissen West New Guinea: — BW Kuswata & Kochummen Toroes Borneo: H. macrocoma H. parviflora (Roxb.) Schram 4446; Kostermans Malaya: Moluccas: & Kadim bin Sinclair si 12325; Sumatra: irya (Gaertn.) Warb.; iryaghedhi (Gaertn.) Warb.; Sinclair; Rahmal paniculatai (A. DC.) zippeliana (Miq.) Sinclair; var. Guinea: G. Haviland 7241, 13696; Rahmat si Roepa (Warb.) Sinclair; — borneensis var. 5326, 5363; Sumatra: Sumatra: coll. 6591. King's 44045 ; Kostermans 17514; crassinervis var. Tassim 1839\ Sinclair; 10437. van — H. .H. Royen 4510; Soepadmo 3; Pleyle KEP80627; van (Miq.) 379. superba (Hook. /: & SFN 40047. ashtonii; 23314. — Borneo: K. Sibal ak attenuate Luang Ashton A4139, Ashton S BRUN 18294. 5587; Malaya: — S (Hook, / conferta (King) Warb.; Singapore: Borneo: Sinclair Borneo: Jacobs et al. SAN Sinclair 5749; Junghuhn 230. West New Ashlon <6 FRI Th.) Warb.; — Warb.; griffithii(Warb.) Sinclair; var. H. sabulosa Sinclair ashtonii Sinclair Murlhv cell guard measured with were magnification bancana; Singapore: var. Galau S Miranda bb 3081 /; Moluccas: Th.) Warb.; Singapore: Knema pole to 400 x G. contract Superficially studied: H. fulva (King) Borneo: at 11044. — 3908; Schram Kostermans 767. Gjellerup 407\ vision Sinclair Sleenis 5274. West New Guinea: sylvestris of forbesii; Malaya: glabra (Bl.) Warb.; Moluccas: pole per leaf. The number of basal parts of hairs G. forbesii (King) Warb. — paniculata; Philippines: Java: or The wax. any follows: The width of the out as from length eugeniifolia (A. DC.) Horsfieldia crassifolia (Hook./ Warb.; to remove per leaf for both midrib and areolae 9420; coll. 6622. Royen 5101; ultrasonically and the lamina between midrib and leaf (Miq.) Malaya; King's H. shaken were leaf. For the lamina thickness out Bujang S Whitmore FRI — of leaves studied. bancana 8035; Malaya: 59. acetic acid. The glacial glycerin-jelly. 1000 magnification in the free hand sections. Ten x field a counts 9437; Anderson 134; Zehnder var. per carried half least 7 Gymnacranthera 3393; at with were chloroform or boiled in water, alcohol or carried and the the midrib Specimens Billiton: and out specimens gold. counts were part of all measured, the tips of the papillae excepted. For both characters surface a ether, alcohol carried were petiole safranin- a for the cuticle thickness the thickest parts of the cuticle between the measurements were with mounted in microscopy fragments micrometer piece tests preparations hydrogen peroxyde 30% coated with were lignin acid. Cuticular stained with were rinsed in fresh dissolvent in order subsequently fragments pairs specimens some stained with Sudan IV and scanning for half For hydrochloric volumes of using equal cuticles sections. The sections paradermal mixture. haematoxylin phloroglucinol was indumentum K. furfuracea 22841\ & Sinclair Whitmore (Hook./. Th.) var. cinnamomea Warb.; India, SFN 40046. FRI & 12326. Th.) K. — — Warb.; K. de Wilde: Peninsular: curtisii elmeri Malaya: (King) Merr.; Kadim & BLUMEA VOL. 27. - No. 1, 1981 118 — Mahmud 22. K. 2488. K. Kochummen Malaya: Borneo: Chew de (Sinclair) mandarahan K. - oblongata; Borneo: Java: Ja. Mikil — 2502. K. S 18478. — Sumatra: Warb.; 24985; K. Java: — Dransfield 3137F K.K. laurina Witde- de Wilde; linguiformis (Sinclair) Plants Warb.; malayana (Bl.) Sumatra: de Wilde & de Assam: Steenis van — Chai S 34822. K. K. hookeriana — Warb. ssp. kunstleri; (King) SAN FRI 2431: 12308. (Bl.) Warb.; K. kunstleri linifolia (Roxb.) Warb.; India, (Miq.) Bakar SA N — 143; Kochummen 41273; <£ Smitinand K. intermedia Borneo: latifolia Warb.; SAN Hansen 12207. — Cenabra FB 29973. Borneo: Chai Wilde; Thailand: Lorzing Wee Lek K. — K. — Ampuria Paie S 27799. llias lunduensis FRI 2252. Borneo: Duyfjes 14429; Warb.; Sumatra: Philippines: laurina; Malaya: var. 40714. Th.) Warb.; korthalsii i Warb.; — Warb. globularia (Lamk.) & (Wall, ex Hook./. 10092. of Assam — Merr. oblongata K. SFN Singapore: Sinclair K. — 6188. ssp. Chelliah oblongifolia (King)Warb.; Malaya: KEP 98224: Whitmore FRI0111. K. pallens ;de Wilde; Borneo: Meijer 2533: Lantoh SAN 73312. — K. Sinclair percoriacea Sinclair; Wilde; Borneo: Kostermans -K. forma sarawakensis Malaya: Carrick Anderson S 5103; Malaya: rubens(Sinclair) de 13963. 1510. I.: 24236. Houtt. M. fatua Koster Guinea: — 787. Beguin Mikil Corner Othman bin Sinclair 12890. BW 1016. M. M. BRUN Corner 32840). M. — 5365; Warb.; 12547. SURVEY 908. M. subalulata Kostermans Borneo: A — Miq.; Sinclair de Borneo: 9908. 9907, Unyong S New Soepadmo 57; — Sulit M. lepidola Bl.; 1289. East A. New Guinea: New s.n Guinea: 121 Lundquist Sumatra: Warb.; West Smith; Floyd Vriese 37177; Kostermans 4873; Guinea: M. maxima C. New de Sinanggul SAN 39995; 7; Borneo: West New — SAN Markgraf; papuana Moluccas: SAN Brass 15066; Singh globosa Warb.; Papua Kostermans UNESCO M. var. 15023; SAN32924; Aganb. Ambulah I.: Guinea, Fergusson Wood SAN Borneo: New & Havel NGF Brass Guinea: 7446. (=bb Thorenaar M. — & villosa Wood SAN A4747. 6438; OF THE s.n. sphaerosperma M. — plumulosa Warb.; Borneo: Asah anak Sinclair; King; Kusxvata & Scortechini - 5056. rufa K. —.K. psilantha pulchra (Miq.) Warb.; (King) Warb.; Malaya: K. Houtt.; Philippines: Java: AbanGibot 1403. 4197. Moluccas: fragrans FRI Malaya: — MS. Thailand: & Geesink c.s. 4747. van Beusekom —K. maingayi Hook./; Malaya: Maingay (Endert) 55E1P624; Grashoff Versteegh Vogel guatteriifoliaA. DC.; SAN30298; Elmer 20850; Wilde; M. cinnamomea fatua; — retusa S 23645. Luang K. S 29064. entrecasteauxensis var. i King; Malaya: gigantea M. — de —K. Sibat ak Wilde; Malaya: Shah MS. 1312. setosa de Brass 25893. var. Borneo: Haron SFN 29015. Moluccas: chrysophylla 25986; Normanby — Wilde; Wilde; Malaya: Shah (Miq.) Warb.; Myristica Carr 14373; K. tenuinerviar de Wilde ssp. tomentella de K. pseudolaurinai de — LEAF ANATOMICAL CHARACTERS IN THE ASIATIC MYRISTICACEAE The indumentum (figs. Hairs present, are Horsfieldia, Knema, abaxial surface frequent on Myristica (in at and on frequent most in both 1 —10, 19, 21, 25) the adaxial and abaxial surface on examined of and sides in the on leaves, species most Gymnacranthera the midrib , 1—3, photos least in young some or Myristica species), Horsfieldia lamina in Myristica) and only usually Knema and Gymnacranthera on and (in the most adaxially in abaxially in Myristica. The indumentum hair type: words, each, a a hair is length in In composed two, one or excepted. appears very rarely of Gymnacranthera The appearance of a the and uniseriate consisting of a few cells consisting of numerous The cells, having the cells have may and the length tall a 'body' hair, which which will be discussed are occur. sometimes hairs as or two of the one arm seen and is basic arms from same (fig. 2,3; above, the Knema, the walls variable. Short to filiform, In Knema the separately. a In other of each cell very small; this may (most typical for Gymnacranthera) cells, one two arms, not Horsfieldia of the 1895). the so-called stalk cells, and Knema have give In recognize one can (Warburg, epidermis, Horsfieldia arms oblique nature. to the (e.g. photo 7). 250 /tm per cell in Knema, may forms, Myristica the arrangement of stellate hair of short nearest the cells in between the cells may be very obscure cells per genus, but uniseriate trichome one row more Gymnacranthera ; photo 4, 9). different sympodially branched, with species hairs, extremely long hairs, a show length of a about range of hair In older leaves the hairs have often been J. Koster & P. Baas: Fig. 1. deciduous part Diagram and of the lucida Fig. hair); drawing 3. g See = Myristicaceae in of the text the hair. — hair; e = a surface view different for definitions = arm; cutinized unspecialized epidermal of hair Diagram derivations. of persistent part of cells. b = ring; — body; f = Fig. types of different in Knema. types. — c = stalk basal cells 2. (e cell; of arms Arrows d = line and f form Gymnacranthera showing unequal length hair 119 Leaf anatomy of Asiatic Myristicaceae of hair indicate separating together the contracta. cells, x — the basal Camera 400. tentative phylogenetic 120 shed, but of this basal a small cuticular wall elevated. to cells' for epidermal The hair index specimen in or even but variable, studied. It is at of fact (in it is essential know When necessary not. or low power with magnification cuticle After structured 11 —18, (photos in boiling this has An identical structure, referred Winteraceae In the Asiatic Bongers (1973). species examined of material usually the overlying it is only Knema slightly present and stomatal species a this to positively with acetolysis it of both acid IV. the leaf warm material, chloroform, the and the cells the on cells Horsfieldia sylvestris sunken alveolar by of the but in Knema present in 12); cells. The have studied most When present the guard cells, margin, (photo 28), (photo a has been recorded thoroughly was only conspicuously stomata not cells in subsidiary material (see under 'The (photo and From sulfuric acid. (used the porous 16). structured species 14) layer is or in Besides the alveolar not as cuticle (but present in some 21) and a erect warts present Asiatic (in material, most Horsfieldia and 15) 11), in 13), press) for of Myristica but also present the abaxial surface in the and to this cutinaceous. (photo are stained moreover solvents organic 9:1 was species a it is dense more with rod- in this genus. An Horsfieldia iryaghedhi a more elaborate description of the Myristicaceae. there may layer overlying species, is a alveolar material and Knema is composed occur (photo on to plates (photo plates The palynology) resistance composition with See Kosterand Baas alveolar material in the and Knema has been boiled in Gymnacranthera concluded that the irregularly structures (photo is in the abaxial side in many on Myristica species. alveolar material in Gymnacranthera irregular, the by studying because the absence this material is absent in some subsidiary do treatment coarse variable in appearance: (photo it anhydride Sudan was reticulum of like are fully complex'). mixture of acetic The the on alveolar epidermis except cells of the usually Myristica One leafof resistent and in plumulosa subsidiary in have persisted been checked (1944), and Myristicaceae the whole Knema the on rough a leaves fully developed shaking or to as Nast midrib, the major veins and intermedia and bordering and Horsfieldia covers use. per surface unit is microtechnical artefacts. to ether, or for the Bailey slightly patterns of nevertheless gives that the cuticle proper remains species. by small one 21, 28) water, alcohol layer, overlying hairs) whether hairs of hairs from anatomical slides may be due The and 'oval dissecting microscope a this Myristica these patterns with the hairs. determine and density, provided to and oval, subtend basal cells may be basal parts of the to epidermis and is therefore of little taxonomic 1965), easy the may be very variable in leaves of one density remaining more course leaves developed leaves leaf(Stace, one not connect on (in Horsfieldia or and described 'circular' did of hair ring to numerous fig. 1). These have epidermis cells. The upright walls cutinized a Gymnacranthera 25 and position (photo of the difference in hair impression in Usually as the nearest characteristic circle a Armstrong (1975) The number of hairs stalk cells Two (fig. 1). in cells, arranged a measure 1981 1, unspecialized epidermal Horsfieldia, but as No. of the hair remain (photo 21). circular cell takes this 27, one or two have the as of the hair has been shed rest Knema) mainly sunken left: always walls, proximal part most after the is part cutinized strongly also VOL. BLUMEA be the cuticle abaxially in faintly adaxially in conspicuously sylvestris, other types of sculpturing of the proper). Striations (fine ridges) Horsfieldia some are crassifolia Knema species. J. Kostkr Grooves corresponding present in & P. Baas: with the anticlinal walls Horsfieldia, Knema sculpturings of the cuticle are that only clearly distinguish ridges, one can electron scanning adaxially. way The surface In this because of the The cuticle although a The 1 in or been studied. the Only of the vein is course In epidermal cells). are the on is very major midrib and the areas veins indeed thickness all less in striking straight, proved over. in nearly all are are perpendicular rather than perceptible to square the are thin some the cell (photos in fairly study, view. of the veins. course and straight to they on Adaxially they have not is when (that the minor veins epidermis only; are in Knema adaxially. curved are usually lower focus at less sinuous. Thin are cells long the overlying pattern of Myristica the adaxial surface slightly The with the Venation patterns of the abaxial visible faintly constant, 13, 16, 20, 22, 30, 34) surface rectangular, to rectangular. pattern flanges areas are areas of species where In present. of the cuticle areas areas of more or (photo 22); of cuticle of of the cuticular flanges frequencies veins and the and leaf the midrib and the transverse the only: the even on some and Gymnacranthera thin but in low major this (>Gymnacranthera species of minor veins has been recorded veins be thin Pitting in present in the loops of the undulations. This character species so. is species in the macerations in the and major light occur. On the abaxial surface the cuticular specimens, the midrib, on to these inconspicuously to Gymnacranthera (photos 22,30), prominent In sinuous. found veins major prominence varying conspicuousness the sections. considered one cells focus and high at of specimens polygonal prominent abaxially in obscures this feature in transverse been not cells epidermal the abaxial surface, granular appear present, epidermal In the macerations the cuticular less sinuous has Gymnacranthera, Horsfieldia visible in the slightly cells may when The thickness per two epidermal polygonal on of the alveolar material, varies between 2—18 /rm of 9 /im has been some be to with the sculpturings of this material. abaxially. yum 1965) material, feature, nature the midrib and the tend 18). It should be noted have been examined this specimens be examined in the must difference between parallel they 11 — unspecialized usually the it important ephemeral abaxially) overlying axis case an unspecialized The Stace, thickness, exclusive and contracta, (sensu convex of the cuticle. sculpturings layer although adaxially all rarely are Adaxial 17). and other grooves not cells epidermal (photo present in Knema rufa (photo smooth. The alveolar or view. Wax, of the Myristica Moreover, alveolar material, when present cuticular microscope and microscope. However, may obscure other 121 Leaf anatomy of Asiatic Myristicaceae outer flanges are mostly Horsfieldia they cuticle be may (cf. Bongers, where the same more or are slightly present, 1973) least inconspicuous, margin, sections these wall has the but is present in inconspicuous flanges are most straight. In to transverse rectangular, The abaxial (photo 34). the sections the adaxial on unspecialized conferta they unspecialized epidermal are epidermal The cuticle of these cells is measurements Abaxial the midrib often cells are slightly usually thickened more on and Knema occur in linifolia), many in species of Knema Horsfieldia iryaghedhi in are less or usually square Gymnacranthera). dome-shaped the top of the dome. For of the cuticle thickness these thickened papillae cells dome-shaped (not parts have been taken. in Knema (most striking and in Myristica gigantea 122 and Myristica maingayi (photos and Knema, where the papillae the Myristica cuticle may Often and in and surrounded occur on veins the adaxial and is or properties, In the less are and are However, wall has whereas in strongly thickened the (photos 28—31) hole in or a sections to on the cells. They the appear may major large as situated in extent (in are of the centre midrib, the cells large a the about circular in epidermal mostly cells epidermal only relatively small, restricted surface a this case the the cell is cells. In view of surrounding epidermal present of several cells, are thin cuticular relatively narrow to are they epidermal cells, and short cuticular are sharply delimited, not flanges (photo 30). From connected with this feature. The in Myristica adaxially, which are large flattened, having flanges (photo 31). and Gymnacranthera The areas Horsfieldia absent in Knema. each specimen examined, but usually in small numbers. present in distributed and irregularly more or distribution, in the Knema Wilde refers to these cork Knema sabulosa, less circular structures, identical frequent most are their origin is shaped, Horsfieldia Horsfieldia glabra, structures Horsfieldia dome-shaped strongly, a four genera oriented radially transverse adjacent and abaxially, kunstleri abound in These with in present warts are they areas idioblasts are vary have features all the sunken and by forms of probably secretory. are thin cuticle and relatively mentioned Since outer covered partly the cells deeply and crystalliferous Cork In not cells the abaxial surface, margin. The cell does usually are mentioned of 20). sections it appears that druses transverse adaxially as species thin cuticle a 1981 as extreme epidermal less epidermis, macerations many staining more or well as mesophyll (photo 29). pear-shaped) by the leaf along empty idioblasts in these single over with These cells (photo 28). shape or I, In the epidermal distributed species thin cuticle present with a very cell surface one top of the papillae (photo miscellaneous Some No. be considered mentioned the species the on VOL. 27. 13, 16, 20). cells, the thickness of the cuticle of a - BLUMEA the on species mostly 1979) and and Knema regular to surface, cork having a warts. regular the midrib and the veins. De along and on (De Wilde, warts abaxial traumatic. probably conferta uses them as a lens character of basal cells separate a group of species from others. In Horsfieldia glabra groups to of hairs hairs on the probably are in the leaf early of origin ontogeny, of the cork most but basal parts the adaxial surface the groups of basal cells small sometimes cells; surface relatively abound. Perhaps development frequently some and occurs The bancana veins a are are var. basal finally on a cells cork wart cell is become a to the the leaf shows abnormalities. a is is not from 13, species the whereas cork course Why they are leaf this suberization understood. Moreover there is a are stomatal Sometimes complex. recognizable in a cork wart. 23—27) epidermis; abundant on the If warts of the however, in Gymnacranthera very small number is also present in the adaxial margin. leaves numerous On the abaxial recognizable. in looses its fully developed and consist of suberized. are part of it) (photos This rarely present, subdivided only abaxial large are wart are developed pair (or complex confined borneensis along cork the walls of these cells randomly distributed, and to a the abaxial surface guard stomatal Stomata They the evidence that the outlines of transition a small basal parts of hairs warts. remain. In the areolae, scanty on the epidermis. midrib, the present here, the stomatal complex J. Koster & P. Several authors variable within Baas, 1973). The 39 f.im for the but the cell guard and a of type in little variation in some not are range from 8 and a is paracytic Knema Therefore, the genus. specimens of Gymnacranthera study. 27). short single one leaf stomata be up can to are to 8 the average values show data for the stomatal sizes per have been recorded for and Gymnacranthera forbesii contracta The (in Myristica relatively relatively stomata water or (photo 21 fim for the width and from 15 to length respectively. However, in tables. Giant given extremely been determined in this not The variation in occurs. be may 1973; Van Staveren & different size range a Gymnacranthera large overlap stomatal index examined species pairs and 12 /<m for the width and specimen the 123 Myristicaceae 1973; Jansen & Baas, The genera have length. stomata fairly that out Asiatic oj anatomy Stomatal indices have therefore stomatal present), pointed Lea] species (Bongers, a dimensionsof the long have Baas: var. forbesii only. The guard cells cells subsidiary fully Gymnacranthera often embedded in the are and Gymnacranthera in Knema; here, enclose the guard cells, such sections also show subsidiary cells, section a the Outer rectangular. stomatal ledges are the to the pore, In protruding subsidiary lowly dome-shaped well in conspicuously excepted (photo 24). sometimes are fairly sides poral pointed, cells, whereas the unspecialized epidermal cells most perpendicular in developed or square to species of complex in some Horsfieldia only. A peculiar Knema and Knema character is formed Myristica. species the stomatal level than the other cells transverse cells characteristic cells. A sections show the cuticle, overarched by epidermal more or (photo form. structure' is formed 24); Siddiqi 11 to varying in stated As less height per species, the elsewhere, directed (1975c) cells. This papillae papillae to on a by (photo 26). less In surface above the stomatal unspecialized epidermal of the (photo cells in a thin bordering the star its 'star-shaped the be incorrect. In to more or lower epidermal 6 stoma give that proposed appears overarched ring 4 by of the position nipple-shaped cells form a left sunken stomatal complex with horizontally partly bordering epidermal indicates the Wilson by hypodermal observe, lying to star-shaped opening, strongly these and the sunken stomatal complex is the 4 the stomatal surrounding itself is difficult complex epidermal the stomatal complex, overarching The 23). the cells by In the cuticular macerations and the free hand sections of the Myristica upright papillae view these of papillae, complex (photo 13,25). most species are without papillae. The hypodermis An adaxial the epidermis, some only Knema In has and developed, with the Horsfieldia as a continuous been recorded for species locally intergrades (photo 32) hypodermis specimens especially are cells of the adjacent to tissue Myristica also of and Myristica species. Myristica guatteriifolia near the midrib the occur, hypodermis occur in the and have the periclinal hypodermis, the adaxial and sometimes elevations of the leaf surface ground of of translucent cells, distinct from Horsfieldia the hypodermis major In is veins, and palisade parenchyma. some genus swollen cells with thin walls these cells layer some (photo 32), and/or midrib and sometimes the petiole, probably are to or the abaxial present where in the these of traumatic division walls. In this few-layered periphery cells origin. groups of epidermis, in in of the Knema and BLUMEA 124 An abaxial inconspicuous 27, No. VOL. 1. 1981 is present in hypodermis Knema some and in species Myristica sphaerosperma. The mesophyll The cells of the virtually four adaxial to with the intergrades in range from a have i.e. lignified; one sclerified Either the cells possibilities. walls birefringent spongy tissue is (in some of Knema and Gymnacranthera, (secondary in the only of walls in the adjacent to N o t Myristica) the cell sectioning thickness because epidermis, This characters thickness of robust. The Horsfieldia, there is species midrib a with is abaxially Myristica The vascular system is phloem often an sections (in the whole or species some In of cells have palisade parenchyma Knema mandarahan large pits (in strongly birefringent the leaf margin the cells, the some in soon, and for the transverse domestic bleach of bad to be lamina quality thickness reservation domestic are damaged made and the thickness not in (and or and the in the study of midrib of to Horsfieldia specimens only, bleach). of lamina 3 and less: quite remarkable. abaxial arc-shaped by (in some in the and adaxially and there is a two are: more: raised in Myristica species and some the adaxial surface. on sometimes in more or less straight joined together. One layers. to The numerous tissue between the main bundles, ground elements xylem Generally bundle, ratio 6 Gymnacranthera to concave often strands, Horsfieldia but in less flat more or interspersed are accompanied well fine; and raised, prominently as ridge adaxially), in separate arranged bundles range of studied is fine to robust; criteria for these characters the midrib is adaxial bundle and is whole a and Knema) the dried leaves break leaves; behaviour raised central Horsfieldia species phloem in without cells with expand completely in midrib/thickness Knema and only and (figs. 4—7) The midrib of the ratio the Horsfieldia). consequence, swelling layer few leaves demonstrated to a mesophyll with such cells or even in have to be taken with in layer isobilateral leaf. The an width ratio per to three, to abaxial have sclerified walls. the transverse difficult. In of lamina ratio midrib caused to walls in the extents species have brittle walls cannot be made of the variation The often Horsfieldia species process. anatomical also are mesophyll (particularly the Most e: sections There pallens). two most abaxial layers of the spongy tissue have thick, strongly birefringent walls, supplied and Knema length Gymnacranthera variable to sabulosa has different a occurs species interspersed the five. to cells) mostly palisade parenchyma, Horsfieldia of cell walls Strong birefringence be tissue. spongy of with dorsiventral, always layers palisade parenchyma species, per is mesophyll sometimes up (fig. 4); these situated centrally phloem bundles have been mentioned before by Worsdell (1908) for Myristica fragrans, by Siddiqi and Wilson Horsfieldia species. and In Myristica. pith (fig. 5); in (1975c) for some some strongly interrupted, sclerenchyma (figs. often 4 — 7); even Knema or there (fig. 7). fibres, there is species are a and by Armstrong is present in complete which centre phloem often occur of the (1975) Horsjieldia, for Knema collateral bundle in the the adaxial bundle is several small bundles (fig. 6). In Gymnacranthera The whole fibre groups also in the species Horsfieldia and Myristica specimens the adaxial bundle is absent, but described above Knema This type of vascular system bundles occur vascular system are associated in the phloem centre bundles in the centre of the midrib is surrounded with peripheral by groups phloem as of groups of the midrib, associated with and (figs. 4 — 7). J. Koster & Figs. 8: 7. 4 Gymnacranthera basal part The Baas: Leal 9. Vascular patterns in midrib and — — 4. Knema 16. x P. of laurina, midrib. — Lines: — is often black: of Asiatic Myristicaceae Camera lucida midrib. lunduensis, 8. Knema xylem; tissue ground collenchymatous, petiole. 5. Knema contracta, midrib. petiole. anatomy from centre interspersed to white: 4, 6, 7, and 9: x 40; 5 and Horsfieldia glabra, midrib. — drawings. 6. — linifolia, distal part sclerenchyma; of petiole. — 9. Myristica lepidota, phloem. periphery to various 125 extent parenchymatous cells with with walls, especially in Gymnacranthera and Knema, in the former mostly abaxially; these genera there midrib. Often the one- to of collenchyma Gymnacranthera sclerified The The flat often several or cells, veins major or the if and midrib, layers of these cells at the sclerified cells appear several-layered hypodermis continuous in or are on periphery the as a in rarely situated abaxially Knema of the in of the locally developed the adaxial side of the midrib. In Myristica, to sclerified adaxial specimens chlorenchyma , collenchyma or is the layers of present. (photos 33, 34) veins are adaxially raised; abaxially they either are slightly depressed (not always raised. in Gymnacranthera), 126 The veins veins and a caps by constituting one number of Knema sheath, which abaxial more less or well-developed two of row bundle sheath. be continuous and A sclerenchyma fibres often have vertically either the adaxial birefrigent or walls and the veins, transcurrent in are caps each at thus side, differentiated usually poorly sclerenchyma. the abaxial the cells of these bundle sheath extensions epidermis have Myristica (little developed In a fairly well-developed parenchymatous bundle a to major intermediate between that of Horsfieldia continuous there is species may (photo 33); from these the Knema the sclerenchyma fibres; bundle sheath surrounds the bundle and the parenchymatous both 1. 1981 system, Gymnacranthera, slerenchymatous a No. the adaxial and abaxial side of the bundle In Gymnacranthera). connected veins. at vascular complex more minor sclerenchyma VOL. 27. with collateral bundles and supplied are have may midrib or - BLUMEA veins are are epidermis adjacent or to the adaxial to probably lignified. Apart embedded in usually the mesophyll. Another character recorded for Knema outline, fibres sclerenchyma these and position, in the places distribution sometimes only; transverse section homologous with minor veins. The (figs. 8, 9) petiole The vascular system vascular system of the Myristica) the of the midrib, adaxial the petiole phloem consists of three bundles is sclerenchyma Brachy- more or Crystals idioblasts, in the a have and Knema fig. second and the are crystal tissue 10g), of found of consisting is may with comparable mostly the Knema and three to up at be considered (in Horsfieldia, bundles, large vascular system of the basal end of collateral bundles, also with free 9); the the abaxial sides. Sometimes to in (not these close and Gymnacranthera), transverse interspersed in the mentioned sections ground were not from tissue of the petiole. occur of the in the by midrib (i.e. 'Some mesophyll restricted in enlarged miscellaneous of species, most the to palisade ordinary parenchyma midrib and in is occurrence (fig. 10a). Metcalfe and Chalk exists spindle-shaped are druses epidermal cells of cells; these Gymnacranthera. have been found in the is occurrence may tissue type of frequent the Large under been found in the their (fig. lOh) ground before They particles also present in the (1950). epidermal are cells represented in cells of the bundle sheath of the veins. Most mentioned that more in the leaves studied. specimens they when druses structures petiole usually minute, particles, grouped parenchymatous crystals of probably often been found in not spindle-shaped, ground of the bundles, few fibres may be present petiole. Myristica Small druses mesophyll A These less arc-shaped been large diversity 31, photo druses have or a (photo 31, fig. 10) show parenchyma. vein the appearance of bicollateralbundles (fig. epidermal features', have frequently but be mentioned: structures, in to confined for the greater part usually astrosclereids Crystals (photo 34). (fig. 8). Generally the basal end of the to one row distal end adaxially, giving small collateral bundles extreme has but often the abaxial and adaxial to the main bundles. Most the only consists part bundles for the abaxial part only unlike the not are most probably In as some by more mesophyll abundant these are or less and the in the the acicular Knema and Myristica well, but here the possibillity fragments of epidermis. Relatively small druses, especially large spindle- to needle- J. Koster & P. Fig. 10. Crystal types. cell of bundle sheath (Myristica fatua fragrans). fragrans). in crystals — — e. h. small Flattened, specimens more in species of to a. spindle-shaped crystals needle-shaped crystals in prismatic crystal (corroded?) edges epidermal cells cells of abaxial tissue some druse in big in parenchyma in basal cells of a hair cell epidermal in abaxial (Myristica epidermal cells (Myristica (Myristicafragrans). g. — 127 Myrislkaceae — f. mesophyll irregular, idioblast small (Knema (Knema hookeriana). Myristica specimens, especially in 10b). prismatic crystals bancana and var. the cells crystals, in abaxial occur Myristica have crystals with four to smaller than the solitary makes, together (fig. lOd). 630. cell of spongy ordinary less or x of Asiatic of the in occur several parenchymatous cells epidermal Myristica species in or, bundle sheath, in some mostly 10c). small Irregular abaxial epidermal bancana all spindle- (Knema hookeriana). (fig. of Knema, in solitary (fig. clustered, cell druse in occur Gymnacranthera in crystals the basal cells of hairs b. flattened c. anatomy drawings, with rounded crystals epidermal shaped crystals — fatua). var. clustered — lucida Leaf — ( Knema hookeriana). d. flattened abaxial hookeriana). Camera Baas: six rounded study if edges; Often ones. edges, rounded grouped, cells in Knema hookeriana peculiar crystals also with the rounded Further with usually epidermal more they edges, in the epidermal than have a one in a sometimes lOf). cells: cell, they Some flattened usually are corroded appearance, which composition a (fig. of calcium oxalate doubtful would be necessary to determine the exact nature of these crystals. Druse-like composed clustered of crystals relatively few, Intermediate forms have been edges and have been prismatic crystals. reported Secretory The large, to enable or have been found in the less in Myristica fragrans, irregularly found of clustered shaped sex distinction of but crystals unlike particles crystals, crystals The different types of structures more occur large, in (fig. druses lOe). with rounded Myristica fragrans seedlings by Nayar et al. (1977). (photos 34, 35) spherical mesophyll oil cells, and the as mentioned in the ground older literature, tissue of the midrib in most of the 128 BLUMEA in great specimens studied, usually There Warburg's remark indications (colour to contrary are of contents these West mucilaginous. large cells idioblasts of the 1981 1, that and they also in staining tannin-like that properties) (in or in occurring and for mutually the irregular confirmed in this in strictly that fact, some and not somewhat study. studied leaves peculiar (photo sometimes be Metcalfe in the protruding even the so-called and tanniniferous Chalk In the not S been c le r e id Sclereids in Knema and they often present in Myristica as the to which Astrosclereids varying they occur is in amounts as clustered are the The traces...' for study, been of the vein epidermis, structures must according latter some these found to describes Horsfieldia have structures for astrosclereids (rarely tissue of the midrib, (photo 37). recorded tissue of the findings and for Gymnacranthera, midrib, most in two Horsfieldia Horsfieldia superba frequent in the spongy tissue of Rao and Chin abundantly fulva to ground high frequency with the Horsfieldia finding a brachy- in the have been ground with contrast have accordance with the in mesophyll, examined: superficially vascular caps the veins (1975). examined in this However, Myristica: adaxial these be cell followed and specimens. the situated in are brachysclereids only) sometimes frequently (photo 38), towards not identify of Knema and specimens Armstrong rarely branched sclereids and in the adaxially, staining technique herbarium could to observed. mostly abaxially; where literature, (photos 36—38) s are Filiform, some which mucilage epidermal features'; impossible epidermal layer (photo 35); Horsfieldia specimens clearly even species. per abaxially of the sclerenchyma and (1950) dried 'branching' sacs, '...tanniferous tubules... adaxial species. in and sometimes 34), On the whole it is were structures occur fairly thick-walled cells, adaxially bundles the cases Annonaceae he noted constant like this, because of the study a all some iryaghedhi) related species. The empty idioblasts in The presence of tanniniferous sacs, also mentioned in the contents in Horsfieldia also recorded tannin-like substances instead of oil in the Myristicaceae, their distribution is unambiguously Gymnacranthera, absent in this genus. probably are have been described under 'Some miscellaneous epidermis probably No. 27. frequencies (photo 34); (1895), are (1969) some cells instead of oil cells VOL. - as well (1966). Horsfieldia superba. of Rao and Chin species, This is (1966) in and Armstrong (1975). Infrequent, they thick filiform, branched sclereids abundant in Knema are bundle sheaths of the veins mostly to GENERIC Explanatory The generic expedient in individual the diagnostic species in some from the Knema species; slerenchymatous epidermis (photo 36). DESCRIPTIONS value only or descriptions specimens are infrageneric variation descriptions. adopted procedure present notes leaf anatomical several characters the such the are extending percoriacea, the For some generic very detailed. If mentioned in the presented given relevant and descriptions. For in tables and referred to variable characters range is makes individual is are of limited without further or specification. specific descriptions superfluous. as without The J. Koster & P. Baas: GYMNACRANTHERA WARB. Asiatic Leaf anatomy of Photos figs. 2,7; — 129 Myristicaceae 22,27,30,31,37,38; figs. 2,7; table I 6 5 species; studied. species surface In frequent on the stalk view: Hairs present the midrib, cells (up length, spread out decussate arrangement (fig. sometimes up oriented horizontally; epidermal cells and reticulum of cells Unspecialized epidermal focus and high thin conspicuous Abaxial straight but walls, cuticle on but rows sinuous in abaxial to the tending on near be in rarely present (in G. 6591). Cork origin. Small cells surrounded In 1). pattern of abaxial bancana 23 — x on 12—16 /<m, absent length with a thin sometimes cuticle in in present oriented the the epidermal section: to rectangular. to layer, G. bancana of about the cells overlying same size midrib usually small and erect, sometimes adaxial epidermal thin cells areas of cuticle epidermal cells flat lumen side. as to cells; protruding above subsidiary stomatal the in ledges centre of the cell and/or adaxial absent x 10—19 2.3, water average stomata coll. s of traumatic surface, probably abaxial 130 excepted. /un 410 — thick epidermis, on the outer Outer abaxially on periclinal periclinal sometimes cells. thick (table often cells slightly cells, sometimes lower; lumen side, Subsidiary surrounding epidermal nm (table I), usually Unspecialized epidermal wall complex complex slightly cells abaxial surface wall corresponding slightly sunken; guard absent. — or 26 — to adaxial cells. convex, cells 1.0 or probably the adaxial epidermal in surface view; slightly G. forbesii var. forbesii, King dome-shaped. mostly slightly Stomata and subsidiary as 15 pairs Giant sometimes erect, with indentations seen of cuticular present width ratio infrequent, or arranged Stomata confined adaxial surface often erect, on major parallel Pitting scarcely of without midrib and axis and areas abaxially as long Guard cell at anticlinal adaxial cells also Lamina dorsiventral, rectangular, usually flattened, dome-shaped, epidermal cuticular the inconspicuous. or contracta, Zehnder 9437 and by radially granular square focus abaxial surface. Minor veins paracytic. a (photo 22). high at abundant in areolae, scanty and aberrant T-pieces transverse a undulations overlying with borneensis var. as margin. lower focus and with at well as Adaxial cuticle 5—18 /<m thick, abaxial cuticle 2—11 with veins and leaf epidermis only. variously present, usually warts cells, rather than conspicuous stomata 18 adaxially square 1, radially of the hair. shedding slightly curved to veins; corresponding polygonal G. 1.2— 1.9. Polar secretory straight in to mostly less more or anticlinal walls Abaxial cells to veins and leaf margin, /<m, average values values of the most cell midrib; midrib, major G. bancana rectangular course to epidermal epidermis, epidermis with anticlinal walls. rows, flanges usually present, visible in or cells midrib, major veins and leaf margin with prominent thin arranged perpendicular in of cuticle, areas sinuous same with sinuous anticlinal walls curved anticlinal walls faintly anticlinal walls. along conspicuously veins cells with thin inconspicuous adaxially of the each other with alveolar material midrib, major on least 2 arms, mostly not to Basal after ring of cuticle between anticlinal wall areas epidermal cutinized a polygonal, usually parallel oval, surrounded by or is absent slightly to abaxial surface, to walls. granular, abaxially to plates, which erect thin with subtending smooth adaxially nearly Cuticle circle a cell same of the whole hair Arms 2). 6, arranged in to arms confined of flattened cells with Arms of the 4) excepted. to (table I), uniseriate, composed of with of abaxial indented on the cells embedded in pointed, Hypodermis as a sometimes continuous BLUMEA 130 Table Sizes in ++ µm; of because abundant; = + or = present; 27, No. characters I. Some leaf anatomical preparation poor VOL. ± = other reasons; ad. ± scarce; = of 1981 1, Gymnacranthera species = very adaxial; ab. = ? scarce; = character not observed abaxial. t/1 "a> 1 o o CO c (J "3 W 1 (/3 1 E W £ C3 c >> U >N o 1 o c — o 1) <D 1- c*_ o x> CT3 E O o T3 tA C ■s ■J u t/i 55 S c C/3 cL'B a E GA 1 3 g •3 "3 3 'C E IE Species and f— studied specimens ad. G. bancana var. bancana G. bancana var. borneensis al — "o Z ad. 8 30 18 10 *3 c C/5 .3 C/5 SI ab. 14 260 410 £ 3 — ab. _ + 50 + + + + + ± G. contracta Jacobs SAN 9437 Kostermans Haviland 13696 9420 134 Kostermans 7241 11 7 5 ± + 220 5 2 15 ± + 260 7 6 5 ± + 300 8 8 15 - 15 - - 14970 350 11 11 15 Jacobs 5326 240 5 5 10 210 6 2 15 Rossum 63 - + - - ? - - + - + ± + - + ± ± + ± - eugeniifolia var. eugeniifolia 250 8 5 5 ± + -H- 260 8 6 5 ± + ++ Rahmat si Toroes 3944 200 8 4 10 ± ++ ++ coll. 6622 240 9 5 5 ± ++ ± Rahmat si Boeea 8035 eugeniifolia var. griffithii King's forbesii Jacobs var. crassinervis 5259 Rosli 3393 G. 270 8 FRI 3908 G. 15 10 van G. 15 8 260 Galau G. 6 310 17514 Anderson 5 10 200 5363 Zehnder forbesii var. 5 5 20 5 2 15 coll. 6591 210 5 2 20 220 6 2 20 G. paniculata var. paniculata 130 5 2 15 G. paniculata 210 7 5 15 layer var. absent. cells, which loose zippeliana Mesophyll composed of are spongy conspicuously Rosli 3393, square so rarely with to 3 times, tissue. Abaxial in G. some either with sclerified supplied 2 — 3 adaxial sometimes up layers to 7 + ++ ++ ± 7 of spongy cells in the centre in layers an abaxial flat to shallowly arc-shaped and with several + - - + - + as long as wide, and fairly G. forbesii var. to the phloem var. or bancana abaxially prominently bundle with the to numerous margin epidermis (table I), in G. bancana concave, most crassinervis, of the spongy tissue sclerified. Leaf parenchyma cells, often adjacent with 7 + - tissue sometimes sclerified, collenchyma. Midrib fine to intermediate, raised, supplied - layers of palisade parenchyma 4 times eugeniifolia,, locally present robust, adaxially approximately groups in 2 + - forbesii FRI 5749 King's 180 190 phloem in separate bundles situated adaxially J. Koster of the collateral bundle, of Groups with the parenchymatous which (table phloem hypodermis and on the with, bundles. Ground tissue from phloem of the scarcely present collenchyma the or elements xylem some 131 (fig. 7). of the collateral bundle, associated sclerified the adaxial side. Ground tissue also and as in epidermis locally a even periphery between the the adaxial near cells to parenchyma cells, vascular system absent to and often centre with sclerified collenchymatous, interspersed groups, often I); by abaxially abundantly present abaxially are central to accompanied Myristicaceae groups, and in the centre, associated the adaxial of, rarely Asiatic Leaf anatomy of fibres present sclerenchyma peripheral phloem centre P. Baas: & developed with interspersed brachy- to astrosclereids; adaxial chlorenchyma continuous in the midrib (not in G. bancana), situated abaxially of the flat adaxially raised. Veins abaxially with small adaxially at in G. (veins eugeniifolia with collateral bundles, embedded in and abaxially caps with very paniculata with 3 bundles usually little less more or adaxially. brachy- astrosclereids. to idioblasts, of several Crystals which then less deeply parenchymatous tissue var. and/or in of midrib. usually be in cells most Minute, grouped sometimes 50 species; In 8 usually species studied, view; in older leaves some hairs usually excepted. on 2 in the basal end at the mostly Arms extended abaxial walls between cells with one hairs upwards arms very robust hairs with thick walls also thin a to especially present more Large, usually a thin Sclereids the ground abaxially; in G. less or tissue spongy cuticle, probably usually tissue filiform, ground in in sometimes in cells. scarce, as spindle-shaped sometimes tissue, and the — Photos Hairs present the less or in var. present of the as midrib rarely branched tissue of the midrib, 38). of cells with midrib eugeniifolia prismatic crystals epidermis. veins, G. flattened and have idioblasts with mesophyll druses large as (in more epidermal on 16, 21, 32; fig. 6; table II examined adaxial and abaxial surface of mainly shed, especially on cells palisade parenchyma, less or species superficially persistent uniseriate, composed hair, in spongy Empty major in the adaxially (photo surface leaves, Minute, abaxial HORSFIELDIA WIILD. C. a xylem, visible in cuticular macerations astrosclereids in to of the frequently present numerous more in in present brachy- sometimes 37), cells. sheaths, rarely tissue of midrib. sometimes clustered sclereids by also with free bundles, frequent extremely invariably present, frequent ground secretory, (photo bundle bancana, oil cells spherical fibres epidermal to may staining crystals usually present, grouped ground in the interruptions collateral cuticle and thin and short cuticular flanges, also bancana surrounded sclerenchyma), sclerenchyma mostly adjacent griffithii only abaxially) the fibres sclerenchyma bundle sheath. Petiole with vascular arc-shaped of Groups veins raised), mesophyll, supplied few a Major never the abaxial side of the bundles. Ground tissue abundantly interspersed to mesophyll areas eugeniifolia var. the distal end similar to that of midrib, but with at confined with the sclerified parenchyma cells. or G. (in differentiatedparenchymatous the basal end phloem raised sclerenchyma mostly poorly system collenchyma slightly to in surface, arm longer (cells oblique, in glabra and particular with arms directions, arms on H. as the adaxially but the midrib. Hairs 1 — 3 — 6 per stalk cells in stellate hairs. Cell with thin walls but in H. Basal cells young sabulosa, 2—11, mostly 3 than elsewhere), in different present. H. 1 — sylvestris 13, mostly 4 or 8, BLUMEA 132 VOL. 27, No. 1, 1981 epidermis curved; adjacent cells, to the ground midrib of tissue slightly in to astrosclereids straight + + + + + ++ ++ + ++ ± + ++ + ++ + Brachy- crystals Spindle-shaped to + Swollen ± ± ++ + + + + + + flattened extremely idioblasts, druse to adjacent cells, epidermal Some = st midrib of pith in Xylem + + + + + ± s + + focus; species <1 areas cuticle of Thin ± ± + s st s st 8 high at ab. st st s(h) ad. s(h) c.st s(h) s(h) ab. 2 2 ad. 8 8 s s st s(h) s(h) st St St s(h) s(h) 4 2 2 4 2 = Horsfielda Anticlinal sinuous; of . * * ? 2 2 ? 3 2,4 4,6 ? 1.5 1.5 = s char cters walls (h) 1.5; only) hair numbers per cells common basal (most 16-43* 18-31* of Number and 4,6,8 1 antomical leaf betwe n areolae of the in parts 2 mm hairs/0.1 basal of Number II. ad. 1.5 1.5 1.5 1 3 2.5 <1 <1 6 9 1 1 8 9 2 3 1 ? 1 < < 1.5 1 > 1 > 1 = 1 > Some <1* <1* 10 10 ab. ab. 3 1 5 3 3 3 10 3 6 4 3 5 13 1; cuticle of Thickness ad. and 4 0 Table betwe n = table very at also see 200 490 250 280 59 230 220 210 310 110 170 250 460 focus. high I. lamina of Thickness sinuous legend only = For c.st studied specimens and Species 5101 5274 12325 en Ste nis cras ifola Theunisen SAR glabra Junghu nIbid. irya van van H. H. H. Roy 4031 2796 iryaghedhi macro ma Mer il parviflora sabulosa BW H. H. H. H. J. Koster _ + + + + + _ + + + _ _ _ _ _ + ± ± ± ± + + + + + + + 7 + + — — — — ± + 7 & P. Baas: + + — _ _— — —_ _ _ — warts. s s s s s s s s s cork C.St -> St 2 C.St s(h) C.St 1 2 1 St 2 s(h) s(h) C.St 2 1,2 2 into parts basal 4 4,8 4 4 4 4 5,6,7 4,8 4 many of 5 <1 3 1 15 1 3 10 3 <1 <1 <1 4 2 4 2 5 4 5 2 5 3 5 10 5 1.5 1.5 1.5 5 4 5 4 7 transfomatin of 6 6 7 because 240 160 240 200 270 180 280 270 310 species, other 3 1123 4510 en sylvestri H. Soepadmo 235 407 4446 & 1 Roy 3081 BW van bb with 379 Gjel rup Robinson Kostermans Kuswat Pleyte 1839 BW compar ble range. Hardly * * Full Leaf anatomy of Asiatic Myristicaceae 133 BLUMEA 134 1 abaxially most — 1 15, mostly 2 or VOL. 27. in arranged a circle or subtending cells and cutinized a probably surrounded oval, in H. (table II), of the groups of basal cells by after ring No. 1981 1. glabra adaxially 16-43, abaxially into cork changed less more or of the hair. shedding Basal cells warts. oriented epidermal radially Usually basal most parts of hairs per surface unit (table II) on the midrib. In H. crassifolia abaxially 2 or more but basal in H. often parts joined together. smooth glabra then absent iryaghedhi only, and at sinuous, least at cells high some thickened. Abaxial midrib and overlying axis long parallel cells also major in arranged rows but the to epidermal cell epidermis, abundant in and striations of abaxial pattern leaf sometimes (photo 21), the to faintly — 1.2 35 x8—19 — pairs /OTI, average average values 3.3, Polar inconspicuous. 1.6 wide: relatively are Subsidiary in H. values 25 2.8; in H. 15—19 nm. frequent probably only, Smalt cells with probably glabra In and H. grooves — 110 — section: 490 pm thick 10 /im thick Cork (table II), cells of cuticle iryaghedhi; complex or not; overlying in H. circular, from stomatal of the cell surface adaxial abaxial and/or in frequent most to midrib often wall cells. flattened, H. than of sunken (in cells in absent H. surface on iryaghedhi partly view. midrib, and slightly embedded in the adaxial epidermal Abaxial major epidermis and on abaxial side more or crassifolia subsidiary cells, but with leaf to epidermal cells; erect, cells cells slightly side, corresponding veins in H. sabulosa, epidermal sometimes dome-shaped and/or complex H. sylvestris with in H. Unspecialized abaxially lower in or, crassifolia and usually epidermal higher periclinal seen as papillae guard variously extent cuticle 2—13 /im thick, abaxial sometimes with shallow indentations on the lumen areas the dorsiventral Lamina in H. of usually dome-shaped, small. Outer usually in warts and centre cell guard sometimes distributed and hairs pairs width ratio to less to a epidermal cells, (table II). Adaxial anticlinal walls rectangular, conspicuously epidermal of parts present oriented relatively sabulosa. above to square sometimes sylvestris cells ledges present, regularly basal cuticular H. and H. sabulosa the thin cuticle, sometimes in the by radially transverse 1 surface, modified a secretory, surrounded isobilateral, cuticle abaxial from developed complexes. epidermis, the on and length inconspicuous. in abaxial midrib, major on prominent of traumatic origin, but in H. glabra and present, probably sabulosa visible to then elsewhere. Guard cell 10—17 /im, parviflora or adaxial rectangular. slightly subsidiary on Outer stomatal absent T-pieces 32x absent with then Abaxial cells to square with the crassifolia H. granular — or some thin, but iryaghedhi. Minor veins cuticle glabra; very Stomata confined cells stomatal pore in so — only. areas sinuous, rather then polygonal curved thin veins; corresponding inconspicuous. thin and, when striations are absent, less 20 be epidermis paracytic. perpendicular to H. crassifolia slightly to when walls in H. of the in H. inconspicuous specimens scanty and aberrant areolae, margin, straight rectangular rows, course tending few papillae in arranged of cuticular flanges present, but Pitting veins veins a in margin, irregularly sylvestris faintly abaxially. (table II); in cells with epidermal perpendicular or leaf sometimes with straight. Cuticular flanges are H. granular, obviously present and anticlinal walls (table II), of cuticle between anticlinal wall undulations division walls conspicuously to 16). Cuticular striations present polygonal; focus veins and sometimes in faintly adaxially, Unspecialized epidermal or faintly midrib, major on structured and distributed (photo abaxially Cuticle Alveolar material abaxially. papillae margin. and H. hardly less flat, with thin in H. Stomatal sylvestris p.p.) so in H. glabra J. Koster & P. H. and as and outer macrocoma; Hypodermis continuous a Baas: Leaf inner layer translucent cells, sometimes with epidermal cells, 2 — of Theunissen crassifolia, of square layer flattened to sometimes broader than division walls. periclinal often present. probably in H. adaxial one higher, to ledges (absent 135 Myristicaceae adjacent Mesophyll composed of 3, in H. crassifolia, H. irya, Van Royen 5101 and H. sabulosa 3—4 adaxial layers cells palisade parenchyma which present), loose fairly raised to square ridge cells, present) in or, in H. abaxial shaped glabra, bundle, H. and parviflora a less more or concave, adaxial straight in 2 usually only abaxially often pith, or (sometimes only sabulosa, H. margin epidermis, adaxial bundle; with the rarely joined together bundles in the phloem to numerous raised robust, adaxially with the to also are wide, and as tissue. Leaf compact spongy adjacent long as bundle, with 2 small adaxial bundles in H. parviflora, and or in small separate groups, phloem palisade layers 5 times to up usually Midrib fine to abaxially prominently raised, supplied interrupted less more or abaxial some times, sometimes 4 parenchyma collenchyma. central H. sabulosa (in sylvestris) H. sclerified with supplied are (in or either with a lower Asiatic stomatal often present 59, H. glabra and H. sabulosa), consisting of rectangular of anatomy an arc- with the and with several layers, accompanied by xylem groups (table II; fig. 6). Groups of sclerenchyma fibres present, surrounding the vascular system, associated with fibres present in the pith, tissue from with the tissue groups, and peripheral phloem associated with and often centre to collenchyma in the even periphery parenchymatous as a are usually present in the pith, Veins with collateral differentiatedparenchymatous poorly the distal end similar that of to collateral bundles, also with free fibres the at basal end Ground tissue druses in often and to macerations as areas often present in the crystals frequent spongy the tissue, the to scarce, ground tissue sometimes most abundantly, peripheral ground the H. a frequent as and H. major veins, abaxially sclereids to astrosclereids. close to and the glabra sometimes abundantly present sylvestris, (in as H. to with supplied by a usually with 3 arc-shaped sometimes with small Groups of sclerenchyma main bundles. Crystals frequent the adaxial and abaxial also II), few, mainly a mesophyll, parenchymatous ground visible in less more or more or most a thin cuticular small druses spindle-shaped frequently in the bundle sheath, and in less tissue of the probably secretory, spherical midrib, oil cells often in the sometimes present in and H. sabulosa. Sclereids often present, clustered brachy- to astrosclereids (table II), usually abaxially, rarely bb large as epidermis, flattened and have extremely flanges (table in the end abaxial side of the in cells in the tissue of the midrib parviflora basal the main bundles. thin cuticle, in H. mesophyll, adaxially, (table II). Large, mesophyll brachy- with slightly raised, abaxially to deeply staining cells; grouped of the midrib in the the at bundles which may be especially ground parenchyma and surrounded tissue of the midrib. Minute, idioblasts with epidermis, pith (in to cuticular of several less ground sometimes usually abundant pith. Empty short sclerified bundle sheath. Petiole with vascular system brachy- epidermal cells, thin and sometimes the adaxial side; interspersed in fibres, confined to the with interspersed on some depressed few midrib, mesophyll idioblasts, mostly adjacent cuticle mostly a phloem collateral bundles close and adaxial with embedded bundles, caps, sometimes only with sclerenchyma bundles. Ground phloem collenchymatous, to and often astrosclereids. Major veins adaxially slightly raised. of the locally developed hypodermis sometimes, mostly abaxially, interspersed cells which at sclerenchyma centre 30811), crassifolia, sometimes in the H. astrosclereids in the glabra and superficially H. in also in the ground tissue of sabulosa), examined and H. fulva BLUMEA VOL. 27. No. and H. superba, solitary in or at least in the groups of the periphery LOUR. KNEMA Swollen cells with thin walls often present, mesophyll. if present, hypodermis, rarely adjacent or sometimes in elevations of the leaf surface epidermis (table II), the in the 1981 1, 136 to the abaxial in (photo 32), and/or tissue of the midrib. ground 1—8, 11,12,18, 23, 24, 28, 29, 33—36; figs. 3—5, 8, 10; — Photos table III 83 31 species; studied species view: surface In leaves often shed midrib and on with the arms 1—3 to many; with upwards arms short; 1—8, very fig. 3, 1 mostly surrounded by ring (table III) or midrib, major is plumulosa mainly adaxially; as with rectangular to cells or not in epidermal horizontally papillae Guard cell to usually on a 17 cell pairs are inconspicuous. traumatic to 1 adaxially in circle a 14, — or oval, subtending a granular, abaxially plates, which is absent long axis and parallel or arranged rarely rufa (photo in K. at sometimes 18). at lower focus and often in arranged perpendicular in rows but Abaxial straight. walls, and veins major epidermal to tending the to cell pattern of abaxial epidermis, epidermis. with rows, course be of the polygonal Stomatal nipple-shaped papillae x 8 — complex of the 4 10—16 — 6 26 — 30 /rm; Polar variously conferta and absent — T-pieces more or less complex (photo 23). 27x9— 14 some sometimes usually K. kunstleri abaxial midrib, major by guard the cell present, infrequent, length im, oblongifolia.FRI0111, in K. lunduensis present, on to bordering epidermal cells; above the stomatal 16 /im, average values 20 pm. warts but in K. or overarched visible slightly Stomata confined in areolae, scanty and aberrant 30 cells epidermal of cuticle between anticlinal wall areas midrib Cuticular complexes. with sinuous anticlinal walls slightly curved anticlinal star-shaped opening Cork origin, relatively Pitting of cuticular flanges present, but inconspicuous. in relatively long: wide: to curved anticlinal walls cell pattern of adaxial — sculpturings convex width ratio 1.3 —2.8, average values 1.7 —2.4; in K. relatively lobed stomatal midrib less sinuous, adaxial cells also directed leave bordering slightly margin, paracytic. pairs are Arms (table III, photos cells and almost smooth polygonal, adaxially overlying rectangular. veins and leaf epidermal with anticlinal walls of inconspicuous very square with the epidermis, abundant the straight prominent Minor veins cells to corresponding cells veins; corresponding rather than hairs arms. may be arms structure Basal cells below). reticulum of erect, anticlinal walls Abaxial papillae. in (table III), arranged adaxially Cuticle cells with straight epidermal have not (cells elsewhere), when excepted; where the varying oriented adaxial cuticle with focus and undulations; cells than longer persistent one arm of the hair. Most basal parts of hairs per surface unit restricted sometimes thin, radially a Unspecialized epidermal (very) high less hairs of cells with in stellate hairs. Cell walls between cells midrib, 1 —4 surface, in older some veins and leaf margin; the alveolar material in K. intermediaand K. striations and grooves present mostly shedding with alveolar material on 1 —8, the midrib. on as the classification of types more after usually distal cells do most Hairs thick walls. to also on stalk 4) to the in different directions, oblique, especially see but the midrib the hairs on hookeriana) 11, abaxially — cutinized sometimes in K. with thin arms adaxial and abaxial on uniseriate, composed midrib sometimes up (on extremely long (as extended veins. Hairs major one Hairs present (table III), especially adaxially, to guard pairs are usually probably of abundant, especially abaxially, J. Koster & and and/or abaxial to usually Outer of leaves, especially most cuticular lower erect, K. absent linifolia papillae often with midrib, major on papillae. Subsidiary translucent cells, continuous major veins, composed high as layer but as parenchyma 2 — cells, 3, are up to square (or than wide (in K. curtisii and K. K. pallens and K. B RUN 5587, K. tissue; cells of abaxial layers sclerified palisade margin and K. laurina, spongy spongy (table III); supplied with a straight ashtonii and K. kunstleri, and with the adaxial bundle; especially abaxially, and accompanied by xylem straight present to arc-shaped in the surrounding an 3 lignified cells at the periphery, side; ground or often abundant the sclerified cells tissue also often as a or the main of 2 in the large, var. peripheral phloem to often less even parenchyma brachy- or are present groups, and in the centre periphery parenchymatous and also present with pith, more or fibres to cells, especially in the locally developed hypodermis interspersed the (very) to abaxial bundle, sclerenchyma with sclerified (table III) Leaf to in K. ashtonii associated with and often bundles. Ground tissue from centre collenchyma as Groups of the usually interspersed large pits. adjacent rarely joined together phloem bundles sometimes 1 fibres present in the collenchymatous, with the only many cells of separate groups, usually in 2 layers, associated with pith, pallens (in curtisii, less compact sometimes interrupted abaxial bundle, sclerenchyma phloem more or 5 times K. abaxially prominently raised, so, oriented 5). to (in to Midrib of intermediate size in palisade scarcely longer to sometimes up supplied never Mesophyll of layers sclerified, adaxial bundle, phloem fig. the vascular system, adaxial times, walls prominently xylem bundles, layer rectangular, the midrib and the (table III), usually one to numerous III; 4 pseudolaurina) groups (table III; fig. 4); pith (table near slightly flattened) or to arc-shaped with the with only cells present, adaxial one wide, and fairly loose collenchyma. arc-shaped directed material; guard of in and Stomatal margin. horizontally in K. mandarahan and K. sometimes to as parenchyma with robust, adaxially raised, even occasionally tissue have thick supplied or tissue seen as conferta palisade parenchyma. 3 — or and K. oblongifolia spongy often with sclerified epidermis, long as in K. flattened of sometimes 4, linguiformis) percoriacea) of cuticle leaf less consisting layer) with the intergrading sometimes which or less shallow adjacent epidermal cells, adaxially locally developed, and sometimes of lower than or areas striking small. usually stomatal ledges sometimes outer abaxial inconspicuous one more sometimes present, Hypodermis of most midrib overlying more or thin cuticle, without alveolar a inner and/or subsidiary cells; inconspicuous. a cells with cells with veins and cells with complex sunken; bordering epidermal cells to square dome-shaped, abaxial side on often (table III), adaxially cells epidermal papillae, thick conspicuously with thin side, corresponding epidermis (sometimes cells; veins and cells. 300 jum thick (table — 9//m — to partly in adaxial along midrib, epidermal of adaxial epidermal complex surface view. Abaxial embedded in slightly adaxial side often dome-shaped, wall periclinal abaxially than adaxial epidermal on indentations on the lumen as oriented by radially and on layer. Unspecialized epidermal sometimes erect, rectangular, and circular, midrib, thin cuticle, sometimes in the a cuticle 2—17 /im thick, abaxial cuticle 1 granular higher the along Small cells with section: Lamina dorsiventral, 80 transverse III). Adaxial a veins and on 137 Myristicaceae only, probably secretory, occasionally present epidermis often surrounded margin, In Leaf anatomy of Asiatic from stomatal complexes. of the cell surface centre with Baas: regularly distributed, mostly developed leaf P. pith; on (mostly) often the the adaxial brachy- to 138 BLUMEA midrib of tissue - VOL. 27, No. astrosclereids to 1981 _ — ± ± — — — — + + ± + + + ++ ± + - — — — — + - ground + in 1, — ± + + + + + — - Brachy- midrib of tissue cells ground peripheral parenchyma + in + 1 midrib of pith in margin cells parenchyma Xylem species Knema bundles. parenchyma leaf + ± 1 + — - + in ± ± ± ± ± — — — — ± ± — — — — + ± a in — — — — — - a + Sclerified text) (see type large + + - Sclerified tissue spongy cells ++ Sclerified 8a Hair 2 7+8b 8a 8b + 3 7 3 6 3+7+8c 7 ++ — ± — — ± ± 1 2 2 2 2 2 2 4 2 2 4 3 3 3 5 1.5 1.5 3 8b 8b 7 a + 7 = 1 of layers; char cters abaxial abaxially with only) hair covered numbers per cells leaves common basal hairs ++ + + 9 — 9 ± Adult ab. 1 1 1 2 1 2 ad. 2 2,4 2 4 2 4 2 ab. 15 40 5 3 5 3 20 30 ad. 3 5 3 3 3 1.5 5 5 ab. 2 2 3 4 3 4 2 2 9 2 ad. 6 6 6 9 4 6 4 4 12 5 90 120 1 1 1 (most of Number = 2 2 8,9 a antomical II. and areolae I of leaf tables Some also III. see Table legend the parts in 10 10 3 3 3 3 <1 8 3 2 6 3 12 9 7 13 4 mm2 hairs/0.1 basal of Number cuticle of Thickness lamina of Thickness 170 140 140 230 120 110 230 160 140 180 160 220 120 For studied specimens and Species ashtoni cin amo ea var. var. kunstleri curtisi var. 5587 A4139 ashtoni ashtoni atenuat conferta curtisi BRUN FRI123 6 elmeri SAN K. K. K. K. K. K. ssp. 18294 S fur ace globulari hokerian interm dia korthalsi kunstleri latifolia K. K. K. K. K. K. K. J. Roster & ++ + + + + + - + — — + + - + - ± - ++ P. Baas: ++ Lea) ++ ++ + + + + ++ ++ + + + - + + + — + ± anatomy ± - ++ 1 - + - — - - - + - - — - - - ++ ++ 1 1 1 1 ? + + + + 2 2 2 4 2 8,9 8 4,6 11 4 5 3 3 3 3 1 1 3 3 5 5 1.5 1.5 1.5 3 5 3 4 100 110 120 140 laurina Chew 2 - 2 2 2 4 1.5 1.5 10 - a + 7+8b + 4 - — — - + - + ± 8c 8a 8a • ++ ++ + 8b 8b ++ — ± 3 ? - - 2 2 1 2 2 2 2 4 6,7 8 2 2 4 4 5 20 10 3 3 1 3 3 1.5 15 1.5 1.5 2 3 3 2 3 5 3 3 2 2 6 3 7 8 5 4 14 7 8 2 2 8 100 180 210 150 300 170 210 3 1.5 1.5 5 80 140 120 SAN — ++ ++ ++ - - + + 1 - +(1) + — - — - + + 7 - + - — ++ ++ - + + + + ± + + + ++ + + - - - ± + - - 1 + + - - a 3 - 2 1,2 3 6 - 8a 5 - 8a 8a 7 — - 2 1,2 2,3 2 1 4 4 5 4 4,5 2 2,4 3 3 3 15 3 1.5 1.5 15 2 2 1 3 4 3 4 4 1.5 10 5 <1 5 9 2 9 17 4 1.5 — 150 160 - SAN 3482 S K. 982 4 2533 73 12 pal ens Meijer SAN percoiace plum losa pseudolarin psilantha K. K. K. K. K. 11 01 + + 1 5 KEP FRI S S K. - - + 4 8b - + 2 2 2 2,3,4 4 2 4 35 20 3 3 1.5 3 10 5 3 3 4 3 2 3 3 7 9 3 8 12 6 1 2 1.5 1.5 1 180 250 150 150 200 150 150 180 pulchra Kostermans ++ ++ 2 1 3 + setosa 5103 14373 29064 + 8b 8b 8b 8c + forma linguiformis linfolia lundensi mal yan mandran oblongat oblngifola K. K. K. K. K. K. + percoiace 24985 Ja - — - 230 300 260 160 130 ssp. 2502 + — a ++ ± 1 — oblongat Wilde(-Duyfjes) 41273 FRI2431 de 1 — 5 ± + 5 143 var. Wee - 4 + ++ ++ ± — 1.5 1.5 14 29 laurina Lek 8a + + + 1 + + 1 K. - - 139 of Asiatic Myristicaceae 9908 9907 29015 SFN retusa Sinclair K. ssp. Sinclair rubens rufa tenui ervia toment la K. K. K. K. 140 BLUMEA astrosclereids then situated adaxial of the No. chlorenchyma collenchyma. bundle sheath, sclerenchymatous layers the adaxial and on abaxial 1981 I, continuous in rarely Major veins the midrib, adaxially slightly depressed transcurrent often entirely bundles K. with fibres sclerenchyma often with epidermis adjacent latter condition sometimes a in in the interruptions Minor veins few- many-celled to in K. laurina, K. xylem, also bundles, with at more less or small of Groups brachy- to in (when large collateral bundles close fibres sclerenchyma astrosclereids. in the idioblasts) the midrib, in the and to the basal at the basal end with at free epidermal crystals frequent to parenchymatous bundle scarce, present in epidermal cells. from grouped cells. in cells in the Flattened, cells epidermal usually tissue of the ground also in the in present the tanniniferous idioblasts with epidermis sacs, in the rarely also sometimes prismatic crystals usually solitary. Irregular sometimes clustered, present in (fig. lOf). Large, less more or mesophyll in small druses spherical in and/or oil peripheral in K. elmeri, K. kunstleri and K. retusa, Sinclair 9908, midrib, pith. Empty grouped, abundant, sometimes tissue of less spindle-shaped tissue of the midrib, less or or the druses large to ground large; (very) more bundle sheath, cells of K. hookeriana present, small to interspersed mesophyll (most frequent ground more with rounded edges, crystals, usually and frequent most and in the sheath), as the main confined mostly often also present in the Usually minute, present in cells of the parenchymatous abaxial bundles phloem adaxially end Crystals usually frequent mesophyll, palisade parenchyma often also in present small oblongata, the distal end abaxial side of the main bundles. Ground tissue usually abundantly with sheath epidermis. small, especially poorly vertically to bundle through the to Petiole with vascular system collateral arc-shaped by sometimes with small, collateral bundles between the main bundles, and adaxially, bundles. of (the with the lateral sides, of several cell on surrounded abaxial and/or pseudolaurina). midrib, less or rows and K. and sides, sometimes sclerified composed that of to more adaxial are vertical or percoriacea similar the to which extensions, one-layered bundle sheath, embeddedin mesophyll differentiated parenchymatous 3 27. raised, abaxially prominently raised. Veins with collateralbundles, supplied to a (table III); abaxially VOL. (photo often present a 29). thin cuticle, probably secretory, More and adaxially less or thick-walled abaxially of the sometimes cells, probably sclerenchymatous bundle sheaths of the veins, sometimes 'branching' towards the adaxial epidermis, sometimes even protruding in the present, sometimes abundant, 12326) or as tissue of the pith, in K. Thick sometimes clustered oblongata filiform, veins Hair For to the chlorenchyma of adaxial of the preparations damaged during studied at in to (in K. curtisii astrosclereids in some ground specimens curtisii, FRI var. peripheral from the tissue of the major veins morphology were percoriacea, 36). of the hairs, of individual hairs for left for the study magnification so transverse light microscopy, have been used. the microtechnical process, low power in the (table III). bundle sheaths of sclerenchymatous epidermis (photo ground of K. laurina also Knema study views obtained with the S.E.M. photographs sclereids brachy- S. 34822 also in the the, mostly detailed sections and brachy branched sclereids sometimes abundant in K. present, types a Sclereids often epidermal layer (photos 34, 35). midrib, mostly abaxially, extending through the as Unfortunately a paradermal well as the hairs surface are often that for certain leaves only S.E.M. of the hairs. The entire with and as tomentum dissecting microscope. In has also been fully developed J. K.OSTER & leaves the hairs cell size, Hairs with diversity wall thickness arms short very other using such arms, total etc. of the leaf. have on young leaves have been studied. (very) case 141 Myristicaceae of the hairs of Knema have been classified the midrib and the on Asiatic Leaf anatomy of type of stalk cells, relative size of the body of the cells with as remaining part present Baas: often shed, in that are The morphological characters P. veins major They bodies) and have of the leaf, places compact (which more short relatively have different from those usually are longer, are been not into account the on the cells These hairs, arms. taken means: rarely in this classification. The and photos indispensable part also a If key. CLASSIFICATION A. One or Type 1. more stalk Long OF 2. Long one or two; some 1, then try type 2 AREOLAE AND distinctly longer cells MINOR considered as an way the classification is etc. VEINS OF KNEMA LEAVES. than wide: part with short and compact; arms arms with thick walls. See 3. photo 1, fig. Type HAIRS ON cells stalk fit type not should be 3) fig. of hair types. In descriptions hair does a 1—8, drawings (photos of the stalk cells up to with three; part short and compact; arms arms with thin walls. See fig. 3. Type 3. Long stalk cells B. Stalk cells Type Type 4. 5. long; walls. See Hairs long; thin Type 6. Hairs with Hairs short to thick walls: Type Type 7. 8a. Hairs short; are thick See Hairs b. Hairs with arms Note: in the case MYRISTICA C. 72 species; In (table 13 L. 2 to obviously — surface of about equal distance more in the 3. proximal regions; arms with thick pointed tip, a and with fairly the is arms considerable); arms thin, 3. at the distal end 100 surface view pm relatively short, and less); with thin to with arms thin to 3. view 65 — hairs with the 250 pm. When a diameter of hairs more with than a diameter of 100 pm also 100 pm occur. Arms 3. photo lengths 8, 3. fig. of hairs of type 8a and c; arms with fairly thin to 3. difficult; some species show short with body, between or long a gradual transition from Still this subdivision may help one to subtype identify a hairs. 10, 13—15, 17, 19, 20, 25, 26; figs. 9, 10; table IV studied view: Hairs present, on sometimes adaxial surface almost (table IV), uniseriate, composed midrib and the the near give precise lengths. Photos 9, IV), in older leaves (on long (in particular arms photo 2, fig. photo 4, fig. walls. See between 8 is species sometimes shed or with thick type of in (fairly) long, photo 6, fig. photo 7, fig. of broad (diameter in surface it is useless The subdivision another. Therefore specimen See in See (see type 7), length a thick walls. to diameter very arms and small present long; longer than long (the photo 5, fig. walls. See c. See 3. walls. of cells small; short very and less of cells thin to thick walls. with photo 3, fig. long; body long; body Hairs fairly short, arms thick (very) long; arms 3. See fairly with part thin to thick walls. at the distal end arms fig. walls. with than wide: longer not Hairs two; one or proximal region), major veins often length, spread out more) to always, on abaxial and at excepted. angles Arms of with surface abaxial surface of flattened cells with 2 stalk cells horizontally, confined arms arms, a of the 1 single cell adjoining 142 BLUMEA VOL. 27, No. 1, 1981 mesophyll crystals; ocurenc ++ cells epidermal abaxial in Crystals epidc adaxial in Crystals cells = of •a s + ++ + (p)-c r-p-c r-p-c p,s,n r-p P.s p,s s,n s,n s,n,h s,n,h (s) b-a sc-b-a r spindle-shaped infrequent + + crystals Spindle-shaped c s druses; Types ground midrib of tissue OJ periph in midrib of cells sc,b-a sc-b Sclerified = hypenatd. crystals; d species Myristca astrocleids. = a cluster d hypodermis forms X<D "a i o CJ stomatal = c char cters brachyslerids; crystals; in - Xylem •3 the bordering Intermdiate X epidermalof ct f, — "a. E oai stomatal the cells, only. - - Continuous 00 1 ■, of pith Number 1 SO OS + + bordering on ++ ++ Papillae abaxially + 1 + 6-10 5-9 6-10 4-6 1 t-rt 1 1 6-10 6-9 + + + 1 1 6-9 6-10 + + antomical cells; prismatic hairs wi covered leaves 6-8 + hairs ++ ++ ++ ++ .c + + Adult = b = lamina of c p leaf Some parenchyma edges; rounded IV. Table sclerifed with = sc I—I I. crystals = tables r only: types parts mm2 hairs/0.1 ab. 5 15 40 30 15 basal of Number ad. in 5 1 1 + + ab. 6 6 6 ad. 9 12 8 material Alveolar basal 1 1 4 + 2 15 15 50 5 3 25 1.5 15 + + + + + 3 3 3 3 5 3 5 6 7 8 8 1 cuticle of Thickness = h crystals; to = n 8 4 4 260 190 220 230 240 140 230 200 210 230 310 lamina of Thickness studied nedl-shaped legends crystal spindleFor For on cells also see the brackets. specimens and betwe n Species fatua 25986 25893 chrysophyla Brass cinamo ea Brass M. M. 1506 24236 pa uan s.n. 15023 var. var. SAN SAN fatua fatua 787 Vriese fragrans Sulit Beguin gi antea globosa M. M. M. de M. M. J. Koster - (P),d,(s) — - + - - - - d,(s) - — — - — - + + + s* - (P),d (s) d,(s) — — - - - - +/± - - + + - + - + ++ - - - + - - - - s (p),s - - - s,h - - - — 5 9 — 10 — 5 - 3 - d r,P - (P),d s,n - b-a b-a b-a sc** sc sc + + - - - sc - - — - - - - - - + — - - - - - 7 6-11 6-9 + + — — - — 45 40 50 50 45 50 50 50 40 20 5 - (P),d (s) ++ ++ ++ ++ ++ ++ ++ ++ ++ 3 ++ d,(s) (s) 3 - 5 — 5 - Leaf anatomy of + 4-7 4-7 5-8 5-8 4-7 5-8 4-8 4-7 4-8 — & P. Baas: 3 1.5 3 1.5 — + + 3 4 5 5 6 8 11 9 8 9 10 9 11 10 9 11 10 5 4-8 5-8 + + + + 7-11 6-10 + + + + • + + + 10 10 15 15 + - - - - 4-6 4-6 - - ++ ++ 30 35 1.5 5 5 10 + + + + + + 5 4 4 6 3 5 7 7 7 11 7 9 8 270 190 220 190 260 290 310 260 310 180 200 1 7 170 200 320 220 Asiatic 5 280 350 origin. traumatic of 7 UNESCO 624 39 95 guaterifola Kostermans Sina g ul M. 77 30298 20850 5365 32924 371 Mihil Elmer BRUN SAN Ambulah 55 ** 908 Kostermans FRI50 6 lepidota maingayi maxima Thorenar Grashof sphaerosp ma subalu ta vilosa Kostermans M. M. M. probably 6438 E1P 4873 M. M. M. A4747 SAN minute; very * Myristicaceae 143 144 cells. Arms with thin walls In M. villosa with arranged in a circle less very frequent smooth material absent on cuticle between in M. wall midrib and or major veins in rows cuticular flanges but epidermis, abundant in veins and leaf 25). to of Guard to guard cell cell areolae, pairs are 1.3 surface origin. be — — Cork average and on or Adaxial overlying long absent on axis cells also of Pitting visible slightly Stomata confined slightly to rectangular. walls in abaxial to midrib, major overarched by a ring of papillae (table IV, photos 13, 15, values 26 average values 1.6 — 2.8; 27—39 gm. Polar in M. — 34x11 — 18 m, sphaerosperma the T-pieces sometimes present, variously present, usually infrequent, probably warts Small cells with especially 21 straight of areas maingayi. square with the complex partly without or thin division straight Minor veins M. with curved anticlinal slightly rather than only. in adaxially veins; corresponding adaxial epidermis Stomatal shallow grooves Very and M. to inconspicuous. ceils with 3.4, of the of variable cells. Abaxial cells epidermal most Cuticle almost plates; alveolar erect cells with by usually polygonal, adaxially to 4, after ring cell one (table IV) present with rectangular — M. fatua, if present, inconspicuous, scanty and aberrant a thin cuticle, sometimes in the centre sometimes present in the adaxial only, probably secretory, epidermis, straight gigantea polygonal but relatively long: usually inconspicuous. of traumatic to 21—39x8 pairs rows, course margin, paracytic. width ratio in abaxial bordering epidermal length the tending cell pattern 4—11 in M. papillae usually present, epidermal cells fragrans. Abaxial epidermal arranged perpendicular cells cells undulations; cells often smaller than abaxial epidermal margin. sometimes very usually with, anticlinal common focus and (very) high at leaf epidermal Unspecialized epidermal curved anticlinal walls and with arranged veins and of the areolae. reticulum of a in cutinized adaxially, often with alveolar material as a composed in frequent most midrib, major lower focus and if surface, or, basal cells surrounded subtending Basal parts of hairs sometimes 13—15), sinuous anticlinal walls parallel abaxial midrib, abaxially gigantea and M. maingayi. especially on 1 mostly row; also are leaves. Basal cells 1 not on a walls and oblique these hairs 10); mostly 2; cells and with anticlinal walls of corresponding at adaxial surface in M. villosa. (photos 4 in - taller than elsewhere. usually (photo bud scales etc., on epidermal granular, abaxially to appearance walls on 1981 of cells with body arms sometimes with 3 oval, basal parts hair; the on but species, I. midrib on with thick-walled oriented small, especially 27, No. Hairs (photo 9). M. maxima radially of the shedding or VOL. extremely long, parallel, other some lepidotaand more or hairs some almost short, present in M. - BLUMEA along the midrib, the veins and of the cell and/or the leaf abaxial margin, surrounded by radially oriented epidermal cells. In transverse IV). Adaxial cuticle 4 with a wall cells complex the lumen Abaxial erect to overlying midrib higher on epidermal side, corresponding in M. higher in M. than abaxial surface with thin gigantea lower to cells with — 350 thick cells thick (table IV), adaxially on same with as as the seen papillae midrib, major height often square to square to Outer cells; periclinal less shallow indentations on of cuticle maingayi to (table adaxial epidermal often small. more or areas and M. Basal cells of hairs abaxially of about the but 140 rarely dome-shaped, abaxially dome-shaped thickened cuticle on the tips; papillae absent cells, dorsiventral, layer. Unspecialized epidermal flattened, of adaxial epidermis Lamina 1 2 pm thick, abaxial cuticle 2—11 often erect, rectangular, epidermal section: cuticular granular rectangular, — in surface with a view. strongly veins and leaf margin. unspecialized epidermal cinnamomea, M. guatteriifolia and M. villosa (photo 19). P. J. Koster & of stomatal complex guard stomatal cells sometimes ledges continuous layer sometimes present sphaerosperma translucent in cells, the also some specimens and parenchyma. 4 — occasionally usually supplied with adjacent the to fine, adaxially adaxial bundle and pith, sometimes with of the developed sometimes IV); to with supplied through bundle the at at tissue with often which adaxially veins bundles that of to adaxially, tending in M. less mostly confined to druses often adjacent (in with M. villosa to an as to be in the situated to pith, are and (table abaxially mesophyll, differentiated vertically transcurrent Petiole with vascular to fibres sclerenchyma astrosclereids. at to the basal usually Crystals usually frequent in idioblasts in the small) in interruptions collateral bundles, also sometimes with small collateralbundles close brachy- they locally slightly raised, poorly sometimes with arc-shaped a interspersed embedded in lepidota. periphery astrosclereids midrib, often in the even to collenchyma depressed with associated centre to of fig. 4). Groups the abaxial side of the main bundles. Ground tissue abundantly interspersed large in usually present by and adaxially from the main bundles. Groups of end as separate bundles in the with and often mostly midrib, more or abaxially in phloem tissue sometimes slightly surrounded caps and the basal end with 3 robust, rarely to system, from the are with collateral bundles, extensions, especially margin parenchyma present, phloem (IV, vascular associated bundle sheath, sometimes sheath phloem the side; ground (table IV), ridge with the sometimes continuous in the Major Veins central groups table pith, sclerified brachysclereids and/or brachy- the distal end similar xylem, with free cells with sclerenchyma parenchymatous system the adaxial chlorenchyma raised. abaxially on to square M. villosa. Leaf some and with 2 —9 Ground collenchymatous, parenchyma collenchyma. the bundles. are less straight, sometimes interrupted or surrounding in groups, and interspersed adaxial of the more xylem near palisade sphaerosperma which of intermediate size raised a in so abaxial bundle, with present, hypodermis with sclerified arc-shaped phloem parenchymatous a M. the wide, and fairly loose spongy tissue; cells sometimes with only a M. rectangular, with guatteriifolia, sclerified, frequently accompanied peripheral phloem centre as layers, especially abaxially, fibres sclerenchyma long as sometimes an as layer (in adjacent epidermal intergrading in M. —3, epidermis. Midrib usually prominently raised, supplied than palisade parenchyma cells, collenchyma, raised, groups, often in 2 sometimes of 2 of layers of spongy tissue cells and 5 times to inner and/or Hypodermis flattened to square rarely higher the cells often guatteriifolia only locally developed, especially veins, 4 adaxial sometimes up times, of M. major Mesophyll composed and M. villosa 3 of layer) upright thin cuticle, without of one adaxial (table IV), consisting broader and lower, a subsidiary cells; inconspicuous. present, abaxial one cells, usually midrib less embedded in more or less cell, separating bordering epidermal sometimes slender, with 26). Subsidiary cells, 145 more or small sunken a both lateral sides of the on one or alveolar material; outer cells often with (table IV); varying conspicuousness present (photo of Asiatic Myristicaceae anatomy sunken; bordering epidermal Stomatal complex papillae Baas: Leal as palisade parenchyma, adaxial epidermal cells, which then may be flattened and have a thin cuticle and thin and short cuticular flanges, also visible in cuticularmacerations as areas of several, epidermal ground in cells cells less deeply staining (table IV) and/or tissue of the midrib. in the mesophyll, parenchymatous in Minute, most cells; small druses more or less in the frequent bundle sheath, and in the sometimes cells of the parenchyma present in mesophyll and the spindle-shaped crystals, grouped spongy ground tissue, especially in the tissue of the midrib, sometimes BLUMEA 146 also present in VOL. - where epidermal cells, they shaped, especially in basal cells of hairs prismatic crystals sometimes present in Flattened 1 crystals with sometimes in Od) Clustered of occur Large, more in abaxial prismatic less or and in mesophyll maxima and M. epidermis, few oil cells in the even In the pith, generic descriptions and been recorded. It is essential species (or variety) characters have as in M. More in thin a M. cuticle, less thick-walled abaxially of the towards the adaxial Sclereids present in epidermal layer. brachysclereids and/or tissue of the sphaerosperma edges. abundant, guatteriifolia, or and 'branching' peripheral ground in M. with rounded idioblasts with most Gymnacranthera, a THE also in sometimes as midrib, usually ground tissue of the CHARACTERS in the tables a very great number of characters has analyse a which of these characters and genus valuefor which is list are in not; are species (or variety), or even relationship for on for constant are other words: a possible are listed to a limited represented following those characters by their are extent because many genus. This is the species for mentioned, species (and, specimens two or more constancy species which or not are a which and of the characters the determination of the constancy for varieties) The characters first for diagnostic level. For the or to OF for the determinationof patterns of important species (or variety) In (fig. smaller in size. sometimes adaxially into the abundant, VALUE genus less Sometimes intermediate midrib; epidermis. sometimes protruding astrosclereids in to grouped crystals present, Empty occur sacs, bundles, sometimes needle- veins (table IV). major also usually and pith. sometimes to more or corroded appearance a when M. fragrans. crystals sometimes present in the in M. maxima in the abaxially, cells of villosa also specimens only, clustered brachy- often with tissue of the caps of the vein spindle- epidermal cells, mostly solitary (table IV). and clustered spherical and larger relatively few, large, irregularly shaped particles epidermal tanniniferous probably sclerenchyma a of 1981 be peripheral ground probably secretory, cells, may 1, adaxially (table IV). Flattened, edges, 6 rounded — No. epidermal cells, mostly solitary, crystals, composed (fig. lOe) present forms 4 27. in this a for study. genera. constant for a species. List Pitting of cuticular Presence of Shape of abaxial of cells in (papillae excepted) ledges prominence of interruptions bundle of the *) Does not apply midrib in the adaxial midrib*) for Number of of phloem groups in Gymnacranthera. phloem bundles in the pith of the midrib Presence the adaxially*) Presence of layers the collateral bundles of the midrib epidermal section Presence of stomatal Degree Number of flanges polar T-pieces transverse 1 of locally developed dermis in the midrib Frequency of druses Frequency of oil cells Presence of tanniniferous sacs hypo- J. K.OSTER & The characters, P. Baas: Leaf mentioned in list anatomy I do 147 of Asiatic Myristicaceae not have diagnostic any or taxonomic value. The next for the characters applicable are to one or two and genera only, not constant are species (or varieties). List 2 Basal cells of hairs in a row Minor veins (Myristica) Difference in size of adaxial and abaxial cells epidermal Presence of adaxial Presence of (Myristica) in stomata adaxial the of giant bundles stomata (Gymnacranthera) The characters of list 2 do the value, especially the frequent one or two but for genera a diagnostic they walls in adaxial the and midrib abaxial (Horsjleldia, occurrence or are few within the applicable to taxonomic value on the these 'prominence genera are species of the minor veins', concerned. each genus and genera of the indicative circumstantial some species (or varieties) they especially, transcurrence (Knema, Myristica) may have latter features and the two The characters of list 3 species, have not genus level because of their of the of in of vertically veins on the ( Knema) Knema) Degree level either, but not periclinal division Connection water or in prominent or hypodermis (Horsfieldia) the epidermis (Gymnacranthera) Presence slightly epidermis are not constant for most When the latter applies are. to mentioned. List 3 Cuticular layer granular or Presence ( Gym- not midrib nacranthera, Horsfieldia) Sizes of guard cell pairs (Gymnacran- of in the of pith the of in chlorenchyma the midrib cells secretory (Hors- Shape fieldia) of veins adaxially (Gymnacran- thera) Length/width chyma xylem (Myristica) Continuity thera, Horsfieldia) Frequency of cells ratio of palisade Size of druses paren- Position of (Knema). Relative size of midrib the (Myristica) brachy- midrib and to astrosclereids in major veins (My- ristica) These characters have species or In the to variety following species. are The latter characters, without either acquisition (very little) diagnostic some particular list those characters each genus, and which for all some level in them or constant species having are for may be on the genera. mentioned, most of the thought become fixed gradual loss). and taxonomic value to in the which are usually applicable species (or varieties), show a 'tendency' speciation process but not for these (through BLUMEA 148 VOL. 27, No. 1. 1981 List 4 Number of basal of parts hairs Degree of covering of the adult leaves Leaf by walls Anticlinal Thin grooves of the cuticle cells epidermal or Thickness of the lamina of of layers of palisade Presence paren- The characters of list 4 have the characters are by ecological factors, Myristicaceae not of oil thought to cells in the of the pith of diagnostic and taxonomic very strong, however: and some most of them of them may be brachy- value can astrosclereids to on the possibly age-dependent species level; be influenced in the persistent leaves. In list 5 characters for and spindle-shaped midrib Frequency constant epidermis, to not prismatic crystals cells chyma flattened, when druse adjacent are Frequency Thickness of the cuticle Number cells Epidermal not or peripheral tissue of the midrib idioblasts of cuticle present not or of sclerification of the ground not or areas in sclerified margin Degree Presence of central spongy tissue the hairs sinuous of sclerification of the Degree per surface unit most show are of the a mentioned, applicable The species. tendency species, to one or two for which genera only, which character is a inconstant, are are for that character. List 5 Presence of cuticular striations Hors- ( Presence of fieldia, Knema) Number matal of pith cells, complex the bordering the (forming Minor veins sto- of 'ring'; on sunken slightly Presence the level of the of a {Horsfieldia, Myristica-, Myristica is Presence of Presence locally bundles as fibres peculiar cells of sclereids hypodermis the as rows or epidermal or ( Horsfieldia) continuous developed hypodermis Spongy epidermis xylem sclerenchyma Presence Myristica) Stomata large in the of the midrib (Knema) small bundles (Knema) crystals in the (Myristica, Knema) relatively thick filiform cells in (Knema)*) of swollen the subepidermal layer (Horsfieldia) in Knema and highly inconstant) tissue compact or not {Horsfieldia, Knema) Abaxial of layers rified or not spongy tissue scle- (Gymnacranthera, Knema) These characters also have level, *) but only some diagnostic and taxonomic value in certain genera. The value is low The thin filiform sclereids in Gymnacranthera are on the not meant here on genus level. the species J. Koster & The characters following constant for the P. species, Baas: anatomy applicable are but Leaf of Asiatic to one for the genera in not 149 Myristicuceae two or genera and only, are question. List 6 Type of hairs with cells with one Presence arm (Horsfieldia, Knema) Basal parts of hairs often joined or stomatal Basal parts of hairs, when cell, very small one cells of hairs unspecialized cork regular many of composed or not epidermal not warts than cells or or chyma not or (Knema) with veins Major not (Myristica) conspicuous sclerified and with parenchyma large pits the the cells, bordering complex, (Myristica) Palisade (Myristica) higher of Upright papillae not (Horsfieldia) Basal of (Horsfieldia, Knema) little very with or scleren- (Gymna- caps cranthera) The characters of list 6 have certain genera; The species on few characters next and for diagnostic and taxonomic value on the level in species the genus level the value is low. most are applicable to each genus and are for the constant of the genera. The genus for which the character is inconstant is mentioned. List 7 Basal Alveolar not of hairs parts present or not adaxial surface on material cells obviously present and (Horsfieldia of Degree (Myristica) being in embedded of the the guard cells subsidiary (Horsfieldia) or Stomata sunken below Myristica) epidermis Midrib or not the level of the (Horsfieldia) raised adaxially not or (Horsfieldia) These characters have genera, and taxonomic value on the diagnostic for which the characters are inconstant; species level for the the characters also have and taxonomic value on the genus level, but this value is not of the same diagnostic degree for each genus. In the constant following list those characters for each genus, but are are which mentioned, inconstant for the are applicable to and family. List 8 Basal of parts frequent on Arms of hairs the latter hairs the most midrib one or two case, of the or least abaxially size or of bordering per cell, and, in same Structure not Major or veins with caps (and Position of The characters of list 8 have diagnostic stomatal complex, the cells included big sclerenchyma the size of the druses in the and taxonomic value on sheaths caps) mesophyll the genus level. BLUMEA 150 In the last list characters some and which only, are No. 1, 1981 mentioned, which applicable are to one genus for that genus. constant are VOL. 27, List 9 vascular bundle in Adaxial the midrib Filiform, absent ( Gymnacranthera) Phloem groups midrib small of collateral The classification of characters, noted that far as Dicotyledons for possibilities Rijksherbarium (Baas, 1975), the taxonomic general Icacinaceae (Van the genus not strict. the midrib & Baas, Baas, and 1973) Den 1973; more that likely the throughout confirms proposed Vliet & is the different levels. It may be occurrence & Baas, level. When it studied, in the evergreen limitations and studies earlier of widely differentfamilies such (Jansen Staveren on analysis as purposes representatives Celastraceae the of on is would have been above 1978), Crypteroniaceae (Van Baas, tissue outlined above, be inconstant to concerned, on sclereids and the and taxonomic value as characters of as are branched mesophyll ground species more characters would appear more rarely in (Gymnacranthera) diagnostic and species per bundles in ( Horsfieldia) These characters have specimens present Hartog-Van Ter Tholen & 1975), Guttiferae (Baas, Winteraceae the in Aquifoliaceae as 1970), 1973; Vink, (Bongers, 1977). important Some Knema' and have Horsfieldia studied, papillae in Sinclair examined have reasons are to at most moreover papillae papillae species in dome-shaped full range of the occur occurrence of Horsfieldia, not unfortunately only M. gigantea and unspecialized M. one in cells. species. to species Myristica species epidermal cells specimen conspicuous maingayi, the other for these are constant have such astrosclereids is unknown. The very slightly dome-shaped do of lists. closely related, whereas as suppose, that the different: the genus; (1968) the foregoing of which species the constancy for the so Myristica, however, regarded by and included in the have been recorded in Papillae has been epidermal papillae (in species respectively), been not as the type and distribution of the alveolar material (variable in Knema and Myristica characters such Myristica), papillae pronounced shapes This In Knema as in this occur in gives things Myristica. These facts make the constancy of this character in Knema doubtful. The constancy for Horsfieldia papillae reflecting The of the alveolar studied with only one specimen of this character for over the leafis restricted plumulosa, placed these for constant C. In to a a these species In scanning they (sub)species character electron species per (a but were into regarded taxa variable in microscopy Baas, as in of lower rank. Myristica) only; however, in this has been examined. Therefore the constancy the stomatal study by series. This Winteraceae alveolar material is be can with the is uncertain. The distribution of the alveolar material bordering one 1970) possibly character variable in Knema. The presence of the material is the cells, represented species. material species Cratoxylum (Guttiferae, formosum, further differentiation of the a type thoroughly S.E.M. is unknown. iryaghedhi not are (Bongers, variable in some one complexes specimen each; suggests 1973) species. a mainly in K. intermedia and De Wilde (1979) K. has constancy of this character for the distribution and the type of the J. Koster & P. Baas: Lea/ Astrosclereids have been recorded for superba; only according be specimen one Sinclair to for these constant in Knema have either a leaf are on a constant as far and H. species are, expected some The and of Specimens numbers, cells, one whereas frequently most Myristica species, trend for Knema inconstant species. of the cuticle in Knema sculpturings to (not and Knema Horsfieldia occur. smallnumber of basal a Gymnacranthera is rufa only specimen examined of this species, while been observed in other not These the constancy is concerned: The as Horsfieldia species. for convex because it is the unknown, number or high and reflect Horsfieldia species The constancy of the has studied. with small numbers also species this character is inconstant for for been The character 'number of basal cells of the hairs' species. but in these genera species, found has numbers of basal cells per hair have been counted in highest 151 Myristicaceae related. Therefore this character is (1958) closely mentioned in the lists either) is difficult, species Asiatic species only: Horsfieldia fulva two species per of anatomy simply the feature species. The feature 'isobilateral leaf' (recorded for Horsfieldia sabulosa) will be discussed elsewhere. TAXONOMIC Introductory In the on an remarks following discussions, suggestions evaluation of the species, IMPLICATIONS genera shared and and families. With our anatomy, this phenetic approach is cladistic methods matics in the (cf. sense Bremer & of whether certain character types in Knema are states perhaps will be further significance feasable than an attempt towards plea, Wanntorp's mainly based because there is and their this, to applying 1978, for phylogenetic should be considered derved exception an only leaf anatomical characters of present stage of knowledge of systematic leaf more This is Hennig). about natural affinity will be distinguishing or no way of syste- telling ancestral. The hair possible phylogenetic explored. In view of the limitationsof the research materials, the taxonomic suggestions can be tentative. only Below the genus level Gymnacranthera This genus is this because of of hairs some, per the fairly homogeneous: G. bancana may be context mostly epidermal surface unit, species regarded as features: cuticular layer differ taking a only relatively large not granular, cuticle, anticlinal walls of unspecialized epidermal cells and the chlorenchyma recognized the by of the ground tissue features of G. are bancana var. findings as midrib, sometimes also forbesii These the forbesii are most other, especially continuous not high degree G. and the in the a a in few parenchyma G. forbesii, specimens agreement cells but of G. and G. forbesii. thick G. less adaxial sinuous conspicuously eugeniifolia can be tissue, the peripheral margin. extent. contracta are (1958a, b); distinct species, and the remaining species eugeniifolia position number of basal parts of the leaf to a with Sinclair few characters. In relatively not midrib. a less isolated of sclerification of the spongy present and partly in in a more or as One These specimen very similar. he regarded very close to G. each 152 BLUMEA Where all Malesian to give a key be 1. 1981 and their varieties have been examined, it species and varieites of species and species leaf separated No. enable identification on leaf anatomical characters. A to the Malesian to paper. All VOL. 27, - varieties, except is Gymnacranthera G. and contracta given seems useful synoptical key the end of this at G. forbesii var.forbesii, can anatomically. Horsfieldia It is possible surface unit H. to distinguish features epidermal H. of hairs smooth) and papillose and stomatal are distributed species, which H. the grooves, H. a many more unit all abaxially; abaxial sylveslris mesophyll. epidermis has some, for too tissue spongy surface), and abaxial cuticle more less or Other characters less reticulate pattern, but there or characters in important are common: share the presence of cuticular striations and sylvestris sunken slightly in species and has isobilateral on of hairs per the abaxial on adaxially H. anatomically: parts clearly present, often robust stomatal cuticular striations and the prominent surface (hairs relatively and crassifolia material only species as the over have joined together complex distinctly sunken); features unique H. sabulosa compact); basal numerous of many cells composed iryaghedhi (alveolar H. examined of Horsfieldia species and basal parts often adaxially glabra (basal parts Horsfieldia most crassifolia (relatively with complex relatively epidermal the characters, parts recorded not cells subsidiary basal many with of hairs for the per other Horsfieldia species examined. H. in crassifolia as common, and H. sabulosa also have the H. sabulosa and (and unique of adult for leaves, some relatively thick H. share another glabra Horsfieldia) many are regular (in view our unimportant) features cuticle and lamina. epidermal: cork warts of set hairs characters; abaxially and the important most absent from the lamina completely many secretory cells in the epidermis. H. and H. fulva the other by Sinclair according H. not examined a 1975), to Sinclair, are placed are on concerned. The one Bivalves crassifolia, Section H. H. irya H. sylvestris H. iryaghedhi H. macrocoma H. sabulosa special relationship relationship and H. the by crassifolia sabulosa, species Sinclair. and H. leaf anatomy. H. between H. fulva and H. between as pairs Our results by this mentioned do not superba and crassifolia suggested of the irya. H. H. study, were, support a is supported by sylvestris, has not however, is a species with a and been as the species, placed deviating not this study. between recognized special relationship Also the division into the sections does iryaghedhi far as related superba H. The close closely Trivalves fulva, glabra section mentioned recorded for line. H. Sinclair; not given here, more parviflora glabra characters character this classification is H. A the on mesophyll, results, outlined above, with the classification of Horsfieldia, (1958a, 1974, examined leaf anatomically species Section (species Horsfieldia species. To compare the made superba both have many astrosclereids in the above) in the H. by same between H. appear from the inflorescence (Sinclair, J. K.OSTER & P. it has 1975). Although the other the species, Asiatic Leaf anatomy of of majority the 153 Myristicaceae leaf antomical characters, important some Horsfieldia Baas: recorded for not characters is in range of the Horsfieldia. key No species the to species of is Horsfieldia small number of because of the given, examined. Knema Only a few of the Knema examined species K. laurina have unique has peculiar crystals the mesophyll and Some other K. point (the distribution of kunstleri to a with Most other the these characters with diagram the alveolar cork less (c. 20, largely compartments with many features in formed: Wilde's Group K. K. K. K. linifolia has percoriacea for Knema many unique and hair type 6); mandarahan and and K. (The characters constant each character gives K. K. sclereids in features tenuinervia and conferta and type 4); (hair for each roman — distributed in 6 of the point species. to a are K. behind the close species listed in there appear to be relationship, few features. In this of way groups numbers But when refer to the 1 Group 2 Group whereas series 3 K. ashtonii II K. malayana XI latifolia V K. attenuata X K. oblongifolia I linguiformis VI K. elmeri II K. globularia IV K. laurina I K. IV oblongata K. pseudolaurina I K. pulchra IV K. retusa II K. tomentella X Group 4 K. rubens Group 5 a species species XI 6 K. intermedia IX K. conferta XII K. furfuracea III K. korlhalsii IX K. kunstleri XII K. hookeriana III plumulosa IX K. K. reticulate pattern a previous chapter) possible species pair which may none or are VI Group (which plumulosa classification). curtisii the pallens (sclerified another classification of the from lists 3 common, other combinations share been K. couple of, material on adaxial cuticle. for instance: K. intermedia and K. warts); K. recognized be cinnamomea, K. korthalsii and combination of types, K. hookeriana unique sculptured, a var. psilantha (hair type 5). more or species: a epidermis, one, or a large pits); lunduensis and K. over or special relationship), (many regular mesophyll has rufa share species may hair types in the abaxial individually can basis of leaf anatomical characters: K. ashtonii K. linifolia I K. lunduensis III K. mandarahan IV K. pallens III K. percoriacea III K. psilantha III K. rufa IV K. tenuinervia IV have of De BLUMEA 154 The first group contains lamina thin leaf margin the linguiformis 1. conspicuously not 1981 which have the characters: hair type 7, 8a species midrib sclerified, not No. adaxial cuticle less than /an), than 5 pm thick, adult leaves and VOL. 27, weakly cells palisade parenchyma 10 pm covered with hairs, sclerified. In K. spongy tissue curtisii than wide, scarcely longer are 8b, or abaxial cuticle less thick, and K. unique a feature in Knema. The second group contains This group exhibits the lamina usually intermediate <5 margin adult pm, leaves sclerified weakly important characters, shared < pm, 200 each other in midrib not, with usually cuticle hairs, are: 8b, thickness of or 10 /im, abaxial < tissue spongy and sclerified. moderately of this group by species reticulate pattern. a 1, 2, 7, 8a /on), adaxial covered usually or to features: hair type 120 (> cuticle linked species, following relatively numerous basal parts of hairs per surface unit abaxially, spongy tissue compact, anastomosing as rows of the midrib. K. elmeri and K. pith similar. The second group shares The third group contains thickness of lamina not covered such The fourth group contains and/or 8c, < characters: is rather group hair type 7 abaxial cuticle 4 — weakly sclerified, 6 /mi leaf group, K. kunstleri in the and K. compact spongy covered with The sixth group contains which have the characters: adaxial cuticle usually (strongly) sclerified, shares > leaf species, and 7 spongy tissue and leaf which species have >5 /mi), adaxial cuticle or the /mi, not, spongy tissue not, midrib moderately sclerified. This or linked 3, 6, 10 pm, abaxial cuticle < contains (>200 characters with some hair types have tissue. hairs, the sclerification of the abaxial as characters particular sclerified. thick sclerified adult <5m, some laurina in adult leaves covered with hairs thick, and/or 8b, which exhibit the features: hair type heterogeneous particular, the sixth group, such Other as species, margin group. sclerified. This group shares 8a, lamina or appear sclerified, leaf margin spongy tissue oblongifolia conspicuously not particular of the first adaxial and abaxial cuticle hairs, sclerified, midrib (moderately) not fifth The species thickness of lamina intermediate, adaxial cuticle 5 pin, adult leaves margin with moderately features, common in retusa characters with which have the characters: hair type 7 of the second group; K. species certain species some usually intermediate, conspicuously not sclerified and midrib with veins of sclerenchyma fibres, large xylem bundles in the pith of the midrib and oil cells present in the leaves leaf Other each other in to 5 3, 4, 5 /mi, adult leaves or not of the spongy tissue. reticulate pattern, and a 8c, lamina intermediate covered with margin usually sclerified, important characters, shared by species of the third and species layers midrib of this group hairs, (moderately) are: to thick, spongy tissue sclerified. sclerification of the abaxial layers of the spongy tissue and large xylem bundles in the pith of the midrib. K. furfuracea, in K. long pallens as wide, characters in species K. a K. hookeriana, and K. percoriacea Some of the fourth group, De Wilde (1979), as of series VII and VIII refers to the and K. far were as species especially are with by us. in particular cells and K. group share are up rufa some to are similar; five times as also have many characters with K. korthalsii. below with the classification examined in this studied leaf anatomical group. linifolia of this compared species, not percoriacea palisade parenchyma feature in Knema. K. unique common. The results outlined above by pallens the study are proposed concerned. The number behind each Species species name J. Koster & 1: Series K. K. K. P. Baas: Asiatic 155 Myristicaceae Series Sericeae II: Laurinae Leaf anatomy of III: Series Lamellariae linifolia 6 K. elmeri 2 K. hookeriana 6 oblongifolia 3 K. retusa 2 K. furfuracea 6 pseudolaurina 2 K. ashtonii 2 K. pallens K. laurina 2 K. K. IV: Series Obovoideae V: Series Latifoliae 2 1 K. oblongata 2 K. mandarahan 6 VI: Series K. tenuinervia 6 K. linguiformis 1 K. rufa 6 K. curtisii 1 K. globularia 2 latifolia XI: percoriacea 6 IX: Series pulchra K. 6 lunduensis K. K. 6 Mamillatae plumulosa 4 K. intermedia 4 K. korthalsii 4 K. Curtisianae 6 psilantha XII: Series X: Series Glomeratae K. tomentella 2 K. rubens 3 K. kunstleri 5 K. attenuata 2 K. malayana 3 K. conferta 5 The classification based Wilde's classification. discussed, (1958a, this Wilde describes K. a by to notes linifolia K. and K. oblongata common supported oblongata) together classification with K. will be The classification of Sinclair (1979). with K. with K. close laurina, study. and K. K. retusa, K. resembling (according oblongifolia leaf anatomical common globularia K. placed and K. rubens, in the species of close vegetative parts, attenuata to to species having as globularia According one. with K. these tomentella and other K. curtisii in the has De a note tenuinervia. in series IV placed were Sinclair K. laurina, the to and K. tenuinervia. In linifolia pulchra whereas leaf isolated position, an rufa De Wilde considered K. this by has characters in oblongifolia resemblances whereas in leaf anatomical aspects series) affinities with his series X and XI, and is De supports generally particular in series I, albeit in resemblance between K. leaf anatomical group 2. which and De Wilde together goes heterogeneous have much in the K. species a globularia, De Wilde differences Punctatae will also be cited, if this is relevant. De Wilde placed anatomically leaf anatomical characters on Some with reference 1961) Series Glaucae in (and one De K. group, Wilde, same manner as a K. in leaf anatomy. De Wilde created a series for K. in accordance with this Not half the number of examined in this leaf anatomical study. latifolia, considering of Knema, included species Therefore it is useless groups, however, may help parallelism of the classifications based on characters, there is De Wilde species as related chance that the to a examined. For De Wilde's series of Knema is A a tentative at the to to give a by key as restrict the De Wilde to the related which is macromorphological species, not it may be A key the In view of the study, regarded by fits the group of the that K. lamellariade Wilde the leaf anatomical group 6. A been to and leafanatomical examined in this hoped (1979) has species. possibilities. examined leaf anatomically, species, example, III) fits given series VI study. synoptical key to the (from groups end of this paper. phylogeny The indumentumof Knema can of the hair types be classified in types with in Knema (very) long hairs (type 3, 4, 5, 6 and 8c) and types with shorter hairs (fig. 3). Leaves with long hairs loose 156 BLUMEA these hairs when they retain them usually least Between type 8a and 7 27, No. the adult stage, come to (at VOL. 1, 1981 whereas leaves with shorter hairs partially). there is (short hairs) hairs of type 8a small hairs also distinction. On leaves with sharp no The classification into these types is artificial occur. and made for identification purposes. Hair type 8b and least specialized structures types, having (as long stalk cells, long then occurrence) rare hairs with been of the indicated in with part arms. c) through peculiar through The types between are the 'forerunners' of & laurina hookeriana, 3 and are, latter and 1 and 2 ashtonii K. korthalsii, K. according to K. var. pallens De Wilde however, has species, and K. hairs belonging one to series It is remarkable that the mandarahan and K. the (type 8c, species groups with type 8c. The species group type 8b) these presumably one to are very K. The 6. belonging K. types. belonging types each to excepted. K. type The and linifolia K. remaining These three a hairs all long of group particular just mentioned), species both have with K. korthalsii. species but without long hairs, 4). type 5 and have to together with the derived hair types with and Only furfuracea, intermedia and K. plumulosa in De Wilde's series IX, 4 3. fig. relationships derived hair most K. korthalsii three both have hairs This supports the of group without species (or variety) have hairs ancestral form of the hair types groups and their mutual within are these species have been discussed type 4; to the hair types of K. korthalsii (of group ones The belonging also have rufa, putative, are is outlined in here, percoriacea De Wilde. K. by type 6 and leaf anatomical 7 and 8a for the determination of (1979), closely related, hairs psilantha in short hairs cinnamomea, respectively). before. K. lunduensis and K. placed these (the types c represented by are belonging the proposed as tenuinervia both have hairs been and types cells. types, Wanntorp, 1978). 1, 2, 3, 4, 5 and 6. Type (K. no and 2 may also have been derived from the types 7 and 8a 1 of the hair (Bremer taxa that 8b and type derived characters should be used uniquely stalk cells) (long derivations of these possible are may have been derived from the types 8b (7 and 8a) extension of the stalk phylogeny The Other relatively have may also species each) one a There c. 5 and 6. The other 3, 4, through specialization reduction; another possibility is structures are common!). (b and) the as special 3. fig. The types with short hairs (and c) regarded hairs with long the distal end etc., types with at 1 and 2, recorded for (type type 8b (and be may have been derived from type 8 may stalk cells long derived from reduction arms c The structures. propose evolution lines between the types to argumentes peculiar no moreover natural character of these are 6, in K. structure like are one the of only leaf anatomical relationship. 1,2,3 and 5 have shorter hairs, belonging 2 the derived hair types 1 and 2. Hair type 7 and 8a in different groups, which may be an to different types, occur (together argument for the primitive with nature of types. Myristica The have species some, leaves, basal in mesophyll abaxial examined of for Myristica are fairly Myristica unique features, cells of not hairs, if solitary, large) and hypodermis present, M. very e.g. distinct small; sphaerosperma sclereids in anatomically. M. villosa major Some (remarkable species hairs on the sclerified and druses spongy tissue (guard veins cell and pairs relatively long, druses in the abaxial J. Koster & M. epidermis); fragrans, P. Baas: Lea) only as anatomy has species, 157 of Asiatic Myristicaceae clustered in crystals the abaxial epidermis. M. and M. gigantea maingayi are Most other more or less characters constant question reticulate pattern. The characters in adult on cells epidermal on etc. sclerification of the spindle-shaped characters diagram, cells As in the done for Knema, it appears that 'character own none or complement'; of a crystals a in the the hypodermis, frequency of the mentioned. When these previous chapter) some in thickness of the unit, presence be to species presence of hairs epidermal: per surface 7 of the — epidermis). distributed over the often have mesophyll from lists 3 features in rare tissue of the midrib, and the peripheral ground crystals some abaxial characters the presence of non-epidermal whereas others share common, are are bordering the stomatal complex, (c. 25, largely as papillose a number of basal parts of hairs leaves, cuticle, papillae has its similar and have very (as grooves in the adaxial cuticle and common species have many few features. Of the former nevertheless groups of listed are each species species can in a features in pair be formed. Not each character involved will be mentioned in the discussion of the groups, but those which only are more or M. chrysophylla, M. fatua, globosa, M. and M. subalulata have many lepidota, features, in particular: M. chrysophylla and M. lepidota, common lepidota; M. fatua and M. M. subalulata. The crystals in crystals play important an and M. lepidota; important for with sphaerosperma M. (such adult M. leaves) fatua. features in It is the are M. as and M. villosa have many basal cells mm abaxially striking. cinnamomea and The M. two higher too has to than study are I: series Maximae or sphaerosperma M. chrysophylla series Fatuae M. M. lepidota fatua villosa far as the series Cinnamomeae series Littorales M. 9: maingayi M. as Myristica 8: Maingavae gigantea series Fuscae M. some, M. cinnamomea Section II: mainly common features; and many hairs present have also guatteriifolia series M. 10: more given here, 7: M. maxima M. 30 concerned. Section series of on mainly epidermal (1958a; with the reticulate distribution of the characters in this M. M. the unspecialized epidermal results with Sinclair's revision compare these struggled and the series. His classification is 4. couple again latter Matures have also been found in maxima species 1: a and species. common. interesting Sinclair most this group of shares 2 cells, basal parts of hairs/0.1 globosa of this group, and species and M. M. fatua and M. chrysophylla. guatteriifolia, the hairs concerning are common role. characters with M. M. cinnamomea. M. globosa particular has many features in epidermal M. globosa; fragrans those less characteristic for the groups. M. Fragrantes fragrans II: Fatua 14: series Cimiciferae M. 15: globosa series Subalulatae M. subalulata species, 1968). over the examined BLUMEA 158 The presence of M. this by also study. some deviating and M. it is distinguish this position, notes; he so globosa M. from the Apart sections is common anatomical species to that the series while case 9 as he has this M. to and fragrans varieties examined of M. to the species of section villosa) alone will species together in one from the differing others in those of to present results. the division into villosa, the two to series 8, series 10 is globosa and M. nearest is globosa series 11, M. to also similar to The supported by this study. are M. two rather different, which is in agreement with the are characters. macromorphological key fatua M. a with M. some extent. to Sinclair series 7 is close has resemblances M. has leaves somewhat similar in agreement with the study, common within the sections and series distinct series, a M. M. fatua but M. villosa takes than with these placed has some from deviates features in I) chrysophylla nevertheless some comment: characters, relationships subalulata in many ways; all these remarks No has species more of section M. fragrans of M. has strongly supported and M. vegetative features (of fatua, this is lepidota; supported by According lepidota this too respect his ideas about regarded and sphaerosperma while this from M. given section I in many ways, M. M. guatteriifolia (species species series. Sinclair has sphaerosperma support. Sinclair stated Although series is one of M. ecological have many lepidota in maingayi 1981 1. mountain species. The series 11 needs a cinnamomea and M. in in characters; somewhat distinct II. relationship leaf anatomical chrysophylla: and M. gigantea The close No. VOL. 27, species of is Myristica because of the small number of species given, examined. On the The level genus Asiatic characters genera of the Myristicaceae, unique This applies with cells with midrib in table V. two arms to Gymnacranthera. of different Horsfieldia genus has flat absent, major veins to with is the least distinctive; for the Asiatic part of the The some, important length, subsidiary shallowly the family cells of concave, small (very) branched sclereids present in the unique distinct. leaf anatomically are Each for the The Asiatic features. particular adaxially approximately in the midrib rarely Myristicaceae summarized in are characters stomata are: hairs protruding, adaxial vascular bundle sclerenchyma filiform, caps and mesophyll. only character, constant is the small size of the for the genus and phloem in the groups midrib. Knema has sclerenchyma For length Myristica and the papillae) characters the unique as sheaths the unique 'ring' above the in the of major characters bordering stomatal Knema. two groups: Regarding grouping. According to with supported by the most this characters, Sinclair (1958a) a primitive more or arms conspicuous, less close leaf anatomy and another and concerning Myristica there appear the to and of equal upright be four genera the hairs separate 2) Horsfieldia no reasons are close to as to it provides the most and no Knema motives advanced. last. to This indicate and for such a each other; series of increasing advancement: first Gymnacranthera study; hairs with cells with two (with each genus, those to furthermore he arranged them in Horsfieldia are: cells 1) Gymnacranthera the other the complex. Of the characters, applicable the genera in stomatal structure' and 'star-shaped veins. Myristica, view one is not genus as J. Koster & P. Table V. Baas: Lea)anatomy of Differential leaf anatomical characters + present; — = absent; —/+ = of in = Asiatic Horsfieldia, Gymnacranthera, some 159 Myristicaceae species absent, in others Knema and Myristica present 03 t— <D I 03 o 2 03 O 03 C3 c characters Important Basal of hairs parts Most basal More than 6 Number of basal arms than more one cells surface per basal per X - unit abaxial side of the midrib on - of hair part o Stomata characters Important Arms sunken spongy in of hair cells of different Arms of hair bordering adaxially + + complex flat (to - shallowly + concave) + Phloem groups in midrib small with small Major Thin, veins with filiform, recorded Characters, branched for one Large xylem bundles cells in the Garrat (1933) characters for number in of and presented here, do be the the can is leaf be used family more mentioned as recent stem & Wilson a as (Wilson, and node pollen (Siddiqi - + - + - + - - - - - - - - - - - - + - - - - - - - - - recorded (1980) (1968) found and Wilson ± - ± ± some - + - - - wood deviating deviating chromosome (1978) regarded the anatomy a somewhat different from those of the other genera. not support a distinct position somewhat isolated. A for Horsfieldia; only grouping of certain with the present results. synoptical key to the four genera. level literature the the closest allies of the have been regarded androecium as - - - anatomically supported + - genus: fibres Ehrendorferet al. as cannot Table V In Armstrong Horsfieldia Gymnacranthera On sclerenchyma Horsfieldia species; Armstrong The results, for that subepidermal layer Horsfieldia; of the flowers of genera constant epidermis of the midrib pith Anastomosing veins, composed Swollen - - + and not genus in the - - - sclereids of minor veins in the Prominence + caps sheaths sclerenchyma rarely - - + - sclerenchyma -/+ - - approximately 2 + - - stomatal Adaxial vascular bundle in midrib absent Major veins + - dome-shaped cells above - 1 only: stomatal structure Star-shaped Midrib genera length equal length of stomata cells Subsidiary Ring of cells of single - + (+)- + -/+ "/+ + (+)- - tissue occurring + 1 + distinctly druses in + + 2 Alveolar material Large •c + (-) - hair cell per £ genus: adaxial surface on of hairs parts in occurring a c 52 the family Annonaceae and Myristieaceae. most closely related, 1966; Wilson & Maculans, (Wilson, 1965; Siddiqi & Wilson, & Wilson, 1975b). Based on the Canellaceae have The Canellaceae in based 1967), on been particular the anatomy of the the anatomy of the young 1975a) and the pollen morphology structure the of the Annonaceae BLUMEA 160 have also been mentioned as the (Joshi, Sinclair 1946). To short a family, and ochocoa, Scyphocephalium (Compsoneura ulei, platyspermum, Virola The anatomy does to allied with the strongly this compare Staudtia is family V. of the African Pycnanthus the Iryanthera deviate from that of the on indispensable. specimens and gabonensis) V. loretensis and calophylla, based the leafanatomy of the three cuneatum, lehmanni, Myristicaceae relationship, leaves of Coelocaryon Dialyanthera not 1981 advocated Myristicaceae, acuminata, species (Brochoneura I. survey of characters of each general the survey of the complete most also (1958a) characters. In order macromorphological families, VOL. 27, No. angolensis, American ulei, species Osteophloeum have been examined. pavonis) Asiatic genera in the characters mentioned here. The variation in the presence of the alveolar material and in the vascular of the midrib, system found in the African described recently shows a similar well as found in the Bernardi and by variation Asiatic Myristicaceae in the American genera. The as in as for Spichinger (1980) the Asiatic confidence that the leafanatomical characteristic of the The comprehensive. have genera their implications closest the on relatives on genus other i.e. the level; American genera, give below is possibility would continents, been of the hairs similarities family given of African and American material, and is detailed study some These genera. has also , morphology that require a us fairly some more the scope of this beyond study. The characteristics of the Myristicaceae when hairs have been shed, present. Stomata embeddedin the collateral phloem bundle, cells. with often caps or more also with free and spindle-shaped particles phloem Sclereids the fragrans, complement in the to mesophyll dinisii, the less shaped guard the mesophyll the ground as tissue epidermis. Tanniniferous filiform, rarely a ground of cells large to sacs mannii) epidermis. absent. Sclereids also the basal at small druses the sacs midrib and often present of tissue species not present in the in midrib, has been Cinnamosma examined are: Hairs to not Midrib with an to arc- in groups. Veins with collateral as at spindle-shaped Oil cells present in the cells sometimes usually the (Canella alba, Petiole with three collateral bundles small druses and sometimes characters, bundles, with only abaxially present, paracytic phloem apparently caps. the bundles in the to macranthum, embedded. Leaf dorsiventral. not less branched elements. number of Warburgia tissue of the midrib; secretory Most of the ground targe or arc-shaped the mesophyll and the ground tissue of present in and and sometimes in the Tanniniferous the mesophyll, as arms; mostly adaxially, three collateralbundles mostly astrosclereids in the as phloem more an with collateral groups. Veins leaf anatomy of collateral bundle, the Crystals with free scanty data in the literature. The characteristics bundles, with sclerenchyma end. cells collateralbundle straight recorded. Alveolar material absent. Stomata bicyclocytic; Alveolar material dorsiventral. Midrib with cinnamodendron corticosum. C. madagascariensis, the visible. adaxially. Crystals mainly Oil cells brachy- Canellaceae, C. Hairs uniseriate, with cells with layers; always cells present in the as sometimes also in the Capsicodendron bundles epidermis. midrib; secretory the veins. or two sheaths. Petiole with end, Of a Leaf usually accompanied by xylem centre, sometimes sometimes in the are: remain usually only abaxially present, paracytic; guard subsidiary in separate groups, often in sclerenchyma the basal parts present the basal particles mesophyll in the in and epidermis. absent. Myristicaceae (stomata only abaxially, J. Koster & bundles; crystals have present) Asiatic Leaf anatomy oj wide a druses as and in the 1950). Important characters, present alveolar material, bundles in the Canellaceae. guard the of the pith Therefore, midrib and close a particles; Dicotyledons the the between & Chalk, the peculiar hairs, subsidiary cells, tanniniferous the relationship cells secretory (Metcalfe Myristicaceae (as embedded in cells with sclerenchyma; petiole spindle-shaped distribution among the 161 Myristicaceae leaf dorsiventral; collateral vein bundles with paracytic; three P. Baas: sacs) phloem the absent are these families in the be hardly can supported. The characteristics of the Annonaceae Jovet-Ast to Blunden et al. not Stomata collateral vascular system with free several Petiole with druses and mesophyll sclereids the brachy- mesophyll and/or to the the centre); in the at with the mucilage or same too lack are as absent. sacs astrosclereids Sclereids or as resemblance, sclereids. close Magnoliales (taken candidates for to it between other in close the to large sometimes brachy- as broad A the and hairs (the superficial a the filiform indications of of the flowers in and comparison of versa only groups no less more or close a Myristicaceae. Myristicaceae sense with are are The Annonaceae and vice show phloem there morphology the Myristicaceae Myristicaceae. characters that say families). affinity present the with Cronquist, Myristicaceae. other does 1968) Yet did particular, Annonaceae or families not the of the yield Myristicaceae not other clearly this order through their leaf anatomical features like oil cells and paracytic The embedded alveolar material cells in the guard of the subsidiary Winteraceae (Bongers, cells of the Lauraceae the branched hairs and sympodially vascular system of petiole and midrib, that the take an the additional Magnoliales. separate phloem Myristicaceae and 1973) provide circumstantial evidence of links with individual families of the through small tissue of the midrib and and cavities present in the Myristicaceae the bundles, to the (1972), studying (among stomata. medullary suffice relationship Canellaceae belong in the Some instance). between the Annonaceae and the Also Nair a the as Therefore relationship see for as in sheaths. or thin, branched, filiform sclereids in the those between the Canellaceae and the different, or (not groups caps Crystals ground cells bundles, tissue of the midrib. ground important characters, present very large sclerenchyma the basal end. mesophyll, not an arc- medullary part (not a often in phloem The resemblances between the Annonaceaeand the the cells dorsiventral. Midrib with abaxial and are, and (1969) Alveolar material peltate. or anatomy, West tissue of the midrib; oil cells sometimes in the abaxial ground Tanniniferous or as leaf (1950), consisted of several small arc adaxial, an Oil cells epidermis. and the epidermis. with three vascular strands mostly prismatic particles in especially complex, Veins with collateral bundles, layers). Chalk stellate isobilateral, mostly bundles in the phloem simple, sometimes the bundle, more and usually only abaxially present, paracytic; guard embedded. Leaf sometimes shaped if present, (1973): Hairs, recorded. Metcalfe (1942), the concerning , according It is groups of the isolated position within the order. REMARKS The majority forests (Sinclair, of the 1958a), Myristicaceae a ON are ECOLOGY confined to the lowland very humid habitat. Therefore it is tropical surprising to rain find so 162 BLUMEA characters in xeromorphic many and adult dense a (characters, usually No. 198! 1, A thick cuticle, tomentum sunken stomata, overarched leaves, alveolar material 27, species. most tissue of the midrib, ground VOL. of the by papillae for the constant the These characters do press). lack these particular apparently are adaptations different. Some not conditions to subjected, quite semi-arid a vesselles for phase). now surface unit cells appear Another (if leaf, same to are at is most characters, least the this there is the isobilateral leaf. The any) for important (an factor) call for appears basal many an of hairs for on the Myristica. never are of them is. isobilateral leaf (Esau, 1960), (1974), have the leaves in a more to serve as explanation an of the leaves with different only Sinclair cannot position not parts high papillae not constant Myristicaceae be to for passing similar relic a The three characters one less distichous arrangement. This fact, however, sun i.e. were presumably primitive, species has, according other relics; are Myristicaceae features.) some extent. but character whereas rows, and in species (cf. Doyle, 1978, might (in agreement with complexes each other the perhaps the the alveolar material and the Horsfieldia sabulosa; leaves in several or on xeromorphic for adult leaves on areolae), replace to present together recorded the mesic habitats as species differentiationof angiosperms markedly xeromorphic occupying the stomatal bordering They on of habitat, present early ecological The characters and 1963, Koster & Baas, circumstances of the ancestors the leaf anatomical explaining The dense tomentum per the on Winteraceae, hypothesis the cells regarded may be species (Horsfieldia ecological xeromorphic which to in all occur different from those of the interesting hypothesis through not whereas the features), mesophyll bordering species) features, restricting the transpiration (Burstrom & Odhnoff, in sclerified the abaxial surface of the on regard from that of the to the other Myristicaceae (De Wilde, personal communication). In the conclusion, seems defy to features all the to Myristicaceae provide generalizations prevailing situation in the genus xeromorphic interpretation not even equally a is if in to they to of an (cf. of a Baas, earlier stretches It leaves the of the example plant group which the question for 1975, a similar adaptive significance limits unanswered of of the functional why at present grows in semi-arid habitats. However, family it is visualize the development and retention of so many xeromorphic were research needed hypothesis Myristicaceae speculation. portion another adaptability of various leaf anatomical conditions ecological The at on all enable stages in evolution totally meaningless the actual role of the anatomical householdand gas water badly and difficult characters, the features small Physiological Ilex). the on exchange more in these and other fruitful discussions on structures tropical ecological for survival. concerned in rain forest plants and functional leaf anatomy. SYNOPTICAL KEY TO THE MALESIAN GYMNACRANTHERA a: Number of basal parts of hairs abaxially (average G. 14-22: — 55: the values of about five 2—11: 28 on SPECIES AND (p.p. = pro VARIETIES OF parte) areolae and minor veins counts): contracta p.p., G. eugeniifolia G. contracta p.p., G. forbesii, G. G. bancana paniculata / 0.1 2 mm J. b: Koster & P. Baas: Leaf anatomy of Asiatic Thickness of lamina in /im (This character has when other leaves than examined here < 200: S: 200, G. forbesii <400: G. forbesii >400: c: G. forbesii; G. var. bancana G. panicuiata < 12: >12: paniculata Abaxial Abaxial paniculata eugeniifolia; G. zippeliana var. borneensis var. layers of contracta p.p., G. forbesii; G. contracta p.p., G. eugeniifolia G. bancana layers tissue sclerified, spongy of spongy tissue at G. forbesii eugeniifolia; G. least var. paniculata locally: crassinervis p.p. sclerified: not G. bancana, G. contracta, G. forbesii forbesii Major veins with well G. G. Druse idioblasts developed Druse idioblasts G. var. crassinervis p.p., G. paniculata sclerenchyma caps: forbesii sclerenchyma: paniculata bordering G. forbesii, var. bancana, G. contracta, G. eugeniifolia, G. veins with little Major f: reservation; quite possible) var. contracta; G. G. G. e: some are Thickness of adaxial cuticle in /im: >7, d: bancana; var. be used with G. crassinervis; var. bancana to studied, overlaps are 163 Myristicaceae the abaxial on eugeniifolia bordering var. epidermis only: griffithii the abaxial and adaxial on G. bancana, G. contracta, G. epidermis: eugeniifolia var. eugeniifolia, G. eugeniifolia, G. forbesii, G. paniculata g: Minute, prismatic crystals present in the abaxial epidermis: G. bancana var. bancana Minute, prismatic crystals absent: G. bancana G. forbesii, SYNOPTICAL (the a: Adult leaves groups densely 2 Adult leaves not KEY borneensis, TO THE specified are 5 very G. contracta, G. GROUPS OF KNEMA on p. covered with hairs p.p., (or var. paniculata 153) (p.p. = SPECIES pro parte) abaxially: p.p. sparsely) covered with hairs: 1, 2 p.p., 3, 4, 5 p.p., 6 b: Hair type (see page 141): 1: 2 p.p. 2: 2 6: 4 p.p. 7: 1 p.p., 2 p.p., 3: 4 p.p., 6 p.p. 8a: 1 p.p., 2 p.p., 5 p.p. 4: 8b: 1 p.p., 2 p.p., 3 p.p. 6 p.p. 5: 6 p.p. c: Number of basal parts abaxially (average < 12: > 12: 8c: 3 p.p., 4 p.p., 5 p.p. 4 p.p., 6 p.p. of hairs on the values of about five counts): 1, 2 p.p., 3 p.p., 4, 5, 6 2 p.p., 3 p.p. areolae and minor veins / 0.1 2 mm BLUMEA 164 d: Thickness of lamina in /im character has (This when other leaves than examined here :g 120: > 120, No. VOL. 27, are 1981 1. to be used with studied, overlaps some reservation; quite possible): are 1, 2 p.p., 3 p.p. <200: 2 p.p., 3 p.p., 4, 6 p.p. 2 p.p., 5, 6 p.p. Thickness of adaxial cuticle in /mi: e: >5, 10: < 10: 1 p.p., 2 p.p., 3, 1 p.p., 2 p.p., 4 p.p., 5 4 p.p., 6 p.p. p.p., 6 p.p. 5 p.p., 6 p.p. Thickness of abaxial cuticle in /mi: f: <5: 1, 2, 3, 4, 5 p.p., 6 p.p. 5 p.p., 6 p.p. tissue Spongy g: sclerified: obviously 2 p.p., tissue Spongy 3, 5 p.p., 6 sclerified: (obviously) not 1, 2 p.p., 4, 5 p.p. h: Leaf margin sclerified: Leaf margin not 2 p.p., 5 p.p., 6 p.p. sclerified: 1, 2 p.p., 3, 4, 5 p.p., 6 p.p. tissue of midrib Peripheral ground i: 2 p.p., 3, tissue of midrib Peripheral ground bundles in Xylem sclerified: weakly sclerified not: or 2 p.p. 1, j: strongly 6 4, 5, pith of midrib large (see fig. 5): 2 p.p., 6 p.p. bundles in Xylem pith Regular cork k: warts of midrib 2 p.p., 3, 1, not large or absent: 4, 5, 6 p.p. abundantly present: 5 cork Regular warts absent: 1, 2, 3, 4, 6 ACKNOWLEDGEMENTS We wish to thank on Knema Horsfieldia. in the text Dr. W. to their prior Mr. M. figures J. J. O. de Wilde publication. Dr. Groeneveld, and for J. E. Mr. B. N. photoplates. Special givingvaluable taxonomic advice and Armstrong kindly provided Kieft and Mr. C. L. Marks thanks are due to Dr. are D. F. with us access to acknowledged for Cutler his results his thesis copy of a for testing on their share the alveolar material. REFERENCES ARMSTRONG, J. E. 1975. Thesis. Miami & T. K. The comparative University, Oxford, 1978. WILSON, Floral anatomy and morphology of Horsfieldia (Myristicaceae). Ohio. morphology of Horsfieldia (Myristicaceae). Amer. J. Bot. 65: 441 -449. BAAS, & — P. 1970. Anatomical contributions to from 1975. 1980. Wood Madagascar Anatomy and 22: Horsfieldia (Myristicaceae). Cratoxylum Vegetative anatomy Oncotheca. Blumea of and taxonomy affinities of IAWA I. Floral and from Indo-Malesia the 311 -407. plant Bull. N.S. I: 121 vegetative anatomy (Guttiferae). Blumea — 129. of Eliaea 18: 369 — 391. Aquifoliaceae, Sphenoslemon, Phelline and , J. K.OSTER & 1. W. &C. G. BAILEY, and & L., 133- J. Am. Asiatic of anatomy comparative morphology of Arb. 25: 1980. SPICHIGER, R. Leaf 165 Myristicaceae the Winteraceae V. Foliar epidermis 342-348. Las Miristicaceas del Jenaro Herrera. Arboretum Candollea 35: 182. BI.UNDEN, G., AYE KYI & K. macrophyllus, H. G., BURSTROM, comparativeleaf and G. velutinus. leaf characters 1978. WANNTORP, & C. 1973. The JEWERS, malayanus Epidermal E. & H. BREMER, K., G. 1973. M. J. BONGERS, 1944. The NAST, sclerenchyma. BERNARDI, P. Baas: anatomy of Goniothamus Linn. Soc. 67: Blumea in botany. Vegetative 21: G. 381 -411. Taxon 27: 317 — Alb. plants. Stockholm, of anatomy andersonii, 361 —376. of the Winteraceae. Phylogenetic systematics 1963. ODHNOFF, Bot. J. 329. Bonniers Boktryckeri. A. CRONQUIST, 1968. The evolution of and classification Mifflin flowering plants. Boston, Houghton Company. DOYLE, J. A. 1978. Fossil evidence P. systems (Ed. EHRENDORFER, F., F. KRENDL, E. K. ESAU, 1960. G. Anatomy A. HARTOG-VAN TER lato. Acta W. JANSEN, Bot. Neerl. P. & T., 1946. affinities A 1942. on J., & P. METCALFE, C. N. C. (in R., certain S. Murty living in 1978. of Woods Myristicaceae. Trop. Epidermal characters 35: 6 — of the Celastraceae 48. sensu 388. — leaf Comparative of anatomy Kokoona and Lophopetalum 153— 178. the development of pollen J. les sur national of Bot. Indian Myristica fragrans Soc. 25: d'histoire van Houtten and the 139- 143. d'lndochine. - Annonacees du Museum 1981. Alveolar material the Anatomie naturelle; - foliaire. Repartition fascicule serie; XVI, nouvelle & L. of nutmeg sex A. N., & W. Y. Flora Jena 1975b. Pollen Leaf J. 1958b. Curr. Sc. 46: malabarica Myristica In: Advances in Press. Lamk. with a discussion Y. plant morphology (Ed. simple morphological technique for distinguishing 156— 157. sclereids in certain 1975a. Stem members of Annonaceae anatomy and nodal anatomy of the and Myristicaceae. genus Knema. Pak. J. The whitmorei The Bull. Bull. genus — XX. - 23: XXXI. 18: Sing. precursores Sing. 102 — — spp. The Gard. genus Bull. 21: 167— Sing. 16: 175. 205-472. Gymnacranthera (Myristicaceae) in nov. The genus Knema (Myristicaceae) in Malaysia and 327. XLII. Gard. Bull. The in Sing. Horsfieldia (Myristicaceae) genus 197-200. in Myristica genus Malesia and outside 1—540. Horsfieldia (Myristicaceae) J. Sinclair 7: 17: 96—120. precursores Gard. Pak. J. Bot. (Myristicaceae). Biologia Malayan Myristicaceae. precursores Sing. Malesianae Gard. (Myristicaceae). genus Knema of the Malesianae Gard. Bull. Florae Knema of the revision Malaysia. Malesia. 1975. genus Florae Malesianae 1974. of Myristicaceae. 1977. A VATSALA, 1966. Foliar WILSON, of the Florae outside 1968. of Clarendon Dicotyledons. Oxford, of the Anatomy systematics seedlings. anatomy 1958a. A Malaysia. 1961. Bot. J. 67-76. 1975c. SINCLAIR, Suppl. 156: 220-231. M. R. & T. K. Bot. 7: Cuticle. 277. — CHIN, B. In: The Plant Myristicaceae. morphology and embryology 264 al.). 1950. CHALK, aspects of the et in press). B. K., R. RAJENDRA & P. SIDDIQI, as trees Univ. Press. & Sons. London, Wiley BAAS, 1973. Recherches 1972. Floral on the RAO, Tropical Cambridge 1968. Chromosome numbers and evolution SAUER, den, & family Myristicaceae. BAAS, Linn. Soc. NAYAR, 3-30. 353. New York, P. and forests. In: trees 125-308. KOSTER, NAIR, — of the wood 21: geographique.Memoires 3: plants. 355 BAAS, note of the S. JOVET-AST, 27: Blumea & W. 337 tropical Zimmermann) anatomy R. M. THOLEN, (Celastraceae). JOSHI, A. C. of seed 17: of origin HABELER, Systematic 1933. the & M. H. Taxon primitive angiosperms. GARRAT, on B. Tomlinson in and 27: and outside Malesia I: H. sabulosa and H. Sing. 28: 133-141. outside Malesia II. Gard. Bull. 1-181. SOLEREDER, H. 1899 & 1908. Systematische Anatomie der Dicotyledonen & Erganzungsband. Stuttgart, Enke. STACE, C. A. STAVEREN, 1965. Cuticular M. G. C. Neerl. 22: VAN & studies P. BAAS, as an 1973. aid to plant VLIET, Bull. Brit. Mus. of the Malesian (Bot.) Vol. Icacinaceae. 4 no. 1. Acta Bot. 329-359. VINK, W. 1977. The Winteraceae of the old world II. 23: taxonomy. Epidermal characters Zygogvnum — Morphology and taxonomy. Blumea 219-250. G. J. C. 173-195. M. VAN & P. BAAS, 1975. Comparative anatomy of Crypteroniaceae s.l. Blumea 22: BLUMEA 166 1897. Monographie WEST, W. C. WILDE, 1969. WILSON, T. K. 1966. The Amer. WORSDELL, 1 Photos 170. hair type 5, hair type 7, C. 52: 369 — 1967. The Internal 4. K. Photos 7—12. types 8b, x 350. material as a hairs, and x Scanning 140. — phloem in 3, material note tall on warty abaxial rods (heavily restricted to stomatal a alveolar corresponding surface with complex, warty layer. complex (s), (‘spongy’) electron infested x material, with convex with Note 800. anticlinal CI. 96: Blumea 329 — 25: of the Anatomy — 82. 344. 321 -478. young stem morphology and conclusions. Ann. Bot. Lond. types in Knema. note tall bodies cuticle, alveolar x type 8c, of hair 820. — material, tall 90. x — hairs, the x 1. — K. Myristica fragrans x 9. material laurina, x — 80. — oblongata, hair type 1, 3. K. lunduensis, 5. K. globularia, hair type 8a, note 170. material. — (± 11. Knema surface as in — 7. Knema retusa, cinnamomea. thin-walled Myristica 150. 527. 85. x cells, K. entire abaxial intermedia,alveolar 526 lying hairs, alveolar types and of hair plates covering 22: — alveolar oblongata, (exept margin,midrib photo 11) restricted to cells x 410. epidermis covered above stomatal bot. I. Flowers of Myristicaceae. of hair with micrographs 12. Knema — II. IV. Floral stalk cells of free abaxial mandarahan, hair of erect, lobed 13 — 18. Scanning Photos opening electron of the Myristica. plumulosa, hair type 6, complex, Torrey 1 —680. 54: 214-220. micrographs hair type Knema 1730. x stomatal Bull. 13: 78 68: (Myristicaceae). the Canellaceae. the Canellaceae. Myristica villosa, thick-walled, reticulum major veins), overlying papillose 10. — Deutsch. Bot. Ges. Leop.-Carol. Ranales. genus Knema morphology J. Bot. electron also alveolar note woody d. Ber. Acta Acad. 378. dome-shaped unspecialized epidermal cells hair the of the 1981 1. 343. Amer. percoriacea, — — 336 1908. 170. x in comparative morphology of — 6. Scanning 2. K. — Bot. MACULANS, W. of oil cells Nova comparative morphology of M. malabarica. and x The Amer. J. J. Bot. 53: & L. M. Myristicaceen. 1979. New account DE, 1965. and node. der Ontogeny W. J. J. O. No. dieHaarbildungderMyristicaceen. O. 1895. Ueber WARBURG, VOL. 27. x fungal hyphae), 16. 270. walls; sculpturings material of erect features. plates. part of hair x 1660. (arrow) 15. — and — Note alveolar 820. — 14. Myristica globosa, x basal — of miscellaneous micrographs with alveolar of cells Myristica maingayi, of cells material Myristica ring 13. ring villosa, overlying as a alveolar and Horsfteldia iryaghedhi, abaxial, papillate epidermis — 17. Myristica maingayi, note basal of cuticle part which of hair are adaxial in centre; much smaller x surface with 400. than — cells, of material surrounding with grooves 18. Knema individual surrounding reticulum rufa, x porous in cuticle adaxial 1470. J. Koster & P. Baas: Leaf anatomy of Asiatic Myristicaceae 167 168 BLUMEA VOL. 27, No. 1. 1981 J. Koster & P. Baas: Leaf anatomy of Asiatic Myristicaceae 169 BLUMEA 170 Photos with tall 19 — basal thick cuticle 22. Light micrographs cell, on x tips 600. of x 20. papillae; prominent striations, and cuticle with — VOL. of various 27, No. I, features. epidermal Myrislica maingayi, transverse x 240. basal part undulating anticlinal — at five cells, high focus x 19. — Myristica guatteriifolia, hair papillate epidermis. Note abaxial cuticular maceration with section 21. Horsfieldia crassifolia, ofhair with flanges 1981 with 380. — 22. Gymnacranihera forbesii,: adaxial and thin areas of cuticle in loops of undulations, 600. Photos 23 of abaxial stomatal Knema — 27. complex laurina, complex Light micrographs of stomatal complexes. — 23. Knema laurina. epidermis showing star-shaped opening (arrows) overlying and in focus; transverse leaving a inset: section star-shaped structure as seen showing ± horizontally star-shaped opening as shown in stomatal in cuticular directed photo 23, paradermal section complex; maceration, papillae overarching x 600. — 25. x at bottom 600. — 24. the stomatal Myristica. — a. M. J. Koster & P. Baas: chrysophylla, paradermal section showing rings — x b. M 380. cinnamomea, ibid, — 26. Myrislica chrysophylla. cells, forming ring cells, x 600. in cuticular — as shown 27. in photo Gymnacranthera 171 Leaf anatomy of Asiatic Myristicaceae of cells overlying maceration. Note transverse 25. Note section of stomatal also small cells contracta, and bordering also small basal paradermal complex (arrows) section parts overarched between with the stomatal of hairs both by ± papillate subsidiary paracytic complex. (arrows), cells and stomata, x ring 600. 172 BLUMEA Photos abaxial absent 28 — 32. cuticle near Light micrographs near leaf leaf margin. margin (right), x cells — 29. with flanges x 380. in — contracta, centre, 31. maceration corresponding Ibid., transverse Horsfieldia sylvestris, adaxial with section location hypodermis I, features. 198! — thin cuticle very thin cuticle with of adaxial with very No. 28. Knema pallens, (some arrowed); maceration alveolar Ibid., transverse section, showing globular cells (or mesophyll cells intruding into epidermis?) Gymnacranthera 27, of miscellaneous Note 380. VOL. shown in cuticle, showing region of large crystal subepidermal with as locally druse swollen idioblast with in idioblast, cells relatively in x epidermis 240. — 30. thin anticlinal subepidermal mesophyll, cf. (arrow), photo 28; of material photo 30; x 240. x 380. — 32. J. Photos vertically 33 — 38. Roster & P. Light micrographs transcurrent bundle phloem, entirely composed (arrows), palisade x 240. very sclerenchyma brachy- to — 35. Knema cap of vein astrosclereid, in fibres; cross x pallens, 380. x with a. 380. section 38. 240. — 34. tanniniferous 36. Knema epidermis, x features. Knema note oil cell in — adaxial — x 173 Leaf anatomy of Asiatic Myristicaceae of miscellaneous sheath, of thick-walled, sclereids, mostly Baas: x 33. tubule running percoriacea, 240. 150. — 37. Knema minor mesophyll (arrow); Gymnacranthera (some arrowed), — laurina, veins devoid into filiform vein with xylem or epidermis (arrow); sclereid Gymnacranthera forbesii, contracta, of b. note tanniniferous tubules from vein thick malayana, mesophyll with connecting midrib numerous with filiform