Anatomical contributionsto I.

Floral and vegetative anatomy of Eliaea from Madagascar and

from Indo-Malesia (Guttiferae)

P. Baas

Contents

Summary 369

Introduction 370

Techniques 371

Materials 371

Results and Discussions

Anatomy of the flower in Cratoxylum 372

Anatomy of the flower in Eliaea 373

of Vegetative anatomy Cratoxylum 373 of Eliaea Vegetative anatomy 381 Occurrence of papillae in Cratoxylum 383 Floral and xylem specialization and related problems 383

Sectional delimitation and in the 385 anatomy Cratoxylum and Eliaea distinct Cratoxylum as genera 387 Acknowledgements 387

References 387

Summary

floral and ofthe allied and The vegetative anatomy closely guttiferous genera Cratoxylum (Indo-Malesia) have in Eliaea (Madagascar) is described. Although the two taxa many features common, differences are sub- stantial them distinct from the enough to recognize as genera, as appears following survey:

Cratoxylum Eliaea

Ovary with 12 or more ovules Ovary with 6(—8) ovules

with with three Ovary 3(or 4) incomplete true septa Ovary 6(or 8) incomplete septa,

being true, three false Styles and filaments glabrous Styles and filaments mostly villous Secondary xylem with scarcely pitted fibres Secondary xylem with densely pitted fibre-

tracheids

Vessels solitary and in radial multiples Vessels almost exclusively solitary

Silica bodies in cells ofwood Silica bodies absent present ray

insertion in view The vascularization and of the hypogynous scales both genera favours the that these structures may be interpreted as staminodial derivatives.

In into rank is the some species of Cratoxylum a further differentiation taxa of lower suggested by presence or absence of abaxial epidermal papillae on the leaves; this is related to geographical distribution.

Floral and xylem specialization of Cratoxylum and Eliaea are discussed. The xylem structure of Eliaea than but flowers be shows more primitive features that of Cratoxylum , the seem to more specialized. The possible implications of these findings for plant geography are discussed.

This is the first of series in which the, of of *) paper a mainly vegetative, anatomy dicotyledonous taxa different ranks will be described and used discuss their affinities. (families, genera,species) to Anatomical studies on Carlemanniaceae,Dirachmaceae, Huaccae, Lophopyxis, Oncotheca, Phelline, and Sphenostemon are in progress.

369 370 BLUMEA VOL. XVIII, No. 2. 1970

section In Cratoxylum the Isopterygium (C. arborescens and C. glaucum) is very different in its wood anatomy

from the other representatives of Cratoxylum. This difference (concerning parenchyma distribution) is

habit in section sections correlated with the evergreen the Isopterygium. The other of Cratoxylum are decid- with data from literature about other with deciduous and uous. A comparison some groups evergreen

members is made.

Introduction

the Blume In his monograph of genus Cratoxylum (Guttiferae, Hypericoideae, Cratoxy-

leae) Gogelein (1968: 457) expressed his views on the relationship of this genus with the

monotypic Eliaea Camb. from Madagascar: 'If I would have studied material of Eliaea before I whether I would have of that genus was described, doubt made a separate genus it, but would certainly have given it a distinction as a section or even subgenus close to sect. Tridesmos, most closest to C. maingayi’. He listed the distinctive characters of the

two genera as follows:

Cratoxylum Eliaea

Ovary with at least 12 ovules Ovary with 6 ovules

Ovary incompletely 3-celled Ovary incompletely 6-celled Styles and fdaments glabrous Styles andfdaments villous ('cottonny hairy')

Fruit with three valves, often partly Fruit with finally 6 semipersistent caducous valves

doubt about the nature ofthe Gogelein expressed some true ovary ofEliaea, because of conflicting evidence given by Engler (1925) and Perrier de la Bathie (1951). The former the and fruit three-celled and the latter in his called ovary reported key 'loges biovulees', refer three-celled but his which would to the condition, in generic description the ovary is said to have '6(—8) loges incompletes uniovulees'. Gogelein himself suggested on the

observations that the of Eliaea three and ground of his limited ovary possesses true three thus confirmed the of Cambesscdes false, incomplete septa. He original description (1830).

for a further detaded about the nature of the finds Gogelein's plea study ovary its response this detaded of the of both in paper. Moreover, a study vegetative anatomy genera was undertakenin order to establish the mutual affinities with more certainty. The distribution Indo-Malesia and several pattern Madagascar occurs in plant genera (e.g. Nepenthes, A of within in Allanthospermum). better understanding the relationships such groups, be able this distribution considered be order to to interpret pattern, is to sufficiently important to justify an elaborate investigation of this kind. substantial of anatomical information Although a amount on Cratoxylum and some the Eliaea have been of notes on secretory cavities in published, a comprehensive account these Accu- the anatomy of genera would add to our knowledge of Guttiferae anatomy. of of families becomes mulation of descriptive accounts representatives large more and

if anatomists use their in more urgent, comparative want to knowledge successfully discussing the affinities of enigmatic small families. Solereder (1899, 1908) and Metcalfe and Chalk (1950) comprehensively summarized the Additional available information on Hypericoideae anatomy. information on structure and apphcations of the timber can be found in Brazier and Franklin (1961), Chowdhury and Balan Schofield and Ghosh (1958), Menon, (1956, 1967). (1968) gave particulars on nodal and petiole anatomy. Chemotaxonomical data on Cratoxylum were summarized

al. by Hegnauer (1966). Jacquemin et (1969) investigated some chemical compounds of Eliaea. P. Baas: Anatomy of Eliaea and Cratoxyhm 371

The anatomical descriptions in this paper were made according to a more or less standardized scheme largely following Metcalfe and Chalk's practice. Accounts ofmature

detailed than those of the of secondary xylem are more secondary xylem young twigs.

TECHNIQUES

For the study of the floral anatomy, series ofpickled young buds and flowers, and in

in the case of Eliaea also fruits, were embedded paraffin wax; serial sections from flower stalk to stigma were stained with auramin, safranin and astrablue according to Maacz and

cleared Vautier Vagas (1961). In addition, other flowers and buds were according to

to which added of (1949) at 40° C in chloral-lactophenol were some drops 30 % hydrogen peroxide. Leaf studied the middle of the lamina anatomy was in transverse sections through and distal and and the basal part of the petiole. Epidermal characters were studied in scrapes of volumes of cuticular preparations (using a mixture equal 20 % hydrogen peroxide and concentrated glacial acetic acid, at 60° C.)

and wood studied in radial and Twig anatomy were transverse, tangential sections. features such Quantitative as vessel member length and fibre length were measured in macerated material. The macerations made in the same the cuticular were way as prep- arations.

the For wood elements means are based on 25 measurements. Other means (stomatal made diameter etc.) are based on 3 —10 measurements. Illustrations were using a Leitz

Orthoplan microscope with drawing tube.

MATERIALS

about In the following list of specimens, particulars are given the investigations carried them. All material studied derived from herbarium and wood out on was specimens present at Leiden, unless stated ortherwise. Institutional wood collections from which specimens were obtained are abbreviated according to Stern (1967). Slides for reference are available at the Rijksherbarium; duplicates of some of them have been deposited in the Kew slide collection.

line (Vahl) Blume. Pickled material from Kepong Arboretum (tree 253, 5) (clearing For. and sectioning of flowers); Malaya, Kadim & Noor 155; Sumatra, Serv. bb 33096 (leaves and twigs);

S. H. SAN Fuchs and Borneo, 25136; 21290 21346 (wood samples). Hallier C. cochinchinense (Lour.) Blume. Hongkong, s.n. (clearing of flowers); Sumatra, Alston 14359 (leaves and twigs); Borneo, Meyer SAN 25128 (wood sample).

C. formosum (Jack) Dyer ssp. formosum. Pickled material from Botanic Gardens (clearing and sectioning of flowers); Java, Cult. Hort. Bog. VI. C, 52a; Borneo, Buwalda 7979; Sumatra, Forbes 3298;

Borneo, Meyer 53506 (leaves and twigs).

C. Pickled material from Bot. Gardens and formosum (Jack) Dyer ssp. pruniflorum. Singapore (clearing Hort. XV. A. XII. sectioning of flowers); Java, Cult. Bog. J. 3 (leaves and twigs). C. formosum (Jack) Dyer. Borneo, SAN 19.6.61 (wood sample).

C. glaucumKcrth. Borneo, Philip. Nat. Herb. 87634 (clearing offlowers); Borneo, Boschb. Proefst. bb 14411 (leaves and twigs) Boschb. Proefst. Bogor Number wood coll. 6468 (= bb 6770 of herbarium series) (wood sample ex RTIw).

C. maingayi Dyer. Singapore, Burkiii HMB 2607 (clearing of flowers); Sumatra, Grashoff 991 (leaves and

WT twigs); Malaya, 6732 (wood sample ex KEPw).

C. sumatranum (Jack) Blume ssp. sumatranum. Borneo, Darnton 577 (clearing of flowers); Java, Neth. Ind.

For. Serv. bb 3939 (leaves and twigs). S.H. C. sumatranum (Jack) Blume. , F.P.R.I., 321 (wood sample ex CLPw). Eliaea articulata Camb. Pickled material of flowers and fruits collected by Dr. Capuron at Fort Dauphin, 372 BLUMEA VOL. XVIII, No. 2, 1970

Madagascar, 1967; Madagascar, d'Alleizette s.n. Aug. 1906 and Nov. 1906. Material from these 3 collections used in flowers. The first for serial The latter for Mada- was clearing sectioning. two vegetative anatomy. TEFw SF gascar, 14577 (wood sample ex TEFw).

RESULTS AND DISCUSSIONS

Anatomy of the flower in Cratoxylum (fig. 1a, 2)

The following account is based on serial sections of buds and flowers of C. arborescens and of C. and cleared material of all of formosum on species Cratoxylum. The arrangement

of floral in transverse section is illustrated in Additional data parts as seen fig. ia. are given below. The the hypogynous scales are inserted at same level as the phalanges and receive

the several vestigial vascular bundles each from the same region of the stelar system as view phalanges. This seems to favour the that they may be regarded as staminodial struc- The of celled. The tures (cf. Engler, 1925: 183). ovary Cratoxylum is incompletely 3 (or 4) of coherent septa are fused at the very base the gynoecium and are (in C. formosum) or fused (in arborescens) below the styles. Placentation there ovules C. is axile; are 3—18 per cell wich are attached the the basal of the (cf. Gogelein, 1968), to placentae at part septa in an ascendant anatropous way. Vascularization in the ovary of is shown loculi bundles in fig. 2. Below the three major vascular depart from a fragmented branches stelar cylinder. Each bundle over a very short distance (fig. 2a, b) into three bundles of which the median one forms the dorsal bundle and each of the other two bundles the attachment of of the vascular runs along one septa ('lateral' bundles). Minor bundles connected with these three carpellary bundles occur in varying numbers. The placentae are supplied with vascular bundles which are more or less linearly continuous with the stelar cylinder. Some vascular tissue extends to above the level of ovule insertion.

Transverse Fig. 1. sections throughflower buds. a. Cratoxylumformosum, X 12.b. Eliaea articulata, X 12. h.s.: hypogenous scale; o: ovule; p: petal; p.s.: petal scale; ph: phalange; s: sepal. Loculi shaded; vascular bundles indicated where present. P. Baas: Anatomy of Eliaea and Cratoxylum 373

orientation difficult of The of vascular bundles was very to determine because the very low number of differentiated elements before the those xylem and at flowering stage. In cases where orientation could be observed all peripheral vascular bundles (dorsal and lateral bundles found have normal plus derivatives) were to arrangement with xylem poles facing the loculi. In the placental vascular supply the situationis much more com-

and is is the plicated obscured because there no distinct boundary between individual arborescens the vascular bundles. In C. bundles were found to be centric (amphicribral)

the the above level of ovule attachment. In same species some minor vascular bundles with their xylem poles facing the axis of the ovary were observed running as a median line These other derived the through the septa. bundles, not foundin the species, are from The observable vascular system of the placentae. course of vascular bundles was hardly cleared material due the of differentiated elements. The in to scanty presence xylem fusion of lateral vascular bundles from different carpels opposite the septa could be observed only in C. maingayi, but not in the other species. The dorsal and lateral bundles enter the The dorsal bundles to the where branch styles. continue up stigmas they pro- fusely.

ofthe Secretory cavities and canals are abundantin the periphery ovary wall and occur in in in various numbers also the style. They are also abundantly present the sepals, occur the in petals (cf. Gogelein 1968), but are absent or very rare in the androecium (apart from special connective glands in C. formosum ssp. pruniflorum).

Anatomy of the flower in Eliaea (fig. 1b, 3)

The arrangement of floral parts in Eliaea is identical to that of Cratoxylum apart from

details the of sections for some in gynoecium (see fig. ib) Fig. 3 gives a series transverse with those of The also comparison Cratoxylum. ovary is incompletely 3 (or 4) celled, the distance the free is There three although over which septa are very short. are conspic-

false of tissue situated to the uous septa, composed a bulky parenchymatous abaxially dorsal vascular bundle. These of the of septa give a superficial impression presence 6(—8) loculiinstead of the There ovules As in 3 (or 4) actually present. are 6(—8) in one ovary. the the Cratoxylum they are attached to the placentae at basal part of septa in an ascendant

The vascularization the in that anatropous way. of ovary Eliaea is fundamentally similar to

There The not of Cratoxylum. are some minor differences, however. stelar cylinder is

arbo- broken up just below the level ofinsertion of carpellary bundles as in Cratoxylum bundles fused rescens and formosum. The lateral vascular of two carpels are over some die distal of The the distance (see fig. 3d—f) in part the ovary. vascular strands in placentae

be double above of The appear to or centric the level ovule attachment (see fig. 3e). lateral carpellary bundles do not continue into the styles. The dorsal bundles branch just below the stigmas. The distribution ofsecretory canals is similar to that in Cratoxylum.

Spherical glands (i for each stamen) occur in the distal end of the connectives of the stamens. These are similar in structure to the two connective glands foundin each stamen of C. formosum ssp. pruniflorum (cf. Gogelein, 1968, fig. 5k). The vascularization of the scales and the of scales in the hypogynous presence petal other characters corolla of Eliaea are among the shared with Cratoxylum.

Vegetative Anatomy of Cratoxylum

Leaf (fig. 4d—k)

In surface view: Adaxial epidermis glabrous in all material apart from C. formosum

Cells in outline with curved to anticlinal walls. ssp. pruniflorum. polygonal straight BLUMEA VOL. No. 374 XVIII, 2, 1970

Some sections Fig. 2. transverse through the gynoecium of a flower of Cratoxylum formosum ssp. for-

all Vascular of central of b—i. Sections mosum, from bottom (a) to top; x 29. a. supply part receptacle.

through ovary and styles. d.b.: dorsal bundle; l.b.: lateral bundle; o: ovule; st: style; st.c.: style canal; v.b.: Loculi shaded. vascular bundle; x: differentiated xylem element(s) (solid black). P. Baas: Anatomy of EHaea and Cratoxylum 375

of of Eliaea articulata from bottom (a) Fig. 3. Some transverse sections through the gynoecium a flower

all For 2. false septum. to top; X 29. legends see Fig. ƒ.s.: BLUMEA No. 376 VOL. XVIII, 2, 1970

Leaf of Eliaea and Eliaea articulata distal Fig. 4. anatomy Cratoxylum. a—c. a. petiole through end,

X 12. b. lamina throughmidrib, X 12. c. abaxial epidermis, X 200. d. C. sumatranum, abaxial epidermis, abaxial f. X 200.e. C. maingayi, epidermis, X 200. C. arborescens, abaxial epidermis, x 200. g. C. formosum

abaxial x 200.h. Cult. Hort. abaxial ssp.pruniflorum , epidermis, C. formosum ssp. formosum, Bog., epidermis,

X 200.i. C. formosum ssp. formosum, Buwalda 7979, abaxial epidermis, X 200.j. C. formosum ssp. formosum,

Cult. Hort. transverse k. Buwalda h: secre- Bog., lamina, section, X 200. ibid., 7979. hair; p: papillae; s.c. :

cavity or canal; v.b.: in tory vascular bundle. Xylem shaded; phloem dotted; sclerenchyma a and b solid black. P. Baas: Anatomy of Eliaea and Cratoxylum 377

Abaxial epidermis with indumentum of mainly uniseriate hairs with broad basal cells formosum present in C. ssp. pruniflorum (fig. 4g). In this sub-species hairs also infre- and low quently present on adaxial leafsurface bluntly conical papillae present on abaxial

in of surface. Tall to short ± cylindrical papillae present on abaxial surface part the

material of C. cochinchinense, C. formosum ssp. formosum, C. glaucum, C. maingayi, and sumatranum Table and discussion confined C. (see fig. 4h, j, III on p. 383). Papillae

to unspecialized epidermal cells and absent from guard cells and subsidiary cells of stomata cells and from elongated overlying midrib and (if conspicuous on epidermis) major veins. Abaxial epidermis glabrous in remainder of material. Unspecialized cells polygonal in outline with almost straight to sinuous anticlinal walls (see fig. 4d—i). Cell outline often obscured by presence of papillae. Stomata confined to abaxial surface apart from some found adaxial surface midrib on overlying the in I specimen of C. formosum ssp. for-

mosum; paracytic with low dome-shaped guard cells and subsidiary cells of various

shapes. Outer anticlinal walls of subsidiary cells curved to sinous (see figs. 4d —i). Di-

of to the random or ameter stomata perpendicular pore 10—20 fj, (mean 14//); orientation

tending to coincide with direction of major veins. thick. cells In transverse section: Cuticle 1—3 p, Epidermal square to flattened, those of abaxial surface smaller than those of adaxial surface. Hypodermis (derived from

protoderm and thereforerepresenting the innerlayer(s) of the multiple epidermis) of I—3

irregularly arranged cell layers, adaxially present in C. arborescens and C. glaucum, absent from other material. Mesophyll composed of I or 2 layers of tall adaxial palisade

cells; abaxial chlorenchyma cells of C. cochinchinense also tending to be palisade-like;

in remainder of material abaxial tissue compact to fairly loose, composed of cubical to

lobed or rounded chlorenchyma cells. Leafmargins pointed downwardsto various extents.

Mesophyll of margins composed of small amount of collenchyma. Midrib with shallow V—U shaped adaxial groove and prominently raised abaxial surface, supplied with single crescentiform vascular strand with of thick broadly V-shaped to sclerenchymatous caps walled fibres interspersed with thin-walledcells. Ground tissue ofmidrib collenchymatous

in Veins and at periphery to parenchymatous more central parts. with or without ad- Sheath of cells abaxial schlerenchyma caps. translucent parenchyma present around

vascular bundles and also including caps, if present. Vertical bundle sheath extensions associated with all from major veins present in species apart C. maingayi. Petiole ± semi-

circular to bean-shaped in T. S. through distal end, with concave to flat adaxial surface and abaxial surface. Vascular of convex system composed single ± crescentiform strand, withoutincurved margin in material from the sections Cratoxylum and Tridesmos. additional latero-dorsal bundles the arborescens 2(—4) present in section Isopterygium (C. and C. glaucum). Ground tissue of petiole composed of collenchyma and thick walled parenchyma. Crystals present in varying amounts; druses predominant in abaxial part of

mesophyll (often in cells which are larger than surrounding cells); clusters predominan.

in ground tissues of petiole and midrib and in mesophyll. Prismatic crystals less commont

Secretory cavities of (sub) spherical shape, varying greatly in frequency confined to palisade of in the or spongy part mesophyll or present both. In a few cases anticlinal and/or peri- clinal cell walls of the epidermis thinner in places opposite secretory cavities than in remainder of canals of small diameter of epidermis. Secretory present in phloem

midrib and petiole. Broader canals present in ground tissue of petiole and midrib. of canals Epithelium secretory cavities and badly preserved in most herbariumspecimens.

Axis of (twigs c. 3—5 mm diameter; see also table I)

cortex and caducous in all which Epidermis, perivascular fibres twigs are 3 mm or more BLUMEA No. 378 VOL. XVIII, 2, 1970

wood — Fig. 5. Transverse sections of from Cratoxylum and Eliaea; all X 30. a. C. sumatranum.

C. SAN C. f. Eliaea articulala. r: b. C. cochinchinense. c. C. formosum. d. arborescens, 26136. e. glaucum. ray;

shaded. f: tracheid; v: vessel. Parenchyma P. Baas: Anatomy of Eliaea and Cratoxylum 379

thick. These a of arborescens. Cork tissues were only seen in very young twig C. origina- ting in pericycle between perivascular fibre sheath and phloem; composed of layers of

approximately square to flattened cells with adaxially lignified ± U-shaped and pitted

wall thickenings; these layers are occasionally interspersed with layers of thin-walledcells.

Cork cells ± hexagonal in tangential view. Secondary phloem often badly preserved in

herbarium specimens; composed of sieve-tubes, companion cells, parenchyma, secretory canals cell-elon- and rays with very slightly dilatating peripheral parts (due to tangential the formation of radial gation and/or an extra wall). Secondary xylem: Growth rings

poorly defined or absent (varying intraspecifically). Vessels diffuse, solitary and in short

oblique to radial multiples of 2 or 3 (—6); round to oval in T.S., mean tangential diameter

vessel member Perforations in the to 35 —31 [x, mean length 255—363 p.. simple oblique Lateral wall transitional alternate slightly oblique end walls. pits in to arrangement,

small and round-oval — and (diameter 2 3p.) or larger slightly polygonal (diameter 3—4[i).

Vessel—ray pits polygonal or flattened and unilaterally compound, resulting in ± reticulate wall of the Solid pattern. Tyloses present in some specimens investigated. brown vessels amorphous contents present in some of most specimens. Tracheids

paratracheally arranged, very sparse to rather frequent ('vascular tracheids'). Fibres

of arranged in radial rows, constituting ground tissue secondary xylem. Walls varying in and bordered thickness very sparsely provided with simple to minutely pits. fibre Average length 420—740 p.. Parenchyma present in varying amounts, predomin- antly in irregular, mainly apotracheal, bands in C. cochinchinense, C. formosum. c.

maingavi and C. sumatranum ; mainly paratracheal or confluent in C. arborescens and c.

— of glaucum. Rays 15 25/mm at periphery of secondary xylem twigs investigated, uni-

seriate or — cells of biseriate, (1 —) 3 many high, heterogeneous, composed square to cells. with radial upright Primary xylem forming a continuous cylinder conspicuous

arrangement of vessels. Node unilacunar with a single leaf trace. Pith forming a flattened of cells which and cylinder, composed parenchyma are rounded in T.S. square to rectan-

in number of gular L.S., interspersed with a varying secretory canals. Crystals present as clusters and abundant prismatic types in pith. Prismatic types in phloem. Amount and of proportion of different crystal types varying infraspecifically Silica bodies, globose or in of irregular shapes, noted ray parenchyma cells secondary xylem ofpart of the material canals with investigated. Secretory I or 2 layered epithelium; present in phloem and pith,

the canals in the phloem being narrower than those in the pith.

Bark

Material: sample from bole of C. arborescens, Fuchs 21290

General features. Colour greyish brown or dark brown with yellow and reddish brown shallow tones. 4 —5 mm thick, with coarse longitudinal fissures, which are more than 2 cm apart. Shedding in thin layers. In T.S. with numerous yellow tangential bands.

For field characters see Foxworthy (1927).

Microscopic features: Ground tissue composed of sieve tubes, companion

strands. cells and (crystalliferous) parenchyma Rays 1—4 seriate. Secretory canals numerous

in tissue. bands of cubical tliick-walled ground Tangential (2 —3 mm thick) composed ± and densely pitted stone cells ('phelloids'); occurring at various intervals (their distance from few microns due the rather of varying a to 0.7 mm to wavy course the bands). The

discontinuous bands accompanied on both sides by layers of flattened suberized cells.

Bands formed by cork cambia on their inner sides. BLUMEA 380 VOL. XVIII, No. 2, 1970

Wood (See also Table II, fig. 5, 6) of A. Section Cratoxylum and section Tridesmos (fig. 5a—c) Material investigated: samples wood from bolesof mature C. sumatranum, C. cochinchinense, C. formosum and C.I maingayi. Wood General features: heavy to very heavy (specific gravity exceeding 1 in C.

sumatranum, C. formosum and C. maingayi). Colour dark brown in C. maingayi, light yellowish to pinkish brown in other three species. Texture fine.

Microscopic features: Growth rings absent to fairly conspicuous, Vessels

2 diffuse, solitary and in radial to oblique multiples of 2 —4(—6), forming an 7—18/mm ,

oblique pattern in C. cochinchinenseand C. maingayi, round to oval (radially elongated) in mean diameter radial diameter walls T.S., tangetial 90—iiOfx, up to 140—1909.,

— — vessel member Perforations 3 4( 6)(j. thick, length 290—710(2 (mean 460—6oo;x).

simple in the oblique to horizontal end walls. Lateral wall pits alternate. Inter-vessel pits oval diameter predominantly round, sometimes to polygonal, 3—5(—6) fx. Apertures slit-like, horizontal to oblique. Vessel-ray pits similar but half-bordered; sometimes Vessel—fibre and minute. Thin-walled unilaterally compound. pits very sparse tyloses

and solid amorphous and granular brown contents noted in some heartwood vessels of C.

cochinchinense and C. maingayi (also recorded for C. sumatranum and C. formosum by Vascular tracheids Moll and Janssonius, 1906). accompanying someofthe vessels, sometimes

arranged in radial 'tails' together with vessels ofnarrow diameter. Ground tissue composed

of with bordered with and fibres very few simple pits or pits strongly reduced borders slit-

like vertical Fibres in disturbed radial in ± apertures. arranged variously rows, hexagonal diameter walls T.S., 10—32[i, 2—8/u thick; length 780—1690[i (mean 1080—1320/t). of cells Parenchyma predominantly apotracheal in tangential bands I—4(—6) wide. betweenbands Paratracheal Distance 100—400//. parenchyma scanty. Diffuse parenchyma

6—8 cells absent or very scanty. Parenchyma strands of cells. Rays io—13/mm, I—many

not exceeding 1.1 other high, height mm, I—2 seriatein C. maingayi, I—3(or 4) seriate in

species, tending to be of two distinct sizes in C. cochinchinense, heterogeneous or tending of to homogeneous, mainly composed procumbent cells, but squarish cells occurring in low uniseriates and in tails of multiseriates. Crytals absent. Silica bodies with rough surface

and of in cells. irregular shapes present ray

B. Section Isopterygium (fig. jd, e, 6)

Material investigated: samples of mature wood of C. arborescens and C. glaucum.

General features: Wood rather light. Colour light pink to brick red. Texture

fairly fine, but conspicuously coarser than in the other two sections.

Microscopic features: Growth rings not seen in material examined. Vessels

2 diffuse, 2 solitary and in radial multiples of 2—4( —5), oval in T.S. (radially —5/mm , diameter radial diameter elongated), mean tangential, 125—200fx, up to 210—280/1,

walls vessel member —660 Perfora- 3 —4.(—6) [i thick, length 300—780 fx (mean 520 /x).

tions simple in the oblique to horizontal end walls. Lateral wall pits alternate. Inter-vessel

round oval with slit-like In the pits to (diameter 3—6/u) oblique apertures. C. glaucum

slits sometimes confluent with others from neighbouring pits. Pits to contiguous paren-

chyma cells (rays and axial parenchyma) half-bordered, similar to inter-vessel pits or and occasionnally unilaterally compound then two pits on the parenchyma cell wall. and brown Thin-walled tyloses solid amorphous contents noted in some heartwood vessels.

Vascular tracheids accompanying some of the vessels, sometimes arranged in radial 'tails'

together with vessels of narrow diameter. Ground tissue composed of fibres with very

Fibres in few simple pits. more or less arranged in radial rows, rounded to rectangular P. Baas: Anatomy of Eiiaea and Cratoxylum 381

diameter — walls T.S., 20—38(2 in C. arborescens, 16 21 (J, in C. glaucum; 2—4( —6) (2

thick. —1210 Parenchyma Length 780—1560 [2 (mean 1070 (2). predominantly paratracheal,

in layers around the vessels to in C. arborescens tending to be aliform con- i—3 scanty , in few continuous in Fucbs fluent some places. Very bands, only seen 21346. In C. aliform. absent. glaucum parenchyma mainly Diffuse parenchyma very scanty or Paren-

of —8 — chyma strands 5 cells. Rays 10 Ii/mm, I to many cells high, height not exceeding

i.i mm, i—4 seriate and tending to be of two distinct sizes in C. arborescens, I or 2

(or 3) seriate in C. glaucum, heterogeneous or tending to homogeneous, mainly composed

cells in uniseriate and in the tails of procumbent cells, squarish present most rays of bodies and of multiseriate rays. Crystals absent. Silica with rough surface irregular shapes present in ray cells.

6. sections wood x C. b. C. Eliaea articulata. Fig. Tangential through rays; 77. a. glaucum. formosum. c.

Vegetative Anatomy of Eliaea

Leaf (fig. 4a—c)

In surface view: Adaxial and abaxial epidermis glabrous. Epidermal cells of adaxial surface and unspecialized cells of abaxial surface polygonal widi straight anti-

clinal walls. Cells overlying midrib and major veins slightly elongated. Stomata confined

to abaxial surface, paracytic with high dome-shaped subsidiary cells. Subsidiary cells often divided to the Diameter to the 10—18 parallel pore. perpendicular pore p. (mean 15p); orientation random.

transverse Cuticle thick. Adaxial cells than In section: 2—5 p epidermal larger abaxial with ones. Multiple epidermis irregular collenchyma-like wall thickenings present in D'Alleizette, August 1906, absent in specimen D'Alleizette, Nov. 1906. Mesophyll of of cells tissue composed 1 or 2 layers palisade and spongy of inconspicuously lobed cells. Leaf margins pointed downwards. Mesophyll parenchymatous at margins. Midrib

with sligh ly raised (almost flat) adaxial surface and convex abaxial surface, supplied with a single shallowly V-shaped vascular strand with or without incurved margin.

Vascular strand with abaxial and adaxial sclerenchymatous caps, composed of thick- walled fibres interspersed with thin-walled cells. Ground tissue of midrib collenchyma- tous at abaxial central and adaxial periphery to parenchymatous in more parts. Veins 382 BLUMEA VOL. XVIII, No. 2, 1970

with which are provided sclerenchyma caps, most prominent opposite xylem, minor of cells veins embedded in mesophyll. Sheath translucent parenchyma present around

vascular bundles and Bundle sheath extensions absent. Petiole sclerenchyma caps.

semi-circular in T.S. through distal end, with concave to flat adaxial surface and convex of crescentiform with abaxial surface. Vascular system composed single strand, or Ground without additional incurved margins. tissue collenchymatous. Crystals as in

of in Cratoxylum. Secretory cavities (sub)spherical shape present both palisade and spongy

tissue. Adaxial epidermal cells opposite secretory cavities with conspicuously thinner

anticlinal and periclinal walls than remainder ofepidermal cells. Secretory canals abundant of midrib and with a two-layered epithelium of large cells in ground tissue and petiole.

Secretory canals in phloem of midrib and petiole narrow and very inconspicuous.

also Axis (twigs of c. 3 mm diameter; see table I) Growth absent. Vessels Peripheral tissues as in Cratoxylum. Secondary xylem: rings diffuse, almost round diameter vessel member exclusively solitary; in T.S., mean 35—36 fx, mean Perforations in the slightly oblique end walls. Lateral wall length 390—450 (X. simple alternate Vessel-fibre diameter pits in arrangement. pits round-oval, c. 3 p.. Vessel-paren- chyma and vessel-ray pits sometimes flattened and unilaterally compound. Tyloses and radial vessel contents not seen. Vascular tracheids rare. Fibre-tracheids arranged in rows,

Walls thick with constituting ground tissue of secondary xylem. and provided numerous bordered fibre pits. Average length 670—780 u. Parenchyma mainly irregularly banded; aliform confluent. Paratracheal bands apotracheal or parenchyma scanty. Raysc. 22/mm at of node periphery secondary xylem, further as in Cratoxylum. Primary xylem, and pith

in and clusters in as in Cratoxylum. Crystals present as elongated primatic types phloem as

pith. Silica bodies absent. Secretory canals as in Cratoxylum.

Wood (see also Table II, fig. 5f, 6e) of of diameter. Material investigated: outer part stem 5 cm features: General Wood very heavy (specific gravity exceeding 1). Colour

dark brown. Texture fine.

2 features: Growth absent. Vessels diffuse, c. almost Microscopic rings 12/mm , roundin diameter walls vessel member exclusively solitary, T.S., mean 95 ft; 4—7 fx thick;

—1200 Perforations simple in the oblique to horizontal end length 450 (j. (mean 780 fx). walls. Lateral wall pits to contiguous fibre-tracheids in alternate arrangement, diameter

and round or slit-like. and vessel—parenchyma 3 —4 fx, apertures minute Vessel-ray pits similar but half-bordered or unilaterally compound. Vessels in heartwood with

solid amorphous brown-yellow contents. Tyloses absent. Vascular tracheids not seen.

in radial with Ground tissue composed of fibre-tracheids, arranged rows, numerous with almost vertical diameter of borders Fibres bordered pits slit-like, apertures; c. 3 p..

to diameter 20 walls —11 round hexagonal in T.S., —30p, 4 p thick; length 900 —1700 p in bands of (mean 1290 p). Parenchyma predominantly apotracheal irregular tangential cells i—3 wide at various distances. Diffuse and paratracheal parenchyma scanty.

cells Parenchyma strands of 5 —8 cells. Rays c. I i/mm, i to many high (height not

exceeding 0.8 mm), uni—biseriate, markedly heterogeneous, composed of procumbent cells. of axial cells. to upright Crystals prismatic types infrequently present in parenchyma Silica bodies absent. P. Baas: Anatomy of Eliaea and Cratoxylum 383

Occurrence of papillae in Cratoxylum

In two out of four leafsamples of C. formosum ssp. formosum sectioned very conspicuous,

tall epidermal papillae were notedon the abaxial leafsurface (fig. 4h, j). The leaves of the

other two of (fig. k). The specimens were completely devoid papillae 4i, very striking

differences in the abaxial epidermides of one subspecies led me to study all material of

the in Cratoxylum present in Rijksherbarium with a stereomicroscope (magn. X 60) whether order to establish the presence or absence of papillae. My aim was to know this character distribution of the showed any relationship to geographical specimens

concerned and whether it could be used to trace taxa of infraspecific rank.

hsted the mark Results are listed in table III. Specimens under question represent cases where be but where the surface smooth no conspicuous papillae could seen, was not leaves found either; when examined in transverse section such were to exhibit an inter- condition with becomes evident that the mediate very low papillae. From table III it

is character for of presence of epidermal papillae not a reliable diagnostic species Cratoxy- but that of exists for the of lum, in some species a kind geographical pattern occurrence is obvious in C. here find papillae. Such a pattern very formosum ssp. formosum; we a high proportion of specimens with papillae in material from Java and Sumatra; in the material

from Borneo and the Philippines this proportion is remarkably lower, whilst all

specimens from Celebes and S.E. Asia lack papillae. It is tentatively suggested here that

in this taxon the occurrence of papillae reflects a further differentiation into taxa of be lower rank. If infraspecific variation patterns were to investigated more extensively this character definitely deserves further attention. The comparatively high proportion of the mark could be indicationthat specimens of C. cochinchinense listed under question an

differentiation of subspecific taxa has not progressed as far as in C. formosum.

The different the of different under patterns for occurrence papillae in species growing

similar conditions (cf. Gogelein, 1968) suggests that the presence or absence of papillae is

not determined by the environment.

Floral and xylem specialization and related problems

criteria for trends of flower In applying the general phylogenetic specialization (see e.g. has the of Eliaea be Takhtajan 1959) one to regard gynoecium to more specialized than

that of of of Cratoxylum because of a reduced number ovules in the former. The presence

in Eliaea also rather than false septa seems to represent a specialization, some vestigial character. with other would for A comparison Hypericoideae be necessary a good under- floral but data from literature standing of trends in specialization, comprehensive are

not available, as far as known to me. be found and Detailed data on xylem specialization are to in Bailey Tupper (1918), and Chalk and Frost (1930), Tippo (1938), Metcalfe (1950), Carlquist (1961). For our suffices the trends: purpose it to summarize following

1) long fusiform elements (particularly vessel members) are more primitive than short

ones; tracheids than libriform 2) fibre are more primitive fibres; with diffuse than 3) xylem solitary vessels is more primitive xylem with vessel multiples.

Bailey (1957) and Carlquist (1961) have pointed out the dangers of using these trends,

which have been established for the higher categories of land to judge the degree of within small of allied In these specialization groups closely taxa. my opinion trends are, valuable for both and related however, equally closely distantly groups, perhaps even BLUMEA No. 384 VOL. XVIII, 2, 1970

more readily interpretable for small groups. In the higher categories such as families and

much orders, parallelism and different speeds of speciahzation may play a more confusing

and members of different often not their part groups are comparable in secondary xylem These less anatomy. factors are apparently important within a group of which we know

on other grounds that its representatives are closely related. Of course we must not member had been ancestral the expect linear sequences, as if the more primitive to other The result is species. only we may hope for to find some representatives which are more others. like a possible common ancestor than Returning to Cratoxylum and Eliaea, we Eliaea the characters find that shows more primitive in the longer fusiform elements

(tables I & II), in fibre tracheids with conspicuously bordered pits and in exclusively

solitary vessels. Fibre length in Eliaea, though comparatively great, is equalled by c.

cochinchinense (in twigs and mature wood) C. formosum (in twigs) and C. maingayi (in that fibre index twigs and mature wood). It is known, however, length is not an accurate

for xylem specialization since considerable elongation of these cambial products takes

place.

it should also be stressed that wood from stem of diameter Here only a sample a 5 cm

was at our disposal in the case of Eliaea. Bole-samples as in Cratoxylum are not available branch wood of from since Eliaea is a shrub. It might well be that in a sinnlar diameter the values lower material Cratoxylum quantitative would be than in the at my disposal;

this would make the differences between Cratoxylum and Eliaea even more pronounced. tables Quantitative xylem characters as listed in I and II are not suitable for diagnostic because purposes, of a high intraspecific variation (e.g. table I: C. formosum; table II: c.

sufficient of values arborescens). For our problem it is to know that in a wide range Eliaea the maximal the that occupies extremes. From above we can tentatively conclude the xylem of Eliaea resembles that of a possible common ancestor more closely than that of

Cratoxylum. It is difficult to reconcile the contradictory conclusions from floral and wood

It should be that the of mixture of anatomy. mentioned, however, occurrence a obviously

advanced and primitive characters in plant taxa is by no means a rare phenomenon (see The that floral and and e.g. Cronquist 1968). suggestion xylem anatomy in Cra oxylum

Eliaea show differently diverging lines implies that Cratoxylum and Eliaea are not so

from closely related as one might expect gross morphology. Within the Hypericoideae

related than the main they are, however, more closely mutually to any other genus. For differences either from the other in gross morphology or vegetative anatomy hypericoid the genera: Ascyrcum, Haronga, Hypericum, Psorospermum or Vismia reader is referred to

and Metcalfe For either Engler (1925) and Chalk (1950). each of these genera there are flower and fruit characters which them rather far from the or xylem put apart Cratoxyleae (Cratoxylum and Eliaea). shrub found the of dunes Eliaea is a in narrow strip along the east coast of Madagascar. la Bathie 'Ce il Perrier de (194.8: 114) stated: genre monotypique est endcmique, mais est affine des Cratoxylon d'Asie ct d'Oceanie. C' est done un type d'origine orientale,' etc.

He tentatively ascribed its occurrence along the east coast to dispersal by see of the winged The Eliaea the seeds. derivation of from Cratoxylum is, however, most unlikely since

of Eliaea more than thatof If the of Eliaea on xylem is primitive Cratoxylum. presence

Madagascar must be explained by oriental origin it, or its ancestor, must have arrived

the at a time that Cratoxyleae in Indo-Malesia still showed primitive xylem characters now found in Eliaea. An alternative hypothesis, already used by Danser (1928) for

explaining the occurrence of primitive Nepenthes species on Madagascar, can also be

used to understand the present distribution of Eliaea. This hypothesis implies that the

distribution back time that connections pattern Madagascar—Indo-Malesia goes to the land P. Baas: Anatomy of Eliaea and Cratoxytum 385

between the two areas were still present and that a common ancestor of Cratoxylum and Eliaea the whole also Schuchert Willis occupied range (see 1932; 1932; Van Steenis 1962 later of Gondwana made and Florin 1963). A fragmenting this continent an independant evolution ofboth and Eliaea this retained Cratoxylum possible. During process Cratoxylum primitive flower characters and Eliaea retained primitive xylem characters. Dapania resembles but grandifolia (Oxal), an endemic form from Madagascar, two Dapania species , also Sarcotheca, all from Malesia (Veldkamp 1968) and might call for a similar explanation. model It is emphatically stressed here that the above mentioned hypothesis is merely a in which the data known fit well. factual evidence available at present seem to fairly No is to much research with similar support it. Obviously, more into plant groups a distribution before ofthe offered. pattern is needed it is possible to ascertain the probability explanation

Great care must also be observed in attaching too much weight to thephylogenetic trends have been of xylem specialization. The general trends of xylem specialization very ele- their has gantly established, but an absolute certainty of irreversability only been claimed

(Baily 1944) and was never proved.

Sectional delimitationand the anatomy in genus Cratoxylum

the wood Den Berger (1926) suggested in a general paper that, on grounds of anatomy, from those Cratoxylum should be split into two sections, different recognized by Englcr he did list the decisive (1925). Unfortunately 70t any specific names or mention anatom- ical characters involved. Desch (1941) emphasized that C. arborescens differs wood- anatomically from the other species investigated by him (C. formosum and C. ligustrinum of distribution. = C. cochinchinense) in having another type parenchyma My own obser- vations, together with wood descriptions from literature (Solereder 1899, Moll and confirm Janssonius 1906, Metcalfe and Chalk 1950 and the papers cited therein), the fact that wood two can be in anatomically only grouwps distinguished Cratoxylum. One group is represented by section Tridesmos (C. formosum and C. maingayi) and by section Cratoxylum and The these (C. sumatranum C. cochinchinense). species of sections possess comparatively narrow vessels, abundantpredominantly banded parenchyma (see fig. 5a—c), and generally fairly thick-walled fibres (high specific gravity). The other group is represented by C. arborescens and C. glaucum constituting the section Isopterygium. Wood of these species is characterized by wide vessels, less abundant paratracheal parenchyma (fig. sd, e), and thin-walled fibres (low specific gravity).

be of The fact that wood-anatomically only two groups can distinguished instead the three should not be as a between wood and sections interpreted disagrement anatomy That the sections Tridesmos and be gross morphology. Cratoxylum cannot separated on the basis ofwood does that have sound basis. anatomy not mean they no morphological

In fact, the great differences between the wood anatomy of section Isopterygium and that of the than lack ofsuch differences between oftheremainder genus are more surprising the the two other sections. It is most unlikely that, in the absence of flowering material, a wood sample of C. arborescens or C. glaucum would be related to other members of the genus. This indicates a rather isolated position of the section Isopterygium within the genus.

The distinction in wood anatomical structure coincides with another important of the of feature: the representatives section Isopterygium are evergreen trees; those the

It would be attractive the other two sections are deciduous. very to interpret anatomical the differences between the two groups in terms of a functional adaptation to behaviour of the increased facilities the abundant cells of foliage. Initially, storage in parenchyma for the deciduous group, useful energy requirements during leafless periods, were consid-

[i No. BLUMEA VOL. XVIII, 2, 1970 386

find collected about comparable cases in order to ered. Some data from literature were whether there is is the of a rule or no out whether the situation in Cratoxylum example distribution and the or deciduous general relationship between parenchyma evergreen of the Notho- described the wood anatomy genus habit at all. Dadswell and Ingle (1954) be abundant in the section from fagus and found wood parenchyma to very evergreen and small in the deciduous sections in a evergreen New Guinea, whilst it was very sparse in Zealand. This reversal, therefore, of what we found Cratoxylum. section from New is a wood-anatomical evidence and Muller tried to find Williams (1939) and Tilson (1942) substantial difference They found no for subgeneric segregation of American Quercus. of and of deciduous oaks, from a not surprisingly between the wood evergreen apart above mentioned in the former of species. The more pronounced ring-porousness group ground for accepting a functional explanation examples, though very few, remove any

on such topics require for the differences in wood anatomy in Cratoxylum. Suggestions involved and such of the and cambial activity of the trees more knowledge autecology studies. knowledge is only obtainable in field the of it should be noted that great Returning to the xylem anatomy Cratoxylum, those exist in the xylem are not parallelled by m differences found to mature secondary and C. of In twigs of C. arborescens the juvenile secondary xylem young twigs. young in the other banded almost the same extent as species may be to glaucumi parenchyma show deviations from the remainder ofthe and vessel diameter and frequency do not and of the mature xylem of C. arborescens C. From this might conclude that genus. one that the first formed secondary xylem a derived condition, considering glaucum, represents characters Bailey 1944)- Such a conclusion seems to often retains the more primitive (see members of the listed in table II where the vessel disagree with the quantitative features the in the of C. arborescens and C. glaucum belong to longest mature secondary xylem condition for the former two species. This would imply a more primitive genus. again note that both C. glaucum and C. the for we However, if we consider data young twigs has beenknown the limitsof individual variability ofC. formosum. It arborescens are within the fusiform element length increase from from Sanio's time onwards (1872) that may later formed mature' secondary xylem. This first formed secondary xylem to the much and 1918: from to (see Bailey Tupper, increase in length, which varies species species e.g. related of vessel member length as to phylogenetic 186) could interfere with our theories the in case of Cratoxylum. Considering specialization, and complicate the picture as the be in C. arborescens and C. glaucum one would wide vessels and paratracheal parenchyma be derived rather than as compared inclined to regard the section Isopterygium to primitive for trends of xylem with the other sections of Cratoxylum (cf. Carlquist, 1961, speciali-

zation). the conclusions about The considerations offered above do not affect on page 384 than Cratoxylum since the characters employed Eliaea having a more primitive xylem

for both and mature secondary xylem. there are consistent young for the rather isolated position Another anatomical character provided an argument the distal end of the of within the as whole: in petiole of the Isopterygium species genus a absent in the latero-dorsal bundles occur which are C. arborescens and C. glaucum 2( —4) of of of a epidermis in leaves other species of Cratoxylum. The presence multiple species the character to distinguish them from the section Isopterygium would offer yet another of varies, however, quite The or absence a multiple epidermis other species. presence 381), andthis character could well often infraspecifically (e.g. in Eliaea articulata, seepage feature if material is examined. break down in Cratoxylum as a diagnostic more P. Baas: Anatomy of Eliaea and Cratoxylum 387

Cratoxylum and Eliaea as distinct genera

follows that there characters both From the descriptive part of this paper it are many in

floral and vegetative structure shared by Cratoxylum and Eliaea. For instance petal scales,

hypogynous glands, and stamens in phalanges; the general histology of leaf and axis is similar. number of characters also essentially However, quite a distinguishing are present All characters could be confirmed and three as well. as listed by Gogelein (see p. 370) more

diagnostic characters could be added from vegetative anatomy: fibre-tracheids with bordered almost and the absence ofsilica the numerous pits, exclusively solitary vessels, in fibres and the wood of Eliaea, as contrasted with scarcely pitted common occurrence of of vessel multiples and silica bodies in rays in the representatives Cratoxylum. substantial than between The differences between Cratoxylum and Eliaea are more the of The differ section Isopterygium and the other two sections Cratoxylum. two genera in and differential characters within both floral wood characters, whilst the Cratoxylum are distribution the The latter character mainly based on parenchyma in secondary xylem.

be as less than vessel and may, moreover, regarded taxonomically important grouping

of fibres. both floral and I see no grounds for type Considering vegetative anatomy com-

bining Eliaea and Cratoxylum to form one genus. that C. is I agree with Gogelein within the genus Cratoxylum, maingayi probably most of closely related to Eliaea. Anatomically this view is supported by the fusion lateral the extensions in bundles in the ovary and absence of vertical bundle sheath the veins of

the leaves in both Eliaea and C. maingayi. These two features were not encountered in the of fibres with that other species Cratoxylum. Moreover, C. maingayi possesses xylem pits

are most numerous and of which the borders are least reduced as compared with those These fibres different from the of the other Cratoxylum species. are yet very densely pitted fibre-tracheids in Eliaea, however.

ACKNOWLEDGEMENTS

after the This study became possible revision of Cratoxylum and was suggested by

ofDr. R. ofTananarive Prof. Dr. C. G. G. J. van Steenis. Through the courtesy Capuron

he obtained excellent pickled material ofEliaea; later Dr. Capuron also provided a wood Th. and of sample. Ir. J. Wassink (Amsterdam) Dr. Meng (Kepong) provided some the wood samples of Cratoxylum.

I have to thank Dr. W. van Heel and Mr. A. J. F. Gogelein for their keen interest and received the critical suggestions. During the initial stages of this study I hospitality in wish thank Jodrell Laboratory, Royal Botanic Gardens, Kew. I to Dr. C. R. Metcalfe and the members of the their in Miss anatomy staff for help many ways. M. Gregory with references anatomical and (Kew) very kindly provided me on literature, polished

the english text.

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and

length 320 510510 pp. —320 diameter, 340 340 310 370 in mean 310—430310—430 280— 410— Tracheid 280 Eliaea vessel for and % only. 820820 850 900 620 720720 960960 35 —620 range 650—850650— 470— 350—850 490—900490— 300300— 400— 420—

length p[A Cratoxylum length, measurements inin Fibre 510510 740 of 10 mean 600 600—730 670 420— 560 670—780 member on xylem based vessel 8484 55 5858 5252 for are rangeränge 16— 31—68 21—84 21—63 15—55 16— 26— 3 % secondary diameter pEr- 25 length inin

36 36

Vessel 42 42 40 formed to tracheid mean 45 37—5137—51 35—36 35—36 first amount of of Means length 440470—215—440 68 16— 1— 485220—485400— 16— They range 215— 175—540175—540 175—445 185—325265—270420—35— 220— 185—32515—225—650420—26—410—35— 255—435 table. p.F- characters length. member in this 270 in fibre 10—20 mean 310 315 345 340 Vessel 255—365 265— 390—450 for

Quantitative recorded %

I: not 15—10— specimens) are specimens) specimens) TABLE (2 (2 deviations (5 Cratoxylum: Tridesmos: Isopterygium:Isopterygium: articulata sumatranum cochinchinense formosum maingayi arborescens glaucum Standard Cratoxylum Section Section Section C.C. C. C. C. C. C. Eliaea VOL. 390 BLUMEA XVIII, No. 2, 1970

wall IX 8 8 3 4 3 4 IX (A}i 5 6 3—6 3-8 3-8 2—3 2—4 2—3 2—4 diameter, inin range 2—5 4— Fibre thicknessthickness vessel 20 2222 38 30 28 3030 [Afj. 32 21

for Fibre in range diameter in 16— 10—15 16— 14— 20—38 20—30 20— 16— 20— Eliaea % 35 16901690 13001300 1610 1430 1430 15601560 1430 17001700 and 15601560 range 910— 1040— 780— rioo— 910—1430 780—1430 1040— 780—1430 900— length, length (iJA inm Cratoxylum Fibre member mean 1190 1260 1080 1320 11201120 1070 1210 1120 1290 of xylem vessel 125 145145 160160 230230 270270 315 160160 140140 for length. range 55—140 30—125 20— 30— 60— 60— 50—315 50— 50— % vessel

24 fibre [A. secondary inji. diameter in for to Tangential % mean 90 90 90 no 155155 180 200 125 95 mature 17 amount of 11— 550 710 1200 They and 600 365—5603—30— 1040— 780— 3—290—55020—20 16— 60——79020—2—400—790910— 780—60—20—2—300—715 20—50—2—390—84528 780——78050—2— 21 1040— —120016—50—900—4—20— length 910—290—6002— 400—71030—3— 1100—14—1610 range 290— 365—560 290— 400— 300—715 390—845 325—780 450— characters 400 325 450 table. [A

inji,m this member

in Vessel mean 470 460 470 600 620 520 660 620 780780 Quantitative recorded

II: not

are25— 13— 25136 TABLE 21290 21346 SAN deviations F F Cratoxylum: Tridesmos Isopterygium:Isopterygium: articulata sumatranum cochinchinense formosum maingayi arborescens arborescens arborescens glaucum Standard Cratoxylum Section Section Section C. C. C. C. C. C. C. C. Eliaea P, BAAS: Anatomy of Eliaea and Cratoxylum 391

(see

1 0 ?? 00 Phillipines glaucum 2 0 papillae - — — 421 — — the 4 C. + 24 stereomicroscope, from conspicuous 0 0 0 0 ? ? — — specimen posses of 8 i 7 0 maingayi — 180 — — one 210 070 500 not 2 I 0 5 C. + — — Only magnification does

0 o i 6 0 0 ?? specimens). 81 pruniflorum formosum 4 5 8 8 19 Cratoxylum — 30 formosum 040 050 800 080 5190 (7 ssp. 0 0 8 5 using + 37 19 0 in C. ssp. blancoi 376). when ssp. papillae formosum p. 9 5 4 0 0 on present ? ? — C. 6 0 4 — 19 42 — sumatranum papillae cochinchinensecochinchinense 669 305 0420 040 6 3 33 0 papillae. 0 Epidermal + — C. description C. doubtfully — and III: epidermalpapillae showed are

?? TABLE 12 specimens) papillae sumatranum nereifolium 212 — specimens) epidermal — — — — — — (43 2 conspicuous + C. ssp. without which arborescens (170 with in

C.

in sumatranum specimens specimens 60specimensx —— —— ——— ——— ——— ——— —— found ssp. of of of =— (mainland) were number number number sumatranum AsiaAsia papillae = = Locality MalayaMalaya Sumatra Borneo Celebes Philippines C. + ? S.E. Java No of