BLUMEA 21 (1973) 193—258
Thewood anatomicalrange in Ilex (Aquifoliaceae)
and its ecological and phylogenetic significance
P. Baas
Rijksherbarium,Leiden
Contents
Summary 193
Introduction 194
Materials and techniques 195
Reliability and comparabilityof the material studied 195
Results and Discussions
Generic description of the wood ofIlex 196
Ontogenetic changes in vessel member length and number of bars 197
Diagnostic value of the characters used 199
Discussion of the individual characters
Growth rings 199
Vessel distribution 200
Vessel frequency and vessel diameter 200
Vessel member length and number ofbars 201
Vessel wall pitting 211
Spiral thickenings 212
Fibre-tracheids 216
Parenchyma 217
Rays 217
Crystals 218
Wood and wood 218 phylogeny ecological anatomy Wood and the classification of anatomy Ilex 221 Specific wood anatomical descriptions 225 Acknowledgements 248
References 249
Addendum 251
Summary
The wood of of Ilex is described detail. The wood anatomical encountered anatomy 81 species in range is presented in a generic description (p. 196). Data on ontogenetic changes in vessel member length and number of bars for three per perforation are given species. The variationin in is great amountof mainly quantitativebut also some qualitativefeatures hardly or not related with subgeneric classification but with latitudinal and altitudinal distribution. In both the northern
and in New southern hemisphere and both the Old and World, temperate and subtropical species are members characterized by conspicuous growth rings, numerousnarrow vessels, relatively short vessel and few bars both vessel and fibre and the fibre- per perforation plate, conspicuous spiral thickenings on walls, bordered wall tracheids are frequently provided with rather numerous conspicuously tangential pits. In tropical lowland species growthrings are absent or less marked, the vessels are scanty and wide, the vessel members are long and the number of bars per perforation plate is high. Spirals are lacking or faint, oroccur only in minor of the axial elements. The fibre-tracheids have few with reduced a part usually pits more borders the walls. resemble the on tangential Tropical montane species temperate ones to a great extent, but this does not apply to growth rings, spiral thickenings, and frequency and size offibre-tracheid pits.
The only wood sample of a climbing Ilex species from the tropics studied deviates from the general trend BLUMEA VOL. No. 194 XXI, 2, 1973
in few bars The I. and I. verticillata having per perforation plate. two temperate species serrata are exceptional in In lacking spiralthickenings. all wood characters they resemble the genus Nemopanthus (also Aquifoliaceae)
very closely.
Comparisons with data from literature and original observations on Prunus, Symplocos, Vaccinium. and also the that Viburnum, to some extent on Hydrangea, support view the gradual differences between
and Ilex conform to trend also in other Therefore temperate tropical species a general present taxa. a major influence climatic on wood structure is indicated. This is discussed with reference to the major trends of
phylogeneticwood specialization. The fact that within Ilex and Symplocos the tropical lowland species have
with the numerous bars be with perforations most cannot broughtin agreement the generalphenomenon of of scalariform in a rare occurrence perforations tropical lowland floras.
Other items such the between the absence of and entire as parallel spiral thickenings the presence of leaf
the lack of a clear taxonomic in the wood anatomical variation and observations margins, pattern in Ilex, former students of by Ilex wood anatomy are also discussed.
Introduction
Whilein search for relatives of the and possible genera Sphenostemon Oncotheca, which had been be allied several the felt thought to to Aquifoliaceae by taxonomists in past, I the of need for a more comprehensive study the wood anatomical range within that family.
The of the of genus Ilex, with a roughly estimated number 400 species, constitutes bulk the from family. The other genera are Nemopanthus North America, and Phelline from
New Caledonia.
Ilex in all into occurs tropical areas of the world and extends temperate regions up to and 65° N (America, Eurasia) 35° S (America, Africa). In the tropics a number of species
is montane but a fair number occurs in the lowlandforests. Because of this wide latitudinal
and and of the wood altitudinal, consequently climatic, range of habitats, a survey ana- tomical within this became all the order range genus more attractive in to analyse any
that between and wood am aware that for relationship might exist provenance anatomy. I
conclusive evidence oncorrelations between climate or other ecological factors and wood the of wood well defined structure, study many samples belonging to a single species
should be carried out in the first place in order to be more certain about genetical homo- the material geneity in used for comparison. Preferably such a study should include natural afford specimens cultivated outside their geographic range. Sufficient material to
such an analysis was, however, not available for this study.
This will of 126 paper report on the results a study of wood samples belonging to 78 ofIlex fromall distribution identified species and 3 unidentified, but different, species major and from of areas most the important taxonomic groups recognized by Loesener (1901, A the affinities of Phelline 1908, 1942). subsequent paper will deal with Ilex, Nemopanthus,
(Aquifoliaceae) Oncotheca, and Sphenostemon in the light of the whole range in characters
from both wood and leaf.
In previous studies some common species have been described wood anatomically by several of detail. elaborate authors with a varying amount The most study is by Penning- of ton (1953), whose thesis is, however, not easily accessible. She studied 54 species Ilex. material Aquifoliaceae, 51 of which belonged to Her was derived from specimens 'evidencing secondary growth' and from her quantitative data it becomes clear that she used for small twigs for most tropical species and both twigs and mature wood samples the
temperate species. In her discussion Pennington did not take these differences in sampling
methodinto account, and all her painstaking measurements and conclusions based on these
therefore (and in complete disagreement with my results) are unfortunately without value. Solereder (1899 and 1908) and Metcalfe and Chalk (1950) comprehensively summarized the older literature. Additional and more recent publications are scanty fragmentary. Hu
(1967) reported on evolutionary trends in the xylem of Ilex. Ingle and Dadswell (1961) P. BAAS: Wood anatomy ofIlex 195
dataof Ilex in their discussion of the affinities of Nouhuysia included quantitative 5 species (= Sphenostemon). More general publications by Abbate (1970), Barghoorn (1940),
Brazier and Franklin (1961), Chattaway (1956), Greguss (1959), Jane (1970), Panshin, De Zeeuw and Brown (1964), Purkayastha (1963), Sarlin (1954), Sudo (1959), and Versteegh
also (1968) contain data on several species of Ilex. The results of these previous studies will
is for the only be referred to in the discussion as far as relevant present study.
MATERIALS AND TECHNIQUES
from numberof institutional collections from Wood samples were obtained a wood or the wood collection of the Rijksherbarium. These institutions will be referred to using each The of detail Stern's (1967) abbreviations, at the end of specific description. amount collection data in appeared to be very variable in the material received, and the methodof listing specimens can therefore not be uniform. collection numbers Herbarium vouchers are listed when known; only referring to wood between brackets. wood specimens are given Mature samples (diam. of main stem least without immature or branch at 5 cm; seep. 197) are listed special indications, samples
(diam. of main stem or branch between 1.5 and 5 cm) are marked with an asterisk.
Woodanatomical in features were studied transverse, radial, and tangential sections, and obtained Franklin's method macerations. The latter were using (Anonymous, 1968) and mounted in glycerin jelly. made for vessel member For each sample 25 measurements were length, vessel diameter, of r and fibre Vessel number bars per per oration plate, length. member length was mea- sured the and including tails. Data on vessel frequency ray frequency are based on at least
in each in of and distances of 3 counts specimen areas 1 square mm, tangential 1 mm vessel due of respectively. For frequency attention was paid to include equal portions
in Pit sizes in early and late wood temperate species. were measured a few pits of the most and of frequently occurring size class, pits some extreme sizes. of for several Presence or absence spiral thickenings was confirmed species with the well scanning electron microscope. Clean cut tangential, transverse and oblique surfaces as as radially split surfaces were examined after coating with carbon and gold.
Reliability and comparability of the material studied
The and most recent monograph of the genus Ilex was published in 1901 1908 by who distributed Loesener, recognized 279 species, over 4 subgenera, numerous sections,
and he added a fifth Much material has subsections, series. Later, in 1942, subgenus. more become available since Loesener's time, many new species have been published, and at the needof This that the present genus is in a comprehensive new revision. implies naming of the woodspecimens I used for this study may be partly incorrect, which is all the more consisted of unvouchered serious since a rather high proportion specimens. Therefore, that the studied one should be aware names of the specimens are not always entirely reliable, though no doubt can exist about the provenance and the fact that all material described here belongs to the genus Ilex. On wood specimens data on localities are usually extremely undetailed.In assigning a degree of latitude to a specimen from 'Japan',
'China' or 'New Guinea' one therefore has to take an arbitrary decision, which will always be incorrect to some extent. However, such mistakes cannot have influenced the results Altitudinal data the general seriously. are usually lacking, and analysis of a direct or 196 BLUMEA VOL. XXI, No. 2, 1973
effect ofaltitude wood indirect on structure has therefore been limited to those specimens and species of which reliable data on altitude were available. All Ilex but the whole from species are woody, range small shrubs to tall trees exists.
Wood have been collected from main from specimens may stems or branches, a part high up the tree or close to the soil. Such data are usually lacking for the wood specimens well known from studied, yet it is that distance the pith and height in the tree (or shrub)
influence wood anatomical may characters, particularly quantitative ones, to a great the and extent. Of these, distance from the pith is most important factor, to increase the of data from both shrubs with comparability on specimens and trees, only material a of diameter of 5 cm or more was considered in the comparisons quantitative features. Such material is referred to as 'mature', and it is assumed that beyond this diameter the increase in cambial initial length and changes in cambial products are negligible for this study.
to substantiate this will be Arguments given on p. 197—199.
RESULTS AND DISCUSSIONS
Generic description of the wood of Ilex
Qualitative features based on 81 species. Quantitative values are given for mature wood for samples only. Only the ranges of means individual samples are given.
General features: Wood light and soft to fairly heavy and hard. Colour white
brown. Growth absent to distinct. Vessels visible thenaked in to greyish rings to eye some the naked tropical lowland species only. Broad rays visible to eye, usually prominent.
Heartwood not differentiated.
Wood Growth Microscopic features: diffuse to semi-ring porous. rings
2 absent to distinct. Vessels and in radial from 13—22o/mm , solitary multiples ranging short vessels oval to very long (over ten vessels), percentageofsolitary 8—70, angular to in
tangential diameter mean vessel member length 600 T.S., mean 30—113 /im, —1920 fim. others Inter-vessel pits usually opposite, but in some species tending to alternate, in
sometimes transitional to scalariform; pits round to rounded rectangular or strongly
diameter to with minute circular slit-like flattened, tangential 5 over 20/mi, to apertures, enclosed within the pit borders, in few species seemingly vestured. Vessel—ray and vessel scalariform half —parenchyma pits to alternate, fully bordered, bordered or tending to sometimes in large and simple, unilaterally compound. Perforations scalariform very end walls with bars. Helical well oblique 13—58 thickenings ('spirals') developed, faint,
or absent. Warts notedin two species only. Tyloses and vessel contents (probably traumatic)
noted in a few specimens only. Ground tissue composed of thin- to thick-walled flbre-
tracheids with minute to fairly large bordered pits. Pits frequent on radial walls, infrequent diameter slit-like and to frequent on tangential walls, average 3 —7 fim, pit apertures
confined within the pit borders to extending several microns beyond the pit borders.
Mean fibre-tracheid length 890—2830/im. Helical or annular thickenings ('spirals') well
developed, faint, or absent. Fibre-tracheids rarely gelatinous. Parenchyma diffuse and/or
diffuse in aggregates, very scanty to fairly abundant. Parenchyma strands of 4—13, of cells. mostly 7 or 8 Rays mostly heterogeneous II, rarely heterogeneous I or III, uniseri-
cells tallest —6.0 ate rays 6—16/mm; broad rays I —7/mm, 2—25 wide, rays 0.8 mm high,
often with sheath cells.
and chambered idio- Crystals present or absent, usually solitary in ordinary, erect, or absent. blastic thick-walled and/or bulging ray cells; rarely irregularly clustered. Silica
Pith flecks (traumatic) occasionally present. P. BAAS: Wood anatomy ofIlex 197
Ontogenetic changes in vessel memberlength and number of bars
Three of different stems widely species from different geographical origin were ana- for the vessel member number lysed variation in length and ofbars per perforation plate with This respect to the distance from the pith. was important in order to establish at which diameter these characters become less stable and stem more or consequently when
we can speak of 'mature' wood. Figures I and 2 give the curves for vessel member
Fig. 1. Vessel member (in in relation distance from I. I. length µm) to the pith in cymosa, dumosa, and I. aquifolium. 198 BLUMEA VOL. XXI, No. 2, 1973
of length and number bars respectively. For vessel member length the classical curves,
and fibres frequently reported for both vessels are obvious: a strong increase in length in first and the the centimetre from the pith curve leveling off towards the periphery.
In I. aquifolium (UN 163, cult. Holland) the most important increase has been completed within the from the vessel member but first 7.5 mm pith; length gradually, very slightly
increases beyond this point. In I. dumosa (Lindeman & Hecking 1916, Brazil, Parana) the
I. length attains a maximum at 15 mm from the pith and fluctuates beyond here. In
is a increase to mm from the cymosa (Ridley s.n., Singapore) strong up 22.5 pith, beyond
fluctuation. If mature which there is a very strong we define wood as secondary xylem
from of the where with those parts stem, there is no important increase in length respect further distance from the conclude that for these Ilex the to pith, we may species mature
is less reached in of which the is and stage more or stems xylem cylinder just over 1.5, 3.0,
2. Number of bars in relation to distance from the in I. I. Fig. per perforation plate pith cymosa, dumosa, and I. aquifolium. Wood Ilex P. BAAS: anatomy of 199
in diameter. With the chosen for wood in of 4.5 cm 5 cmlimit, arbitrarily mature samples
Ilex (cf. p. 195) I am therefore on safe grounds.
The number in of bars per perforation plate is virtually constant regard to distance the both and dumosa. from pith in I. aquifolium I. In I. cymosa there is a considerable in-
the first from the and rather fluctuation data crease in centimetre pith a strong beyond. No from from literature are available for comparison, and it is impossible to conclude the
whether for the rule or those present data the curve I. cymosa is an exception to not. In of which studied both and trend species I mature immature samples no consistent in number ofbars couldbe five differences in per perforation found. In species the immature
wood sample had slightly more or about the same number ofbars per perforation as the
mature wood sample(s); in three species this number was considerably lower for the im- of mature specimens. In most cases exact diameters of the stems the samples were un- known, which decreases compatibility.
It is, however, justified to conclude from our graphs that xylem of samples more than diameter also with number of bars Further 5 cm in is mature respect to per perforation. in study is necessary to establish whether any general trends the ontogenetic changes of
numberof bars per perforation plate exist in Ilex and other woody plants.
Diagnostic value ofthe characters used
From the that generic description it appears only very few wood anatomical characters
are diagnostic at the genus level. Values for size and frequency of the different elements
them for show an enormous range, which makes useless diagnostic purposes. Qualitative features such vessel as growth rings, distribution, pitting of the vessel walls, spiral thick- also variable Ilex. The characters enings and crystals are within only which are constant
at the genus level are: vessel perforations scalariform, ground tissue composed of fibre-
tracheids, parenchyma diffuse or diffuse in aggregates, rays heterogeneous, broad and
This character is in number narrow. complex also present a fairly great of other families with wood primitive structure (cf. Metcalfe and Chalk, 1950).
At the level values also in species quantitative exhibit a considerable range those species of which I studied sufficient material (cf. I. cymosa, I. mitis and I. spec. from New Guinea).
the of there Accordingly in 63 species which mature wood samples were available is a anatomical great amount of overlap, so that species are not separable on quantitative wood of the features. For a number qualitative features same applies. Semi-ring porosity is not varies constant within e.g. I. integra, inter-vessel pit arrangement usually considerably of the within a single tangential section (Plate II, 6 & 7), and different specimens same if species may show a different range (e.g. I. anomala). Spiral thickenings, well developed faint the are constant within the species, but in tropical species with only some spirals in fibre tips, spirals may be entirely lacking as well (e.g. I. cymosa). Presence or absence of
crystals haslong beenknown to be variable at the species level and Ilex makes no exception
to this rule.
the fact within of of From that even the restricted number species which I studied ofanatomical features be more than one wood specimen the majority proved to variable,
one may conclude that it is impossible to separate Ilex species using wood anatomy.
Discussion of the individualcharacters
Growth rings
Virtually all temperateand subtropical species of Ilex show distinct growth rings; most have lack these In this the tropical species faint growth rings or entirely. respect genus 200 BLUMEA VOL. XXI, No. 2, 1973
Ilex follows well known rule. where less a general In some tropical species more or distinct growth ring boundaries occur this is probably due to a seasonal climate with pronounced dry periods (e.g. I. crenata and I. quercetorum). In other species (e.g. I. curranii and climatic for the of distinct I. volkensiana) no apparent reason occurrence growth rings can be given.
Vessel distribution (Plate I)
in Most Ilex species possess diffuse porous wood, but a few temperate and subtropical species the wood is semi-ring porous. This feature is not restricted to the deciduous sub- but also numberof Prinus as one might expect, occurs in a evergreen species (e.g. genus , there I. dipyrena, I. ficoidea, I. integra, I. latifolia, and I.perado). In some species is a transition between the diffuse-porous and semi-ring-porous condition (e.g. I. aquifolium).
There is a considerable range of variation in the extent to which the vessels are arranged
radial This the where in multiples. is only very incompletely covered in descriptions per-
of vessels The of radial consider- centages solitary are given. length multiples may vary the ably and strongly alter impression thewood gives as seen in transverse section. It would be to such differences the very time consuming express quantitatively. Moreover, intra- is considerable: for instance in anomala the in specific variability very I. range percentage of solitary vessels in mature specimens is 31 —64; in the specimen with the lowest per- the radial but centageof solitary vessels, multiples are not only more frequent, on average other There between also longer than in the specimens. is no clear relationship taxonomic subdivision of the Ilex and vessel there a correlation genus grouping, nor is very strong with all the latitudinal or altitudinal distribution. In distribution areas range is very large.
The Ilex species with the strongest tendency for vessel grouping (few solitary vessels, radial from the those with the shortest long multiples) are, however, temperateregions, and and of from multiples highest percentage solitary vessels are the tropics.
Vessel frequency and vessel diameter (Plate I)
The vessel within the Ilex great range of variation in frequency genus was found not to be related to its taxonomic subdivision. The same applies to vessel diameter. Both characters are inversely correlated, and are both related to latitudinal distribution (see fig.
3, 4, and 5). Temperate species are characterized by numerous narrow vessels, tropical species are characterized by few relatively wide vessels per unit surface area of the trans-
tend to verse section. All intermediates occur. Within the tropics, the lowland species have the wider and vessels, the montane species have narrower vessels. Vessel frequency in tropical montane species does not differ much from that of the lowland species (fig. 6), and is much lower than in temperate species growing under comparable temperature regimes.
variation in in Intraspecific vessel frequency and diameter may be considerable (e.g.
I. cymosa, I. havilandii, and I. integra).
For the latitudinal trends in vessel frequency and vessel diameter Ilex follows a more or less wood of has vessels general rule. In general temperate diffuse-porous trees narrower than that of tropical trees.
Versteegh (1968) has provided evidence that tropical montane species are frequently whilst their lowland relatives be This that the microporous, may megaporous. suggests Ilex for vessel diameter follows trend. Ab- altitudinal trend in tropical also a general bassova (1969) showed that both vessel diameter and vessel frequency decreased with
altitude in vessel is increasing Quercus. This does not apply to Ilex as far as frequency concerned. P. BAAS: Wood anatomy of Ilex 201
in in Fig. 3. Vessel frequency and latitude 64 species Fig. 4. Vessel diameter (inµm) and latitude 64 of Ilex (I. crenata plotted at 17° and 52°). species of Ilex (I. crenata plotted at 17° and 52°).
Vessel member length and number ofbars
(Plate II, I—5; III, 5 & 6)
the correlation between vessel member and latitude of In fig. 7 length provenance is shown wood of for mature samples 64 species of Ilex. Although there is a considerable variation in all latitudinal zones, the trend is evident: the tropical species have considerably vessel members than the The numberofbars longer temperate ones. per perforation plate is also inversely correlated with latitude (fig. 8). Accordingly, vessel member length and number ofbars are positively correlated (fig. 9). The vessel member lumping of data on length and number ofbars of all species as done in & is is figs. 7, 8, 9 justified. There no obvious correlation of thesefeatures with taxonom- ic subdivision, and the general correlations apply equally well in the southern and northern hemisphere, and in the Old and the New World. 202 BLUMEA VOL. XXI, No. 2, 1973
Fig. 5. Vessel frequency and vessel diameter (inµm) in 64 species ofIlex.
in lowland Fig. 6. Vessel diameter (inµm) and vessel frequency tropical montane(o) and tropical species (•). Ilex P. BAAS: Wood anatomy of 203
Fig. 7. Vessel member length(inµm) and latitude in 64 species of Ilex (I. crenata plotted at 17° and 52°).
Within the tropics the montane species studied have shorter vessel members and less 'alti- bars per perforation plate than the lowland species (fig. 10). For the analysis of the those and taken which tudinal' effect only specimens species were into account on reliable dataon altitude were available. Of the species in fig. 10, eight are represented by specimens from from sea level to well below iooo m, one is an altitude between iooo and 2000 m, and the remaining seven species are from altidudes over 2000 m. There is only little overlap in the values for vessel member length, and the number of bars is consistently 204 BLUMEA VOL. XXI, No. 2, 1973
ofbars and latitude in ofIlex and Fig. 8. Number per perforationplate 64 species (I. crenata plotted at 17° 52°). BAAS: P. Wood anatomy ofIlex 205
Vessel member and number ofbars in of Ilex. Fig. 9. length (in per perforation plate 64 species
2 206 BLUMEA VOL. XXI, No. 2, 1973
Fig. 10. Vessel member length(in µm) and number of bars in tropical lowland and tropical montane
= species of Ilex; — o = montane species (altitude over 2000 m); — x from altitude between 1000 and
2000 m; — • = lowland species (altitude below 1000m). P. BAAS : Wood anatomy ofIlex 207
lower in the montane than the lowland species from both the Old and New World tropics.
Fig. II gives a comparison of microtherm tropical montane species (wood samples from altitudes than with microtherm latitudes higher 1600 m) temperate species (from
0 These be considered to under over 35 ). species groups can grow comparable temperature regimes (not taking into account seasonal influences), according to the altitude—latitude scheme proposed by Van Steenis (1962). It is clear that the tropical montane species tend have to longer vessel members than theirtemperate counterparts, and that the numberof bars does differ in the and not considerably two categories (26 23 on average respectively).
In order to explore whether the correlation trends of vessel member length and number of bars per perforation plate with latitude and altitude in Ilex is an example of a general rule or not, two methods can be applied. One possibility is to compare data for 'whole' woody floras from the to different latitudes and altitudes; other is compare species from different latitudes and within other altitudes eurytherm genera. As the for first possibility one must bear in mind that vessel member length is a character which be much less variable for taxa such as and families than for in- may higher genera stance vessel frequency and vessel diameter. Therefore it is not justified to compare our data Ilex data for whole floras and the on with woody available for e.g. Japan, Formosa,
Fig. 11. Vessel member length (in µm) and number of bars per perforation plate in microtherm species from and from sites 1600 temperate regions (latitude over 35°: •), tropical montane (altitude over m: o). 208 BLUMEA VOL. XXI, No. 2, 1973
Philippines (Kanehira 1921 a & b, 1924). Making a frequency distribution for maximum member classes of the described finds vessel length species by Kanehira, one hardly any
diiference between tropical and subtropical to temperate floras. However, the frequency
distribution for the Philippines is highly influenced by the comparatively high proportion
of Dipterocarpaceae, Guttiferae, Leguminosae, and Meliaceae, which are characterized by
rather short vessel members. These families and are very poorly represented in Formosa On the other hand, families with members like Japan. comparatively long vessel e.g.
Magnoliaceae and Theaceae are better represented in the materials studied by Kanehira. If
the of those of which material from all one compares species genera Kanehira studied that three countries it appears in Albizia, Cinnamomum, Litsea, and Quercus the 'more have the tropical' species longer vessel members. For Ficus this is not very evident, and for
Melia and Diospyros no clear differences can be found in Kanehira's data for the different
localities.
results Of Prunus I studied 12 species from widely different regions (Table I). The
clearly demonstrate that in Prunus too tropical species tend to have the longer vessel and the shorter table similar members temperatespecies ones. In II a phenomenon is shown
for Viburnum, Vaccinium, Symplocos, and Hydrangea. For Prunus and Symplocos table I and evidence II also containsome that withinthe tropics montane species have shorter vessel
members than the lowland species. the literature found for the idea that within with In I some more support taxa represen-
tatives inboth the tropics and subtropical to temperateregions the tropical species have the members. Schweitzer longer vessel (1971) showed this to hold for Celtis (Ulmaceae),
Carlquist (1966) for woody Compositae. For Nothofagus, Dadswell and Ingle (1954)
showed that species belonging to the temperate sections had the shorter vessel members those the subsection. If vessel member than belonging to single tropical one equates
length with fusiform cambial initial length, as is generally accepted to be valid, one may
draw an interesting parallel with data on conifers. Dinwoodie(1963) showed that tracheid
in Sitka decreased with latitude of length Spruce (Picea sitchensis) increasing provenance cultures he could also establish that tracheid (41 —63° N). In seedling length was genet- ically fixed in the different populations from different latitudinal origin. He tentatively ascribed these differences effect' of cambialinitial to a 'long term temperature on length.
Reviewing all these data on latitudinal variation in vessel member length, it seems within the fully justified to accept as a general rule that taxa tropical representatives have
the longer vessel members and the species from higher latitudes have the shorter ones.
The same probably applies to altitudinal variation in vessel member length.
Data on number of bars per perforation plate in relation to latitude and altitude are hardly available in the literature. One may cite my own results onyoung twigs ofPlatanus species (Baas, 1969), and interpret the higher percentage of scalariform perforation plates ofbars the and the higher number in tropical species from Laos and Mexico as comparable the latitudinal Ilex. for to trends observed in Novruzova (1968) compared some species specimens from different altitudes in Soviet Azerbaijan, and found similar altitudinal found Ilex: thenumber ofbars trends as I in per perforation plate decreased withincreasing altitude. observations and Viburnum also Original on Symplocos (Table II) are suggestive Ilex that the latitudinal variation of number of bars per perforation plate as found in is a general phenomenon. The two species ofHydrangea studied seem to contradict this, but here deal with wood from climbers which be modified that we may so strongly they be this The from Ilex cannot compared in respect. only wood specimen an climber I studied, also does not follow the latitudinal trends Tabata's data (see p. 248). (1964) on P. BAAS: Wood anatomy of Ilex 209
faint well well faint or developed developed developed developed spiral thickenings fine fine very very absent absent very absent very absent absent well fairly well fairly
length means) Prunus 650 910 770 700 780 660 590 1030 510 440 640 380 & of member range in 480—310— 710—390— 580—290— 530—310— 580—360— 460—340— 400—270— 710—410— 270—390— 270—360— 470—330— 290—190— species vessel (full
12
m m in m m m 35 altitude ? ? 2780 100 ? 3600 50 o low low low thickenings 21598) studied 327 (IF 24491) spiral 5173 s.n. Balgooy 053) 9680 (MADw 9217 229 1918 Tervuren and specimen Kalkman s.n.) s.n.) Lewalle BW BW Van BW (Vienna length and Stahel (Kw (Kw cult. Guinea, Pakistan, Guinea, Guinea, Guinea, Guinea, member Origin Belgium, Burundi, New East New New Surinam, New New Austria, England, England, Vessel Kalkman Kalkman I. Kalkman f.) Miq. Kalkman Table Kalkman (Koehne) Muell.) Urb. SPECIES (Hook, (Wt.) SPECIES L. L. (Bl.) Bl. (Koehne) (C. L. L. TROPICAL africanum arborea ceylanica grisea javanica myrtifolia pullei schlechteri TEMPERATE avium cerasus lusitanica virginiana
A. P. P. P. P. P. P. P. P. B. P. P. P. P. 210 BLUMEA VOL. XXI, No. 2, 1973
&.
8 bars 26 25 60 11* 13* 38 34 64 54 34
m of range fi 18— 17— means 6—2— 7—1— 2—4— genera in 13—19— 11— 28—39— 24—15— 11—19— 21—34— 35—19— 7— number ■full
miscellaneous length means) -1440 -1100 -2270 670- 860- -1450 -1340 -1800 -1750 980- -1240 & member ,um 990— 860— 1690— 550— 640— 1040— 1090— 1320— 1490— 720— 1100— some range in 590— 450— 1190— 440— 490— 720— 720— 790— 1130— 480— 1000— in vessel (full plate m m m altitude ? ? 2000 ? 2400 7 ? 3140 low low ? perforation 23539 (climber) per 6521) 6080) studied Reitz (climber) bars (U 30653) 18849) (CTFT 276) 7435 7026 4870 3407 of 588, (USw (UN specimen (USw Catharina, Sarlin number Jacobs Jacobs Kalkman cult. and Barkorda 19399) Breteler and Maryland, Maryland Santa Origin (USw Guinea, Caledonia, length Portugal, Philippines, Philippines, Netherlands, Venezuela, U.S.A., Texas U.S.A., Brazil, New New
member Zucc. Gris
Vessel Merr. simple L. Vidal l'Hërit. Brand. Sieb.et Marsh. Brongn.et Briq. II. Species
L. Table prunifolium glaberrimum arboreum barandanum tinctoria tenuifolia arborea petiolaris durifolia predominantly Viburnum tinus Vaccinium Symplocos spec. Hydrangea V. V. V. V. V. S. S. S. S. H. H. * P. BAAS : Wood anatomy ofIlex 211
Ilex and Japanese species of Betula also seem to be contradictory to those obtained on other found that the lowest number of bars occurred in some genera. He per perforation
species with a preference for wet sites in the valleys. For some high-montane species, partly also from higher latitudes, he recorded much higher numbers of bars. One has to
be cautious with Tabata's data since he obtained them from twigs 5 mm in diameter, and from his clear whether he studied the of the paper itis not peripheral part secondary xylem
or all of the xylem formed. He also did not give exact data on altitude, and his general associated with with conclusion that a low number of bars is species growing in sites a results high moisture content seems to be contradicted by his own on Betula platyphylla
sites var. japonica. In this variety he found the specimens from marshy to have more bars per perforation plate than the specimens from well-drained sites. If that is one nevertheless accepts within taxa with scalariform perforations there a
of decrease with latitude general trend for the number bars per perforation to increasing and would the lowland habitat be a favourable altitude, one expect tropical to environment to foster species with elaborate scalariform perforation plates and the montane tropical better suited for with reduced scalariform and temperate environment to be species perforations viz. simple perforations. An analysis of the abundanceof species with simple scalariform this idea. or perforations in different woody floras does not support Kanehira showed that houses much of with (1921a) Japan a higher percentage (32.6%) genera scalariform perforations than the Philippines (5.7%), Formosa being intermediate with
19.4%. A comparison of the European woods with woods from Java (Greguss, 1959, and also of with scalariform Janssonius, 1906—1936) gives a higher percentage genera per- forations for than for temperate and subtropical Europe (23%) tropical Java (13%). In a comparison of the woody montane flora with the lowland woody flora of Indonesia,
Versteegh (1968) showed that scalariform perforation plates were of more common oc- currence in the mountains than in the lowland. The low frequency of taxa with scalari- form perforation plates in tropical lowland forests is well known to wood anatomists. for Therefore the trends reported Ilex, Symplocos, Viburnum, and for Platanus and some of species from Azerbaijan seem to be contradictory to the general distribution species with scalariform and simple perforations. This will be further discussed in relation to
See also Addendum phylogenetic implications (p. 218). on p. 251.
Vessel wall pitting (Plate II, 6— 13; III, 1, 3—9)
in size wall within is The range type and of vessel pitting Ilex considerable. In many also within each species there is a considerable range individual specimen, restricting the character. diagnostic value of this Particularly the ranges transitional to opposite, and opposite to alternate inter-vessel pits frequently occur within one sample. Other species show from may even the whole range scalariform to alternate (e.g. I. aquifolium and I.
the berteroi). Usually one type predominates, however. Of species studied, I. anomala,
I. berteroi, I. casiquiariensis, I. chinensis, I. coriacea, I. cornuta, I. decidua, I. glabra, I. longipes, have These and I. volkensiana a fairly high proportion of scalariform inter-vessel pits. from all three of species are subgenera Ilex studied, and are distributed haphazardly over the various series and sections recognized by Loesener. Surprisingly enough, there is there- fore no correlation betweentype of inter-vessel pits and subgeneric classification. A corre- lation of of with latitude altitude of also type pits or provenance is absent, nor is there with vessel member numberof bars This any relationship length or per perforation plate. what would is contrary to one expect, since in general analyses it has been established for whole that of correlated with vessel Dicotyledons as a type pitting is strongly member 212 BLUMEA VOL. XXI, No. 2, 1973
vessel length: the scalariform type of pitting being far more frequent in woods with long
members, and opposite to alternate pitting being more in evidence in woods with short vessel members (Metcalfe and Chalk, 1950: XLIII—XLV).
the inter-vessel Often the Vessel—ray pits are usually similarly arranged as pits. arrange-
ment is slightly disturbed, however. Unilaterally compound pits are of frequent occur-
than bordered rence within the genus. Sometimes one ray pit corresponds with more two
vessel wall pits, which are usually in horizontal rows. Occasionally the pits on the vessel
wall side are small clusters of or arranged in 3 4 ('complex unilaterally compound', Plate The which the and III, 6). extent to pits on both vessel- ray-side are overarched by scalariform borders varies also considerably within the genus. In some species with to
transitional vessel—ray pits the borders are much reduced, resulting in almost simple
pits.
Spiral thickenings (Plate III, 2; IV, V; 1 & 2)
Plate IV shows the range of vessel and fibre wall sculpturing. All possibilities from
realized within Ilex. absence of spirals to spirals well developed are In some species only
faint spirals occur in a number of fibre-tracheid tips. In others the vessels are devoid of
spirals but most fibre-tracheids have fairly well developed spirals. Yet another possibility
the of faint the walls of both vessels and fibre- is presence very spiral thickenings on is and in tracheids. Very rare the combination of spirals faint infrequent the fibre-tracheids and well the vessels. and The less developed in (I. curranii I. dumosa). more or continuous
well the range from developed spirals to very faint or no spirals includes at least two of
types recognized by Parham & Kaustinen (1973). Therefore, and probably also because of the small number of different woods their classification of of studied, types spiral thickenings is rather artificial.
Fig. 12 shows the proportional distribution of species with or without well developed well intermediate different latitudinal Almost spiral thickenings, as as species over zones. all without the species spiral thickenings appear to be confined to tropical zonebetween The I2°30' S and N. two temperate species without spiral thickenings are exceptional with both and (see p. 225). Most species well developed spiral thickenings in vessels fibre-
tracheids to be found the and Those with are in subtropics temperate regions. species
0 0 well from the latitudinal between and are fact developed spirals zones 13 25 N or S in
mostly from the transitional zones between tropics and subtropics (20°—25°). Species with faint vessels and with well one spirals in fibres, or spirals developed in only category of these elements the Of are mainly confined to tropics. these the species growing closest the have the faintest confined few fibre-tracheid to equator spirals, usually wholly to a This tips. condition is therefore much closer to 'spirals absent' than to 'spirals well devel-
oped'. The temperate I. montana with its faint spirals in the vessels is exceptional for the
temperate regions, where development of spiral thickenings is usually most conspicuous. be Summarizing, there is a very obvious tendency of spiral thickenings to rare in the the tropics and to become very common and conspicuous in subtropics and temperate regions. the Ilex further made the For tropical species a attempt was to analyse variation en- countered. Table III gives the distribution of species with well developed spirals, no and intermediate altitude and spirals, species in relation to seasonal or everwet climate. Total of be absence spiral thickenings seems to more common in montane species than in lowland species. The climatic condition as expressed in seasonal or everwet also seems to have bearing the matter. All Ilex without some on tropical species spirals grow in P. BAAS: Wood anatomy ofIlex 213
in Fig. 12. Occurrence of spiral thickenings 81 species of Ilex , and latitudinal distribution ( Ilex crenata
— = — = faint restricted ofvessels plotted at 17° and 52°). white spirals absent; hatched spirals or to part
= and/or fibre-tracheids; — black spirals well-developed.
in climate with short habitats with an everwet climate or a a very dry season. Climatic data used for this analysis were taken fromWalther and Lieth(i96o). Because the number of for data recorded those stations which meteorological are is restricted, only species couldbe ofwhich data suitable with Walther and considered locality were for comparison Lieth's data.
From the fact that in exceptional cases well developed spiral thickenings occur both
conclude that and everwet and montane habitats, one must rain fall temperature cannot
Table III. The occurrence of spiral thickenings in the wood of lowland and montane
tropical Ilex species in relation to climate and altitude
Number of species with:
well developed spirals faint or rare spirals no spirals
CLIMATE
everwet 1 10 7
wet with short dry season - 7 1
with pronounced dry season 1 9 -
ALTITUDE
lowland (below 1000 m) - 12 2
montane (over 1600 m) 1 4 6 BLUMEA VOL. No. 214 XXI, 2, 1973
be considered as entirely responsible for the presence or absence of spiral thickenings.
of Because this one must assume that thecharacter 'spiral thickenings' has to a great extent fixed the absence of become genetically in species involved. The spirals in the temperate species I. serrata and I. verticillata also points in this direction. Moreover, in general,
presence or absence of spiral thickenings remains a valid diagnostic feature within tem-
floras. For instance Viburnum characterized perate woody some species are by spiral whilst others thickenings, lack them (Greguss, 1959), also if they grow under the same
held for either conditions. Temperaturecan certainly notbe responsible favouring presence absence of the cool habitats absence of or spirals. In tropical montane spirals is the most condition of latitudes with frequent in Ilex; in regions higher comparable temperature absence of spirals that regimes is very rare. Novruzova (1968) reported spiral thickenings
were of more common occurrence in mesic species than in xeric species in Soviet Azer-
baijan. Whatever the climatic correlations found, however, from the foregoing it is clear
absence that the environmental conditions cannot fully explain presence or of spiral
thickenings, and one has therefore to assume that the evolutionary background plays
an important role in the distribution of this character over the different species of Ilex.
However, Ihave been unabletofind a taxonomicpatternbehind the variationencountered. of The more frequent occurrence spiral thickenings in fibres and/or vessels in plants
growing in subtropical and temperate regions as compared with tropical plants is a before general phenomenon, recognized (Kanehira, 1921a; Janssonius 1906—1936;
Panshin, De Zeeuw & Brown, 1970), though it has not received much attention by students of wood The trend becomes clear if the anatomy. general one compares per- of with in the flora and in the centages genera spiral thickenings European (Greguss, 1959) index of Javanese flora0anssonius, 1936, p. 31). In European trees and shrubs 91 generaout
studied have that is of all For out of 153 spiral thickenings, 60% genera. Java only 54 377 these genera (15%) have spirals. Moreover, 54 genera have usually faint or infrequent of spiral thickenings occurring in only a minority the species. Kanehira (1921a) reported of with of and for and the percentages genera spirals 40.7, 22.3, 4.7 Japan, Formosa, Philippines respectively. For the United States Record (1919) noted spiral thickenings in
of also in In both 47 genera out 122 studied, resulting a fairly high percentage: 38.5.
tropical America and Africa the occurrence of spiral thickenings is a rare phenomenon
(Mennega and Dechamps, private communications). The trend of be less towards the also general spiral thickenings to common tropics can be within demonstrated a number of genera. Celtis (Schweitzer, 1971), Gleditsia and
& and Elaeocarpus (Kanehira, 1921a b, 1924), Euonymus (Moll Janssonius, 1911), Prunus and and results in table Koelreuteria and (Moll Janssonius, 1914, my own I), Symplocos
(original observations) are a few examples. There between is an interesting parallel the diminishing frequency of spiral thickenings the towards the tropics and the higher percentage of species with entire leaf margins in with floras. tropics as compared temperate Sinnott with entire in Bailey and (1916) gave percentages of species leaf margins and floras. discussed the tropical temperate Ferguson (1971) gave further percentages and value of the percentage of species with entire margins in assessing the climatic conditions
of the past in paleobotanical studies.
Within leaf similar of Ilex too, margin characters behave as presence or absence spiral Ilex thickenings (see table IV). From the table it appears that in leaf margin characters are related to latitude to a considerable extent, but that the occurrence of spiral thickenings is even more strongly related to latitude. One cannot conclude from this parallel between spiral thickenings and leaf margin characters, that sculpturing of the vessel and/or fibre P. BAAS: Wood anatomy of Ilex 215
regions speciesspecies Ilex temperate 5 2 3 of 10 26 36 81
and the species of 3.
77 be 81 Subtropics for in should
S) here 8 available sculpturing N number
23°30' 5 wall 20 11 12 26 Tropics 2(3)** the and descriptions (between species temperate specificspecific 77 with fibres and in or later margins with and/or spirals tropical fromfrom characters margins margins or leaf and both leaf leaf spirals vessels developed margin under spirals of entire subentire dentate monographmonograph crenulate) faint part well Leaf treated with with with without with to with IV. or Loesener's is Table confined species species dentate species species species species crenata fromfrom MARGIN* of of of SCULPTURING of of of (weakly spines spirals studied. I. Number Number Number *Data*Data If LEAF Number Number Number WALL **** 216 BLUMEA VOL. XXI, No. 2, 1973
wall has the of climatic conditions of the fossil same potentiality predicting past, using
woods. In doing so one would ignore the diversity in climatic conditions in the tropics due in of the wood in to differences altitude. Moreover, comprehensive surveys anatomy
different floras would be needed in order to have firmer statistical basis to work many a with.
Fibre-tracheids (Plate V)
In Ilex fibre-tracheid length shows the same relationships with latitude and altitude as
vessel member length. It decreases with increasing latitude and/or altitude. The ratio the fibre length/vessel member length ranges from 1.2 to 2.2 (on average 1.6), most
1.8. ofintrusive frequent range is between 1.4 and This means that the amount growth of
the differentiating xylem elements is fairly constant within Ilex. Part of the variation, for the also ascribed differences individual possibly responsible extremes, may be to within
growth rings of temperate species. Many authors have shown fibre length to vary con-
siderably from early to late wood within one growth ring, the vessel members being
rather constant in length throughout the ring (e.g. Suss, 1967). No special attention was
devoted to sampling chips for macerations from special parts of the growth rings in this
it is that of the indeed the study, so that likely part variability was caused by different positions within the rings.
No relation exists between the elongation factor and vessel member length (regarded as cambial initial had more or less identical with length) in Ilex. Chattaway (1936) shown
that the shorter the vessel members the the of the dif- in general stronger elongation elements into for cambial ferentiating xylem developing fibres. However, the range of of the of recorded initial length 600—1900 /um (which is range Ilex) the differences she in
the of fibre to cambial initial are small and between ratios length length only range 1.3 which in and 1.8, is almost the same as Ilex.
Usually the bordered pits of the fibre-tracheids are more numerous on the radial than
but the latter from on the tangential walls, on they may vary very infrequent to abundant
and from having conspicuous borders enclosing the slit-like apertures to having much reduced borders with the slit-like far apertures extending beyond. There the is no clear correlation between diversity in fibre-tracheid pitting and the for classification by Loesener. In looking a correlation with latitudinal zonation a con-
siderable in size and is found in all latitudinal Yet there is range pit pit frequency zones. for the be small an obvious tendency tropical species to characterized by bordered pits in
low frequencies, whilst the bordered pits are larger and more numerous in subtropical and the restricted number of of which sufficient temperate species (fig. 13). In tropical species
I find trend data on altitude were available could not any of correlation in pit size and frequency with altitudinal distribution.
No from from adequate data other plant groups or the literaturewere at my disposal to establish whether the latitudinaltrend in fibre characters found in Ilex is a general one.
Two references may be significant here. Novruzova (1968) concluded that primitive
fibre elements (fibre-tracheids) were more frequently observed in plants growing under
conditions of adequate water supply, and that specialization towards libriform fibres
depends on atmospheric dryness. Versteegh (1968) concluded that fibres with bordered
pits were more in evidence in the tropical mountainsthanin the tropical lowland. My data insufficient are to support or reject either view.
Gelatinous fibres in irregular groups or partial concentric bands are present in I. laurina,
I. myricoides, and I. rimbachii. They have hitherto not been recorded for Ilex (Hoster and
Liese, 1966). P. BAAS: Wood anatomy of Ilex 217
in Ilex Fig. 13. a. Frequency of bordered pits on tangential walls of fibre-tracheids 64 species of and — = white = latitudinal distribution. — hatched pits infrequent; pits rather infrequent to rather frequent;
— black = pits frequent.
b. Diameter of bordered on tangential walls of fibre-tracheids in 64 species of Ilex and latitudinal pits ,
= — in — in distribution. white = borders less than black borders or more pit 5 µm diameter; pit 5 µm diameter.
in I. I. Hu (1967) reported the presence of septate fibres hawaiensis and cymosa. She must for because have mistaken the thin-walled parenchyma strands these, septate fibres are
material and absent from all I studied of I. cymosa from the other Ilex material, except for fibre-tracheids and Plate some infrequent septate-like in I. martiniana (see p. 240, V, 3). The fibres she for hawaiensis strands. septate pictured I. are clearly parenchyma
Parenchyma
The range in parenchyma abundance from very scanty to fairly abundant shows a
have diffuse sub- correlation with latitude. Temperate species very scanty parenchyma, which both diffuse and tropical and tropical species have more abundant parenchyma is This diffuse in aggregates. applies to lowland as well as to montane species.
Rays (Plate VI, I—6)
There tremendousvariation in size and within Ilex. Since the variation is a ray frequency be is continuous no relationships could established with taxonomic grouping, and an of the distribution size classes in analysis latitudinal of different of rays only resulted a
for the be abundant in the and the lower vague trend higher rays to more tropics rays to be evidence the width does not show correlations more in in temperate regions. Ray any with in and width has been latitude. Only the case of I. verticillata I. serrata, narrow ray used additional character their mutual close affinities as an supporting (p. 225). 218 BLUMEA VOL. XXI, No. 2, 1973
Crystals (Plate VI, 7—10)
all recorded here for Ilex to the the Although virtually crystals belong solitary type, than realized variation is larger previously (cf. Chattaway, 1956). Crystals in idioblasts are
reported for the first time. About30% of the species studied lack crystals. Of the crystallif- the be found the erous species, majority is to in tropics. clustered for few The irregularly crystals reported species may possibly be regarded as fragmented solitary crystals.
Wood and wood also phylogeny ecological anatomy (see p. 251)
the variation in in Ilex in Much of wood anatomy and some miscellaneous genera which reported above concerns characters are regarded as important indicators ofphylo-
In the context of this the trends genetic specialization. paper following specialization are for Frost's relevant (see Carlquist, 1961, a general summary of mainly Bailey's and work).
1. Short vessel members are derived from long vessel members.
with from with 2. Perforation plates few bars are derived perforation plates numerous bars.
of and of bordered fibre walls 3. Decrease size frequency pits on means specialization. and Trend 1 2 in particular have often beenregarded to represent irreversible evolution-
Of interest here is also the view that increase vessel ary processes. generally accepted in and of vessel in be diameter, a higher degree grouping multiples may interpreted as
specialization, though most authors are aware of the fact that particularly vessel diameter
environmental for may be easily affected by conditions, which opens possibilities reversi- bility of the specialization series.
After all the work that has been done on large and representative samples of Dicotyle- from be from donous woods, and what can deduced the vesselless ancestry of the angio- there be doubt about the ofthe trends of sperms, can no validity general xylem special- established and confirmed wood after him. This ization by Bailey by many anatomists
applies in particular to the three specialization trends listed above. It shouldbe pointed out
that the of trend is and that it ofthe evolution- wording 2 my own, represents only a part
ary pathway from a tracheid with crowded scalariform pits, via a vessel with scalariform
perforations to a vessel with simple perforations. results trends also be deduced: From my own some can
ia. Within taxa (genera) the longer vessel members are found in the tropical lowland
the shorter vessel members in the and certain species, temperate regions, to a extent
also in tropical high montane species.
2a. In trees and shrubs with scalariform perforations the number of bars is highest in the
tropical lowland and decreases with increasing latitude and altitude.
In of Ilex there is for fibre-tracheids have 3a. tropical species some tendency to more reduced and fewerbordered than the pits in temperate regions. The question arises which is the significance of these correlations betweenwood struc- for the which ture and climate general phylogenetic trend, or reversely, part phylogenetic wood specialization has played in these ecological correlations. Because of the evolutionary aspects of these questions the discussion must unfortunately remain highly speculative. vessel member and reduction ofbars Trends i/ia and 2/2a concerning length per per- that within foration plate could be harmonized by stating eurytherm genera the tropical lowland species have retained the most primitive wood structure, and that species of
latitudes altitudes in their wood For Ilex this higher or are more specialized anatomy. Wood Ilex P. BAAS : anatomy of 219
Ilex lowland and South would imply that primitive occurs in tropical Asia, Africa, Ameri-
if Loesener's that thecradle ofIlex is somewhere ca. However, one accepts assumption area the northern and that the under mild climatic conditions in hemisphere, genus originated this ('mildes See-Khma', 1908, p. 246), assumption is not very likely. Moreover, Loesener regarded most of the tropical species as derived rather than as primitive, and a number of series of subgenus species which he regarded as very primitive (belonging to Lioprinus
Euilex) have for Ilex a relatively highly 'specialized' wood anatomy (e.g. I. canariensis, latitudinal I. chinensis, I. montana, I. rotunda, and I. uniflora), in accordance with their Whether Loesener's has cradle distribution. one accepts view or not, one to assume some low area for Ilex, and this makes it hard to understand that there is a similar very special- izationlevel in the tropical lowland ofthree, now widely seperated, continents. Accepting also all the purely phylogenetic interpretation of trends ia and 2a, would imply that
montane as this does not likely tropical species should be regarded specialized; sound very in other features. With these to me, andfinds no support morphological respect to monta- the that ne species, I mentionin passing that the wood anatomical evidence supports idea than they are more closely related to tropical lowland species to temperate or subtropical vessel members than under species. They have longer temperate species growing com- the absence of parable temperature regimes (see p. 207), and they are charactarized by
faint and in the lowland One spirals or by very infrequent spirals, as tropical species. can, the however, not entirely exclude possibility that some tropical montane species are more than lowland closely related to temperate species to tropical ones. trend for fibre-tracheids be less the and The to specialized in temperate subtropical than species in the tropical species calls for opposite reasoning, viz. that the more spe- cialized species are to be found in the tropics and the more primitive ones in subtropical and temperate regions. This argument, derived from fibre-tracheid specialization is, because the correlations with latitude here much however, not a very strong one, are weaker than for vessel specialization. Moreover, mixtures of advanced and primitive characters are of quite common occurrence in the wood and other parts of higher plants.
For Icacinaceae Bailey and Howard (1941) found fibre specialization to parallel vessel specialization in trees and shrubs, but in scrambling and climbing Icacinaceae the fibre- tracheids or even tracheids were provided with numerous large borderedpits in spite of the very high specialization level of the vessel elements. According to Bailey and Howard this indicated that the specialization ofthe imperforate tracheary elements 'lagged behind'.
this well be reversal of the In my opinion phenomenon can just as explained by a general Ilex similar of reversal have specialization series. In a sort might occurred in temperate shrubs and trees.
that the Ilex In passing, I mention Hu (1967) suggested that primitive species are found the Pacific and she in area (N. Borneo, Philippines, Cochinchina, Formosa, Hawaii); her based this conclusionalso on the primitive woodcharacters of those species. Although the Pacific results on xylem specialization in some species of Ilex agree with mine, her From suggestions cannot be upheld by wood anatomy. the study of more species, pre- sented here, it is clear that equally primitive wood is foundin Central and South America.
On balance, I find the interpretation of the correlation of vessel member length and bars withaltitude number of per perforation plate and latitude, in terms of phylogenetic specialization of the species concerned, not very satisfactory. One may also take another view and assume that the latitudinaland altitudinal correlations foundin several eurytherm
climate genera simply mean that has some sort of effect (long term or short term) on vessel member length and number ofbars. This does not necessarily contradict the phylogenetic but climatic influence factor of interpretation puts emphasis on as a primary importance 220 BLUMEA VOL.XXI, No. 2,1973
also that the wood of modified climate It implies anatomy a eurytherm genus is easily by the and that 'independent' specialization trends, if any, can easily be obscured by dominant
effect of climate. For vessel member length this would nicely agree with Dinwoodie's
that the results on tracheid length in Sitka Spruce (see p. 208). Moreover, it implies with
the Ilex of the wanderings of genus over the globe in the past, reversals general speciali-
zation trends have and that the from may occurred, change a subtropical or temperate to the a tropical climate (through major changes in situ, or through extensions of distribution have in the vessel members and increase in the area ofIlex) may resulted elongation of an number of bars This would then be the view per perforation plate. an argument against
strongly upheld by Bailey (1953) that the general trends ofwood specialization are irrevers- be beliefrather than law. the litera- ible, a statement which I consider to a a biological In
correlationsof climateand woodstructure which ture some recent papers also indicate can hardly be incorporated in the general phylogenetic specialization series. These studies showed mainly refer to influence ofwater supply on wood structure. Carlquist (1970) an
almost perfect correlation of rain fall and vessel member length in comparing several from For Hawaiian species of Euphorbia (ranging on average 206—600/urn). Compositae he foundsimilar correlations between vessel member length and mesic or xeric conditions
(1966). Novruzova (1968) analysed the woody flora in Soviet Azerbaijan and found and 'primitive woods' characterized by scalariform or mixed scalariform simple per- forations to be more in evidence in sites with mesic conditions than in xeric habitats. Her
her analysis of 196 species led to the following conclusions which are relevant for this discussion:
elements. 1. Plants in drought areas have specialized structural the 2. Primitive fibres in conducting system are observed in plants with adequate water
supply. Specialization would depend on atmospheric dryness. shorter. 3. Under drought conditions all fusiform elements become much colonize xeric 4. Quantitative and qualitative changes occur when mesophytes conditions,
or when mesic conditions change into xeric.
with scalariform the number of 5. In specimens of the same species perforation plates, bars decreases with increasing altitude. with the altitudinal and latitudinal Though these results are not directly comparable variation in and Novruzova's data correlations between reported this paper, Carlquist's on water supply and wood structure present interesting parallels. Drought conditions seem the effect to have same on fusiform element length as lower temperatures (as extrapolated from fibres found under both conditions and higher altitude or latitude). Specialized dry at tropical latitudes would, however, suggest a similar effect of drought and of the hot,
climate. mostly very wet, tropical
I fmd it impossible to understand all the implications of the relationships existing be- anatomical features. tween climatic conditions and the predominance of certainwood Yet it is obvious that such relationships exist, and the simplest interpretation would be in
Whether climate acts the terms of climatological effects on wood structure. only on influences in phenotypic variation of wood structure or a long term selection process
cannot be evolution so that functionally best fitted anatomical features are favoured, take assessed with the data available now. I would be inclined to the latter view. There
be and latitude can be little doubt that the characters found to correlated with altitude are
extent characters like genetically fixed to a certain (cf. Dinwoodie, 1963). Moreover, role vessel member length and fibre specialization will certainly play an important in the functions and natural selection of water transport and water storage (Braun, 1970) may of favour certaincombinations of anatomical characters for each major type environment. P. BAAS: Wood anatomy of Ilex 221
All of within the possibilities the genome, still witnessing affinities with relatives in the expression of certain wood anatomical features. be Much remains to done on the correlations of wood structure and environmental conditions. have been cited role ofthe Numerous examples above, suggesting a major en-
Other studies, however, do not indicate relation between and vironment. any environment Hawaiian wood structure. Metrosideros polymorpha growing at widely different altitudes and under different conditions of water did show relation of supply not any appreciable wood structure with altitude or rainfall (Sastrapradja and Lamoureux, 1969). Apparently the variational range following environmental conditions also depends on the plant also group. Novruzova (1968) indicated this by concluding that different genera exhibit different of the degrees variability in water conducting system. Though I have abandonedthe purely phylogenetic interpretation of the altitudinal and latitudinal 'effects' wood this does that and on structure, not imply the results onIlex some for other eurytherm genera have no significance wood phylogeny in general. The view
factors have that climatic aninfluence on characters such as vesselmember length, number ofbars and also in fibre indicate that per perforation, specialization, may the same factors have played a role in directing the general phylogenetic specialization trends ofsecondary
for reversals of these trends. If xylem, leaving possibilities open my results really represent examples of a general rule, this would mean that the tropical lowland is the best environ-
for the if of ment retention, or reversals specialization have occurred, the promotion, of primitive vessel characters. This, however, is contradicted by the majority of wood the lowland floras. mentioned before scalariform structure types in tropical As (p. 2x1) in the and of the dominant perforations are rare tropical lowland, many families are characterized rather short vessel members. The these facts by attempt to bring in harmony with the hypothesis that the tropical lowland favours primitive wood structure would involve fantastic like the of evolution outside the speculations, assumption an hot tropical for those families ofthe environment now constituting the bulk tropical flora. Obviously needed reach sensible more facts are to a more synthesis.
Wood anatomy and the classification ofIlex
Ilex The only comprehensive systematic subdivision of the genus is that proposed by and his Loesener (1901, 1908, 1942), and although system may be 'out of date' because of of much material from the availability more areas underexplored up to his time, there is
Loesener's that his no reason to challenge claim subdivision in subgenera, series, sections, natural and subsections reflects grouping and that the arrangement ofthe infrageneric taxa reflects natural affinity. His system rests on the use of character complexes from inflores-
of the floral cence, merousness whorls, ovary, ovules, and leaves and was based on the
of about c. 280 this therefore follow investigation 2000 specimens (for species). In part I his his Loesener's system adopting nomenclature, including intercalation of a rank 'series' between that of subgenus and section. It will appear that most sections, series, and sub- wood those genera are anatomically very heterogeneous, except infrageneric taxa of habitats lowland which the species I studied are from similar (tropical or mountains,
a few the of subtropical or temperate regions). In only sections common occurrence of and certain anatomical features is independent latitude altitude, and seems to point to natural affinity. On the grounds of this lack of correlation between wood anatomical diversity and Loesener's classification, it should not be concluded that his system is weak.
This would be so if wood anatomically one could propose an alternative classification of anatomical based on a consistent combination wood characters for different infrageneric
221 222 BLUMEA VOL. XXI, No. 2,1973
From the continuous variation in all relevant wood anatomical features groups. as re- the follows that corded in descriptive part it such an alternativesystem cannotbe proposed.
Notes the wood Loesener’s and on anatomy of sections, series, subgenera
Subgenus RYBONIA
material this No was available of the opposite-leaved species of subgenus. According to
Merrill Arb. the wood like that of other Ilex (J. Arn. 20, 1939, 222) anatomy is species.
Subgenus BYRONIA
Series A. EUBYRONIA
Species studied: I. anomala, I. cymosa, I. hypoglauca, I. ledermannii, I. sclerophylloides.
The features and most of the wood anatomical characters quantitative qualitative agree well five I. its scalariform very for these species. anomala stands out by and transitional inter-vessel pits and large vessel—ray pits. Except for I. anomala from the Pacific islands
Tahiti lowland of and Hawaii, all other species are from the tropical Malesia.
Series B. MICROCOCCA
Species studied: I. micrococca. from This Japanese species differs the foregoing series in having all features similar to and of wood the subtropical temperate type anatomy. Subgenus Byronia is wood anatomically heterogeneous. The two wood anatomical types encountered coincide with the two series recognized by Loesener; they also coincide with distribution: restricted the the other geographical one type being to tropics, type to a temperate region.
Subgenus YRBONIA
available of the No material was only species I. teratopsis.
Subgenus EUILEX
Series A. LIOPRINUS
Section Excelsae
Species studied: subsection Umbelliformes: I. chinensis, I. godajam, I. rotunda; subsection
Laxae: I. amplifolia, I. kwantungensis, I. laurifolia, I. macfadyenii, I. nayana, I. sebertii.
This section has wide in both the and and a range tropical, subtropical, temperateNew Old Worlds. Generally the large wood anatomical variation follows the latitudinal
that the from zonation. It may be significant three tropical species the Old World (I. I. I. lack in have godajam, laurifolia, and sebertii) spirals their vessels and only very faint spirals in some of the fibre tips, whilst the New World species from the tropics (I. ampli- folia, I. macfadyenii, and I. nayana) have well developed spirals in the fibre-tracheids and weak the I. amplifolia and I. macfadyenii spirals in vessels as well.
Section Cassinoides
Species studied: I. canariensis, I. cassine, I. coriacea, I. glabra, I. opaca, I. quercetorum. for the all from Except tropical-montane Mexican I. quercetorum, species studied are The anatomical variation subtropical and temperate regions on the northern hemisphere. small number from is fairly accordingly, though the average of bars varies 15—36. Of studied the of well I. quercetorum I only an immature specimen; presence fairly developed in it spirals the vessels of this tropical montane species suggests that may be derived from a subtropical or temperate ancestor. P. BAAS: Wood anatomy ofIlex 223
Series B. PALTORIA
Section Rupicolae
Species studied: I. uniflora. of Ilex This tropical montane species follows the altitudinal trend in showing short low number ofbars the vessel membersand a per perforation plate. Presence of spirals in vessels combined with their frequent absence from the fibre-tracheids is unusual for Ilex.
Section Polyphyllae
Species studied: I. crenata, I. dumosa, I. myricoides. The each other. three species differ widely from I. crenata has all characteristics of a from the to subtropical Ilex also in the tropical montane specimen Philippines. temperate ,
I. dumosastands out like I. uniflora in having more conspicuous spirals in the vessels than in the fibre-tracheids, and I. myricoides has a wood anatomical character complex typical lowland of altitude. Thus of the for tropical species, in spite its high part heterogeneity within this section cannot be explained by the latitudinal and altitudinal trends in Ilex.
Series c. AQUIFOLIUM
Section Lemurensis
Species studied: I. fargesii, I. mitis.
The two species differ widely in quantitative features, following the latitudinal variation in Ilex.
Section Aquifolioides
Species studied: subsection Oxydontae: I. aquifolium, I. cornuta, I. dipyrena, I. integra,
I. perado subsection Insignis: I. insignis, I. latifolia. of this all rather similar The species studied section are in exhibiting typically temperate wall to subtropical wood anatomical features. In all species tangential fibre pits are fairly frequent.
Section Microdontae
Species studied: subsection Eumicrodontae: I. brevicuspis, I. krugiana; subsection Repandae:
I. argentina, I. buergeri, I. cinerea, I. ficoidea, I. formosana, I. liukiuensis, I. paraguariensis, I.
repanda, I. toluccana, I. volkensiana; subsection Vomitoriae: I. caroliniana; subsection Sider-
oxyloides: I. divaricata, I. goshiensis, I. sideroxyloides, I. wilsonii.
has and The Section Microdontae a pantropical pansubtropical distribution. species I
studied are mainly from subtropical regions, and those from the tropics were only re- would presented by immature specimens. Anatomically the wood agrees with what one
except in the subtropical species. I. krugiana stands out, however, by the absence of spirals
from the vessels. The tropical species conform to the latitudinal trend in lacking spirals in
their vessels, and have only faint or few spirals in the fibre-tracheids.
Section Daphnophyllae
Species studied: I. caliana, I. laurina. other conform These two species resemble each very closely, and clearly to the altitu- the dinal effect in tropical montane Ilex. They moreover share large gash-like vessel—ray which rather unusual character Ilex. pits, is a in BLUMEA VOL. No. 224 XXI, 2, 1973
Section Megalae
Species studied: subsection Pedicellatae: I. brasiliensis, I. theezans. values and the of well these In quantitative presence developed spirals two species Ilex conform to what one would expect of subtropical species.
Section Micranthae
Species studied: subsection Punctatae: I. petiolaris; subsection Epunctatae: I. berteroi, I.
guianensis, I. inundata, I. jenmanii, I. umbellata. the lowland. wood shows All species studied are from tropical Accordingly the rather the long vessel members and fairly numerous bars per perforation plate, though values lower than for instance for the lowland of are distinctly tropical species subgenus Byronia. The I. berteroi from Cuba stands out with very few bars and short vessel members. oc-
currence of faint to well developed spirals in the fibre-tracheids is shared by all species.
Section Rugosae
Species studied: I. curranii, I. versteeghii. differ the Both species studied grow in the tropical montane habitat. Yet they in absence of presence (I. curranii) or (I. versteeghii) spirals.
Series D. THYRSOPRINUS
Section Racemosae
Species studied: I. havilandii. of wood This species conforms to the tropical montane type Ilex anatomy.
Section Indico-Malaicae
Species studied: I. cissoidea.
The wood anatomy of this species is similar to that of Ilex species from the lowland instance those tropics, as for belonging to subgenus Byronia.
Section Thyrsiflorae
Species studied: I. casiquiariensis, I. martiniana, I. ovalifolia. The three studied all from the lowland of South and species are tropical America are similar wood also resemble other of the lowland very anatomically. They species tropical and absence (cf. section Indico-Malaicae) with their long vessel members, numerous bars
of spirals. Euilex Subgenus is, in this study, the best represented one: 61 species from all major
variation encountered distribution areas of the genus. Accordingly the wood anatomical
within this is almost within the whole the variation subgenus as large as genus. Frequently altitude related with latitude and is also very considerable within the individual sections of based or even subsections. In some sections one finds an indication systematic affinity for and on wood characters, as instance in sections Daphnophyllae Aquifolioides. Yet, in other sections (Rugosae and Polyphyllae) the woodanatomical diversity is considerable and cannot be entirely correlated with the general latitudinal and altitudinal trends.
Subgenus PRINUS
Series A. EUPRINUS
verticillata. Species studied: I. serrata, I.
The two species resemble each other very closely in their semi-ring-porousness, short vessel members, few bars per perforation, rather narrow rays and the — for temperate Wood Ilex P. BAAS: anatomy of 225
of from the vessel- and fibre walls. This whole character species unique — absence spirals rather isolated withinthe According complex put these species together in a position genus. series closest the North American to Loesener (1901, p. 501) the species of Euprinus are to
The wood of is identical to that ofI. serrata genus Nemopanthus. anatomy Nemopanthus ± verticillata and Loesener's and I. very strongly supports suggestion.
Series B. PRINOIDES
Species studies: I. chapaensis, I. decidua, I. dubia, I. longipes, I. montana. the in The three former species share presence of numerous tangential wall pits the fibre-tracheids, whereas I. montana has infrequent fibre-wall pits. Other characters vary latitudinal trend considerably, partly in relation to the (20° —41 °N).
based wood Series Subgenus Prinus can be split into two groups, as on anatomy.
Euprinus stands out by its absence of spirals, and series Prinoides simply follows latitudinal trends with its fairly faint spirals in the tropical I. chapaensis and the well developed spirals of deciduous in the subtropical to temperatespecies. Subgenus Prinus is composed species, The in contrast to the remaining evergreen subgenera. heterogeneity of this deciduous that anatomical characters and subgenus indicates there is no relationship between wood the in Ilex. deciduous or evergreen habit
Specific wood anatomical descriptions
on Explanatory notes. — The specific descriptions are as short as possible. Data average classes of and maximal numberof vessels per radial multiple, frequency of different size strands radial vessel multiples, abundance of parenchyma, length of parenchyma etc. are of below the not included in the specific descriptions because the great variation species level of the vessel distribution characters, and the time consuming nature of measuring de- parenchyma frequency. For these characters the general statements in the generic scription and in the discussion of the characters must suffice. Abbrevations and very con- densed sentences have been used to shorten the descriptions even further. The general descriptive scheme adopted is as follows: Growth Vessel distribution. rings. Average vessel frequency(ies), average percentage(s)
in oftwo of solitary vessels, tangential vessel diameter (full range and range of means; case the specimens studied both means are given as e.g. 800 & 1000, with more specimens inter-vessel extreme means are connected by a hyphen), vessel member length (ibid.), number of pits and vessel—ray pits (arrangement and size), bars per perforation plate (full and range range of means), helical and annular thickenings (spirals). Fibre-tracheid length
and size helical and (full range and range of means), frequency of tangential wall pits, of broad and annular thickenings (spirals). Average frequencies narrow rays, ray type
(following Kribs), maximum height and width of the broad rays, sheath cells. Crystals. Peculiarities (if any).
The abbreviations used are: alt.=alternate; diam. =tangential diameter; freq.=frequent; het.=heterogeneous; infreq.=infrequent; length=vessel member length; occ.=occasion- ally; opp.=opposite; scal.=scalariform; sol.=solitary; spirals=helical or annular thicke-
—r nings of the cell wall; tang. =tangential; unil. comp.=unilaterally compound; v pits= vessel—ray pits; v—v pits=inter-vessel pits. in that the total Ray frequency is expressed two figures, e.g. 3+8/mm. This means BLUMEA 226 VOL. XXI, No. 2, 1973
of multiseriate and number rays per millimetre is 11, and that 3 of these are 8 are uniseriate.
the For citation of specimens and meaning of the terms mature and immature, see p. 195
and 196.
Ilex amplifolia Rusby (subg. Euilex, series A, sect. Excelsae, subsect. Laxae)
Latitude in graph 16°S.
Diffuse faint. 2 porous. Growth rings Vessels 43/mm 47% sol., diam. —67—88 , 42 /rm, similar and length 1130—-1720—2270 /um, v—v pits trans, to opp., 7 —10/tm, v—r pits sometimes unil. bars 26 faint comp., —38—52, spirals very to hardly present. Fibre-
tracheids wall 6 well 1260—2500—3420 fim, tang, pits infreq., fim, spirals developed. Rays sheath Sol. 1 (2)+io/mm, het. II, up to 4.1 mm high, to 11 cells wide, cells occ. present.
in crystals ordinary ray cells.
Material studied. BOLIVIA. Krukoff 11198 (SJRw 39912).
of the this Note. The presence well developed spirals in fibre-tracheids in species, which shows the typically tropical character complex in its other wood anatomical fea- and tures, is noteworthy curious.
Ilex anomala Hook. & Arn., inch f. sandwicensis (Endl.) Loes. and f. taitensis (Gray)
Nadaud — Plate (subg. Byronia, series A. Eubyronia) III, 3. and Latitude I7°S 19—22 °N; in graph 21°.
2 Diffuse Growth rather indistinct to absent. Vessels (27—5o)/mm porous. rings 33—48 , diam. 31—64 (28 —62)% sol., 29 —50—76 (55 —68) —97fim, length 1050—1570 —1690 —2160 vestured in (960—1350) fim, v—v pits mainly seal., seemingly some specimens,
to —r similar and and sometimes unil. bars ranging opp., v pits large gash-like, comp., absent. Fibre-tracheids 19—28—35 (29—37) —51, spirals 1740—2i8o—2580 (1450 —1950)
wall infreq. in mature freq. in immature —3230 jum, tang, pits specimens, specimens,
— — absent. het. het. 3 5 (4 6) fim, spirals Rays I—2+8—io/mm, II to I, up to 3.5 mm high, to 8 cells wide, sheath cells often present. Sol. crystals, sometimes slightly elongated,
in cells in 8 of present ordinary ray out 9 specimens studied.
HAWAII al. Material studied. (f.I sandwicensis). (BISHw 24); Bryan Jr et s.n. (BISHw 2758*), alt. 1040 m;
116 MacDaniels 161 MacDaniels 84 (SJRw 2494}*), alt. 750 m; MacDaniels (SJRw 2497 7*), alt. 50 m;
— TAHITI taiten- (SJRw 25019*), alt. 1300 m; Rock & Homer s.n. (BISHw 46); Thrum s.n. (BISHw 140). (f. sis).. Van Balgooy 1689*, alt. 850 m; MacDaniels 653 (SJRw 43839), alt. 1300 m.
this Note. In its qualitative and most of its quantitative wood anatomical features species is rather constant. The scalariform v—v pits and large, gash-like, v—r pits are striking features.
Ilex aquifolium L. (subg. Euilex, series C, sect. Aquifolioides, subsect. Oxydontae) -
Plate IV, 1 & 2.
Latitude 39—54°N, in graph 48 °N.
Growth 2 Diffuse to rings distinct. Vessels 128—192/mm 8—23% sol., semi-ring porous. , diam. trans, to 17—31—38—51 fim, length 520—690—890—1130/tm, v—v pits opp., bars rarely also seal, or alt., v—r pits opp. to alt., very rarely unil. comp., 12—19—21—32, well Fihre-tracheids wall spirals developed. 490—990—1320—1840/um, tang, pits freq.,
well het. to mm 4 —6 jum, spirals developed. Rays 3—4+7—9/mm, II, up 0.9 —2.1 high, to cells sheath cells absent. absent. 5 —7 wide, Crystals P. BAAS: Wood anatomy of Ilex 227
Material studied. GREECE. (MADW 17874). — BELGIUM. (IF 14473). NETHERLANDS, cult. UN 163 (Uw) low alt.
Note. The three samples studied are very similar.
Ilex argentina Lillo, An. Soc. Cient. Argent. 72 (1910) 171 (subg. Euilex, series C, sect. Microdontae, subsect. Repandae)
Latitude c. 27 °S.
2 Diffuse Growth distinct. Vessels 69/mm 26% sol., diam. 38—56—76 porous. rings , jum, similarand length 670—920 —1190 [im, v—v pits trans, to opp., 6—9fim, v—r pits some- times unil. comp., bars 15 —21—34, spirals well developed. Fibre-tracheids II00—1740 —
wall 6 well het. 2290 /urn, tang, pits infreq., [Am, spirals developed. Rays 3+8/nun, II, abundant. in up to 2.0 mm high, to 10 cells wide, sheath cells Sol. crystals infreq. ordinary cells. ray
Material studied. ARGENTINA. (Molfino s.n., presented by Prince of Wales no 392 toFHOw), alt. c. 1200m.
Note. According to Lillo I.e. this species is closely related to I. paraguariensis. I have therefore included it in subsection Repandae.
Ilex berteroi subsect. Loes. ex. Urb. (subg. Euilex, series C, sect. Micranthae, Epunctatae)
Latitude c. 21 °N.
Growth Vessels 2 Diffuse porous. rings faint. 29/mm 52% sol., diam. 38—55 —71 , /urn,
to 6—8 seal, to alt. and length 520—880—1210 fim, v—v pits opp. alt., //m, v—r pits some- Fibre-tracheids times unil. comp., bars 8—15—21, spirals absent. 890—1370—!7oo /urn, wall well het. tang, pits freq., c. 5 jum, spirals fairly developed. Rays 3 + 8/mm, II, up to
1.8 cells Sol. in cells. mm high, to 10 cells wide, sheath present. crystals ordinary ray
Material studied. CUBA. Bucher s.n. (SJRw 19290*).
Note. The number of bars per perforation plate is remarkably low in this tropical species. This can only partly be due to the fart that the specimen studied was immature.
Pedi- Ilex brasiliensis (Spreng.) Loes. (subg. Euilex, series C, sect. Megalae, subsect. cellatae)
Latitude c. 25 °S.
Diffuse Growth rings distinct. Vessels 2,13% sol., diam. —46 porous. 104/mm 25 —59 /um, length —820—1180 v—v to alt., v—r similar and occ. unil. 470 fum, pits opp. 7—9 /um, pits bars well Fibre-tracheids comp., 15—27—43, spirals developed. io8o—1320—1500 ,«111, wall rather well het. tang, pits freq., 6 fim, spirals developed. Rays I—2+9/mm, II, up cells sheath cells in to 1.1 mm high, to 10 wide, rarely present. Sol. crystals ordinary and chambered cells. erect ray
Material studied. BRAZIL. Parana: Lindeman & Hecking 873 (Uw 12773*).
Ilex brevicuspis Reiss (subg. Euilex, series C, sect. Microdontae, subsect. Eumicrodontae)
Latitude c. 28 °S.
2 Diffuse to semi-ring Growth rings distinct. Vessels 105/mm 10% sol., diam. porous. ,
25 —42—59 fim, length 520—800 —1130 fim, v—v pits opp. to alt., 6—9 /im, v—r pits similar unil. bars and sometimes comp., 13—19—30, spirals well developed. Fibre- tracheids wall rather well 740—1100—1370[im, tang, pits infreq., c. 6 /tm, spirals developed. 228 BLUMEA VOL. XXI, No. 2, 1973
het. to cells sheath cells Sol. Rays 2+10/mm, II, up 2.6 mm high, to 14 wide, rare. crystals in ordinary and chambered erect ray cells.
Material studied. BRAZIL. Santa Catharina: Herb. Barb. Rodr. Sant. Cat. 31568*, high alt.
Ilex buergeri Miq. (subg. Euilex, series C, sect. Microdontae, subsect. Repandae) Latitude e in graph 41 N.
Growth Vessels 2 diam. Diffuse porous. rings distinct. 140/mm 30% sol., , 29—42—59fim,
—1120 to 6—8 similar and length 510 —1470/tin, v—v pits opp. alt., jum, v—r pits some-
times unil. bars 18 well Fihre-tracheids — comp., —30—53, spirals developed. 1050 —1970 wall rather well 2550 //m, tang, pits infreq., 5 —7 jam, spirals developed. Rays 3+8/mm, het. cells sheath Sol. idio- II, up to 1.9 mm high, to 8 wide, cells occ. present. crystals in blastic thick-walled and cells. bulging square ray
Material studied. = Govt. Sta. JAPAN. (IF 23288 Jap. Exp. 756, Tervuren).
Ilex caliana Cuatr., n sect. Lloydia (1948) 213 (subg. Euilex, series C, Daphnophyllae) Latitude in graph 4° N.
Growth 2 Diffuse rings faint. Vessels 21/mm 33% sol., diam. —71 porous. , 42 —92 fim,
v—v 6—8 pits similar and unil. length 550—980—1540 fim, pits opp. [im, v—r occ. also and bars absent. Fihre-tracheids comp., tending to large gash-like, 18—23—34, spirals wall absent. 1500—1860—2370fim, tang, pits infreq., 3 —4 fim, spirals Rays 3 +8 /mm, het. Sol. II, up to 2.5 mm high, to 9 cells wide, sheath cells occ. present. crystals abundant cells. in ordinary ray
Cuatrecasas 2160 Material studied. COLOMBIA. 18281, type (SJRw 43277*), alt. m.
Note. This species is closest to I. laurina according to Cuatrecasas, and is therefore
included in section Daphnophyllae here.
The vessel and bars low quantitative values for member length number of per per- of the foration are probably due to both the immature nature specimen and the high
altitude of its habitat.
Ilex — Plate canariensis Poir. (subg. Euilex, series A, sect. Cassinoides ) VI, 6.
Latitude c. 28° N.
2 Diffuse Growth rather Vessels & 45/mm & 63% sol., diam. porous. rings vague. 43 , 47
29—50 & 59—80 fim, length 620—1030 & 1190—1500 /uni, v—v pits trans, to opp.,
similar and sometimes unil. bars 10 & 16 6—10 fim, v—r pits comp., —15 —22, spirals
well developed. Fibre-tracheids —1600 & 2000 wall 6 1130 —2500fim, tang, pits freq., fim,
well absent fibres. & & het. to spirals developed, in some Rays 2 5 + 5 6/mm, II, up
mm to cells sheath cells remainderof cells also bigger 1.5 & 1.7 high, 8 wide, present, ray and clustered than usual. Sol. irregularly crystals in ordinary ray cells.
Material studied. CANARY ISLANDS. Baas & Ridsdale 8, alt. 600 m; Bourgeau s.n. (K-Jw).
and cells Note. The rather vague growth ring boundaries the unusually large ray are
noteworthy in this subtropical species. I also studied an unvoucheredspecimen (IF 18236, and Tervuren) which does not show these peculiarities, moreover differs in quantitative
2 features: Vessels 105/mm 29% sol., v—v pits to alt., 6—7 bars 20—28—36. , opp. fim,
Rays 3+9/mm.This specimen fully agrees with the two specimens ofI. perado I studied,
also from the Canaries. I suspect therefore that IF 18236 is misnamed.
The fart that two species growing together in the same islands in the same habitat show P. BAAS: Wood anatomy of Ilex 229
differences their wood remarkable for where dif- rather clear in anatomy is Ilex, usually similar habitats ferences between unrelated species growing in are very slight or fully absent.
Ilex caroliniana (Lam). Loes.; I. vomitoria Ait. (subg. Euilex, series C, sect. Microdontae,
subsect. Vomitoriae) — Plate II, 4 & 12; HI, 9; VI, 2.
in N. Latitude 15—34° N, graph 30°
Diffuse Growth rings distinct. Vessels 2 sol., diam. porous. 135—22o/mm , 9—16%
—800 21—34—42—50 fim, length 410 —910—1130 fim, v—v pits opp., rarely tending to —8 v occ. unil. bars spirals well alt., 6 fim, —r pits similar, comp., 9—14—16—21, Fihre-tracheids wall developed. 740—n6o—1490—1840 fil11, tang, pits fairly infreq.,
— well het. to 1.6 111m 5 6 /mi, spirals developed. Rays 2 —4+!0—12/mm, II, up —2.7 high, of smallest to 15 cells wide, sheath cells abundant in specimens diameter, rarely present in Sol. cells only. other two specimens. crystals in ordinary ray of MADw 731
Material studied. MEXICO. Chiapas: Page 9643 (MADw 23880*). — U.S.A. Texas: (IF 18236*); (MADw 751)-
Note. The quantitative values for the mature and immature specimens do not differ The from is which widely. tropical specimen Chiapas probably from a montane locality, low values for vessel member and number of bars would explain the length per per- foration plate.
Ilex casiquiariensis Loes. (subg. Euilex, series D, sect. Thyrsiflorae)
Latitude in graph 6° N.
2 Diffuse Growth rings faint. Vessels 64/mm 19% sol., diam. 38—67—80 porous. , fim, seal, similar and length iooo—1260—1700 /urn, v—v pits to opp., 6—8 jum, v—r pits
unil. bars absent. Fibre-tracheids — sometimes comp., 29—40—67, spirals 1390—1840
wall absent. Rays het. to 237° tang, pits infreq., 4 fim, spirals 2+14/mm, II, up 2.5 mm cells sheath cells absent. high, to 8 wide, occ. present. Crystals
Material studied. VENEZUELA. Amazonas: Wurdack & Adderley ? 43197 (SJRw 54122).
Ilex cassine L.; I. myrtifolia Walt. (subg. Euilex, series A, sect. Cassinoides)
Latitude in graph 30° N.
Diffuse Growth distinct. Vessels 62 & 8o/mm2 18 & sol., diam. porous. rings , 20%
& & 21—44 50—71 /mi, length 640—910 1020—1290 fim, v—v pits opp. to alt., 5 —16fim,
—r unil. bars & well v pits similar, rarely comp., 12—24 29—40, spirals developed.
Fibre-tracheids & wall 840—1320 1550—1950 fim, tang, pits infreq. in MADw 2634,
—6 well freq. in MADw 11284, 5 /urn, spirals developed. Rays 3 & 4+8 & 9/mtn, het. II,
het. & to 8 cells sheath cells or II+III (MADw 2634), up to 1.7 1-9 nun high, wide, absent. Crystals absent.
Marts Material studied. U.S.A. Florida: (MADw 2634); 4 (MADw 11284).
Note. in in the fibres in The differences tangential pit frequency and ray histology the between the two specimens, raise the question whether one of samples is misnamed, whether is indeed with or I. cassine (MADw 2634) synonymous I. myrtifolia (MADw 11284). BLUMEA VOL. No. 230 XXI, 2, 1973
Ilex —Plate chapaensis Merr., J. Arn. Arb. 21 (1941) 373 (subg. Prinus,, series B, Prinoides)
III, I & 2; IV, 3-
Latitude c. 20° N.
2 Diffuse Growth faint. Vessels 46/nun sol., diam. — porous. rings very , 28% 50—84
v—v to similar 101 fim, length 880—1380—1660 fim, pits opp. alt., 6—8 fim, v—r pits
and sometimes unil. bars 20 spirals faint, warts noted with SEM. comp., —31 —38 fim., Fibre-tracheids wall rather well !39°—2550—2890[xm, tang, pits freq., 5 jum, spirals fairly
het. to mm to cells sheath cells developed. Rays 2+7/mm, II, up 3.0 high, 7 wide, infreq. Sol. in cells with much thickened walls. crystals enlarged erect ray
Material studied. NORTH VIETNAM. Chevalier 30129 (CTFT 11009, GTFw).
this Note. According to Merrill I.e. species is closets to I. henryi, which Loesener
included in series Prinoides. Therefore I. chapaensis is included in this section here.
Merrill's is from alt. Our trends' in type 1500 m specimen clearly shows some 'tropical vessels member and number of the of length bars, although presence spiral thickenings is line This more in with a subtropical habitat. agrees well with latitudinal and altitudinal distribution of this species.
The small warts on the vessel the inside of the borders are present walls, including
overarching the half-bordered vessel—parenchyma pits (Plate III, 1 & 2).
Ilex chinensis Sims; I. purpurea Hassk. (subg. Euilex, series A, sect. Excelsae, subsect.
Umbelliformes) — Plate II, 9; V, 2.
° Latitude in graph 41 N.
2 Diffuse Growth rings distinct. Vessels 29% sol., diam. porous. 72/mm , 39 —42—55 fim,
pits seal, to —r similar and length 670—990—1490/tm, v—v opp., opp. pits 7—9 jum, v pits often unil. bars well Fibre-tracheids complex comp., 10—14—17, spirals developed. wall well 1000—1760—2240 fim, tang, pits freq., 6—7 fim, spirals developed. Rays in 3 +8/mm, het. II, up to 1.3 mm high, to 8 cells wide, sheath cells infreq. Sol. crystals thick-walled idioblastic often erect ray cells, elongated.
Govt. Sta. Material studied. JAPAN. (IF 17637,Tervuren = Jap. Exp. 2254).
Arb. the reduction of Note. Hu (J. Arn. 30, 1949, 299) proposed I. purpurea to I. chinensis.
Ilex — cinerea Champ, (subg. Euilex, series C, sect. Microdontae, subsect. Repandae)
Plate V, I.
Latitude c. 22° N.
2 Diffuse Growth rings distinct. Vessels 78/mm 50% sol., diam. 38—59—63 porous. , fim,
to 6 —r unil. length 560—880—1280 fim, v—v pits opp. alt., jum, v pits similar, rarely
bars well Fibre-tracheids 1000 —2100 comp., 19—26—33, spirals developed. —1470 fim, wall well het. fairly freq., c. spirals Rays 2+7/mm, II, tang, pits 5 fim, developed. up cells to 1.9 mm high, to 15 wide, sheath cells occ. present. Sol. crystals infreq. in ordinary
ray cells.
HONGKONG. Material studied. Ford s.n. (K-Jw).
Ilex cissoidea Loes. (subg. Euilex, series D, sect. Indico-Malaicae)
Latitude in graph o°.
Diffuse Growth fairly distinct (one in sample studied). Vessels 14/mm 2 porous. ring , 30%
diam. —2160 to 6 sol., 50—92—120 fim, length 1290—1670 fim, v—r pits opp. alt., fim, P. BAAS: Wood anatomy of Ilex 231
v—r pits similar and often unil. comp., bars 22—33 —44, spirals absent. Fibre-tracheids
1760—2550—3160/im, tang,wall pits very infreq., —5 spirals absent. Rays 2+7/mm, 3 /im, . het. cells II, up to 2.0 mm high, to 7 wide, sheath cells occ. present. Crystals absent.
INDONESIA, Material studied. bb. 2119 (RTIw 2564).
The of distinct this Note. presence a growth ring in typically tropical representative
is remarkable. The small size of the wood sample studied leaves the question unanswered whether formed growth rings are periodically or whether the ring boundary in our unusual specimen is due to an dry spell.
Ilex coriacea (Pursh.) Chapm. (subg. Euilex, series A, sect. Cassinoides) Latitude 0 c. 32 N. distinct. Vessels 2 diam. Diffuse Growth rings 140/mm sol., 21 porous. , 15% —34—50jum,
—1060 seal, and length 430—850 /urn, v—v pits to opp., v—r pits opp. sometimes unil.
6 bars well Fibre-tracheids — comp., c. /urn, 17—35—48, spirals developed. 710—1100
wall —6 well het. 1550 fim, tang, pits freq., 5 /«m, spirals developed. Rays i+7/mm, II,
mm cells up to 2.5 high, to 15 wide, sheath cells very rare. Crystals absent.
Material studied. U.S.A. Florida: MADw 4567*.
wood Note. The number ofbars per perforation plate is very high for this immature sample offairly high latitude.
Ilex cornuta Lindl. & Paxt. (subg. Euilex, series C, sect. Aquifolioides, subsect. Oxydontae)
Latitude c. 30° N.
Diffuse Vessels 61 2 & porous. Growth rings distinct. & 97/mm 29 44% sol., diam. ,
& & seal, to 17—30 35 —50 fim, length 420—750 850—1130 fim, v—v pits opp., v—r unil. bars pits opp., rarely comp., 13—19 & 21—29, spirals well developed. Fibre-tra-
cheids 6oo & 1660—2180 wall 6 well devel- —1360 //m, tang, pits fairly freq., //m, spirals
& & & 8 cells oped. Rays 4 5+6 y/mm, het. II, up to 1.2 1.9 mm high, to wide, sheath Sol. cells occ. present. crystals in chambered erect and ordinary ray cells.
Material studied. U.S.A. Florida: NYSCF 8701 (MADw 1108); (IF 22715*).
Note. The immature specimen does not appreciably deviate from the mature wood
sample. I. cornuta is native in China.
Ilex inch f. crenata Thunb., var. typica Loes. luzonica (Rolfe) Loes. (subg. Euilex,
series B, sect. Polyphyllae)
0 0
Latitude — both under and 17 52 N, in graph plotted 17° 52 .
2 Diffuse to semi-ring Growth rings distinct. Vessels 54—82/mm 21—60%, sol., porous. ,
diam. trans, to 21—31—41 —55 /im, length 300 —620—900 —1130 jum, v—v pits opp.,
similar and unil. bars —28 well occ. seal., v—r pits comp., 13—17 —43, spirals developed.
Fibre-tracheids wall rather —8 well 790—11oo—2050 /2m, tang, pits freq., 5 jum, spirals
0.8—2.0 cells developed. Rays 2—4+7—9/mm > het. II, up to mm high, to io wide,
sheath cells occ. present. Crystals absent.
NETHERLANDS, UN — Material studied. cult. 215 (Uw), low alt. CHINA. Makai: (IF 23290, Tervuren),
montane. — PHILIPPINES (var. typica f. luzonica). Mountain Province: (BF 18040,CLPw), high alt.
N ote. The threecollections studied allowfor a comparison ofsamples from widely differ- ent latitudes and altitudes.The montane sample fromthe Philippines exhibits all characteris- BLUMEA No. 232 VOL. XXI, 2, 1973
tics of of Ilex. The from the Netherlands a typically temperate representative specimen
shows the lowest values for number of bars per perforation plate and vessel member
The wide in of sol. vessels is The fact that all length. range % noteworthy. samples the exhibit the same type ofintervascular pitting supports the view that material studied here all belongs to one species.
Ilex curranii Merr., Philip. J. Sci. 17 (1920) 273 (subg. Euilex, series C, sect. Rugosae)
Latitude 17° N.
2
diam. — Diffuse Growth rings fairly distinct. Vessels 69/mm 41% sol., 25—49 porous. , similar 63 fim, length 460—1030—2060 jum, v—v pits opp. to alt., 5 —7 fim, v—r pits and sometimes unil. bars with rather well Fibre- comp., 18—34—50, developed spirals.
tracheids —i8oo wall infreq., —6 faint 1240 —2300 fim, tang, pits 5 fxm, spirals only present
of the fibres. het. to cells sheath in some Rays 2+9/mm, II, up to 2.1 mm high, 10 wide, Sol. in cells. cells occ. present. crystals ordinary ray
Material studied. PHILIPPINES. Luzon, Mountain Province: Stem & Rojo 2290*, high alt.
This in The low member Note. species grows the mossy forest. values for vessel length
and number of bars per perforation are in line with the montane habitat and immature
this well the nature of specimen. The presence of developed spirals in vessels, but less so
in the fibres is noteworthy.
Ilex cymosa Bl.; I. bogoriensis Loes., I. pleiobrachiata Loes. (subg. Byronia, series A.
Eubyronia) — Plate I, 6; II, 8; III, 7; V, 5.
Latitude io° S—12° N, in graph o°.
Diffuse Growth distinct indistinct absent in porous. rings only in CSIRO 4993, or remainder. mm2 Vessels 12—3°/ 17—66% s°l-» diam. 42—76 length > —113—139 /um,
740—1040—1920—2630 /um, v—v pits opp., occ. tending to alt., 5 —6 /um in CSIRO and Elmer in vestured 4993 12806, 7—9 ,um remainder, seemingly in some specimens,
v—r pits opp., occ. unil. comp., bars 19—36—58—101, spirals absent. Fibre-tracheids wall rather freq., in CSIRO 1050—1630—2710—3160 fim, tang, pits 3 —4 fim 4993, and Elmer —6 absent all KEPw 2337 12806, 5 um in remainder, spirals in samples except for faint ones in some Ridley s.n. Rays i—4+6—9/mm, het. II, up to 1.5—5.0 mm high, cells sheath cells few Sol. chambered to 25 wide, only present in specimens. crystals in erect ray cells.
Material studied. PHILIPPINES. Palawan: Elmer 12806. — MALAYA. H 3792 (CSIRO 4993, FPAw); (KEPw
— — alt. 2337). SINGAPORE. Ridley s.n.(K-Jw 39. 1901). BORNEO. Sarawak: S 9789, alt. 3 m; S 13480, ioo m.
— INDONESIA. bb R New Guinea: BW 8494 (Ind. 8082, RTIw). Java: (SFCw 373—12). 339, alt. 100 m; BW BW 611, alt. 50 m; 1067, alt. 10 m.
The material examined shows characters Note. a great deal of variation in quantitative of and The lowest values for vessels, pits, rays. vessel member length were recorded in Elmer and 12806, CSIRO 4993, S 9789.
Ilex deciduaWalt. — (subg. Prinus, series B. Prinoides) Plate I, 3. Latitude c. 33 °N.
2 Diffuse to semi-ring Growth distinct. Vessels 88 & 113/mm & porous. rings , 52 56% sol., diam. 21 length & to —38—50/mi, 440 —780 860—i2io/ Fibre-tracheids —1180 & wall well 870 1260—1840/im, tang, pits freq., 6—7 fim, spirals P. BAAS : Wood anatomy of Ilex 233 & to to 12 cells sheath cells developed. Rays 3 + 10 13/mm, up 1.4 mm high, 7 & wide, in absent. infreq. Sol. crystals infreq. ordinary ray cells or Material studied. U.S.A. North Carolina: E.S. Harrar NYSCF 8575 (MADw 849), Texas: Nogle-Wilson T-40 (MADw 17611). similar wood I could Note. The two specimens are very anatomically. Although not of the the determine the exact diameter stems from which small specimens were taken, the the curving of growth rings suggested almost or just mature wood. Ilex dipyrena Wall. (subg. Euilex, series C, sect. Aquifolioides, subsect. Oxydontae) Latitude 31 °N. 2 diam. Semi-ring Growth rings distinct. Vessels 130/mm 21% sol., 21 porous. , —34—54 unil. length 680—830—1060 v—v to alt., v—r similar, rarely [im, fxm, pits opp. 5 fim, pits bars well Fibre-tracheids comp., 10—18—26, spirals developed. 920—1420—1970fim, wall well het. tang, pits fairly freq., 3 —4 fim, spirals developed. Rays 3+7/mm, II, up to cells 1.3 mm high, to 7 wide, sheath cells abundant. Crystals absent. INDIA. For. Material studied. Natigana Simla (Ind. Dept. 1878, K-Jw), 2300 m alt. Ilex divaricata Mart. (subg. Euilex, series C, sect. Microdontae, subsect. Sideroxyloides) Latitude c. 4°N. Diffuse Growth rings faint. Vessels 55/mm 2 sol., diam. —67 porous. , 18% 34—55 fim, 680 to pits similar and length —1050—1440fim, v—v pits opp. alt., 6—7 fim, v—r some- unil. bars absent. Fibre-tracheids times comp., 16—32—50, spirals 1000—1470—1710fim, rather wall pits infreq., —5 spirals very faint. Rays 2+8/mm, het. II, up to tang,„ . 4 fim, ...... Sol. in 2.6 mm high, to 10 cells wide, sheath cells occ. present. crystals ordinary and chambered cells. erect ray Material studied. VENEZUELA. Amazonas: Wurdack & Alderly (SJRw 7448*), low alt. The for vessel member here due Note. low value average length is probably entirely the the in to immaturenature of specimen (diam. 1.5 cm). The presence ofweak spirals the fibres is noteworthy in this species from the hot tropics. Ilex dubia (Don) Trel. var. macropoda (Miq). Loes. (subg. Prinus, series B. Prinoides) Latitude 36 °N. 2 Diffuse to faintly semi-ring Growth rings distinct. Vessels i86/mm 17% sol., porous. , diam. 21 length —1080 v—v —6 v—r —46—71 fim, 450—790 fim, pits opp., 5 fim, pits similar and unil. bars well Fibre- sometimes comp., 21—32—46, spirals developed. wall rather well tracheids 1000—1310—1630 fim, tang, pits freq., 5 fim, spirals developed. het. to to cells sheath cells Sol. Rays 3+7/mm, II, up 2.5 mm high, 11 wide, occ. present. crystals (occ. slightly elongated) in erect marginal and sheath cells. Material studied. JAPAN. Mt. Fujioka: (MADw 8135). Ilex dumosa Reiss. (subg. Euilex, series B, sect. Polyphyllae) Latitude c. 25 °S. 2 Diffuse Growth rings distinct. Vessels 138/mm 25% sol., diam. porous. , 34—44—59 fim, similar — 6 — and sometimes uni- length 670—910—1080fim, v v pits opp., fim, v r pits bars well Fibre-tracheids — laterally compound, 15—21—25, spirals developed. 1240—1710 No. 234 BLUMEA VOL. XXI, 2, 1973 wall 2ioo pits infreq., spirals absent or faint in fibre 2+io/mm, //m, tang, 5 /zm, tips. Rays to mm to cells sheath cells abundant. absent. het. II, up 2.3 high, 14 wide, Crystals Material studied. BRAZIL. Parana: Lindeman & Hecking 1916 (Uu> 13412). This ilustr. Note. species usually grows at fairly low altitudes (cf. Reitz, Flora Cata- The absence of in the of the fibre tracheids is rinense 1, Aquifol., 1967). spirals majority remarkable. Franch. — Plate Ilex fargesii (subg. Euilex, series C, sect. Lemurensis) I, I. Latitude 52 °N. 2 Diffuse Growth rings distinct. Vessels 99/mm 9% sol., diam. —38—50 porous. > 29 jum, — —r similarand sometimes length 410—660— 1040 fim, v vpits opp. to alt., 6—7/um, v pits unil. bars well Fibre-tracheids comp., 15—19—33, spirals developed. 840—1450—1790 Hm, wall well Rays het. II, to tang, pits fairly freq., 5 /tm, spirals developed. 3 +8/mm, up sheath Sol. in cells. i mm high, to 6 cells wide, cells rare. crystals very infreq. ordinary ray Material studied. NETHERLANDS, cult. UN 162 (Uw). Note. I. fargesii is native in China. Ilex ficoidea Hemsl. (subg. Euilex, series C, sect. Microdontae, subsect. Repandae) Latitude in graph 25 °N. 2 Growth distinct. Vessels sol., diam. — Semi-ring porous. rings 84/mm , 33% 29—54 similar 6774m, length 870—1160—14407401, v—v pits opp. to alt., 674m, v—r pits and unil. bars well Fibre-tracheids — — occ. conip., 20—29—40, spirals developed. 139° I8ÇK> wall well het. 2370 fim, tang, pits infreq., 4—5 /im, spirals developed. Rays 2+6/nim, II, cells cells almost absent. Sol. in up to 2.8 mm high, to 12 wide, sheath crystals chambered and cells. erect ordinary ray Material studied. CHINA. (SJRW 21930). Ilex formosana Maxim, (subg. Euilex, series C, sect. Microdontae, subsect. Repandae) Latitude c. 24 °N. 2 Diffuse Growth rather faint. Vessels sol., diam. — porous. rings 74/mm , 40% 29—55 80 6—8 fim, length 630—1100—1550/tm, v—v pits opp. to alt., //m, v—r pits similar, rarely unil. comp., bars 22—28—39, spirals well developed. Fibre-tracheids ïï8O—2000 wall rather well —2840 /(m, tang, pits infreq., c. 6 fim, spirals fairly developed. Rays het. to to 10 cells sheath cells abundant. Sol. 3 +5/mm, II, up 2.6 mm high, wide, crystals in and cells often with thickened ordinary chambered erect ray cells, crystalliferous bulging walls. Material studied. TAIWAN. (MADw 3266). Ilex glabra (L.) Gray (subg. Euilex, series A, sect. Cassinoides) Latitude & 36 52 °N. 2 Diffuse Growth rings distinct. Vessels 124 & 135/mm & 34% sol., diam. porous. , 29 & —1180 seal, —29 & 34 —42 m length 500—790 830 v—v pits to opp., v—r pits 17 A< > //m, & Fibre- similar and sometimes unil. comp., bars 22—34 36—46, spirals well developed. tracheids & 1000 wall pits well developed. 600—950 —1390/zm, tang, freq., 5 /zm, spirals cells sheath cells Rays 2 & 3 + II & i8/mm, het. II, up to 1.3 & 1.7 mm high, to 14 wide, cells of 2628. absent. Sol. and clustered crystals in ordinary ray MADw BAAS P. : Wood anatomy ofIlex 235 Material studied. U.S.A. North Carolina: MADw 4567*. — NETHERLANDS, cult. UN 374* (Uw). Note. As in I. coriacea, the high number of bars is noteworthy for these immature temperate specimens. Ilex godajam Colebr. (subg. Euilex, series A, sect. Excelsae, subsect. Umbelliformes) Latitude c. 22 °N. 2 diam. Diffuse Growth rings absent. Vessels 15/mm 60% sol., —122 porous. , 59—90 //m, to similar and often length 1130—1620—2370fim, v—v pits opp. alt., 6—7/um, v—r pits unil. comp., bars 23—30—37, spirals absent, warts noted with SEM. Fihre-tracheids wall absent faint 2240—2950 —342° /«ni, tang, pits infreq., 3 —4 [im, spirals or very in the het. cells sheath some of fibre tips. Rays 3+6/mm, II, up to 2.1 mm high, to 7 wide, cells absent. present. Crystals EAST PAKISTAN. & Material studied. Majumdar Islam 35 (MADw 24486). Note. The warts on some of the vessel walls are similar to thoseof I. chapaensis. The number of species studied with SEM is too low to know whether we deal here with a feature and whether a of unique for I. godajam I. chapaensis, of warty layer is more com- mon occurrence in Ilex. Ilex goshiensis Hayata, Mater. Flora Formosa (1911) 54; I. hanceana sensu auct. Jap. non Maxim, (subg. Euilex, series C, sect. Microdontae, subsect. Sideroxyloides) — Plate VI, 7. Latitude c. 36 °N. 2 Diffuse Growth distinct. Vessels & 82/mm 21 & sol., diam. porous. rings 73 , 25% & & to alt., 29 —46 59 —76/mi, length 720—1040 1130—1530/tm, v—v pits opp. bars v—r similar, not unil. & spirals well developed. 6—8 fim, pits comp., 15—23 29—jo, Fihre-tracheids & wall well 950—1760 197°—2210//m, tang, pits infreq., 5 —6jum, spirals & het. & cells sheath developed. Rays 2+6 7/mm, II, up to 1.6 2.2 mm high, to 12 wide, cells Sol. idioblastic thick-walled chambered erect occ. present. crystals in or ray cells, or absent. Material studied. JAPAN. (IF 23291); Mt. Fujioka, (MADw 8333). Note. According to Hayata I.e., this species is closely related to I. championii which belongs to subsection Sideroxyloides. I therefore treat I. goshiensis in this subsection. Ilex O. I. guianensis (Aubl.) Ktze, inch var. macoucoua (Pers.) Loes.; panamensis Standi, (subg. Euilex, series C, sect. Micranthae, subsect. Epunctatae) 8 Latitude 2 —15 °N, in graph °N. & & 2 & Diffuse Growth rings faint. Vessels 20 50 (49 57)/mm 30 (30 39)% sol., porous. , diam. & 86 & & & 42 —54 (32 56) —122 fim, length 740—1220 1440 (840 1420) —2210 trans, to similarand sometimes unil. bars 11—21 & /j,m, v—v pits opp., v—r pits comp., & absent. Fihre-tracheids & 30 (26 32) —45, spirals 1450—2050 2520(1530 &2000)—3100 im wall infreq., 4—5 spirals faint to fairly well developed. Rays / , tang, pits fim, very 1.6 cells sheath cells 2+7 & 12 (7 & 8)/mm, het. II, up to (2.2) mm high, to 9 (10) wide, Sol. cells chambered cells. present. crystals in ordinary ray and/or in erect ray Crystals sometimes sheathed with thick layer (presumably cell wall material). Material studied. HONDURAS. SJRW 7363*. — NICARAGUA. (IF 20780). — SANTA DOMINGO. Jovero: — Lindeman Abott 2521 (SJRw 7102*), var. macoucoua. SURINAM. Lanjouw & 1474 (Uw 1502). 236 BLUMEA VOL. XXI, No. 2, 1973 Note. Loesener treated (1942) I. panamensis as belonging to I. guianensis. Ilex havilandii Loes. (subg. Euilex, series D, sect. Racemosae) Latitude6 °N. Diffuse Growth absent 2 rings or extremely faint. Vessels & 1* & porous. 34 Ji/n" . 29 diam. 61% sol., 30—42 & 46—83 jum, length 310—1050 & 1150—1520 jum, v—v pits seal, to — to seal, and and opp., rarely trans., 5 6/um, v—r pits opp. occ. tending to large gash-like, bars 14—20 & 22—29, spirals absent. Fibre-tracheids ii8o—i8io & 2080— wall rather absent. het. 2370jum, tang, pits infreq., 3 —4 jum, spirals Rays 2+12/mm, I 6 cells and II, up to 2.0 mm high, to wide, sheath cells abundant. Crystals absent. Pith flecks abundant in SAN 76642. studied. SAN & Material NORTH BORNEO. Mt. Kinabalu: 76642 76645 , alt. 2200m. Ilex hypoglauca (Miq.) Loes. (subg. Byronia, series A. Eubyronia) — Plate IV, 6. Latitude c. 4°N. 2 Diffuse Growth rings faint but Vessels 13/mm 31% sol., diam. — porous. present. , 63 80 —113 fxm., length 1130—1730—2370 jum, v—v pits opp. or occ. alt., 6—8 fim, v—r unil. Fibre-tracheids — pits similar, occ. comp., bars 24—48—67, spirals absent. 1710—2240 wall absent. het. 2550 //m, tang, pits infreq., 4—5 jum, spirals Rays 2+io/mm, II, up to 4.2 mm high, to 10 cells wide, sheath cells absent. Sol. crystals in chambered erect ray cells and procumbent ray cells. Material studied. BORNEO. Sarawak: S 30197, low alt. Ilex insignis Hook. f. (subg. Euilex, series C, sect. Aquifolioides, subsect. Insignis) Latitude 27° N. 2 Diffuse porous. Growth rings distinct. Vessels 76/mm 32% sol., diam. —62—-80 , 38 /am, v similar and length 850—1170—1580/urn, v—v pits opp. to alt., 7—8 jttm, —r pits some- times unil. bars well Fibre-tracheids — comp., 17—30—42, spirals developed. ii6O—1840 wall freq., well developed. I—2+8/mm, het. 2370 //m, tang, pits 4—5 //m, spirals Rays mm cells sheath cells absent. II, up to 2.0 high, to II wide, occ. present. Crystals INDIA. For. Material studied. Darjeeling: Ind. Dept. 1878 (K-Jw), alt. 2400 m. subsect. Ilex integra Thunb.; I. othera Spreng. (subg. Euilex, series C, sect. Aquifolioides, Oxydontae) Latitude 36 °N. Diffuse & 2 or semi-ring porous. Growth rings distinct. Vessels 52 93/mm ,25 & 29% sol, diam. & length & v—v trans, to alt., 29—42 46—63 /um, 700—990 1130—1540fim, pits v—r similar and unil. bars & well developed. pits sometimes comp., 1 x—14 25 —39, spirals Fibre-tracheids & wall infreq. or ffeq., 1130—1790 1910—2450//m, tang, pits fairly —6 het. & 5 fim, spirals well developed. Rays 3 & 3+ 5 & 7/mm, II, up to 1.4 1.9 mm to cells sheath cells Sol. chambered and high, 10 wide, occ. present. crystals infreq. in erect ordinary ray cells, or absent. Material studied. JAPAN. (IF 17234)', Mt. Fujioka (SJRw 9662), probably montane. in wood Note. The two specimens are rather different from each other general anatomical appearance, and several quantitative features. Perhaps one of the identifications is erroneous. P. BAAS: Wood anatomy of Hex 237 Ilex inundataPoepp. (subg. Euilex, series C, sect. Micranthae, subsect. Epunctatae) Latitude c. 6°S. Diffuse Growth faint. Vessels 37/mm 2 60% sol., diam. 46—56—96 porous. rings , /urn, —v 6—10 pits similar and sometimes length 980—1390—1700 /an, v pits opp., /.im, v—r unil. bars 16 absent. Fibre-tracheids —2i8o comp., —31—39, spirals 1580 —2700/mi, tang. wall pits rather infreq., spirals fairly well developed. Rays het. II, to 5 [cm., 2+8/mm, up cells sheath cells Sol. and thick- 2.2 mm high, to 12 wide, occ. present. crystals in ordinary walled ray cells. Material studied. BRAZIL. Para: Maguire51823 (MADw 21515*), low alt. Ilex jenmanii Loes. (subg. Euilex, series C, sect. Micranthae, subsect. Epunctatae) Latitude c 4°N. 2 Diffuse Growth faint to absent. Vessels 27(7i)/mm (24)% sol., diam. porous. rings , 33 —100 length (1370)—2160 v—v to alt., c. 50—83 (55) jum, 820—1360 fini, pits opp. 6/cm, bars —r similar and often until, sometimes tending to gash-like, v pits comp., 24—37 absent in but faint in immature (31)—50, spirals mature specimen, present specimen. Fibre-tracheids wall c. spirals absent 1420—2240 (2030) —2890/im, tang, pits infreq., 4/mi, fibre well in immature or confined to tips in mature specimen, developed specimen. cells Rays 2+6 (i2)/mm, het. II, up to 3.2 (2.1) mm high, to 8 cells wide, sheath occ. pre- sent. Sol. crystals in ordinary ray cells. Material studied. SURINAM. Heyligers 704* (Uw 6737); Stahel 185, low alt. The of in the fibre-tracheids of the immature Note. presence conspicuous spirals and in specimen (quantitative values between brackets) of hardly any spirals the mature specimen ofthis tropical lowland species is remarkable. kleinii Ilex Edwin in Reitz, Flora ilustr. Catarinense 1, Aquifol. (1967) 41 (not assigned to a subgenus or section) Latitude 28 °S. 2 diam. Diffuse Growth distinct. Vessels 94/mm sol., 14 porous. rings , 19% —26—38 /cm, alt. to v—r similar and sometimes length 620—940—1290 /mi, v—v pits opp., 7 /mi, pits unil. bars well Fibre-tracheids comp., 14—26—38, spirals developed. 970—1500—1920/cm, wall rather well developed. 2+i2/mm, het. II, tang, pits infreq., 5—67cm, spirals Rays cells sheath cells Sol. in and up to 1.6 mm high, to 12 wide, present. crystals ordinary thick-walled cells. chambered, occ. ray Material studied. BRAZIL. Santa Catarina: Herb. Barb. Rodr. S. C. 30382*, alt. between 450 and 1200 m. Ilex krugiana Loes. (subg. Euilex, series C, sect. Microdontae, subsect. Eumicrodontae) Latitude c. 28 °N. 2 Diffuse Growth faint. Vessels 4o/mm 38% sol., diam. —88 porous. rings very , 42—63 trans, to occ. seal., v—r similar and /im, length 650—1100—1550, v—v pits opp., pits unil. bars absent. Fibre-tracheids 12 sometimes comp., 15—24—39, spirals 70—1950—2310 wall spirals fairly well developed to absent. jum long, tang, pits fairly freq., 5 jum, Rays cells sheath cells Sol. I—2+io/mm, het. II, up to 3.5 mm high, to 12 wide, common. crystals in chambered erect and ordinary ray cells. Material studied. U.S.A. Florida: (SFCw R 114—66). Note. The absence of spirals from the vessels in this subtropical species is noteworthy. The well line with latitude of quantitative values are in provenance. 237 238 BLUMEA VOL. XXI, No. 2, 1973 flex kwantungensis Merr., J. Am. Arb. 8 (1927) 8 (subg. Euilex, series A, ? sect. Excelsae) Latitude in graph 25 °N. Diffuse Growth distinct. Vessels 111112 !b% sol., diam. — porous. rings . 34 4*5—55 /urn, bars length 690—1020—1550fim, v—v pits opp., 6—7/xm, v—r pits similar, 13—18—25, spirals well developed, tending to annular thickenings. Fibre-tracheids 1240 —1700—2550 wall well het. //m, tang, pits infreq., 3—5 jum, spirals developed. Rays 3 -h 9/mm, II, up to 6 cells 1.2 mm high, to wide, sheath cells very infreq. Sol. crystals in ordinary erect ray cells. Material studied. CHINA. (SJRW 21939). Note. Merrill I.e. placed I. kwantungensis in section Excelsae, but added that he was that this the for it. not sure was proper place Thunb. subsect. flex latifolia (subg. Euilex, series C, sect. Aquifolioides, Insignis) — Plate II, 13; III, 6. Latitude c. 41 °N. 2 Semi-ring Growth distinct. Vessels 77/mm sol., diam. 29—50—71 porous. rings , 24% /um, length 720—970—1140/urn, v—v pits opp. to alt., 6—8 fim, v—r pits similar, some- unil. bars —22 well developed. Fibre-tracheids —166o— times comp., 14 —29, spirals 970 wall well 2160 fim, tang, pits fairly freq., 6 /im, spirals developed. Rays 2+9/mm, het. II, up to 1.7 mm high, to 8 cells wide, sheath cells occ. present. Crystals absent. Material studied. JAPAN. (MADW 5033). flex laurifolia Zipp. ex Loes. (subg. Euilex, series A, sect. Excelsae, subsect. Laxae) - Plate VI, 10. Latitude c. o°. 2 Diffuse porous. Growth rings extremely faint. Vessels 48/mm 21% sol., diam. 46— , similar and 68—105 /tm, length 720—1330—1840/um, v—v pits opp., 6—8 /um, v—r pits often unil. bars absent. Fibre-tracheids comp., 28—43—55, spirals 1580—2050—2760jim, wall infreq., spirals typically absent, for weak spirals in tang, pits 3—4 f/m, except some of the fibres. het. II, het. to to cells some Rays 2+9/mm, rarely I, up 1.9 mm high, 7 wide, differentiated. Sol. abundant in sheath cells not clearly crystals extremely ordinary ray cells, occ. in chambered erect ray cells. Material studied. NEW GUINEA. Waigeo Isl.: Van koyeti 5499*, low alt. the with Note. In spite of fact that we deal here an immature specimen (diam. the and 1.5 cm), quantitative values for vessel member length etc. are comparatively high typical for a tropical lowland species. flex lamina H.B.K. (subg. Euilex, series C, sect. Daphnophyllae) — Plate II, 10, Latitude c. 5 °N. 2 Diffuse Growth rings faint. Vessels sol., diam. —118 porous. 18/mm , 50% 25 —85 jim, similar and length 870—1220—1550/urn, v—v pits trans, to opp., 8—11 /tin, v—r pits and bars absent. often unil. comp., tending to large gash-like, 18—25 —34, spirals Fibre- wall tracheids 2030—2310—28io//m, occ. gelatinous, tang, pits infreq., c. 4 //m, spirals absent. het. to mm to cells sheath cells Rays 2+9/mm, II, up 6.0 high, 13 wide, occ. present. Crystals absent. Material studied. MADW VENEZUELA. 21223 (MERw 44). P. BAAS: Wood anatomy of Ilex 239 the wood Note. In its absence of spirals this species conforms to tropical type of and number of bars plate anatomy, the values for vessel member length per perforation low. This is dueto thehigh altitude (2500—2900 m) are, however, comparatively probably which this Bot. at species occurs (Steyermark, Fieldiana, 28, 1957, 958—962). Ilex ledermanniiLoes. (subg. Byronia, series A. Eubyronia) Latitude c. 6°S. 2 diam. — Diffuse Growth faint. Vessels il—15/mm —38% sol., 63—100 porous. rings ,27 120—139//m, length 930—1630—1800—2270/am, v—v pits opp., 6—!8/um, v—r pits unil. bars similar or horizontally more elongated, occ. comp., 32—51 —55—79, spirals Fibre-tracheids wall infreq., 4—6 absent. 1660—2390—2830—3550/um, tang, pits /um, noted fibre of spirals typically absent, but a few spirals in tips 2 specimens. Rays het. to mm high, to 8 cells wide, sheath cells hardly or 2+7—9/mm, II, up 1.3—2.9 Sol. in cells. not differentiated. crystals abundant chamberederect ray BW Material studied. NEW GUINEA. BW 1467, alt. 6 m; 783s, alt. 630 m; NGF 3293. Ilex liukiuensis Loes. (subg. Euilex, series C, sect. Microdontae, subsect. Repandae) — Plate & VI, 8 9. Latitude 25 °N. 2 Diffuse Growth rather faint. Vessels sol., diam. 29—54— porous. rings 32/mm , 63% v 6—8 v—r pits similar, rarely 76fim, length 880—1260—1600/urn, v— pits opp., /um, ' well Fibre-tracheids unil. comp., bars 25 —32—40, spirals developed. 2130—2740—3020/um, het. wall 6 well I II, to tang, pits infreq., c. /um, spirals developed. Rays —2+7/mm, up sheath cells abundant. in thick-walled-idioblasts, 2.5 mm high, to 11 cells wide, Crystals and in cells. chambered erect ray cells with thickened bulging walls, ordinary ray Material studied. JAPAN. Ryukyu Isl.: Iriomote (MADw 12946). Ilex longipes Chapm. (subg. Prinus, series B. Prinoides) Latitude c. 30° N. Vessels 2 diam. Diffuse to Growth distinct. 107/mm 50% sol., semi-ring porous. rings , seal, similarand length 580—860—1180 v—v pits to v—r pits 17—38—55 fjtm, fim, opp., unil. bars spirals well developed. Fibre-tracheids sometimes comp., 11—17—25, 730— wall freq., —6 spirals well developed. Rays +io/mm, 1260—1710/urn, tang, pits 5 fim, 3 sheath cells differentiated. Sol. het. II, up to I.I mm high, to 10 cells wide, hardly crystals cells. in ordinary or enlarged ray Material studied. U.S.A. Texas: (MADw 13271*). Ilex macfadyenii (Walp.) Rehd.; I. montana (Sw.) Griseb. (subg. Euilex, series A, sect. Excelsae, subsect. Laxae) — Plate I, 4; II, II; III, 8. 0 Latitude 18 N. 2 Diffuse Growth rings faint. Vessels 48/mm 60% sol., diam. 38—58—80 porous. , /um, unil. v trans, to v—r pits similar, occ. length 500—970—1290yitm, —v pits opp., comp., wall bars 26 rather faint. Fibre-tracheids —i68o tang, —35—46, spirals 1030 —1970 /urn, well het. II, to mm high, pits infreq., 3 —5 /um, spirals developed. Rays 2+9/mm, up 1.3 absent. to 6 cells wide, sheath cells occ. present. Crystals Material studied. JAMAICA. MADW 20901 (= USw 6112). 240 BLUMEA VOL. XXI, No. 2, 1973 Ilex — Plate martinianaD. Don (subg. Euilex, series D, sect. Thyrsiflorae) III, 5; V, 3; VI, 3. 0 Latitude c. 4 N. Diffuse Growth rings faint. Vessels 32/mm2,22% sol., diam. porous. very 50—70—84fim, v—r similar length 1170—1650—2010/urn, v—v pits opp., 6/tm, pits and sometimes unil. or gash-like, bars absent. Fibre-tracheids comp. 36—52—71, spirals 1550—2310—2890/um, wall absent. — tang, pits infreq., 4 /;m, spirals Rays 2 Il/mm, het. II, up to 2.9 mm high, 6 cells sheath cells Sol. cells. to wide, occ. present. crystals infreq. in ordinary ray Material studied. SURINAM. Lindeman 6340 a (Uw 4466 a). Note. In very few of the fibre-tracheids I noted septa, staining blue with haematoxylin These distances within (Plate V, 3). septa may occur atvery irregular asingle fibre tracheid. and I hesitate to refer to these as septate fibres I am not certain whether the structures involved are artefactual in the sense that they were formed under unusual conditions of the followed (contraction protoplast by the deposition of a thin membrane at the for have noticed fibres other wood of periphery instance). I not septate in any specimen Ilex. Ilex micrococca Maxim, (subg. Byronia, series B. Micrococca) Latitude 36° N. Growth distinct. 2 Diffuse porous to semi-ring rings Vessels 70/mm 2 s°l-> porous. , 4% diam. —61— length 840 —1630 v—v to alt., 6 — 34 70 [mi, —1320 fim, pits opp. fini, v r well Fibre-tracheids pits similar and often unil. comp., bars 10—18—24, spirals developed. wall 1050—1890—263071m, tang, pits infreq., 6/im, spirals well developed. Rays 2+5/111111, wide. het. II, up to 0.9 mm high, to 6 cells Crystals absent. Material studied. JAPAN. Fujioka: (MADut 7419). Ilex mitis (L.) Radlk.; I. capensis Harv. & Sond., I. monticolaTul. (subg. Euilex, series C, sect. Lemurensis) 0 0 Latitude to at 25 S 12° N, in graph plotted 12 . Growth 2 Diffuse rings extremely faint to rather distinct. Vessels 32-48/mm porous. , —50% sol., diam. 38 —76 length 820—11 —2060 v—v 13 —70 —105 fim, 30—1340 fim, to —8 v—r similar and unil. bars — pits opp. alt., 5 fim, pits sometimes comp., 16—25—43 faint almost absent. Fibre-tracheids 52, spirals to 1320—1950—268o—3!6o / het. to to cells sheath cells Sol. II, up 1.8—2.7 mm high, 13 wide, occ. present. crystals in ordinary or chambered erect ray cells, or absent. — Mann Material studied. BURUNDI. Lewalle 3623 (IF 21963, Tervuren). CAMEROONS. s.n. (K-Jw), high alt. — alt.; Letouzey & M. C. 183 (CTFT 14646, CTFw), high MADAGASCAR. Thouvenot 15 (CTFT 1792, CTFw); (CTFT 13474). Note. Considering the fact that I. mitis is a widely distributedand variable species, the wood anatomical variation is rather limited. The species generally occurs in montane and the values well with what one would of habitats, quantitative agree expect a tropical species growing at higher altitudes. In our material, specimens from Madagascar showed the spirals most clearly (almost subtropical latitude!). Abbate (1970) pictured the wood of with an I. mitis specimen from Eritrea (lat. c. 14° N) very conspicuous spirals as one in Abbate's of would expect a subtropical species. drawing a vessel member is misleading and erroneous. However, his photomicrograph leaves no doubt that he has studied an P. BAAS : Wood anatomy ofIlex 241 Ilex. 2 be His values for vessel frequency (63/mm ) and diameter (50 /im) may possibly due to his specimen being immature. flex montana Torr. & Gray (subg. Prinus, series B. Prinoides) Latitude in graph i8° and 41° N. 2 faint distinct. Vessels — Diffuse to Growth rings to 48 46 semi-ring porous. —98/mm , diam. —80 trans, 6o% sol., 17—42—58 /un, length 440—760—970—1130 fim, v—v pits bars rather faint. Fibre- to alt., v—r pits similar, not comp., 23 —31 —39—60, spirals tracheids —i68o wall well 1030—1320 —1970 fim, tang, pits infreq., 4 —5 jum, spirals developed. — mm het. II, to mm high, to 14 cells wide, sheath cells Rays 3+6 9/ » up 1.7 absent or Sol. thick-walled cells occ. present. crystals in erect ray present in SFRw R in other 151—7, absent specimens. U.S.A. Material studied. JAPAN. (R IJJ —7, SFRw). — Pennsylvania: MADw 14603. Note. Loesener treated I. montana Torr & var. mollis as a of I. (1901) . Gray variety dubia Trel. Rehder Arb. maintained the (Don) (J. Arn. 3, 1922, 215), however, species. and have The wood specimens I studied of both I. montana I. dubia, a relatively high in In number of bars per perforation plate common. some qualitative features they differ (see p. 233). flex myricoides H.B.K. var. polyphylla (Benth.) Loes. (subg. Euilex, series B, sect. Polyphyllae) 0 Latitude c. 4 N. 2 Diffuse porous. Growth rings virtually absent. Vessels i6/mm sol., diam. —61— , 70% 46 80 6—8 similar fim, length 660—1470—2130 jum, v—v pits trans, to opp., fim, v—r pits and often unil. comp. with wide slits on parenchyma cells extending over the whole radial bars absent. Fibre-tracheids wall, 15—33—42, spirals i66o—2390—2890 occ. wall absent. gelatinous, tang, pits infreq., c. 5 //m, spirals Rays 2 +io/mm, het. II, up to to cells sheath cells 1.5 mm high, 7 wide, occ. present. Crystals absent. Material studied. COLOMBIA. Cuatrecasas 19166(SJRw 43288), alt. 2700 m. Note. In spite of thehigh altitude of the locality of this specimen, the values for vessel member length and number of bars per perforation plate are high. flex nayana Cuatr., Lloydia 11 (1949) 214 (subg. Euilex, ? series A, ? sect. Excelsae, subsect. Laxae) 0 Latitude 4 N. 2 Diffuse porous. Growth rings faint. Vessels 40/mm 20% sol., diam. —84—114 , 50 fim, and length 790—1080 v—v to alt., similar some- —1390 fim, pits opp. 7—9 fim, v—r pits times unil. bars absent. Fibre-tracheids —1810 —2160 comp., 22—36—49, spirals 1450 fim, wall well tang, pits infreq., 4 —5 fim, spirals fairly developed. Rays 3+6/mm, het. II, to to cells sheath cells Sol. abundant up 2.5 mm high, 7 wide, present. crystals in ordinary and chamberederect ray cells. Material studied. COLOMBIA. Cuatrecasas alt. 14286, type (SJRw 42735*), 3 m. Note. Cuatrecasas I.e. mentioned that this species is closely related to I. amygdifolia (of this section) and to I. laureola (from section Thyrsiflorae of series D of subgenus Euilex). The short vessel membersand fibres should be accounted for by the small diameter ofthe sample. BLUMEA VOL. No. 242 XXI, 2, 1973 Ilex opaca Ait. (subg. Euilex, series A, sect. Cassinoides) Latitude c. 29° N. 2 Diffuse to semi-ring Growth rings distinct. Vessels 128 & /mm 9 & 12% porous. 143 , diam. & 1020 v—v sol., 21—34 & 42—50 /tm, length 720—1010 —1390[im, pits opp. to unil. bars well alt., 5 —7 fim, v—r pits similar, rarely comp., 11—26 & 27—38, spirals developed. Fibre-tracheids & wall to rather 760—1390 1520—2050 fi m, tang, pits infreq. well & & het. to freq., 4—6 [im, spirals developed. Rays 4 5 + 5 6/mm, II to III, up & mm to cells sheath cells absent. absent. i.i 1.4 high, 7 wide, Crystals Material studied. U.S.A. Florida: (MADw 2636); Louisiana: (IF 19728, Tervuren). Ilex ovalifolia Mey. ( sensu Loes). (subg. Euilex, series D, sect. Thyrsiflorae) 0 Latitude c. 4 N. 2 diam. Diffuse porous. Growth rings faint. Vessel 47/mm 38% sol., 29—63 —84 , /um, —8 similar and length 740—1340—1800[im., v—v pits opp., 5 /tm, v—r pits sometimes 18 — unil. comp., bars —33—41, spirals absent. Fibre-tracheids 1230—2450 }i6o jum, tang. absent. het. to wall pits fairly freq., 3—5 fim, spirals Rays 4+7/mm, I to II, up 1.7 mm high, to 5 cells wide, sheath cells abundant. Crystals absent. Material studied. SURINAM. Lindeman 4541 (Uw 3150). Ilex paraguariensis St. Hil. (subg. Euilex, series C, sect. Microdontae, subsect. Repandae) Latitude c. 25° S. 2 Diffuse porous. Growth father faint. Vessels 80/mm 28% sol., diam. 38—55 — rings , length v—v to alt., 6—8 v—r similar and 80 fim, 690—910—1340 /jm, pits opp. fim, pits sometimes well unil. comp., bars 14—20—25, spirals developed. Fibre-tracheids 710 —I8IO wall rather spirals well 2+9/mm, —2370 fini, tang, pits infreq., 4—5 fim, developed. Rays het. cells sheath cells Sol. in chambered II, up to 3.5 mm high, to 15 wide, infreq. crystals cells. erect and ordinary ray Material studied. BRAZIL. Parana: Lindeman & Hecking 2829 (Uw 13942). Ilex perado Ait.; I. platyphylla Webb (subg. Euilex, series C, sect. Aquifolioides, subsect. Oxydontae) — Plate I, 2. Latitude c. 28° N. 2 Growth distinct. Vessels —123/mm sol., diam. Semi-ring porous. rings 89 , 14—28% 21—39—44—63 //m, length 310—690—880— 1240 //m, v—v pits trans, to opp., 6—9 /um, similar and sometimes unil. bars well v—r pits comp., 16—20—25—39, spirals developed. Fibre-tracheids wall rather — 650—1130—1520—iSio/zm, tang, pits infreq. to freq., 4 7/um, well het. to spirals developed. Rays 3—4+7—io/ mm, II, up 0.9—1.4 mm high, to 8 cells wide, sheath cells occ. present. Crystals absent. Baas 600 Material studied. CANARY ISLANDS. (IF 22937); Bourgeau s.n. (K-Jw); & Ridsdale 9, alt. m. Note. This species differs widely from two specimens I studied ofI. canariensis growing in the same habitat; one probably misnamed specimen of I. canariensis is very similar to I. perado. See also the note under I. canariensis. Ilex petiolaris Benth.; I. parviflora Benth. (subg. Euilex, series C, sect. Micranthae, subsect. Punctatae) S. Latitude c. 4° 2 Diffuse Growth rings faint to absent. Vessels 16/mm sol., diam. porous. very , 56% P. BAAS : Wood anatomy ofIlex 243 6—8 55 —8o—ioi fim, length 930—1240—1730 pm, v—v pits opp., /xm, v—r pits sometimes Fibre-tracheids similar and unil. comp., bars 27—36—47, spirals very faint. wall faint. 1390—1790—2160/xm, tang, pits infreq., 5 —6/xm, spirals very Rays 2+8/mm, het. 8 sheath II, up to 2.9 mm high, to cells wide, cells occ. present. Sol. crystals in ordi- nary ray cells. Material studied. BRAZIL. Amazonas: Krukoff 6561 (MADw 12529). Ilex quercetorum Johnston, J. Arn. Arb. 19 (1938) 122 (subg. Euilex, series A, sect, Cassinoides) 0 Latitude c. 16 N. 2 Diffuse Growth rings distinct. Vessels 64/mm 40% sol., diam. —42 porous. , 25 —59 /xm, to similar and length 670—860—1060fim, v—v pits opp. alt., 6—7 jum, v—r pits some- times unil. comp., bars 16—24—35, spirals fairly well developed. Fibre-tracheids 870— wall — but faint. 1420—1920 jum,tang, pits infreq., 5 6 fx m, spirals present Rays 5 + 8/mm, het. to i.i mm to 6 cells sheath cells absent. Sol. in II, up high, wide, crystals rare ordinary ray cells. Material studied. MEXICO. Chiapas: Page 9754 (MADw 23937*), probably high alt. I.e. this Note. According to Johnston species shouldbe included in section Cassinoides. is from The the The type specimen a montane plant i860 m alt. diameter of present have been almost from the of the specimen must 4 cm, judged curving growth rings in the peripheral block I obtained. The low values for vessel member length are probably that only partly due to the fact this specimen was not fully mature, but also to the montane habitat of this species. Ilex repanda Griseb. (subg. Euilex, series C, sect. Microdontae, subsect. Repandae) Latitude c. 21° N. Diffuse Growth faint. Vessels 42/mm2 60% sol., diam. porous. rings , 34—55—67 /um, to 6— similar and often length 670—1110—1490 /urn, v—v pits opp. alt., 7 //m, v—r pits bars — extensively unil. comp., 9—22—33, spirals absent. Fibre-tracheids 1450—!940 wall 6 rather faint. 2550 pits infreq., 5 — spirals and Rays i ( 9/mm, jum, tang, jum, present het. to to cells sheath cells Sol. and II, up 2.7 mm high, 12 wide, freq. crystals in ordinary cells. chambered erect ray Bucher Material studied. CUBA. 243 (SJRw 21418*). Note. The low of member values vessel length and number of bars per perforation for this are dueto the fact that the for plate tropical species probably specimen study was immature. The absence of spirals in the vessels is more in line with latitude. Ilex rimbachii Standley, Publ. Field Mus. Nat. Hist. Chicago, Bot. Ser. 17 (1937) 198. — Plate II, 6 & 7; V, 7 & 8; VI, 5. Latitude c. 1° S. 2 Diffuse Growth rather faint. Vessels sol., diam. —62— porous. rings 34/mm , 68% 30 80 v—v seal, — similar often fim, length 960—1250—1500 //m, pits to opp., v r pits and unil. bars faint. Fibre-tracheids —2ioo comp., 19—32—37, spirals 1630 —2370 jMm, occ. gelatinous, wall infreq., c. faint. het. to tang, pits 6 /xm, spirals Rays 2+9/mm, II, up 5.5 mm high, to 15 cells wide, sheath cells occ. present. Crystals absent. Material studied. ECUADOR. Solis alt. Acosta 6689 (MADw 16614), 2500—3000 m. BLUMEA 244 VOL. XXI, No. 2, 1973 I.e. did indication about the of this with Note. Standley not give any affinity species other Ilex species. Ilex rotunda Thunb. (subg. Euilex, series A, sect. Excelsae, subsect. Umbelliformes) — PlateV, 6. Latitude 0 in graph 41 N. Diffuse Growth rings distinct. Vessels mm2 4°% sol., diam. —61 —86 porous. 33/ > 42 jiva, —1600 v—v 6—8 —r similar and often unil. length 850—1210 fxm, pits opp., [im, v pits bars well Fibre-tracheids comp., 12—20—25, spirals developed. 1840—2290—2760 fim, wall pits fairly ffeq., 6 spirals well developed. Rays + 5/mm het. II, to tang, /tm, 3 > up cells sheath cells in 1.4 mm high, to 6 wide, very rare. Solitary crystals present ordinary ray cells, occ. somewhat elongated. Material studied. JAPAN. ( CSIRO 13253, FPAw). Ilex sclerophylloides Loes. (subg. Byronia, series A. Eubyronia) Latitude c. 4° N. Diffuse Growth faint. Vessels (i6)/mm 2 (54)% sol., diam. porous. rings 13 , 44 59—:113 (1190) —2210 v—v —8 v—r (62)—134 /urn, length 360—1680 fim, pits opp., 7 /um, pits similar and unil. bars often comp., 22—41 (36) —58, spirals absent. Fibre-tracheids mo—- wall to rather absent or 2740 (1940)—3420 /tm, tang, pits infreq. freq., 4—5 //m, spirals and faint S & 8 & het. & rare (in 20855). Rays 2 3 + 9/mm, II, up to 1.9 6.0 mm high, to 9 cells wide, sheath cells present. Sol. crystals abundant in ordinary and chambered erect ray cells. Material studied. BORNEO. Sarawak: S 20855-, ?S 16867 ( CSIRO 50222*, FPAw), low alt. Ilex sebertii Panch. & Seb. (subg. Euilex, series A, sect. Excelsae, subsect. Laxae) ° Latitude c. 21 S. 2 Diffuse Growth rings faint. Vessels 29/mm sol., diam. — porous. very , 48% 38—63 6—8 similar and 84/ Material studied. NEW CALEDONIA. Sarlin 85 (CTFT 6117, CTFw), low alt. Ilex serrata Thunb. (subg. Prinus, series A. Euprinus) — Plate II, 3 Latitude 40° N. 2 Semi-ring Growth distinct. Vessels sol., diam. 21 — porous. rings 115/mm , 30% —38 and length v—v pits trans, to 6—8 v—r 46 fim, 380—600—870[im, alt., fim, pits opp. sometimes unil. comp., bars 10—17—24, spirals absent. Fibre-tracheids 070—1000—1320 wall pits rather freq., spirals absent. 5+9/mm het. II, to , Rays jum tang, 3 —4 jum, > up 0.6 mm high, to 5 cells wide, sheath cells occ. present. Crystals absent. Material studied. U.S.A. cult. Brooklyn Bot. Garden (SJRw 40816). absence Note. This species is native in Japan. The of spirals in this temperate species remarkable also under and discussions is very (see I. verticillata taxonomic on p. 224 and 225). Wood Ilex P. BAAS : anatomy of 245 Ilex sideroxyloides (Sw.) Griseb. (subg. Euilex, series C, sect. Microdontae, subsect. Sideroxyloides) — Plate II, 2. Latitude c. 16° N. Diffuse Growth Vessels 2 diam. porous. faint. (22)/mm 51 (50)% sol., 46 rings 37 , —76 6—8 (59) —105 /on, length 1000—1320 (990) —1700 fim, v—v pits opp., /im, v—r pits similar and sometimes unil. bars absent. Fibre-tracheids comp., 24—36 (23) —56, spirals 1200 (1580) wall rather infreq., spirals faint. Rays —2230 —3140 /tin, tang, pits 4—5 /im, —2 8 het. to to cells sheath cells 1 (2)+ (9)/mm, II, up 3.7 (3.0) mm high, 15 wide, occ. Sol. abundant in and chambered cells. present. crystals ordinary erect ray Material studied. DOMINICA. Stern & Wasshausen 2506 (MADw 24200*). — GUADELOUPE. (HM 2279—55. RTIw) Ilex subrugosa Loes. in Sargent, Plantae Wilsonianae 1 (1911) 80 (subg. Euilex, series C, sect. Microdontae) 0 Latitude in graph 25 N. 2 Diffuse to Growth distinct. Vessels 109/mm sol., diam. semi-ring porous. rings , 43% pits to v—r pits 25 —50—67 fjLm, length 530—960—1340 jum, v—v opp. alt., 5 —7 (J,m, similar and unil. bars well Fibre-trach- sometimes comp., 18—27—34> spirals developed. eids wall well 1210—1810—2520 /im, tang, pits freq., c. 5 /im, spirals developed. Rays het. t5 cells sheath cells Sol. 3+7/mm, II, up 1.2 mm high, to 7 wide, occ. present. crystals in ordinary ray cells. Material studied. CHINA. SJRW 21734. Note. Loesener doubtfully included I. subrugosa in section Microdontae, but he also remarked that it recalls section Rugosae. Wood anatomically it is closer to section Micro- dontae than section but this be due to the fact that of the latter to Rugosae, may entirely examined characters from section I only tropical species and that no special apart those the latitudinaltrend in the wood of I. following general are present subrugosa. Ilex theezans Mart. (subg. Euilex, series C, sect. Megalae, subsect. Pedicellatae) Latitude 0 S. c. 25 Vessels 2 Diffuse Growth rings distinct. 66/mm 20% sol., diam. 34 —63 porous. , —50 /urn, to similar and length 1050—1300—1600 /un, v—v pits opp. alt., 6—7 /un, v—r pits sometimes unil. comp., bars 16—27—37, spirals well developed. Fibre-tracheids IylO—- — wall rather well 2390 276o /urn, tang, pits freq., 5 //m, spirals developed. Rays 1—2 + cells sheath absent. 7/mm, het. II, up to 4.5 mm high, to 20 wide, cells abundant. Crystals BRAZIL. Parana: Material studied. Lindeman & Hecking 1152 (Uw 12923). Ilex toluccana Hemsl. (subg. Euilex, series C, sect. Microdontae, subsect. Repandae) — Plate VI, 4. Latitude 0 in graph 24 N. 2 Diffuse Growth rather faint. Vessels 4-8/mm sol., diam. —52 — porous. rings , 37% 38 6—8 similar 72 fim, length 760—1200—1650/im, v—v pits trans, to opp., /mi, v—r pits and unil. bars sometimes comp., 10—17—22, spirals well developed. Fibre-tracheids wall 6 spirals well developed. Rays 2 1470—2160—2310/tm, tang, pits infreq., /tm, —3 + het. 8 cells sheath cells abundant. absent. 10/mm, II, up to 1.5 mm high, to wide, Crystals Material studied. MEXICO. (IF 19128, Tervuren). 246 BLUMEA VOL. XXI, No. 2, 1973 Ilex series umbellata Kl. var. humirioides (Reiss.) Loes. (subg. Euilex, C, sect. Micranthae, subsect. Epunctatae) — Plate n, I. Latitude c. g° N. 2 Diffuse Growth rings faint. Vessels 24/mm 67% sol., diam. 50—80—101 porous. , fim, —v v—r similar and length 1030—1430—1870 fim, v pits trans, to opp., pits sometimes unil. bars absent. Fihre-tracheids comp., 25 —40—58, spirals 1970 —26oo—3°8O jum, tang. wall 6 faint. I het. II, to to pits infreq., jum, spirals Rays (2)+8/mm, up 5.5 mm high, cells sheath 12 wide, cells occ. present. Sol. crystals in ordinary ray cells. Material studied. BRITISH GUIANA. Forest Dept. 3693 (SJRw 43839), probably low alt. Ilex uniflora Benth. var. panamensis Cuatr., Lloydia 11 (1948) 206 (subg. Euilex, series B. sect. Rupicolae) 0 Latitude in graph 4 N. 2 Diffuse porous. Growth rings very faint. Vessels 28/mm 57% sol., diam. — , 29 —63 v—v trans, similar 88 fim., length 430—1090—1440 /ira, pits to opp., 6—8 fim, v—r pits and sometimes unil. well comp., bars 16—22—30, spirals fairly developed. Fibre-tracheids wall absent or faint. 1260—1920—2390 [Jim, tang, pits infreq., 4 —5 [im, spirals very het. to to cells sheath cells Rays 3 +4/mm, II, up 1.4 mm high, 6 wide, sometimespresent. Sol. in cells. crystals infreq. ordinary ray Material studied. COLOMBIA. Cuatrecasas alt. 20365, type (MADw 17630), 3400 m. The low values for vessel member and number of bars Note. length per perforation with the of in the make this of plate, together presence spirals vessels, species high mon- tane habitat resemble temperate representatives of Ilex. Arb. Ilex versteeghii Merr. & Perry, J. Arn. 22 (1941) 259 (subg. Euilex, series C, sect. Rugosae) Latitude c. 5° S. 2 Diffuse Growth absent. Vessels 26/mm 36% sol., diam. 40—71 —97 porous. rings , /urn, to similar and length 720—1070—1550 fim, v—v pits opp. alt., 6—7/um,1 v—r pits some- unil. bars absent. Fibre-tracheids —2160 times comp., 17—23—30, spirals 1000—1740 fim, wall rather infireq., spirals absent. 2+7/mm, het. II, to tang, pits 3 —4 /um, Rays up mm cells sheath cells absent. I.I high, to 7 wide, occ. present. Crystals Material studied. NEW GUINEA. NGF 9460, alt. 2200 m. be Note. I. versteeghii is said to close to I. archboldiana (Merrill & Perry I.e.) which in turn is said to be close to I. revoluta. The latter species belongs to section Rugosae, and therefore I. versteeghii is treated under this section. Values for vessel member length and number of bars per perforation plate are low in this tropical montane species. This is probably due to the altitudinal effect. Ilex verticillata (L.) Gray (subg. Prinus, series A. Euprinus) — Plate IV, 5; V, 4; VI, 1. Latitude c. 40° N. 2 Semi-ring Growth rings distinct. Vessels 24% sol., diam. —80 porous. 97/min , 25—42 —640—820 v—v trans, to alt., v—r and (im, length 330 fim, pits 6—9 //m, pits opp. bars absent. sometimes unil. comp., 12—21—36, spirals Fibre-tracheids 780—890—ll8o wall absent. het. to i.o (im, tang, pits freq., 6—7//m, spirals Rays 6+io/mm, II, up mm cells high, to 3 (rarely 5) cells wide, sheath absent. Crystals absent. P. BAAS : Wood anatomy ofIlex 247 Material studied. U.S.A. Ohio: (SFCw R 206—14) for detailed description; Michigan: (MADw 2629) for checking of absence ofspirals. values Note. As in I. serrata, the absence of spirals in this temperate species with low and for vessel member length number of bars per perforation plate is very remarkable discussions and (see taxonomic on p. 224 225). Ilex volkensiana Kaneh. & (Loes.) Hatus., Bot. Mag. Tokyo 50 (1936) 607; I. mertensii subsect. Maxim, var. volkensianaLoes. (subg. Euilex, series C, sect. Microdontae, Repandae) 0 Latitude 7 N. 2 Diffuse to Growth fairly distinct. Vessels 31/mm sol., semi-ring porous. rings , 30% v—v seal, diam. 42—59—80 /«m, length 700—1040—1390 fxm, pits to opp., v—r pits similar and unil. bars absent. Fibre-tracheids —2060— comp., 21—28—36, spirals 1450 wall 6 faint. Rays 3+7/mm, het. II, to 2680 fim, tang, pits fairly freq., c. /um, spirals up 2.2 mm high, to xo cells wide, sheath cells abundant. Sol. crystals in ordinary ray cells. Material studied. MICRONESIA. Ponape: Kanehira 802 (SJRw 20384*), alt. 650 m. This the Note. specimen is from summit area of Mt .Tolotom, and the low value for immature of the and the vessel member length may be due to the nature specimen fact that the summits of the mountains of Ponape are exposed to frequent strong winds which results in stunted tree growth (Glassman, Bern. P. Bish. Mus. Bull. 209, 1952, 152 pp.). Ilex wilsonii Loes. (subg. Euilex, series C, sect. Microdontae, subsect. Sideroxyloides) Latitude in graph 25 °N. 2 Diffuse Growth distinct. Vessels 52/mm 38% sol., diam. —42 porous. rings , 25 —63 /um, 6— —r similar and length 550—1060—1320/um, v—v pits opp. to alt., 7/tm, v pits some- unil. bars well Fibre-tracheids 1180—1810— times comp., 13 —23—30, spirals developed. wall rather — well 2210 /um, tang, pits inffeq., 4 6/um, spirals developed. Rays 2+6/mm, cells het. II, up to 1.8 mm high, to 8 wide, sheath cells rare. Sol. crystals infreq. in ordinary chambered or erect ray cells. Material studied. CHINA. Chow s.n. (SJRw 42563). Ilex spec, (unidentified) Latitude 6°S. 2 Diffuse Growth absent. Vessels 17/mm sol., diam. —67—84 porous. rings , 59% 50 /um, & similar length 520—1020 1390—1770/tm, v—v pits trans, to opp., 6—9/ & wall — absent. 2 & 1500 22Ô0—268Ofim, tang, pits infreq., 4 6 /nm, spirals Rays &3 + 5 mm cells sheath cells absent. 7/mm, het. II, up to 1.8 high, to 9 wide, present. Crystals NEW GUINBA. Kalkman alt. Material studied. Kalkman 4863, alt. 3140 m; 5136, 3350 m. in Note. The two specimens differ considerably in quantitative values spite of the from similar altitude. fact that both samples are from mature stems and (very high) Kalkman had the lowest values for vessel member and number of bars 5136 length per perforation plate, but occurs in a similar vegetation (mossy forest) as Kalkman 4863. Ilex spec, (unidentified) Latitude 4°N. 2 Diffuse Growth rings faint to fairly distinct. Vessels sol., diam. porous. 41/mm , 51% 248 BLUMEA VOL. XXI, No. 2, 1973 trans, to seal., v—r 38—55—7 1 jMtn, length 720—1400—196071m, v—v pits opp., occ. pits sometimes bars absent. similar, unil. comp. or tending to gash-like, 20—34—41, spirals Fibre-tracheids wall —6 faint or 1710—2340—2890/im, tang, pits infreq., 5 /j,m, spirals absent. het. to mm to 16 cells wide, sheath cells occ. Rays 2+9/mm, II, up 3.3 high, idioblastic thick-walled present. Sol. crystals in ordinary, chambered erect, and bulging cells. ray studied. SUMATRA. De Wilde alt. Material NORTH 13020, 1300—1500 m. The of this from the montane forest Note. presence growth rings in specimen rain is Both the and the in ofthe fibre-tracheids curious. growth rings presence of spirals some would climate with but the collector suggest a a dry season, according to dry spells are virtually absent in the habitat where this species was found. Ilex — spec, (unidentified) Plate I, 5; II, 5; III, 4. Latitude 15 °N. 2 diam. — Diffuse Growth faint to absent. Vessels 25/mm 44% sol., 48—84 porous. rings , 118 /um, length 470—1090—1440 /urn, v—v pits opp., rarely trans., 7—lOfim, v—r pits similar or irregularly arranged and large and gash-like, occ. unil. comp., bars 4—13—29, spirals absent. Fibre-tracheids E400—1830—2510/mi, tang, wall pits freq., 3—5 //m, spirals faint or absent. het. to cells sheath cells Rays 2+io/mm, II, up to 6.5 mm high, 17 wide, in fairly common. Sol. crystals ordinary ray cells. Material studied. PHILIPPINES. Jacobs 7928*, alt. 600—700 m (climber!). Note. This specimen represents the only climbing species of Ilex available for this such the wood anatomical study. It shows some striking anatomical deviations, as very bars for the low number of per perforation plate (the lowest wholegenus, and very strong- ly deviating from other tropical lowland species, which are all characterized by a high number of bars The bars spaced. The per perforation plate). are, moreover, very widely the parenchyma, which is diffuse in aggregates, is unusually abundant. Although specimen with diameter of was not the minimum its 3.5 cm yet fully mature, according to arbitrary size of stems used as a standard here, thevessel members and fibre-tracheids are remarkably short for lowland Ilex. All here well in with tropical deviations reported are agreement general anatomical tendencies in the wood of climbing plants. The generic identity of this specimen was confirmed by pollen morphology. ACKNOWLEDGEMENTS This study was carried out under supervision of Prof. Dr. C. G. G. J. van Steenis, to whom I wish to express my gratitude for his remarks. Numerous colleagues provided me with wood samples of Ilex. Thanks are due to the Curators of the wood collections of: Forest Prod. Res. and Industr. Development Commission, College, Laguna; Forest Department, Sarawak; Forestry Commission of New South Wales, Sydney; Centre Technique Forestier Tropical, Nogent sur Marne; Bernice P. Bishop Museum, Honolulu; Forest Products Res. Lab., Madison (special thanks to Mr. Miller for providing most materials for this Instituut de Instituut study!); Koninklijk voor Tropen, Amsterdam; voor Systematische Plantkunde, Utrecht; C.S.I.R.O., Melbourne; Royal Botanic Gardens, Kew; Commonwealth Forestry Institute, Oxford: Koninklijk Museum voor Midden Afrika, Tervuren; Pusat Penyelidekan Hutan, Kepong. Mr. Cockburn (Sandakan) is acknowledged for arranging the collecting of wood specimens ofI. havilandii in the field. P. BAAS: Wood anatomy of Ilex 249 the The Director of the Institute for Geology and Mineralogy kindly gave access to of themicrotechnical scanning electron microscope, operated by Mr. H. Kammeraat. Most C. L. Marks and Mr. B. N. Kieft work was skillfully carried out by Mr. W. Star. Mr. prepared the diagrams and plates. read the Prof. Dr. W. K. H. Karstens and Prof. Dr. C. Kalkman critically manuscript. REFERENCES (not including those cited under 'specific descriptions') of the stand in Oak forests in relation to ABBASSOVA, V. S. 1969. Structural specialization components environmental conditions. Izv. AN AzSSR (Ser. Biol. Nauk) (6): 16 20. dei dell' Africa orientale. Cartella VI, No. 67. ABBATE, M. L. E. 1970. Atlante micrographico legni examination. Forest Prod. Res. Lab. Leaflet, ANONYMOUS. 1968. The preparationof wood for microscopic No. 40. Bot. Linn. Soc. 62: BAAS, P. 1969. Comparative anatomy of Platanus kerrii Gagnep. J. 413 421. —8. I. W. ofthe tissue of land Amer. Bot. 40: 4 BAILEY, 1953. Evolution tracheary plants. J. Am. The of the Icacinaceae III. Arb. 22: 432— & R. A. HOWARD. 1941. comparative morphology J. 442. Amer. SINNOTT. distribution of certain of leaves. J. & E. W. 1916. The climatic types angiosperm Bot. 3:24—39. . . . , and of rays in the BARGHOORN, E. S. 1940. The ontogenetic development phylogenetic specialization xylem of dicotyledons I. Amer. J. Bot. 27: 918—928. der sekundaren in W. Zimmermann, P. Ozenda, & BRAUN, H. J. 1970. Funktionelle Histologic Sprossachse; H. D. Wulff (ed.), Handbuch der Pflanzenanatomie IX, 1. Res. Identification of a key. Forest Prod. BRAZIER, J. D., & G. L. FRANKLIN. 1961. Hardwoods, microscope Bull., No. 46. CARLQUIST, S. 1961. Comparative Plant Anatomy. wood of with on factors controlling 1966. Wood anatomy Compositae: a summary, comments evolution. Aliso 6: 25—44. other of Euphorbia; in N. K. B. —— of and species 1970. Wood anatomy Hawaiian, Macaronesian, in Bot. J. Linn. Soc. Robson, D. F. Cutler, & M. Gregory (ed.), New Research Plant Anatomy. 63, suppl. I. fibre and cambial initial in woods. CHATTAWAY, M. M. 1936. Relation between length dicotyledonous Trop. Woods 46: 16—20. Woods 1956. Crystals in woody tissues II. Trop. 104: 100—124. The ofNew Guinea Nothofagus Bl. Austr. J. Bot. DADSWELL, H. E., & H. D. INGLE. 1954. wood anatomy 2: 141—153- sitchensis Carr. Forest Prod. Special DINWOODIE, J. M. 1963. Variation in tracheid lengthin Picea Report 16 D.S.I.R. Kon. Miocene Kreuzau. The leaf-remains. Verh. FERGUSON, D. K. 1971. The flora of Western Germany 1. Ned. Akad. Wet., Afd. Natuurk. II, 60, No. 1. Laubholzer und Straucher. GREGUSS, P. 1959. Holzanatomie der Europaischen fiber das Vorkommen Reaktionsgewebe in Wurzeln und Asten HOSTER, H.-R., & W. LIESE. 1966. von 80— der Dikotyledonen. Holzforschung 20: 90. distribution of the of Aquifoliaceae in the Pacific area (1). J. Jap. Hu, S. 1967. The evolution and species Bot. 42: 13—27. The of the xylem ofSouth West Pacific tree INGLE, H. D., & H. E. DADSWELL. 1961. anatomy secondary Pac. aid their delivered at the 10th Sci. Congr., 12 pp. species as an to taxonomy. Paper Structure of Wood. JANE, F. W. 1970. The des Holzes der auf vorkommenden Baumarten I—VI. JANSSONIUS, H. H. 1906—1936. Mikrographie Java with critical remarks Anatomical Characters and Identification of Formosan Woods, KANEHIRA, R. 1921 a. from the climatic point of view. b. of the Woods anatomical characters. 1921 Identification important Japanese by Woods anatomical characters. 1924. Identification ofPhilippine by I. Nova Acta Abh. Kais. Leop.-Carol. Deutsch. LOESENER, TH. 1901. Monographia Aquifoliacearum Akad. Naturf. 78 1908. Ibid. II. Ibid. 89 (1). in Prantl, Natiirliche Pflanzenfainilien ed. 2, 20 b; 36—86, 1942. Aquifoliaceae; Engler and 250 BLUMEA VOL. XXI, No. 2, 1973 METCALFE, C. R., & L. CHALK. 1950. Anatomy ofthe Dicotyledons. MOLL, J. W., & H. H. JANSSONIUS. See Janssonius 1906 —1936. NOVRUZOVA, Z. A. 1968. The Water-Conducting System of Trees and Shrubs in Relation to Ecology. Izd. Azerb. AN SSR, Baku, 230 pp. A. H. P. BROWN. PANSHIN, J., C. DE ZEEUW, & 1964. Textbook ofWood Technology. PARHAM, R. A., & H. KAUSTINEN. 1973. On the Morphology of Spiral Thickenings. IAWA Bull. 1973/2: 8—17. of PENNINGTON, M. T. 1953. A comparativestudy ofwood anatomy fifty-four species ofthe family Aqui- foliaceae. Thesis Univ. Virginia. Doct. Diss. Ser. Publ. No. 7981. PURKAYASTHA, S. K. 1963. Aquifoliaceae; in Indian Woods II: 175—179. RECORD, J. S. 1919. Identification of the economic woods of the United States. SARLIN, P. 1954. Bois du forets de la Nouvelle Caledonie. SASTRAPRADJA, D. S., & C. LAMOUREUX. 1969. Variations in wood anatomy of Hawaiian Metrosideros Ann. (Myrtaceae). Bogor. 5: 1—83. E. Arn. SCHWEITZER, M. 1971. Comparative anatomy of Ulmaceae. J. Arb. 52: 523—585. SOLEREDER, H. 1899 & 1908. Systematische Anatomie der Dicotyledonen (& Erganzingsband). in STEENIS, C. G. G. J. VAN. 1962. The Land-Bridge Theory Botany. Blumea N: 235 —542. STERN, W. L. 1967. Index Xylariorum. Regnum Vegetabile 49. SUDO, S. 1959. Identification of Japanese Hardwoods. Bull. Govt. Forest Exp. Meguro, Tokyo, 118. Suss, H. 1967. Ober die LSngenanderungen der Parenchymstrange, Holzfasern und Gefassglieder von Laubholzer im Verlauf einer Zuwachsperiode. Holz als Roh- und Werkstoff 25: 369—377. & TABATA, H. 1964. Vessel element of Japanese birches as viewed from ecology and evolution. Physiol. Ecol. (Jap.) 12: 7—16. VERSTEEGH, CHR. 1968. An anatomical study of some woody plants ofthe mountain flora of the tropics (Indonesia).Acta Bot. Neerl. 17: 151—159. WALTHER, H., & H. LIETH. I960. Klimadiagramm-Weltatlas, I. P. BAAS: Wood anatomy oj Ilex 251 ADDENDUM While this the data became available the latitudinal and altitudinal paperwas in press on variation in 19 other widely distributed genera, studied by Mr. N. van der GraafFin our of these the and trends for institute. In most genera latitudinal altitudinal vessel member length are the same as in Ilex, so there are very strong arguments to regard these trends to of in the scalariform be general validity. However, seven genera possessing perforations, the number of bars did not show the same altitudinal and latitudinal trends as in Ilex; on six for the have the contrary: of the genera showed a trend tropical lowland species to the number of bars This fact what would if lowest per perforation. is in one expect, one considers the rare occurrence ofscalariform perforations in the woody flora ofthe tropical and also for and this that the lowland. For Ilex, perhaps Symplocos Viburnum, may imply possibly 'exceptional' latitudinal trends for scalariform perforations are, after all, of great Ilex the then phylogenetic significance within these genera. For wood anatomy would for lowland and derived for suggesta primitive status tropical species a more one tropical and and Mr. der GraafF's data will be the montane subtropical temperate species. Van subject of a separate publication. BLUMEA VOL. No. 252 XXI, 2, 1973 Plate I. Ilex. Vessel frequency, arrangement, distribution,and diameter, all x 75. 1. I. fargesii (UN 162), diffuse radial vessels porous, long multiples; 2. I. perado» (Bourgeau s.tt.), tending to semi-ring-porous, numer- in ous, mostly long radial multiples; 3. I. decidua (MADw 849), semi-ring-porous,high percentage ofsolitary diffuse of Ilex vessels; 4. I. macfadyenii (MADw 2090), porous, high percentage solitary vessels; 5. spec. diffuse 6. I. diffuse vessels wide. (climber, Jacobs 7928), porous; cymosa (KEPw 2337) y porous, 11. Plate Ilex. I —5. vessel perforations, all x 220. 1.I. umbellta (SJRw 43839), maceration, over 50 bars» 2. I. sideroxylodides (HM 2279—53), radial section, 38 bars; 3. I. serrata (SJRw 40816), maceration, 23 bars» I. radial I. radial 4- caroliniana (MADw 751), section, 15 bars; 5.. spec. (climber,Jacobs 7982), section, 12 widely spaced bars. 6—13. inter-vessel pits in tangentialsections. 6 & 7 I. rimbachii (MADw 16614),transitional in 8. I. R — and opposite pits same section, x 550; cymosa (SFCw 575 12), opposite, seemingly vestured pits, x 800; 9. I. chinensis (IF 17637), scalariform to transitional pits, x 350; 10. I. laurina ( MADw 21223), I. I. transitional pits, X 350; 11. macfadyenii (MADw 2090), opposite, small pits, x 350; 12. caroliniana (MADw 751), opposite to alternate, ± polygonal pits, x 350; 13. I. latifolia (MADw $053), opposite to alternate, circular pits, x 350. Plate SEM I. III. Ilex. 1 & 2. photographsin radial surface of chapaensis (CTFT 11009). !• half-bordered pits between parenchyma and vessel, note warts on overarching pit borders from vessel side, x 3300; 2. in vessel wall with and — radial weak spirals warty layer, x 1300. 3—9. vessel—ray pits section; 3—5, I. I. large and gash-like pits; 6—9, opposite and opposite to alternate. 3. anomala (BISHw 140), x 550; 4. I. martiniana 6. I. with spec, (climber,Jacobs 7928), x 350; 5. (LJw 4466 a), x 350; latifolia (MADw 5053), indicated 800; I. R some 'complex' unilaterally compound pits by arrows, x 7. cymosa (SFCw 575—12), x 8. I. macfadyenii (MADw 2090), x I. caroliniana (MADw 75i) x 350; 350; 9. t 350. Plate IV. Ilex. Presence and absence of spiral thickenings. All SEM photographs. 1 & 2. I. aquijolium in (IF 14473), x 310; i. transverse surface, broad spirals vessels; 2. oblique tangential surface, broad spirals in in I. faint vessels and narrow ones fibre-tracheids (arrow); 3. chapaensis (CTFT 11009), radial surface, spirals on vessel walls, x 290; 4. I. guianensis (SJRw 7565), vessel without spirals, fibre-tracheid with faint ~ vessel- spirals (arrow), x 290; 5. I. verticillata (SFCw R 206—14), radial oblique surface, spirals absent from and fibre-tracheids, scalariform perforations with few bars, x 290; 6. I. hypoglauca (S 30Ç17), spirals absent, scalariform perforations with numerousbars, x no. V. I. cinerea Plate Ilex. Fibre-tracheids. 1 —5. tangential sections. 1. (Ford s.n.),helical thickenings, x 800; in fibre- 2. I. chinensis (IF 17637), annular thickenings, x 800; 3. I. martiniana (Uw 4466 a), 'septa' (arrows) I. R tracheid; 4. verticillata (SFCw 206—14), tangential wall pits numerous and with conspicuous borders ~ wall with minute borders visible enclosing pit apertures, x 350; 5. I. cymosa (KEPw 2337), tangential pits (not in 6. I. photograph) and extended slit-like apertures, x 350. —6—8. transverse sections, x 350; rotunda (CSIRO 13255),thin-walled fibre-tracheids; 7 & 8..I. rimbachii (MADw 16614), thick-walled fibre-tracheids in same transverse section, stained with safranin and haematoxylin; 7. normal fibre-tracheids; 8. gelatinous (tension wood) fibre-tracheids. —6. I. I. Plate VI. Ilex. Rays and crystals. 1 tagential sections, x 65. 1. verticillata : (SFCw R 206—14); 2. caroliniatia (MADw 751); 3. I. martiniana (Uw 4466 a), with occasional sheath cells; 4. I. toluccana! (IF 19128), sheath cells and in I. I. canariensis abundant conspicuous ray on the right; 5. rimbachii (MADw 16614); <>• broad abundant. — I. radial section, (Bourgeau s.n.) t rays unusually 7—10. x 26$. 7. goshiensis (IF 2J2Ç1 ), in in chambered crystal thick-walled isioblast; 8 & 9. I. liukiuensis (MADw 12946); 8. radial section, crystals in thick-walled I. bulging ray cells; 9. tagential section, crystal idioblast; 10. laurifolia (Van Royen 3499), adial section, slightly elongate crystals in chambered ray cells. r P. BAAS Wood : anatomy ofIlex 253\I Plate I (for legends of plates sec p. 252) 254\II BLUMEA VOL. XXI, No. 2, 1973 Plate II P. BAAS: Wood anatomy ofIlex 255\III Plate III BLUMEA 256\IV VOL. XXI, No. 2, 1973 Plate IV P. Baas : Wood anatomy of Ilex 257\V Plate V BLUMEA VOL. 258\VI XXI, No. 2, 1973 Plate VI W. A. VAN HEEL: Scaphocalyx spathacea 259 Photo Scaphocalyx spathacea. — 1—3. C.s. of pistil in upward sequence. — Photo 4. Ovule, ± median — — 6 l.s. Photo 5. Ovule, ± transversel.s. Photo & 7. Ovule, subdistal c.s. showing separation of inner integument into four tips. 260 BLUMEA VOL. XXI, No. 2, 1973 8 13 15 9 — 8 & distal of — Photo Scaphocalyx spathacea Photo 9. Ovules, c.s. showing lobing outerintegument. Halved with of 10. ovary five ovules. — Photo 11 & 12. Young ovules. — Photo 13. Micropylarregion seed, showing tips of inner integument. Exostome out off section. — Photo 14. Micropylar region of lobes. Photo seed, showing ectostome and endocarp — 15. Lateral base ofseed, transverse l.s., showing the separation of inner cell layers of testa.