Harley 1991 Pollen Morphology Sapotaceae

The Pollen Morphology of the Sapotaceae
Author(s): Madeline M. Harley

Source: Kew Bulletin, Vol. 46, No. 3 (1991), pp. 379-491
Published by: Springer on behalf of Royal Botanic Gardens, Kew
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The pollen morphology of the Sapotaceae
MADELINE M. HARLEY
Summary.The pollen morphology of 398 species from all but five of the 53 genera of the Sapotaceae
defined by Pennington (1991) in his recent generic review, has been studied using light, scanning
and transmission electron microscopy. Within the family pollen diversity is sufficient for twelve
pollen types to be described, eleven of which have been further subdivided. A key to the pollen
types and further keys to the numerous subtypes that occur within Pollen Types I-VIII are
provided. In two Chrysophyllum species, C. marginatum subsp. marginatum and C. inornatum, the
structure of the pollen wall is anomalous within the Sapotaceae. The infratectum is comprised
of long, slender columellae whereas in all other species examined it is narrow, granular or
comprised of reduced columellae. In the other anomalous species, Englerophytum stelechantha,the
apertures are more or less porate and represent an extreme of the colporate condition observed
in other species in the family. The pollen types are considered in relation to Pennington's revision
of the family and the contribution that pollen morphology has made towards resolving taxonomic
uncertainties. A broad tribal correlation is observed between Mimusopeae with Pollen Types I &
III; Isonandreae with Pollen Type II; Sideroxyleae with Pollen Type VI; Chrysophylleae with Pollen
Type VII and Omphalocarpeae with Pollen Type VIII. There are examples of genera in which only
one pollen type or subtype occurs, although not exclusively: Burckella,Pollen Type I; Sarcosperma,
Pollen Type IV and Omphalocarpum, Pollen Type VIII. A few pollen subtypes are restricted to
only one genus: Delpydorasubtype VIIC; Sarcaulus,subtype XIA; Chromolucuma, subtype XIC and
Diploon, subtype XIIA. Pollen Type IX occurs only in Pouteria,sect. Oxythece.
Aperture configuration and the appearance of the tectum from scanning electron microscopy
have been found to be particularly useful in defining differences between pollen types. The wall
stratification, comprising a very thick tectum and foot layer intersected by a narrow infratectum,
is unusual. Both the functional rationale of this type of pollen wall and the occurrence of a
granular infratectum in both primitive and more highly evolved angiosperm families are con-
sidered. A variable number of (usually 3-4 or 4-5) regularly spaced apertures between pollen
grains within single samples is apparently normal within the Sapotaceae; possible reasons for this
are discussed.
Two major evolutionary trends are postulated, culminating either in 3-4-colporate, prolate
grains with continuous equatorial endexinous thickening and differentiated ornamentation, or
in 4-5-colporate, subprolate grains with discontinuous equatorial endexinous thickening and
undifferentiated ornamentation.
Pollination in the Sapotaceae is not well documented; possible vectors include insects, bats and
wind. Available data is summarized and discussed.
A number of fossil records for the Sapotaceae from the upper Cretaceous onwards are reviewed
and a summary diagram provided. Pollen characteristicsfrom selected, mainly Tertiary, records
for the family from many parts of the world are compared with the pollen types described in the
present paper. Some previously described affinities of fossil pollen with recent sapotaceous pollen
types are considered doubtful, in particular an early record from the upper Cretaceous.
An appendix provides, from published data and personal observations, the main characteristics
of pollen in four families allied or associated with the Sapotaceae:Ebenaceae, Myrsinaceae, Styracaceae
and Symplocaceae. Characteristics of pollen in two other families: Burseraceaeand Meliaceaeless
closely associated with Sapotaceae
but with pollen that often shows similar characteristics, are also
summarized. Features by which the various grains can be distinguished from those of Sapotaceae
are highlighted.
Two tables provide data for; 1. Correlation of pollen types with Pennington's systematic
arrangement and 2. A list of all species examined, arranged systematically and providing full
pollen data in condensed form.
Accepted for publication December 1990.
379

380 KEW BULLETINVOL. 46 (3)
CONTENTS
Introduction 381
Materials and Methods 382
Terminology 383
General Pollen Morphology 384
Pollen Types 385
Key to Pollen Types 385
Descriptions of Pollen Types 386
Pollen Type I-'Mimusops' 386
Pollen Type II-'Palaquium' 389
Pollen Type III-'Manilkara' 391
Pollen Type IV-'Sarcosperma' 392
Pollen Type V-'Pichonia' 394
Pollen Type VI-'Sideroxylon' 396
Pollen Type VII-'Pouteria caimito' 398
Pollen Type VIII-'Pouteria alnifolia' 401
Pollen Type IX-'Oxythece' 403
Pollen Type X-'Elaeoluma' 403
Pollen Type XI-'Sarcaulus' 404
Pollen Type XII-'Diploon' 405
Anomalous Pollen Types 406
Anomalous Pollen Type I 406
Anomalous Pollen Type II 406
Discussion 407
General characteristics of pollen 407
Comparison of pollen within systematic groups 407
Pollen morphology as a taxonomic character 410
Pollen morphological considerations 411
Evolutionary trends 414
Pollination 416
Fossil pollen 417
Acknowledgements 420
Appendix-Pollen of related and/or similar families 420
References 424

THE POLLEN MORPHOLOGY OF THE SAPOTACEAE 381
INTRODUCTION
The Sapotaceae are a fairly large, very natural pantropical family of trees,
shrubs and, rarely, climbers comprising approximately 1100 species. The
flowers may be solitary but in many species are borne in simple fasciculate
inflorescences. The position of the fascicle varies from axillary to ramiflorous
or cauliflorous. They are usually hermaphrodite, regular and generally small.
The corolla is frequently white or cream and sometimes sweetly scented. The
leaves are alternate, simple and almost always entire, and in some species are
clustered in a terminal whorl. The fruits are berries, generally with a thin
outer layer and a soft, juicy or dryish pulp surrounding the seeds. The fruits
are borne on short stalks or, in Omphalocarpum,sessile on the main trunk. The
slashed bark exudes a milky latex, a useful field identification character in
a family whose members are frequently tall forest trees. Hooker (1854),
Harrison et al (1969), Isawumi (1978), Whitmore (1978), Kuruvilla (1989)
and Brficher (1989) discuss and/or summarize economically useful products
from this family.
Within the family, genera are very difficult to circumscribe and since the
family was first described by Endlicher (1836) with twelve genera, there have
been numerous revisions. Subsequent systematists, in particular De Candolle
(1844), Bentham & Hooker (1876), Hartog (1878, 1879), Radlkofer (1887) and
Engler (1890), considered the most significant macromorphological features
for distinguishing genera or groups of genera to be floral; the arrangement of
the sepals, the presence or absence of staminodes, the number and nature of
the stamens, the degree of fusion and position of stamens in relation to the
petals, the number of petals in relation to the number of sepals, and whether
the corolla lobes are simple or divided.
Until the end of the nineteenth century the number of genera of Sapotaceae
described did not exceed thirty-four. In 1892, however, Baillon described
sixty-four genera and also introduced seed characters into the classification of
the family for the first time. Seed scar position and size, and the presence or
absence of endosperm, were particularly important in his classification. In the
present century, Dubard (1912 & 1915), following the unfinished work of
Pierre (1890-1), published yet another classification in which the number of
genera reverted to thirty-four. This treatment was followed by Baehni (1938)
and Lam (1939). Aubreville (1964), devised a system in which he described
one hundred and twenty five genera. Hemsley (1968) argued against exces-
sive subdivision of natural groups. The most recent revision of the family
(Pennington 1991) differs from previous treatments because it uses not only
traditional characters but also features such as chemistry, cytology, wood
anatomy and pollen morphology.
Generic delimitation in the family is difficult because there is so much overlap
between characters. To quote Hemsley (1968), 'The relationship between this
promising array of characters is nonetheless markedly reticulate, so that the
distinction between many genera is often reduced to a single character and,
further, particularly in the number of flower-parts and development of
staminodes or endosperm, varies in some parts of the family between apparently
closely related species and even within a single species'
In the present study the potential of pollen morphology as a useful additional
character for classification has been examined. Almost 400 species of the
family have been studied. However, like all other characters the systematic

382 KEWBULLETINVOL. 46 (3)
contributions of pollen morphology are erratic. Overall, the palynological results
reflect many of Pennington's taxonomic conclusions and there are a number
of instances where pollen morphology is particularly well-correlated with his
taxonomic groupings. For example the pollen morphology of the New World
genera Bumelia, Dipholis and Mastichodendron and that of the pantropical Sideroxylon
is very similar, supporting his decision to combine these taxa. Within the large
genus Pouteria,Pennington includes the genus OxytheceMiq. However, its pollen
morphology is sufficiently distinctive to influence his maintenance of Oxythece
at sectional rank. Pollen morphology has also provided good evidence for
upholding the generic status of Sarcaulus, a genus not clearly distinguishable
from Pouteria on the basis of macromorphological characters alone.
Accounts of the pollen morphology of Sapotaceaeare few (Harley 1986a;
Harley 1990a and in Pennington 1991). A few species are described in regional
pollen floras, for example: Tsukada (1964), a general survey of mainly south
American tropical species from a wide range of families; Correa-Gomes (1966),
Brazilian caatinga; Straka, Simon & Cerceau-Larrival (1967), Madagascar and
the Mascarenes; Bonnefille (1971), Ethiopia; Heusser (1971), Chile; Huang
(1972), Taiwan; Sowunmi (1973), Nigeria; Caratini (1974), tropical Africa;
Gupta & Sharma (1977), India; Markgraf & D'Antoni (1978), Argentina; and
Lieux (1982), south-eastern U.S.A. In addition, a few authors have examined
one or a few sapotaceous species in various contexts such as broad palynological
surveys (Erdtman 1952), pollen morphology of a single species combined with
either anatomy (Ayensu 1972, Barry & Perez de Paz 1979), embryology
(Bhatnagar & Gupta 1970) or taxonomy (Barth 1973).
MATERIALS AND METHODS
Pollen from 398 species (472 specimens), representing all but 5 of the 53
genera comprising the Sapotaceae in Pennington's (1991) revision, have been
examined. Material from the remaining 5 genera (Baillonella Pierre, Labramia
A.DC., LecomteodoxaPierre ex Dub., GluemaAubr. & Pellegrin and Eberhardtia
Lecomte) was unobtainable during the study. The Old World specimens studied
were from the collections in the Herbarium of the Royal Botanic Gardens, Kew
(K), but much of the New World material was from other herbaria (see
Pennington (1990), for details). Mature buds were stored in glacial acetic acid
for a limited period, not exceeding 3 months or briefly (2-3 days) in water
in a refrigerator. Anthers, which are frequently less than 1mm long, were
dissected out of the soaked buds.
To prepare the material for subsequent examination, anthers were broken
open in distilled water to release the pollen. The fragmented anthers and loose
pollen were washed into centrifuge tubes with distilled water, centrifuged
and decanted. Glacial acetic acid was then added and the samples were re-
centrifuged. The acid was then decanted prior to acetolysis using a standard
combination of 10% sulphuric acid and 90% acetic anhydride (Erdtman,
1960). The samples were held in this mixture in a boiling water bath for 2-4
minutes and then thoroughly washed. After the second washing following
acetolysis, samples were passed through suitably graded mesh sieves (50-
100 sm) to remove any extraneous plant material. The prepared samples were
then divided to provide material for light microscopy (LM), scanning electron
microscopy (SEM) and transmission electron microscopy (TEM).
For LM the washed pollen was transferred into 50% glycerol solution,

centrifuged, decanted and mounted in glycerol jelly. Measurements from 10
well-expanded grains were collected using a Nikon L-Ke microscope with a
x 100 oil immersion objective. Photomicrographs were obtained using a Nikon
microflex model AFM automatic photomicrograph attachment. Slide prep-
arations of all specimens examined are deposited in the pollen slide collection
of the Royal Botanic Gardens, Kew.
Pollen for SEM was suspended in 95 % ethanol, pipetted onto clean, polished
aluminium stubs and left to dry at room temperature for c. 24 hours under
an inverted flask. In a few samples there was a tendency for the pollen grains
to collapse when air dried and in some instances improved results were obtained
by dehydrating in a Balzer's Union CPD 010 critical point dryer. The drying
liquid was CO2 and the transfer liquid, absolute ethanol. Following critical
point drying, a fine artist's brush was used to transfer the pollen onto stubs
previously covered with double-sided adhesive tape. Prepared stubs were coated
with c. 20nm of platinum evaporated in a Balzer's Union SCD 030 sputter
coater, and pollen was examined in a Jeol JSM 35 SEM.
For TEM washed pollen was pre-stained and embedded using the techniques
described by Skvarla (1966). Ultrathin sections were cut using a diamond knife
fitted into a Cambridge-Huxley ultramicrotome or, later in the study a Reichert
Ultracut. Sections were collected on 200 hexagonal mesh copper grids and
stained in 5 % aqueous uranyl acetate for 15 minutes, followed by 5 minutes
in lead citrate (Venable & Coggeshall, 1965). Later in the study excellent results
were also obtained with an LKB 2168 Ultrostainer, using LKB stabilised
solutions of uranyl acetate and of lead citrate. The standard timing and
temperature settings specified by the manufacturer were used. A Jeol 100S TEM
and, later in the study, an Hitachi H300 were used for examination. Photo-
graphic negatives of all specimens examined with SEM and TEM are in the
pollen negative collection of the Royal Botanic Gardens, Kew.
Comparisons and descriptions of tectum characteristics are from SEM
micrographs of the mesocolpial area. For pollen wall stratification and charac-
teristics, TEM micrographs of thin sections and SEM micrographs of fractures
from the apocolpial region of the pollen wall were used.
Terminology
In general, terminology in this paper follows that of Faegri & Iversen (1989)
and Erdtman (1952), including the terms 'lolongate' meaning longitudinally
elongated and 'lalongate' meaning transversely elongated. Apertures present
only in the endexine are described as 'endoapertures' by Van Campo
(1958). 'Hemispherical', 'sub-hemispherical' and 'pointed' (Thomson &
Pflug, 1953) are used to distinguish the shape of the polar region in descrip-
tions of some of the subtypes. The terms 'protrudent' or 'semi-protrudent'
(Harley 1990a) were introduced to describe the particular form of the tectum
surrounding the endoaperture in the pollen grains of some of the species in
this family, as no suitable term was found in the existing literature. With rare
exceptions the endexinous thickenings underlying the tectum in the region of
the apertures are not sufficiently abrupt to constitute costae sens. str. The
terms 'costa', 'costae' or 'costate' are used to describe the more localized
endoapertural thickenings in the pollen of a few of the species included in
Pollen Types V and XI only. 'Microfossulae' (see Harley, 1990b) are defined
as 'elongate indentations not penetrating through to the infratectum'. For
some descriptions of surface ornamentation as seen with SEM, the text refers
the reader to plate figures.

GENERAL POLLEN MORPHOLOGY
Number of specimens examined: 472

Number of species: 398
The pollen grains are single, isopolar, angulaperturate or, less frequently
in some of the species with 4-colporate grains, planaperturate (Figs 17: F &
N and 34: E). The amb is more or less circular or modified according to the
number of apertures present. For example, tricolporate grains tend to appear
triangular (Figs 28: F and 40: L), 4-colporate, square (Fig. 22: M) and
5-colporate, pentagonal (Fig. 20: L). Rarely oblate-spheroidal, occasionally
spheroidal, predominantly prolate-spheroidal, subprolate or prolate. Poles
hemispherical (Fig. 5: B & C), sub-hemispherical (Fig. 24: B, C & F) or
pointed (Fig. 14: D & E). 3, 3-4, 4, 4-5 or, rarely, only 5-colporate. Very
rarely some 6-colporate grains occur in samples with mainly 4- 5-colporate
grains. Average polar length, 16 -0 (Pouteria sagotiana)- 92 -2 sm (P. speciosa).
The colpi are usually narrow, occasionally broad, and range in length from
0 -15 to 0 -85 polar length. They are provided with a tough, usually granular,
Fic. 1. To show the two extremes of aperture configuration in Sapotaceae pollen in transverse
section. A. Protrudent tectum overlying a, usually narrow, endoaperture. Particularly associated
with Pollen Type VII (the endexine is continuous in the equatorial region). B. Non-protrudent
tectum, the, usually broad, endoaperture is covered by a layer of endexinous material on which
fragments of ectexine are often apparent (the endexine thins gradually from the endoapertural
region). Slight variations of this aperture form are encountered in most of the pollen types,
excluding Pollen Type VII.

membrane which does not rupture during acetolysis, except in the endo-
apertural area, where there is no underlying endexine. The endoapertures are
narrowly lalongate (polar length less than half equatorial breadth (Fig. 34: B))
or broadly lalongate (polar length more than half equatorial breadth (Fig. 6:
H &J)), less frequently circular (Fig. 10: C & D) or, extremely rarely broadly
lolongate. They are described, in proportion to equatorial width, as 'average'
- -
(c. 0 15 equatorial width or less (Fig. 11: E - F)) or 'large' (c. 0 16 equatorial
width or more (Fig. 8: A - B)). The tectum in many species is protrudent (Fig.
28: L and Fig. 1A) or semi-protrudent; this is usually associated with narrow
endoapertures and vestigial (c. 0 -15 polar length) or short (up to 0-65 polar
length) colpi. The wall thickness at the poles is 1 -0-3 -0 (-5 -0) Jm. At the
-
equator the wall thickness is 1 0- - 5 0 (-8-0) ttm. Throughout the family, with
rare exceptions, the ectexine comprises a thick tectum, a very narrow infra-
tectum and a foot layer similar in thickness or slightly thicker than the tectum.
The infratectum is either granular or with very reduced columellae which
may be interspersed with granules. The slight increase in the width of the
infratectum at the poles correlates with a small reduction in the thickness of
the tectum and/or foot layer, so that a more or less constant ectexine thickness
is usually maintained. The endexine is thickened, particularly in the area
surrounding the endoapertures and underlying the colpi. In some species it
is absent or very thin in the mesocolpial areas (Fig. 10: H -J), while in others
it may be slightly thinner than, or as thick as, the endoapertural area. The
result is a continuously thickened band of endexine in the equatorial region
(Fig. 30: H -J). The thickness of this band is reduced towards the polar
regions where it is completely absent. The tectum is usually sparsely punctate
or perforate (Fig. 7: A). The perforations, which sometimes occur in shallow
pits (Fig. 7: G), may be more dense and/or coarse in either the mesocolpia
or, more frequently, the apocolpia (Fig. 10: A). Microfossulae are recorded
for a few species (Fig. 17: C & G). In some species the entire surface, seen
with SEM, appears psilate (Fig. 37: E), subpsilate, scabrate (Fig. 16: D & E),
finely, and frequently regularly, granular (Fig. 5: G) or coarsely, and often
less regularly, granular (Fig. 5: F), anastomosed granular (Fig. 5: H), finely
striate-rugulate (Fig. 10: L), striate (Fig. 24: A & D), low-relief, angular-
rugulate (Fig. 29: N) or weakly (Fig. 31: L) to coarsely rugulate (Fig. 27: J).
In other species there is clear differentation between the more or less smooth
apocolpia and the weakly or distinctly rugulate mesocolpia (Fig. 27: H & K).
POLLEN TYPES
Twelve pollen types are recognised. Eleven of these have been subdivided
to produce a total of 49 subtypes. Some subtypes are distinguished from
similar forms on only one character. In some instances size or shape have
been found to be useful in picking out a group of species that form sections
within genera.
KEY TO POLLEN TYPES
1. Average polar length not exceeding 35

"0 /m:
2. Supratectal spines, spinules or coarse granules present:
3. Tectum protrudent, spinules or coarse granules present ........ XI
3. Tectum not protrudent, spines present ..................... XII

2. Supratectal spines, spinules or coarse granules absent:
4. Pollen grains 3 -4-colporate, perforate or coarsely perforate and rugulate
or coarsely rugulate, perforations often coarser in apocolpium, up to
35-0 Am average polar length ............................... X
4. 3- 5-colporate, tectum striate, up to 25 0 Am average polar length IX
1. Average polar length usually exceeding 35-0 Mm:
5. Endexine thickened in mesocolpia forming a continuous band in equatorial
region:
6. Apocolpium and mesocolpium differentiated, tectum protrudent VII
6. Apocolpium and mesocolpium not differentiated, tectum not, or semi-
protrudent ........................................ VIII
5. Endexine thin or absent in mesocolpia, not forming a continuous band
in equatorial region:
7. Poles always hemispherical:
8. Occasionally subspheroidal or spheroidal, up to 65 -0 Am average polar
length:
9. Mesocolpial wall thickness up to 3-0, rarely 5-0 Am. Costae not
present, average polar length 0 m .............. IV
9. Mesocolpial walls usually 3- 0 -24"-0-57-0 -0 m. Costae frequently present,
6"Mm ....................... V
average polar length 33-0-65-0
8. Usually prolate-spheroidal or subprolate, up to 85 - 0 Am average polar
length:
10. Entire surface of tectum usually finely or distinctly granular .. I
10. Entire surface of tectum usually finely striate-rugulate ...... II
7. Poles not always hemispherical, colpi frequently up to 0 -85 polar length:
11. Equatorial faces slightly convex, tectum usually psilate, subpsilate or
scabrate, poles frequently pointed ........................ III
11. Equatorial faces usually parallel, tectum frequently striate, poles
frequently sub-hemispherical ............................. VI
DESCRIPTIONS OF POLLEN TYPES
The number of specimens examined for each genus within a pollen type is
given in parentheses after each genus in the generic list that precedes each
pollen type description. Under main pollen type descriptions, genera listed in
bold type are those for which pollen of all species examined falls within that
type.
The description for each pollen type outlines the shared characteristics
of the pollen from all species included in that pollen type. Descriptions for
subtypes generally only include the characters used to distinguish the subtypes
from each other.
POLLENTYPE I. 'MIMUSOPS'.
Number of genera included: 14-Burckella (5), Diploknema(1), Madhuca (25),
Manilkara (13), Mimusops (13), Neolemonniera(1), Northia (1), Palaquium (13),
Payena (8), Pichonia(1), Sideroxylon(1), Synsepalum(2), Tsebona (1), Xantolis (3).
Pollen grains prolate-spheroidal or subprolate, infrequently prolate, rarely
spheroidal. Poles hemispherical, equatorial faces convex. 4, 4-5 (3, 3-4 or
6)-colporate. Average polar length, 31 0 -78 5 Colpi 0- 65-0 85 polar
/m.

length. Endoapertures usually broad occasionally narrow. Wall thickness,

- - - - -
poles: 1 0 - 4 0 (5 0) Am; equator: 2 0 -- 5 0 (6 0) lm. Infratectum narrow or
very narrow, granular or with reduced columellae. Endexine thin or absent in
mesocolpia, thickened in apertural region. Tectum not protrudent, occasionally
coarsely rugulate, apocolpia may be coarsely perforate; surface fine or coarse
granular, distinct or anastomosed or, occasionally psilate or scabrate.
Key to subtypes of Pollen Type I

1. Entire surface of tectum psilate, subpsilate or scabrate ............ IE
1. Entire surface of tectum finely or distinctly granular:
2. Tectum coarsely rugulate:
3. Colpi long, c. 0-85 polar length, grains prolate-spheroidal or sub-
prolate ........................................ IF
3. Colpi short, c. 0-65 polar length, grains + spheroidal ......... IG
2. Tectum not coarsely rugulate:
4. Average polar length not exceeding 60 0 Am:
5. Apocolpia coarsely perforate .............................. ID
5. Apocolpia not coarsely perforate ........................... IA
4. Average polar length up to 75-0 Am:
6. Endoapertures very wide in equatorial direction. Infratectum, granular,
unusually wide ........................................ Ic
6. Endoapertures average width. Infratectum granular or reduced col-
umellate, narrow ........................................ IB
SubtypeIA. (Figs 5: A-L; 6: A-M)

Pollen grains usually prolate-spheroidal or subprolate, infrequently prolate.
4, 4-5, rarely, 3, 3-4 or 6-colporate. Average polar length, 31 -0-57-2 2Am.
Tectum not coarsely rugulate, surface distinct fine or coarse granular or
anastomosed granular.
Species included: Mimusopeae.:Mimusopinae;,Mimusops aedificatoriaMildbr.,
M. angel Chiov., M. caffra E. Mey., M. fasciculata Warb., M. fruticosa Boj., M.
kummel Bruce ex A.DC., M. obovata Sond., M. seychellarum(Oliv.) Hemsley.
Manilkarinae;,Manilkara cuneifolia Dub., M. dawei (Stapf) Chiov., M. hexandra
(Roxb.) Dub., M. jaimiqui (Wright) Dub. subsp. emarginata(L.) Cronq., M.
jaimiqui (Wright) Dub. subsp. haitensis (Cronq.) Cronq., M. letestui Aubrev.
& Pellegr., M. roxburgii (Wight) Parker, M. samoinsis Lam & Meeuse, M.
subsericea(Mart.) Dub., M. vitiensis (Lam & Olden) Meeuse, Northia seychellana
Hook. f. Glueminae; Neolemonniera clitandrifolia (Chev.) Heine. Isonandreae;
Palaquium dasyphyllum (Vriese) Pierre ex Dub., P. ellipticum (Dalz.) Baill.,
P. rioense Lam, P. rivulare Lam, P. walsuraefolium Pierre ex Dub., Madhuca
aristulataLam, M. betis Macbride, M. elmeriMerrill ex Lam, M. korthalsiiLam,
M. malaccensis Lam, M. microphylla(Hook.) Alston, M. mindanaensis Merrill,
M. pubicalyx Ridley, M. sericea Lam, M. spectabilis Royen, M. stipulacea
Fletcher, M. subquincuncialis Lam, M. sp. (Pennington et al. 10234), Payena
lanceolataRidley, P. lucida (Wall. ex Don) Baehni, P. microphyllaBurck, Burckella
banikiensis (Royen) T. D. Penn., B. macropoda(Krause) Lam. Sideroxyleae;
Sideroxylon oxycanthum Baill. Chrysophylleae,;Xantolis assamica (C. B. Clarke)
Royen, X. burmanica(Coll. & Hemsl.), X. embeliifolia (Merrill) Royen.

SubtypeIB. (Figs 7: A-D, F-G; 8: F)
Pollen grains prolate-spheroidal or subprolate, rarely prolate. 4, 4-5, rarely,
3 - 4-colporate. Average polar length, 56 - 2 -72 - 0 tm. Colpi 0 -65, rarely up
to 0 -85, polar length. Tectum not coarsely rugulate, surface distinctly granular
or anastomosed granular. Very similar to sybtype IA, apart from the larger
pollen size.
Species included: Isonandreae;Palaquium burckii Lam, P. grande (Thwaites)
Engl., P. ottolandieriKoord. & Valet., P. sp. (Jayasuriya 1920), P. sp. (Soepadmo
& Mahmud 1115), P. sp. (SAN 34407), Payena dasyphylla (Vriese) Pierre ex
Dub., P. endertii Lam, P. maingayi Clarke, P. obscuraBurck.
SubtypeIC. (Fig. 8: A-B, E)

Pollen grains subprolate. 4-5-colporate. Average polar length, 74-5 Jim.
Colpi long, c. 0-85 polar length. Endoapertures broad, unusually wide.
Tectum not coarsely rugulate, surface finely and irregularly granular. Similar
to subtype IB but colpi longer and endoapertures larger. Furthermore thin
sections show the infratectum to be broader and more densely granular in
Tsebona than in species attributed to subtype IB.
Species included: Omphalocarpeae;Tsebona macranthaCap.
Subtype ID. (Fig. 10: A-F)

Pollen grains prolate-spheroidal or subprolate, less frequently prolate. 4,
4-5 (3-4)-colporate. Average polar length, 33-1-55-0 sm. Colpi 0-65-
0 - 75 polar length. Tectum not coarsely rugulate, apocolpia coarsely perforate;
surface distinctly granular or anastomosed granular, occasionally finely
granular or psilate.
Species included: Mimusopeae:Mimusopinae; Mimusops antongilensisAubrev.,
M. dispar N.E. Br., M. frondosa Hiern., M. marginataN.E. Br. Manilkarinae;
Manilkara valenzuelana(A. Rich.) T. D. Penn. Isonandreae;Madhuca decipiens
Sincl., M. laurifolia Lam, M. leucodermis(Krause) Lamand, M. sp. (Kostermans
10052). Chrysophylleae; (Welwitsch) T. D. Penn., S. gabonense
Synsepalumcerasiferum
(Aubrev. & Pellegr.) T. D. Penn.
Subtype IE. (Figs 7: E; 8: C-D)

Pollen grains prolate-spheroidal, 4-5-colporate. Average polar length,
62 - 3-78 - 5 sm. Tectum not protrudent, surface scabrate or psilate.
Species included: Mimusopeae: Manilkarinae; Manilkara littoralis Dub.

Isonandreae;Madhuca motleyana (Vriese) Baehni, M. pierrei Lam (Haviland &
Hose 3481).
SubtypeIF. (Fig. 9: A-G)

Pollen grains prolate-spheroidal or subprolate. 3-4, 4, 4-5(6)-colporate.
Average polar length, 41 4 Vm.Colpi long, 0" polar length.
"5-61 "4 75-0"85
Tectum coarsely rugulate; surface distinctly granular or anastomosed granular.
Species included: Mimusopeae: Mimusopinae; Mimusops riparia Engl. Manil-
karinae; Manilkara pubicarpa Monachino, M. triflora (Allemao) Monachino.

Isonandreae;PalaquiumpolyanthumEngl., Madhuca bourdilloniiLam, M. burckiana

Lam, M. longifolia (Koenig) Macbride, M. pierrei Lam (Kerr 4352), Payena
acuminata(Blume) Pierre, Burckellafijiensis(Hemsl.) A. C. Smith & Darwin, B.
obovata (Forster) Pierre, B. parviflora A. C. Smith & Darwin, Diploknema
krabiensis(Aubrev.) T. D. Penn. Chrysophylleae;Pichonia calomeris(Baill.) T. D.
Penn.
SubtypeIG. (Fig. 9: A-G)

Pollen grains more or less spheroidal. 4-5 or 5-colporate. Average polar
- 1 m. Colpi c. 0-65 polar length. Tectum coarsely
length, 47 0 and
66"-
rugulate; surface granular or finely striate-rugulate.
Species included: Isonandreae;Palaquium obovatum(Griffith) Engl., Madhuca
neriifolia (Moore) Lam.
POLLEN TYPE II. 'PALAQUIUM'.

Number of genera included: 12-Aulandra (1), Diploknema(1), Inhambanella(1),
Isonandra (4), Labourdonnaisia (2), Letestua (1), Madhuca (10), Manilkara (2),
Palaquium (20), Sideroxylon(4), Synsepalum(2) and Tieghemella(1).
Pollen grains subprolate or prolate-spheroidal, subspheroidal or prolate.
Poles hemispherical, equatorial faces convex. 4, 4-5, rarely 3-colporate.
Average polar length, 27 7 - 84 -7 [m. Colpi 0 -65-0 -75 (0 -85) polar length.
Endoapertures usually broad, occasionally narrow. Wall thickness, poles:
1 -0-4- 0 m; equator: 20- 5 0 (6-0) .tm. Infratectum narrow or very narrow,
granular or with reduced columellae. Endexine thin or absent in mrresocolpia
thickened in apertural region. Tectum rarely protrudent, occasionally distinctly
perforate, rarely finely reticulate; surface finely striate-rugulate, infrequently
psilate or subpsilate. Apocolpia and mesocolpia occasionally show differences
in surface characteristics; apocolpia occasionally coarsely perforate or
foveolate, psilate or subpsilate, mesocolpia infrequently rugulate or low-relief
rugulate.
Key to subtypesof Pollen Type II

1. Average polar length 27 -0-62-0 0m.
1. Average polar length 59-0-85-0 sm .......................... IIB
2. Apocolpia and mesocolpia differentiated.
3. Pollen prolate-spheroidal or subprolate, never prolate, colpi 0 - 65 -0 - 75
polar length ............................................. IIc
3. Pollen prolate or subprolate, never prolate-spheroidal, colpi 0 -75-0 -85
polar length ............................................. IID
2. Apocolpia and mesocolpia not differentiated ................... IIA
SubtypeIIA. (Figs 10: G-L; 11: A-H; 12: C-E)

Pollen grains subprolate or, less frequently prolate-spheroidal, infrequently
prolate, rarely subspheroidal. 4, 4-5, rarely 3-4 or 5-colporate. Average
polar length, 29 2 - 61 7 sm. Colpi 0 65-0 75 (0 85) polar length. Endo-
apertures usually broad. Tectum not protrudent. Endexine thin or absent in
mesocolpia. This subtype is very similar to Pollen Type IA except that the

surface of the tectum is finely striate rugulate.
A few species of Madhuca included in this subtype have coarsely perforate
or foveolate apocolpia and, in one species, M. tubulosa, the tectum is finely
reticulate.
Species included: Mimusopeae:Manilkarinae;Manilkara discolor(Sond.) Hemsl.

(Williams 663), M. staminodella Gilly, Labourdonnaisia calophylloidesBoj., L.
revoluta Boj. Isonandreae;Palaquium calophyllum (Teysm. & Binn.) Pierre, P.
fijiense Pierre ex Dub., P. galactoxylum (F. Muell.) Lam, P. gutta (Hook. f.)
Baill., P. hexandrum(Griffith) Baill., P. hornei (Hartog) Dub., P. leiocarpum
Boerl., P. multiflorumPierre ex Dub., P. philippenseC. B. Robinson, P. pseudo-
rostratumLam, P. rostratum(Miq.) Burck, P. rufolanigerumRoyen, P. sericeum
Lam, P. stipulare Pierre ex Dub., Madhuca coriaceaMerrill, M. costulata Lam,
M. curtisii Ridley (Jacobs 5011), M. kingiana (Brace) Lam, M. orientalis(Assem)
T. D. Penn., M. palustris Royen, M. tubulosa Lam, M. sp. (Pennington et al.
10250), Diploknema butyraceoidesLam (Kanjilal 3137). Chrysophylleae;Synsepalum
dulcificum (Schum. & Thonn.) Dub., S. glycydorum(Engler) T. D. Penn.
SubtypeIIB. (Fig. 12: A-B)

Pollen grains prolate-spheroidal or subprolate. 4-5 or (Palaquium sp.)
- - -
3-colporate. Average polar length, 59 0 - 84 7 Am. Colpi 0 65 -0 75 polar
length. Tectum: apocolopia foveolate or, conversely, mesocolpia foveolate;
surface psilate or subpsilate.
Species included: Isonandreae;Palaquiumcochlearifolium

Royen, P. sp. (Streimann
& Helean NGF 18338), Madhuca obovatifolia Merrill.
SubtypeHIC. (Fig. 12: F-L)

Pollen grains prolate-spheroidal or subprolate. 4-5 or 5-colporate. Average
- - - -
polar length, 42 6 - 53 5 m. Colpi 0 65 - 0 75 polar length. Endoapertures
broad or narrow. Tectum: mesocolpia rugulate, apocolpia psilate.
Species included: Isonandreae;Palaquium formosanum Hayata, P. lobbianum

Burck, P. neo-ebudicumGuill., Madhuca pallida (Burck) Baehni.
Subtype IID. (Fig. 13: A- L)

Pollen grains subprolate or prolate. 4, 4-5, rarely 3-4-colporate. Average
- - - -
polar length, 27 5 -51 7 ~m. Colpi long, 0 75 -0 85 polar length. Tectum
occasionally slightly protrudent, apocolpia psilate or subpsilate, mesocolpia
finely striate-rugulate or fine, low-relief rugulate. In some species tectum
distinctly perforate.
Species included: Mimusopeae: Mimusopinae; Tieghemella africana Pierre.

Manilkarinae; LetestuafloribundaLec. Glueminae;Inhambanellahenriquesii(Engl. &
Warb.). Isonandreae;Palaquium ridleyi King & Gamble, Aulandra longifolia Lam,
Isonandra lanceolataWight, I. perrottetianaA.DC., I. stocksii C. B. Clarke, IL
villosa Wight. Sideroxyleae;Sideroxylonmascatense(A.DC.) T. D. Penn., S. inerme
L., S. leucophyllumWatson, S. wzghtianumHook. & Arn.

POLLEN TYPE III. 'MANILKARA'.

Number of genera included: 17-Chrysophyllum (4), Englerophytum(1), Faucherea
(1), Inhambanella (1), Madhuca (1), Manilkara (17), Mimusops (2), Neohemsleya
(1), Neolemonniera(1), Nesoluma (1), Niemeyera(2), Palaquium (1), Pouteria (9),
Pradosia (2), Sideroxylon(2), Synsepalum(4), Xantolis (1).
Pollen grains prolate-spheroidal, subprolate or prolate, infrequently

spheroidal. Poles hemispherical or pointed, equatorial faces slightly convex.
3, 3-4, 4, 4-5, rarely 6-colporate. Average polar length, 8/m.
Colpi 0-75-0 85 polar length, frequently giving the poles22-9-55"8
of dehydrated
grains a somewhat pointed appearance. Endoapertures broad, frequently
more or less circular or narrow. Wall thickness, poles: 1 -0 -2 -5 (3 -5) /m;
equator: 1 0- 3 -0 (5 m. Infratectum granular or with reduced columel-
lae. Endexine thin or "-0)absent in
mesocolpia, thickened in apertural areas.
Tectum not protrudent; surface psilate, subpsilate, scabrate, less frequently
finely granular, irregularly granular or finely striate-rugulate.
Key to subtypesof Pollen Type III

1. Average polar length 16-0-45-0 jim. Pollen grains may be prolate:
2. Pollen grains 4, 5, rarely 6, never 3-colporate, endoapertures often circular,
never narrow ............................................ IIIA
2. Pollen grains 3, 4, never 5 or 6-colporate, endoapertures narrow or broad,
never circular:
3. Tectum with coarse perforations .......................... IID
3. Tectum without coarse perforations ........................ IIIc
1. Average polar length 29-56 jim. Pollen grains never prolate .... IIIB
Subtype liIA. (Fig. 14: A- L)

Pollen grains prolate-spheroidal or subprolate, 1 species, Manilkara multi-
nervis, prolate. 4, 4-5 or 4- 6-colporate. Average polar length, 24 7-45-0
jim. Colpi long, c. 0-85 polar length. Endoapertures frequently more or less
circular. Tectum, surface scabrate, subpsilate, psilate or, occasionally, finely
granular.
Species included: Mimusopeae:Mimusopinae;Mimusops elengi L.. Manilkarinae;
Manilkara bidentata (A.DC.) Chev. subsp. surinamensis (Miq.) T. D. Penn.
(Ducke 2501 & 1523), M. butugiChiov., M. concolor(Harv. ex Wright) Gerstn.,
M. discolor (Sond.) Hemsl. (Chase 4559 & 4664), M. excelsa (Ducke) Standl.,
M. hoshinoi (Kaneh.) Royen, M. huberi (Ducke) Chev., M. mochisia (Bak.)
Dub., M. multifida T. D. Penn. M. multinervis(Bak.) Dub., M. obovata(Sabine
& Don) Hemsl., M. sulcata (Engl.) Dub. Chrysophylleae;Xantolis tomentosa
(Roxb.) Raf.
SubtypeIIIB. (Figs 14: M-P; 15: A-M)

Pollen grains prolate-spheroidal, subprolate or, infrequently, spheroidal. 4,
-
4 5 (3- 4)-colporate. Average polar length, 29 5 (Englerophytum),34 7 &
/m
35 2 (Synsepalumbrevipesand S. afzelii) or, more commonly, 40 00- 55 88tm.
tsm
Tectum sparsely perforate; surface psilate or scabrate or, less frequently, very

finely striate-rugulate.
Species included: Mimusopeae: Mimusopinae; Mimusops schimperi Hochst.
Manilkarinae; M. maxima T.D. Penn., M. zapota (L.) Royen, Faucherealaciniata
Lec. Glueminae; Inhambanella guereensis (Aubrev. & Pellegr.) T. D. Penn.,
Neolemonnieraogouensis(Pierre ex Dub.) Heine. Isonandreae;Palaquiummicrophyllum
King & Gamble, Madhuca curtisii Ridley (Ridley s.n.). Sideroxyleae;Sideroxylon
discolor A. R. Smith, S. portoricenseUrban subsp. portoricense(Ekman 8737).
Chrysophylleae;Pouteria: sect. Gayella; P. splendens(A.DC.) Kuntze (Reed s.n.),
Synsepalumafzelii (Engler) T. D. Penn., S. brevipes(Baker) T. D. Penn., S. sereti
(De Wild.) T. D. Penn., Englerophytumletestui Aubrev. & Pellegr.
Subtype IIIC. (Fig. 16: A- M)

Pollen grains prolate or, less frequently, subprolate or, rarely, prolate-
spheroidal. 3, 3-4 or 4-colporate. Average polar length, 22-9-35-5 m.
Colpi 0-75-0-85 polar length. Endoapertures broad or narrow. Tectum,
surface psilate, subpsilate or scabrate.
This subtype can usually be differentiated from subtype IIIA by shape, size
(average polar length not greater than c.35 -5 Am), smaller possibly indistinct
endoapertures and an aperture number not greater than 4.
Of the two collections of Chrysophyllumrufum included in this pollen type,
one (Martius 521) has an unusually thick apocolpial region (Fig. 16: M).
Species included: Sideroxyleae;NeohemsleyausambarensisT. D. Penn., Nesoluma

polynesica (Hillebr.) Baill. Chrysophylleae;Pouteria: sect. Rivicoa; P. altissima
(Chev.) Baehni; sect. Franchetella; P. ramiflora (Mart.) Radlk. (Froes 1841;
Irwin & Soderstrom5892; Spruce926), P. reticulata(Engl.) Eyma subsp. reticulata
(FDBG 2413; Krukoff6265), Chrysophyllum:sect. Aneuchrysophyllum;C. imperiale
(Linden) Benth. & Hook.; sect. Chrysophyllum;C. oliviformeL., C. Mart.
,ufum
(Glaziou 11146; Martius 521), Niemeyerachartacea(Bailey) White (Hyland 9167;
Brass 2218), Pradosia beardii (Monachino) T. D. Penn., Synsepalumpassargei
(Engler) T. D. Penn.
Subtype IIID. (Fig. 16: N-R)

Pollen grains subprolate or prolate. 3-colporate. Average polar length, 16 -0 -
28 - 0 Am. Colpi 0 75-0 85 polar length. Endoapertures narrow. Tectum,
coarsely perforate; surface psilate, subpsilate or somewhat irregularly granular.
Similar to subtype IIIC except that tectum coarsely perforate.
Pouteria:sect. Franchetella;P. reticulata(Engl.)
Species included: Chrysophylleae;
Eyma subsp. reticulata(Prance et al. 8142), P. sagotiana (Baill.) Eyma, Pradosia
schomburgkiana(A.DC.) Cronq. subsp. schomburgkiana(Prance & Ramos 23128).
POLLENTYPE IV. 'SARCOSPERMA'.
Number of genera included: 12-Chrysophyllum (1), Diploknema(1), Labourdon-
naisia (1), Manilkara (1), Palaquium (1), Pouteria(2), Pradosia (1), Sarcosperma
(2), Synsepalum
(6), Sideroxylon (1), Xantolis(2).
(2), Tieghemella
Pollen grains spheroidal, prolate-spheroidal or subprolate, rarely prolate.
Poles hemispherical, equatorial faces convex. 4, 4-5 (3)-colporate. Average

- - - - -
polar length, 24 1 - 56 8 um. Colpi 0 65 -0 75 (0 85) polar length. Endo-
apertures broad or very broad. Wall thickness, poles: 1 0-3-0 /Am; equator:
1 - 5- 3 -0 (5 - 0) zm. Infratectum granular or with reduced columellae. Endexine
very thin in mesocolpia. Tectum not protrudent or, occasionally very slightly
protrudent, perforate or with microfossulae; surface psilate, subpsilate,
scabrate or, less frequently, rugulate.
Key to subtypes of Pollen Type IV

1. Average polar length 23-0-35-0 tm:
2. Tectum distinctly perforate and/or with microfossulae, pollen grains 4 or
5, never 3-colporate:
3. Pollen grains prolate-spheroidal, colpi 0 - 65 -0 - 75 polar length . IVA
3. Pollen grains subprolate or prolate, colpi 0-75-0-85 polar length IVB
2. Tectum without distinct perforations or microfossulae, pollen grains 3 or
4, rarely 5-colporate ...................................... IVc
1. Average polar length 38-56 ~/m:
4. Pronounced costae, pollen grains never spheroidal ............. IVE
4. Costae not apparent, grains may be spheroidal ............... IVD
SubtypeIVA. (Fig. 17: A-N)

Pollen grains prolate-spheroidal. 4 or 4-5-colporate. Average polar length,
24 - 1-32 -4 m. Colpi 0 - 65 - 0 - 75 polar length. Endoapertures broad. Tectum,
perforate or with microfossulae; surface, psilate, subpsilate or scabrate.
Species included: Sideroxyleae;SarcospermaarboreumBenth. ex Clarke, S.
griffithii Hook. f. ex Clarke, S. kachinense(King & Prain) Exell., S. laurina
(Benth.) Hook. f. (Ying Hu 12355A), S. paniculatum (King) Stapf & King.
Chrysophylleae;Pouteria: sect. Oxythece;P. rigida (Mart. & Eichl.) Radlk. subsp.
tomentosa(Aubrev.) T. D. Penn., Pradosia brevipes(Pierre) T. D. Penn.
Subtype IVB. (Figs 17: P; 18: A-B)

Pollen grains subprolate or prolate. 4 or 4- 5-colporate. Average polar
- -
length, 29 1 & 30 6 m. Colpi 0 -75-0 -85 polar length. Endoapertures very
broad. Tectum perforate and/or with microfossulae; surface rugulate or
scabrate.
Species included: Chrysophylleae;Xantolis racemosa(Dub.) Royen, X. siamensis

(Fletcher) Royen.
SubtypeIVC. (Fig. 18: C-P)

Pollen grains subprolate. 3, 3-4, 4, infrequently, 4-5-colporate. Average
- - - -
polar length, 23 1 - 34 5 Colpi 0 65 - 0 75 (0 *85) polar length. Tectum
/m.
occasionally slightly protrudent, perforate; surface subpsilate, scabrate or
(Chrysophyllumeximium) rugulate.
Species included: Sideroxyleae; Sideroxylon microphyllum (Elliott) Baehni,
Sarcospermalaurina (Benth.) Hook. f. (Tsang 30726). Chrysophylleae," Pouteria:
sect. Oxythece,P. opposita (Ducke) T. D. Penn., Chrysophyllum:sect. Aneuch-
rysophyllum; C. eximium Ducke, Synsepalum msolo (Engler) T. D. Penn., S.
subverticillatum(Bruce) T. D. Penn.

Subtype IVD. (Fig. 19: A-M)
Pollen grains spheroidal, prolate-spheroidal or subprolate. 4 or 4 -5-
- - -
colporate. Average polar length, 38 0 - 56 8 Am. Colpi c. 0 75 polar length.
Tectum semi-protrudent, densely perforate; surface subpsilate or scabrate.
Species included: Mimusopeae: Manilkarinae; Manilkara bidentata (A.DC.)

Chev. subsp. bidentata, Labourdonnaisia glauca Boj. Isonandreae; Diploknema
butyraceoidesLam (Strachey & Winterbottom 1). Sideroxyleae;Sideroxylon capiri
(A.DC.) Pittier subsp. capiri.
Subtype IVE. (Fig. 20: A-J)

Pollen grains prolate-spheroidal or subprolate. 4 or 4 - 5-colporate. Average
-
polar length, 40 5 and 48 -66Am. Pronounced costae seen with the light micro-
scope. Tectum semi-protrudent, perforate; surface subpsilate.
Species included: Mimusopeae: Mimusopinae; Tieghemella heckelii Pierre ex
Chev. Isonandreae;Palaquium quercifolium(Vriese) Burck.
POLLEN TYPE V. 'PICHONIA'.

Number of genera included: 7-Autranella (2), Elaeoluma (1), Leptostylis (1),
Pichonia (6), Pouteria (11), Pycnandra(6) and Tridesmostemon (1).
Pollen grains subspheroidal, spheroidal or prolate-spheroidal, less frequently
subprolate. Poles hemispherical, equatorial faces convex. 4, 4-5 (3)-colporate.
- - - - - -
Average polar length, 33 8 - 64 5 m. Colpi (0 50) 0 65 -0 75 (0 85) polar
length. Endoapertures usually broad, occasionally more or less circular. Wall
thickness, poles: 1 5-4 Am; equator: 3-6 Am. Infratectum narrow or very
narrow, granular or with reduced columellae. Endexine thin or absent in
mesocolpia, thickened towards or, in some species distinctly costate, around
apertures. Tectum not protrudent, perforate or with coarse slits, weakly or
coarsely rugulate, rarely, coarsely reticulate, broken reticulate or foveolate
with granular lumina; surface psilate, subpsilate, scabrate, weakly striate-
rugulate or low-relief rugulate.
Key to subtypesof Pollen Type V

1. Pollen grains subprolate ..................................... Vc
1. Pollen grains prolate-spheroidal, occasionally subspheroidal:
2. Pollen wall exceptionally thick in relation to grain size:
3. Pollen grains 3 or 4-colporate, never 5, average polar length 39-47 Am,
no obvious costae:
4. Entire surface of tectum psilate ........................... VB
4. Entire surface of tectum finely striate-rugulate or low-relief rugulate
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . VE
........
3. Pollen grains 4 or 5, rarely 3-colporate, average polar length, 33 - 65 Am,
costae frequently pronounced .............................. VA
2. Pollen wall very thin in relation to grain size .................. VD
Subtype VA. (Figs 20: K-M; 21: A-K)

Pollen grains subspheroidal or prolate-spheroidal. 4, 4-5(3-4-Pouteria

laurifolia)-colporate. Average polar length, 33-8 & 37-8, 44-5-64-5 Am.

Colpi 0-65-0-75 polar length. Endoapertures usually broad, often sur-
rounded by pronounced costae (seen with the light microscope). Pollen walls
exceptionally thick, poles: 20-4 -0 /sm; equator: 3 -0- 60 itm. Tectum weakly
or coarsely rugulate or, exceptionally (Pouteriacampechiana),mesocolpia coarsely
reticulate or broken reticulate and apocolpia foveolate or entire; surface
psilate, scabrate or, less frequently, rugulate with microfossulae.
Species included: Mimusopeae:Mimusopinae; Autranella congolensis(De Wild.)

Chev. ex Aubrev. & Pellegr. (Louis 3521). Chrysophylleae;Pouteria: sect.
Oligotheca; P. laurifolia Radlk., sect. Rivicoa; P. campechiana(Kunth) Baehni
(Lundell & Contreras19241; Hinton 5391), P. macrophylla(Lamk.) Eyma; sect.
Antholucuma;P. domingensis(Gaertn.) Baehni subsp. cuprea(Urban & Ekman)
T. D. Penn., P. grandiflora (A.DC.) Baehni, P. multiflora (A.DC.) Eyma, P.
venosa(Mart.) Baehni subsp. amazonicaT. D. Penn. sect. Pouteria;P. gardneriana
(A.DC.) Radlk., Pichonia lauterbachiana(Lam) Baehni (Brass 22061 & 21910;
Womersley19233), P. solomonensisRoyen, Leptostylisgrandifolia Vink.
Subtype VB. (Fig. 21: L-M)

Pollen grains prolate-spheroidal. 4-colporate. Average polar length, 42 - 5 m.
Colpi long, c. 0 -85 polar length. Endoapertures broad. Walls exceptionally
thick in relation to overall grain size, 3 - 5 - 4 -0 tm in both apo- and mesocolpia.
Surface of tectum, psilate.
Species included: Chrysophylleae;Pouteria: sect. Oligotheca;P. keyensis (Lam)

Baehni.
Subtype VC. (Fig. 22: A-J)

Pollen grains subprolate. 4 or 4-5-colporate. Average polar length,
49 - 8 ,m. Colpi 0 - 50 -0 75 polar length. Endoapertures usually broad.34"-0-
Wall
-
thickness, poles: 3 ,m; equator: 2 0 -4-0 m. Tectum perforate or
-0-6"0
with coarse slits and/or rugulate mesocolpium; surface subpsilate. This sub-
type very similar to subtype VA except that shape differs.
Species included: Mimusopeae:Mimusopinae; Autranellacongolensis(De Wild.)
Chev. ex Aubrev. & Pellegr. (Louis 4048). Chrysophylleae;Pouteria: sect. Rivicoa;
P. robusta (Mart. & Eichl.) Eyma, Pichonia deplanchei (Baill.) T. D. Penn.
(McKee 13552; Deplanche 442), Elaeoluma nuda (Baehni) Aubrev. (Philcox et al.
3257).
Subtype VD. (Fig. 23: J-K)

Pollen grains prolate-spheroidal. 4-colporate. Average polar length, 49 0 Pm.
Colpi short, c. 0 -50 polar length. Endoapertures more or less circular. Polar
and equatorial walls very thin, c. 1 -0 ,sm. Tectum, surface subpsilate.
Species included: Omphalocarpoideae;TridesmostemonomphalocarpoidesEngl.
Subtype VE. (Figs 22: K-M; 23: A-H)

Pollen grains prolate-spheroidal. 3-4 or 4-colporate. Average polar length,
1 Wall thickness, poles: pm; equator: 3"0-4"0 ,m.
39"2-46 ksm. 4"0-6"0

Endexine increases in thickness towards and around endoapertures. Tectum,
very finely striate-rugulate or very finely low-relief rugulate. This subtype
similar to subtype VA except for lesser number of apertures and walls that
are exceptionally thick in proportion to overall grain size.
Species included: Chrysophylleae;Pycnandra benthamii Baill., P. controversa

(Guill.) Vink (McPherson 2885), P. kaalainsis Aubrev., P. vieillardii (Baill.)
Vink, P. aff. vieillardii, P. sp. (Pennington & McPherson 10283).
POLLENTYPE VI. 'SIDEROXYLON'.

Number of genera included: 6-Argania (1), Capurodendron (2), Pouteria
(2), Sideroxylon(23), Vitellaria (2) and Vitellariopsis (1).
Pollen grains prolate or subprolate, exceptionally prolate-spheroidal. Poles
sub-hemispherical or hemispherical, equatorial faces parallel or convex. 3,
3-4, 4-5 or 5-colporate. Average polar length, 23 -7-50-8 (61 4) Am. Colpi
(0 -65) 0- 75 -0 85 polar length. Endoapertures broad or narrow. Wall thick-
-0 -
ness, poles: 10-3 (40) Am; equator: 1-5-3 5 (50) m. Infratectum
narrow or very narrow, granular or with reduced columellae. Endexine thin
in mesocolpia. Tectum not or, occasionally, slightly protrudent, occasionally
perforate and/or with microfossulae; surface weakly or distinctly striate-
rugulate or striate, psilate, subpsilate or, rarely, finely granular.
Key to subtypesof Pollen Type VI

1. Entire tectum striate or striate-rugulate:
2. Pollen grains prolate, colpi 0 - 75 -0 -85 polar length:
3. Poles sub-hemispherical .................................. VIA
3. Poles hemispherical ...................................... VIB
2. Pollen grains subprolate, colpi 0-65-0 -75 polar length ........ VIc
1. Entire tectum psilate, subpsilate or very finely granular:
4. Pollen grains 3-colporate ................................... VIF
4. Pollen grains 4 or 5, rarely 3-colporate:
5. Average polar length not exceeding 50 Am, poles hemispherical VIE
5. Average polar length up to 62 Am, poles sub-hemispherical ... VID
Subtype VIA. (Fig. 24: A-G, L)

Pollen grains prolate, poles sub-hemispherical. 4, 4-5, rarely 3 - 4-colporate.
Average polar length, 29 -6- 34 -2 (43 -5) m. Colpi long, 0O75 -0 -85 polar
length. Endoapertures usually broad. Tectum occasionally semi-protrudent;
surface weakly or distinctly striate-rugulate or striate.
Species included: Sideroxyleae;Sideroxyloncelastrinum(Kunth.) T. D. Penn.

(Curtiss 1765), S. eriocarpum(Greenm. & Conz.) T. D. Penn., S. ibarrae
(Lundell) T. D. Penn., S. lycioides L., S. puberulumA.DC.
Subtype VIB. (Fig. 24: H-K, M-Q)

Pollen grains prolate, poles hemispherical. 3, 3-4 or 4-colporate. Average
polar length, 23 7-37 7 tm. Colpi long, 0 polar length. Tectum
75-0"85
occasionally semi-protrudent; surface striate-rugulate or striate.

Species included: Sideroxyleae;SideroxylonborbonicumA.DC., S. celastrinum

(Kunth) T. D. Penn. (Hayes 451), S. confertumWright, S. jubilla (Ekman ex
Urban) T. D. Penn.; Chrysophylleae;Pouteria: sect. Franchetella;P. dictyoneura
(Griseb.) Radlk. subsp. dictyoneura.
Subtype VIC. (Fig. 25: A-K)

Pollen grains subprolate (prolate-spheroidal-Pouteria splendens), poles
hemispherical. 4, 4- 5, rarely 3-colporate. Average polar length, 30- 5- 50- 8
tim. Colpi 0-65-0 75 polar length. Tectum, striate or striate-rugulate.
Pouteria splendensis included here since it is closer to this subtype than to any
other, but both on aperture number (3) and shape it is somewhat out of place.
Species included: Sideroxyleae; Griseb. subsp. floribundum,
Sideroxylonfloribundum
S. galeatum (Hill) Baehni, S. mermulanoLowe, S. occidentale(Hemsl.) T. D.
Penn., Argania spinosa (L.) Skeels. Chrysophylleae;Pouteria: sect. Gayella; P.
splendens (A.DC.) Kuntze (Capt. King s.n.).
Subtype VID. (Figs 25: L-P; 26: A-B, J)

Pollen grains usually prolate, occasionally subprolate, poles sub-hemis-
pherical. 4, 4-5, rarely 3-4-colporate. Average polar length, 36-1-61-4
jim. Colpi long, 0 -75-0-85 polar length. Endoapertures broad or narrow.
Tectum occasionally perforate and/or with microfossulae; surface granular or
subpsilate.
Species included: Mimusopeae:Mimusopinae; Vitellariopsiskirkii (Baker) Dub.,
Vitellariaparadoxa Gaertn. f. (Oldeman 911). Sideroxyleae;Sideroxylonamericanum
(Mill.) T. D. Penn., S. foetidissimum Jacq. subsp. foetidissimum, S. portoricense
Urban subsp. minutiflorum(Pittier) T. D. Penn.
Subtype VIE. (Fig. 26: C-H, K-M)

Pollen grains subprolate or prolate. Poles hemispherical or sub-hemispherical.
4 or 4- 5-colporate. Average polar length, 31-1-49-5 Colpi long, 0-75
- 5/m.
(0 85) polar length. Tectum surface psilate or subpsilate.
Species included: Mimusopeae: Mimusopinae; Vitellaria paradoxa Gaertn. f.
(Coull 1). Sideroxyleae;Sideroxyloncubense(Griseb.) T. D. Penn., S. gerrardianum
(Hook. f.) Aubr6v., S. montanum(Swartz) T. D. Penn., S. obtusifolium(Roem.
& Schultes) T. D. Penn. subsp. buxifolium(Roem. & Schultes) T. D. Penn.,
S. rotundifolium(Swartz) T. D. Penn., S. tenax L..
Subtype VIF. (Fig. 27: A-G)

Pollen grains subprolate, poles hemispherical. 3-colporate. Average polar
length, 35-6-36-6 sm. Colpi c. 0-75 polar length. Wall thickness, poles:
2-0-3 -0 equator: 2-0-3-5 rather thick in proportion to grain size.
/m; /tm,
Tectum surface very finely granular, psilate or subpsilate.
Species included: Sideroxyleae;Sideroxylonfloribundum Griseb. subsp. belizense
(Lundell) T. D. Penn. Chrysophylleae;Capurodendron bakeri(Elliott) Aubr~v., C.
rubrocostatum(Jum. & Perrier) Aubr~v.

POLLEN TYPE VII. 'POUTERIA CAIMITO'.

Number of genera included: 12-Aubregrinia (1), Breviea (1), Chrysophyllum
(38), Delpydora (2), Ecclinusa (4), Englerophytum(1), Isonandra(1), Micropholis
(8), Niemeyera (1), Pouteria (79), Pradosia (2), Sideroxylon(5).
Pollen grains prolate or, less frequently, subprolate. Poles hemispherical,
infrequently sub-hemispherical, equatorial faces usually parallel, occasionally
convex, rarely concave. 3, 3-4, 4 (5)-colporate. Average polar length, 20 -7-
92 - 2 ~m. Colpi 0 - 15 -0 - 50, less frequently 0 - 65 -0 - 75 polar length. Endo-
apertures usually narrow, occasionally broad. Wall thickness, poles: (-5)1 -3
(4) um; equator: 2-5 (8) Lm. Infratectum narrow or very narrow, granular
or reduced columellae. Endexine continuously thickened in mesocolpia.
Tectum usually protrudent, occasionally semi-protrudent. Apocolpia and
mesocolpia usually differentiated: surface of apocolpia psilate, subpsilate,
scabrate, finely striate, very finely rugulate or low-relief rugulate; mesocolpia
weakly or distinct fine or coarse rugulate, rarely corrugate-rugulate in polar
direction, occasionally tectum undifferentiated, finely striate-rugulate or
rugulate.
Key to subtypesof Pollen Type VII

1. Equatorial faces concave ................................... VIIB
1. Equatorial faces usually parallel, occasionally convex:
2. Tectum ornamentation not differentiated .................... VIID
2. Tectum ornamentation differentiated:
3. Pollen grains frequently prolate:
4. Average polar length 30-68
/zm:
5. Apocolpia always psilate, mesocolpia semi-corrugate in polar direction
... ..........................................VIIc
5. Apocolpia may be finely rugulate, low-relief rugulate or psilate,
mesocolpia rugulate:
6. Mesocolpia distinctly or coarsely rugulate .............. VIIA
6. Mesocolpia weakly rugulate .......................... VIIE
4. Average polar length 21-32 ttm ......................... VIIF
3. Pollen grains usually subprolate, rarely prolate, occasionally prolate-
spheroidal ............................................. VIIG
Subtype VIIA. (Figs 27: H-K; 28: A-P)

Pollen grains subprolate (1/3 of species) or prolate (remaining 2/3). Poles
hemispherical. Equatorial faces of grain, parallel. 3, 3-4 or 4 (rarely 4-5)-
- - -
colporate. Average polar length, (20 7), 32 7 - 62 5 ~m. Colpi vestigial--0 50,
less frequently, 0-65-0-75 polar length. Endoapertures usually narrow,
occasionally broad. Tectum protrudent or occasionally semi-protrudent,
apocolpia psilate or with very fine low-relief rugulae, mesocolpia distinctly
rugulate, maybe coarse.
Pouteria: sect. Oligotheca;P. brownlessiana(F.
Species included: Chrysophylleae,;
Muell.) Baehni, P. cotinifolia (A.DC.) Baehni, P. firma (Miq.) Baehni (Kerr
15958), P. maclayana (F. Muell.) Baehni, P. macrantha (Merrill) Baehni,
P. malaccensis(C. B. Clarke) Baehni, P. myrsinoides(Cunn. ex Benth.) Baehni,

P. rubicunda(Pierre) Baehni, P. singuliflora (C. T. White & Francis) Baehni,
P. sp. (McKee 14204), P. sp. (Pennington & McPherson 10282); sect. Gayella; P.
decussata(Ducke) Baehni; sect. Aneulucuma;P. ephedrantha(A. C. Smith) T. D.
Penn.; sect. Pouteria; P. caimito (Ruiz & Pav.) Radlk. (FDBG 3652 & 2320),
P. guianensis Aublet (Sandwith 550, Hart 5982); sect. Oxythece;P. cuspidata
(A.DC.) Baehni subsp. cuspidata (Jenman 924), Aubregriniataiensis (Aubrev. &
Pellegr.) Micropholis:sect. Micropholis;M. melinonianaPierre, M. mensalis(Baehni)
Aubrev., M. resinifera(Ducke) Eyma. Chrysophyllum:sect. Donella; C. welwitschii
Engl. (Jordan 254); sect. Ragala; C. sanguinolentum(Pierre) Baehni subsp. balata
(Ducke) T. D. Penn. (Klug 3699), Ecclinusaguianensis Eyma, E. ramifloraMart.
(Hoehne SP 28590 & s.n.)
Subtype VIIB. (Fig. 29: A-N)

Pollen grains subprolate or, more frequently, prolate. Poles hemispherical.
Equatorial faces concave. 3, or, rarely (Pouteriaspeciosa)4 - 5-colporate. Average
- - - -
polar length, 28 8 -92 2 tm. Colpi 0 50 - 0 65 polar length. Endoapertures
narrow or, less frequently, broad. Endexine massively thickened around
equator. Tectum, apocolpia psilate, scabrate or very finely striate, occasionally
distinctly perforate; mesocolpia weakly or distinctly rugulate. Pouteriaspeciosa
is included here because, in spite of the atypical aperture number and polar
length (av. 92 *2 tm) it is similar in all other respects, particularly the concave
equatorial faces.
Species included: Chrysophylleae;Pouteria: sect. Rivicoa; P. speciosa (Ducke)
Baehni (Ducke 22264), Chrysophyllum:sect. Donella; C. sp. (Donella ogowensis
(A. Chev.) Aubrev. & Pellegr.), C. welwitschii Engl. (Bos 1860); sect. Ragala;
C. sanguinolentum(Pierre) Baehni subsp. balata (Ducke) T. D. Penn. (Schultes
& Lopez 9980); sect. Prieurella; C. amazonicum T. D. Penn., C. cuneifolium
(Rudge) A.DC.; sect. Villocuspis; C. flexuosum Mart.; Ecclinusa lancifolia (Mart.
& Eichl.) Eyma.
Subtype VIIC. (Fig. 30: A-F)

Pollen grains prolate. Poles hemispherical. Equatorial faces more or less
parallel. 4 or 4- 5-colporate. Average polar length, 64 -3 & 68-0 tm. Colpi
c. 0-50 polar length. Endoapertures narrow. Tectum, apocolpia psilate;
mesocolpia rugulate, semi-corrugate in polar direction.
Delpydoragracilis Chev., D. macrophyllaPierre.
Subtype VIID. (Figs 30: G-K; 31: A-C)

Pollen grains prolate. Poles hemispherical or sub-hemispherical. Equatorial
faces parallel. 3, 3 - 4 or 4-colporate. Average polar length, 27 - 1 - 56 -2 Jm.
Colpi 0 -15-0-50 polar length. Endoapertures broad. Tectum very finely
striate-rugulate or rugulate, no differentiation between apo- and mesocolpium.
Specie included: Isonandreae;IsonandrazeylanicaJenken. Sideroxyleae;Sideroxylon
boutonianumA.DC., S. contrerasii(Lundell) T. D. Penn., S. sp. (Bouton s.n.).
Chrysophylleae;Pouteria: sect. Franchetella;P. ucuquiPires & Schultes (Schultes &
Lopez 9563).

Subtype VIlE. (Figs 31: D-M; 32: A-G)
Pollen grains prolate. Poles hemispherical, occasionally sub-hemispherical.
Equatorial faces parallel or slightly convex. 3, 3-4 (4)-colporate. Average
- -
length, 30 0- 37 8 im (2/3 of specimens) or 40 7 - 60 ,m (1/3 of specimens).
- 50 -0 -75
Colpi 0 polar length. Endoapertures narrow, less frequently broad.
Tectum sometimes only semi-protrudent, apocolpia psilate or finely rugulate;
mesocolpia weakly rugulate.
Englerophytumnatalenseis included here although its finely striate surface is
unique within this subtype.
Species included: Chrysophylleae;Pouteria:sect. Oligotheca;P. australis (R. Br.)
Baehni, P. baillonii (Zahlbr.) Baehni, P. baueri (Montr.) Baehni, P. sp.
(Planchonellaglabra (Ridley) Lam), P. nitidd Radlk., P. obovoidea(Lam) Baehni,
P. oppositifolia(Ducke) Baehni, P. pohlmannia(F. Muell.) Baehni, P. sandwicensis
(A. Gray) Baehni, P. sericea (Ait.) Baehni (Brown 2823), P. sp. (FD 1034),
P. sp. (Vodonaivalas.n.), P. toricellensis(K. Schum.) Baehni, P. velutina(Elmer)
P. hochreutineri
Baehni; sect. Pierrisideroxylon; (Lam) Lam, P. kaernbachiana (Engl.)
Baehni, P. vrieseana (Pierre ex Burck) Baehni; sect. Gayella; P. eugeniifolia
(Pierre) Baehni (Cowan & Soderstrom2198A; Silva & Bahid 3093), P. nemorosa
Baehni; sect. Aneulucuma;P. procera(Mart.) T. D. Penn.; sect. Pouteria;P. torta
(Mart.) Radlk., P. torta (Mart.) Radlk. subsp. glabra T. D. Penn., P. torta
(Mart.) Radlk. subsp. torta; sect. Franchetella;P. cladantha Sandwith (Clark &
Maquirino s.n.), P. engleriEyma; Micropholis: sect. Micropholis;M. comptaPierre,
M. venulosa(Mart. & Eichl.) Pierre; Chrysophyllum:sect. Aneuchrysophyllum;C.
boivinianum (Pierre) Baehni (Capuron SF 11403 & SF 2521), C. gonocarpum
(Mart. & Eichl.) Engl. (Hassler 12382), C. pomiferum (Eyma) T. D. Penn.,
C. venezuelanense (Pierre) T. D. Penn. (Ducke 15802), C. sp. (Gambeyaboukokoensis
Aubrev. & Pellegr.); sect. Donella; C. pruniformePierre, C. welwitschii Engl.
(Leeuwenberg 2267); sect. Chrysophyllum; C. argenteumJacq. subsp. auratum
(Miq.) T. D. Penn., Pradosia subverticillata Ducke, Englerophytumnatalense
(Sonder) T. D. Penn.
SubtypeVIIF. (Figs 32: H-L; 33: A-L)

Pollen grains usually prolate, occasionally subprolate. Poles hemispherical.
Equatorial faces parallel or slightly convex. 3, 3-4 (4)-colporate. Average
polar length, 21-3-31-6 /m. Colpi 0-50-0-75 polar length. Tectum
occasionally only slightly protrudent, psilate or subpsilate apocolpia; mesocolpia
weakly rugulate. This subtype is very similar to subtype VIIE except that the
grains are smaller.
Species included: Sideroxyleae;SideroxylonsalicEfolium(L.) Gaertn., S. saxorum
Lec. Chrysophylleae;Pouteria: sect. Olzgotheca; P. linggensis (Burck) Baehni,
P. obovata (R. Br.) Baehni (Procter 3950); sect. Oxythece; P. ambelaniifolia
(Sandwith) T. D. Penn., P. kaieteurensisT. D. Penn.; sect. Franchetella; P.
bangii (Rusby) T. D. Penn. (Ule 9693; BW 6719), P. belizensis (Standl.)
Cronq., P. coriacea(Pierre) Pierre, P. durlandii (Standl.) Baehni, P. durlandii
(Standl.) Baehni subsp. durlandii (Lundell 16759), P. egregia Sandwith, P.
gardneri (Mart. & Miq.) Baehni (Prance & Silva 59467), P. reticulata (Engl.)
Eyma subsp. reticulata(Schipp 1238; Gentle 1208), P. ucuquii Pires & Schultes
(Schultes& Lopez9484). Micropholis:sect. Micropholis;M. gardneriana
(A.DC.)

Pierre, M. guyanensis (A.DC.) Pierre subsp. guyanensis. Chrysophyllum:sect.

Aneuchrysophyllum;C. albidum Don, C. begueiAubr6v., C. gonocarpum(Mart. &
Eichl.) Engl. (Ibbarola 3563), C. lucentifoliumCronq. subsp. pachycarpumPires
& T. D. Penn.; sect. Prieurella; C. colombianum(Aubriv.) T. D. Penn.; sect.
Chrysophyllum;C. argenteumJacq. subsp. argenteum, C. cainito L. (Valeur 677;
Docters&Leeuwenbergs.n.), C. hirsutumCronq., C. mexicanumBrand. (Pennington
& Sarukhdn9126; Contreras2782), C. ovale Rusby; sect. Villocuspis; C. sparsi-
florum Klotsch ex Miq..
Subtype VIIG. (Figs 33: M-P; 34: A-N)

Pollen grains usually subprolate, occasionally prolate-spheroidal, rarely
prolate. Poles hemispherical. 3, 4 (3-4 or 4- 5)-colporate. Average polar
length, 30-0- 49-5 Am, infrequently between 54- 1 and 71 -1 Am. Colpi
0 - 50 -0-065 (0 - 75) polar length. Endoapertures broad or narrow. Tectum,
apocolpia psilate or subpsilate, mesocolpia weakly rugulate. Very similar to
subtype VIIE except for difference in shape.
Species included: Chrysophylleae;Pouteria: sect. Oligotheca;P. amieuanaGuill.,
P. cinerea (Panch. ex Baill.) Baehni, P. firma (Miq.) Baehni (Anta 232), P.
grandifolia (Wall.) Baehni, P. obovata (R. Br.) Baehni (Burkill 2611), P. sarco-
spermoides (Lam) Lam, P. sebertii (Panch.) Baehni, P. unmackiana (Bailey)
Baehni, P. wakere(Panch. et Seb.) Baehni; sect. Pierrisideroxylon;P. multinervis
T. D. Penn. ined., P. thyrsoidea(White) T.D. Penn. sect. Rivicoa; P. speciosa
(Ducke) Baehni (Archer7718); sect. Aneulucuma;P. sapota(Jacq.) H. E. Moore &
Stearn; sect. Pouteria; P. glomerata(Miq.) Radlk. subsp. glomerata, P. gomphiae-
folia (Mart.) Radlk.; sect. Franchetella; P. durlandii (Standl.) Baehni subsp.
durlandii (Gentle 1809), P. gardneri (Mart. & Miq.) Baehni (Hassler 7514), P.
ramiflora (Mart.) Radlk. (Ratter & Ramos 444). Breviea leptosperma(Baehni)
Heine. Micropholis: sect. Exsertistamen;M. maguireiAubr6v. Chrysophyllum:sect.
Aneuchrysophyllum;C. bangweolenseFries, C. giganteum Chev., C. venezuelanense
(Pierre) T. D. Penn., C. viride Mart. & Eichl. ex Miq.; sect. Donella; C.
roxburghiiDon, C. viridifoliumWood & Franks; sect. Villocuspis;C. aff.flexuosum,
C. splendens Sprengel. Niemeyera lissophylla (Pierre ex Baill.) T. D. Penn.,
Pradosia ptychandra(Eyma) T. D. Penn. (Mori & Veyret8982).
POLLEN TYPE VIII. 'POUTERIA ALNIFOLIA'.

Number of genera included: 10-Chrysophyllum (1), Englerophytum (2),
Leptostylis(2), Magodendron (1), Niemeyera(5), Omphalocarpum (4), Pouteria
(5), Pradosia (4), Pycnandra(1) and Sideroxylon(2).
Pollen grains subprolate or prolate-spheroidal, occasionally prolate. Poles
hemispherical, equatorial faces convex. 3, 3-4, 4 or 4-5-colporate. Average
- - - -
polar length, 22 1 - 60 3 m. Colpi 0 65 -0 75 polar length. Endoapertures
broad, occasionally narrow. Wall thickness, poles: 1-0-3-5 Am; equator:
1 0-5 0(7 0) Atm. Infratectum narrow or very narrow, granular or with
reduced columellae. Endexine continuously thickened in mesocolpia. Tectum
semi-protrudent (rarely protrudent) or not, occasionally perforate; psilate,
subpsilate, scabrate or, occasionally, low-relief rugulate, very finely striate or
striate-rugulate, no differentiation between apocolpium and mesocolpium.

Key to subtypesof Pollen Type VIII
1. Pollen grains never prolate:
2. Pollen grains 3 or 4-colporate, average polar length not exceeding 36 m
....................................................... V IIID
2. Pollen grains 4 or 5-colporate, average polar length 43-44 Am VIIIB
1. Pollen grains prolate or subprolate, rarely prolate-spheroidal:
3. Average polar length not usually exceeding 50 m:
4. Tectum not protrudent, psilate, subpsilate or scabrate ...... VIIIA
4. Tectum semi-protrudent, entire surface low-relief rugulate ... VIIIc
3. Average polar length 45-61 Am:
5. Tectum coarsely perforate, protrudent, 3 or 4-colporate ..... VIIIE
5. Tectum not coarsely perforate, semi-protrudent, 4 or 5-colporate VIIIF
Subtype VIIIA. (Fig. 35: A-K)

Pollen grains usually subprolate, occasionally prolate-spheroidal or prolate.
Equatorial faces convex. 3, 3-4, 4 or 4-5-colporate. Average polar length,
23 -6 - 49 - 5 m, (exceptionally 78 - 7 Am). Colpi 0 - 75, occasionally shorter, c.
0 -65 polar length. Endoapertures usually broad. Tectum not protrudent,
perforations apparent in some species; surface psilate, subpsilate or scabrate.
Species included: Chrysophylleae;Pouteria: sect. Oligotheca; P. balanseana
(Pierre ex Baill.) Baehni, P. pinifolia (Baill.) Baehni; sect. Rivicoa; P. alnifolia
(Baker) Roberty. Chrysophyllum:sect. Aneuchrysophyllum; C. lanatumT. D. Penn.
Niemeyera antiloga (F. Muell.) T. D. Penn., N. balansae (Baill.) Aubrev., N.
deplanchei(Baill.) T. D. Penn. (McKee 13654), N. francei (Guill. & Dub.) T.
D. Penn., N. sp. (Pennington & McPherson 10311). Pradosia atroviolaceaDucke,
P. schomburgkiana (A.DC.) Cronq. subsp. schomburgkiana (Stockdale178), Leptostylis
petiolata Vink. Englerophytumoblanceolatum(S. Moore) T. D. Penn., E. magalis-
montanum(Sonder) T. D. Penn., Magodendronvenefici(White & Francis) Vink.
Subtype VIIIB. (Fig. 35: L-N)

Pollen grains prolate-spheroidal or subprolate. 4 or 4 - 5-colporate. Average
polar length, 43-3 & 43-6 Am. Colpi c. 0-65 polar length. Endoapertures
broad. Tectum rugulate, rugulae slightly weaker in apocolpial region.
Species included: Chrysophylleae;Pouteria: sect. Oligotheca;P. sericea (Ait.)
Baehni (Parker 509). Omphalocarpeae;OmphalocarpumbequaertiiDe Wild.
Subtype VIIIC. (Fig. 36: A-E)

Pollen grains subprolate or prolate. 4-colporate. Average polar length, 24 - 0
or 39 - 4 m. Colpi 0 - 65 -0 - 75 polar length. Endoapertures broad or narrow.
Tectum semi-protrudent; low-relief rugulate.
Species included: Sideroxyleae;SideroxylonportoricenseUrban subsp. portoricense
(Harris 5388), S. reclinatumMichaux.
Subtype VIIID. (Fig. 36: F-J)

Pollen grains subprolate. Equatorial faces convex. 3-4-colporate. Average
polar length, 22 1 and Colpi c. polar length. Endoapertures
broad or narrow. Tectum 35"55/m. 0"65
protrudent; surface very finely striate.

Species included: Chrysophylleae;Pradosia cuatrecasasii(Aubrev.) T. D. Penn.,

P. ptychandra(Eyma) T. D. Penn. (SF 7240).
Subtype VIIIE. (Figs 36: K-M; 37: A-D)

Pollen grains subprolate or prolate (Pycnandracontroversa).3-4 or 4-colporate.
- - - -
Average polar length, 45 5 - 60 3 Am. Colpi 0 65 - 0 75 polar length. Endo-
apertures broad. Tectum semi-protrudent, coarse perforate; subpsilate.
Pollen from Pycnandracontroversa(McKee 5100) can be separated from that
of the three species of Omphalocarpumincluded in this subtype by its smaller
size and prolate shape.
Species included: Chrysophylleae;Pycnandracontroversa(Guill.) Vink (McKee

5100). Omphalocarpeae;Omphalocarpumahia Chev., 0. elatum De Wild., O.
mayumbenseGreves.
Subtype VIIIF. (Fig. 37: E-K)

Pollen grains subprolate or prolate. 4-5-colporate. Average polar length,
51 -7 and 52 -9 Am. Colpi c. 0-75 polar length. Endoapertures broad. Tectum
semi-protrudent; surface psilate or finely striate-rugulate.
Species included: Chrysophylleae;Pouteria: sect. Rivicoa; P. sp. (Aningeria

pseudoracemosaJ. H. Hemsley), Leptostylisfilipes Benth.
POLLEN TYPE IX. 'OXYTHECE'.
Number of specimens examined: 8. (Fig. 38: A-L)

Number of genera included: 1-Pouteria.
Pollen grains usually subprolate, less frequently prolate-spheroidal. Poles
hemispherical. Equatorial faces convex. 3, 3-4 (4- 5)-colporate. Average
polar length, 18 -9-24 -6 Am. Colpi long, 0- 75-0 -85 polar length. Endo-
apertures broad or circular. Wall thickness, poles: 1 -0-2-5 m; equator:
1 -0-3-0 0Am. Infratectum narrow with reduced columellae. Endexine thin in
mesocolpia. Tectum not protrudent, in some species apocolpia have coarse
perforations; surface finely or distinctly striate. This pollen type is restricted
to some of the species in Pouteria sect. Oxythece.
Species included: Chrysophylleae;Pouteria: sect. Oxythece;P. cuspidata(A.DC.)

Baehni subsp. cuspidata(Pranceet al. 5541), P. cuspidata(A.DC.) Baehni subsp.
dura (Eyma) T. D. Penn. (Sandwith 328; Maguire et al. 56468), P. cuspidata
(A.DC.) Baehni subsp. robusta (Mart. & Eichl.) T. D. Penn. (BW 6613;
Schunke 2146), P. gabrielensis (Gilly ex Aubrev.) T. D. Penn., P. pallida
(Gaertn. F.) Baehni, P. scrobiculataMonachino.
POLLENTYPE X. 'ELAEOLUMA'.
Number of genera included: l-Elaeoluma.
Pollen grains prolate-spheroidal, subprolate or prolate. Poles hemis-
pherical. Equatorial faces convex. 3, 3-4 or 4-colporate. Average polar
length, 23 - 34-0 m. Colpi 0- 65-0- 75 polar length. Wall thickness, poles:
Mm; equator: (4-0) sm. Infratectum narrow with reduced
2"0-3"0 2"0-3"0

404 KEWBULLETIN
VOL. 46 (3)
columellae. Endexine thin in mesocolpia. Tectum not protrudent, perforate

or coarsely perforate, in some species perforations coarser in apocolpial
region, rugulate or coarsely rugulate with coarse slits; surface subpsilate,
scabrate or indistinctly granular.
SubtypeXA. (Figs 38: M-P; 39: F-S)

Pollen grains prolate-spheroidal, subprolate or prolate. 3, 3-4 or 4-colporate.
- - -
Average polar length, 23 1 - 34 0 Am. Colpi 0 65 -0 75 polar length. Wall
-
thickness, poles: c. 2-0 Am; equator: 20- 3 0 Atm. Tectum perforate or
coarsely perforate, in some species perforations coarser in apocolpial region;
surface subpsilate, scabrate or indistinctly granular.
Species included: Chrysophylleae;Elaeoluma crispa T. D. Penn., E. glabrescens

(Mart. & Eichl.) Aubrev., E. nuda (Baehni) Aubrev. (Maguire et al. 42219;
Maguire 24596), E. schomburgkiana(Miq.) Baill. (Maguire & Fanshawe 32176).
SubtypeXB. (Fig. 39: A-E)

Pollen grains subprolate. 3-4-colporate. Average polar length, 29-0 Jim.
-
Colpi 0-75 polar length. Wall thickness, poles: c. 3-0 Am; equator: c. 3 0
Am. Tectum coarsely fossulate-rugulate; surface subpsilate.
Species included: Chrysophylleae;Elaeoluma schomburgkiana(Miq.) Baill.
(Spruce 1836).
POLLEN TYPE XI. 'SARCAULUS'.

Number of genera included: 3-Chromolucuma (2), Pouteria(5) and Sarcaulus
(3).
Pollen grains spheroidal, prolate-spheroidal, subprolate or prolate. Poles
hemispherical, equatorial faces convex or, occasionally more or less parallel.
3-4 or 4-5-colporate. Average polar length, 17 -0 - 25 - 7 m. Colpi 0 - 15 -
0 - 75 polar length. Endoapertures circular or narrow. Wall thickness, poles:
1 - 0- 2 - 0 m; equator: 1 -0- 3 -0 tm. Infratectum narrow or as wide or wider
than foot layer or tectum, columellate. Endexine thin or absent in mesocolpia,
may be greatly thickened, costate, around endoapertures. Tectum protrudent
or semi-protrudent, finely rugulate or rugulate with coarse, sometimes sparse,
granules or, insulate with spinules or, surface granular.
SubtypeXIA. (Fig. 40: A- L, W- Y)

Pollen grains spheroidal or prolate-spheroidal. 3-colporate, colpi c. 0- 15
-
polar length. Average polar length, 17 - 0 21 0 m. Endoapertures circular.
Endexine absent in mesocolpia. Tectum protrudent, finely rugulate with
coarse granules or insulate with spinules.
Species included: Chrysophylleae;Pouteria: sect. Gayella; Pouteria longifolia

(Mart. & Eichl.) T. D. Penn., Sarcaulus brasiliensis (A.DC.) Eyma subsp.
brasiliensis, S. oblatus T. D. Penn., S. wurdackii Aubrev.
SubtypeXIB. (Fig. 40: M-V)

Pollen grains prolate-spheroidal or subprolate. 3-colporate. Average polar

length, 18- 2- 23 -4 Am. Colpi c. 0 -65 polar length. Endoapertures circular.

Endexine thin in mesocolpia. Tectum sometimes semi-protrudent; surface
irregularly granular.
Species included: Chrysophylleae;Pouteria: sect. Franchetella; P. cladantha
Sandwith (FDBG 2233 and Cid et al. 1945), P. vernicosaT. D. Penn., P. sp.
cf. vernicosa(Pennington 22773).
SubtypeXIC. (Fig. 41: A- F)

Pollen grains subprolate or prolate. 3-4 or 4-5-colporate. Average polar
length, 24-0 or 25-7 tAm. Colpi 0-65-0-75 polar length. Endoapertures
narrow. Endexine thin in mesocolpia. Tectum protrudent, rugulate, sparse
coarse granules or spinulae.
Species included: Chrysophylleae;Chromolucumabaehniana Monachino, C.

rubrifloraDucke.
POLLEN TYPE XII. 'DIPLO(N'.

Number of genera included: 2-Diplo6in (2) and Micropholis (1).
Pollen grains spheroidal, prolate-spheroidal or subprolate. Poles hemis-
pherical, equatorial faces convex. 3-porate or, 3-4 or 4-5-colporate.
- - -
Average polar length, 24-27 Am. Colpi 0 15 or 0 65 -0 75 polar length.
Endoapertures circular. Wall thickness, poles: 1- 0- 2 -0 Mm; equator: 1 -0-
2 -00m. Infratectum more or less same width as tectum and foot layer, or
slightly narrower, columellate. Endexine thin or absent in mesocolpia. Tectum
not protrudent, rugulate or broken reticulate, short or long, supratectal
spines; crooked and blunt or straight and pointed; surface scabrate, finely
rugulate or finely reticulate.
SubtypeXIIA. (Figs 41: G-L; 42: A-D)

Pollen grains spheroidal or prolate-spheroidal. 3-4 or 4-5-colporate.
Average polar length, 24-0-25-7 Am. Colpi 0-65-0-75 polar length.
Tectum not protrudent, rugulate or broken-reticulate with short or long,
blunt 'crooked' supratectal spines; surface scabrate or very finely reticulate.
Species included: Sideroxyleae;Diplo6n cuspidatum(Hoehne) Cronq. (Handro

956; Wurdack& Monachino 39595).
SubtypeXIIB. (Figs 41: M; 42: E-J)

Pollen grains subprolate. Colpi c. 0 -15 polar length or 3-porate. Average
polar length, 27 0m.
0 Tectum, rugulate with long straight, pointed supra-
tectal spines; surface finely rugulate.
Thin sections show the spines on the pollen exine of this species to differ
from those of Diploon in that they contain small cavities whereas those of
Diploon appear solid.
Species examined: Chrysophylleae;Micropholis: sect. Micropholis; M. retusa
(Spruce ex Miq.) Eyma.

ANOMALOUS POLLEN TYPES
Although the pollen of most species within the family can be considered to
have an affinity with one or another of the described pollen types, three
species, from two genera, possess grains with some features anomalous to
pollen in Sapotaceae. All the grains nevertheless display some other typical
sapotaceous characteristics.
ANOMALOUS POLLEN TYPE I
Subtype IA. (Fig. 43: A-G)

Pollen grains prolate. 3, 3-4, or 4-colporate. Average polar length, 22 -0-
29 44tm. Colpi 0-65-0-75 polar length. Endoapertures broad. The wall
stratification and the tectum characteristics are very distinctive and not
observed elsewhere in the family: wall thickness, poles: 2-5-5-0 '4m;
equator: 2-0-3-0 0tm. The infratectum in the apocolpial region is extremely
wide and composed of long, slender columellae. Both tectum and foot layer
are comparatively narrow. The infratectum is narrower in the equatorial
region, although it remains columellate and wider than either the foot layer
or the tectum. Contrary to the situation in Pollen Type VII, although the
endexine is continuously thickened in the mesocolpial region, the total wall
thickness in this area is still markedly less than in the apocolpial region.
Tectum not protrudent, surface finely striate and, in addition, the mesocolpial
tectum is coarsely perforate.
Species included: Chrysophylleae; Chrysophyllum: sect. Chrysophyllum; C.

marginatum(Hooker & Arn.) Radlk. subsp. marginatum(Malme 1394; Heringer
et al. 3446; Regnell s.n.; Reitz & Klein 8070; Morong 701).
Subtype IB. (Fig. 43: H, J-L)

Pollen grains prolate. 3-colporate. Average polar length, 34-4 Colpi
/Jm.
c. 0 -85 polar length. Superficially similar to Chrysophyllummarginatum subsp.
marginatum but average polar length shows c. 17 % increase, the colpi are
longer, the endoapertures are circular and, the mesocolpial tectum has distinct
vertical folding (cf. Delpydora, Pollen Type VIIC). Pollen of this species has
not been thin sectioned, light microscopy indicates that it too, has a columel-
late infratectum, although the walls are not as thick.

Chrysophyllum: C.
sect. Chrysophyllum;
inornatumMart.
POLLENTYPE II
ANOMALOUS
Pollen grains prolate-spheroidal. 3-porate. Average polar length 3xm.
24"-3
Endoapertures broad. The pollen differs from the usual situation within the
Sapotaceaein that the colpi are almost imperceptible. With light microscopy the
pollen grains appear to be triporate. Tectum surface rugulate. The aperture
characteristics in combination with a protrudent tectum and undifferentiated
ornamentation are unique within the Sapotaceae.(Fig. 42: K - Q.)
Species included: Chrysophylleae:EnglerophytumstelechanthaKrause.

DIscuSSION
Generalcharacteristicsof pollen
With few exceptions the pollen morphology of the Sapotaceaeis distinctive,
but correlation of pollen characters with generic limits is less marked.
Previously, Harley (1986a & 1990a) defined two major pollen groups within
the family, A & B, based mainly on presence or absence of continuous
endexinous thickening in the equatorial region, number of apertures, surface
ornamentation and colpus length. Pollen group A comprised most of the
genera of Aubreville's subfamilies Sideroxyloideae(Sideroxyleae + Chrysophylleae
in Pennington 1991) and Omphalocarpoideae (= Omphalocarpeae in Pennington
1991). Ten pollen types were recognised in Group A in these previous studies
(see Column 14 in Table 2). Pollen group B comprised most of the genera of
Aubreville's Mimusopoideae( = Mimusopeaein Pennington 1991) and Madhucoideae
(= Isonandreaein Pennington 1991).
Examination of the pollen of approximately 300 more species from both the
New World and the Old World, has demonstrated that patterns of variation
in the pollen morphology of the family are more complex than previously
thought and twelve pollen types are recognised here. Within these types few
pollen characters show great diversity, but sufficient variation exists for
recognition of numerous subtypes.
The most fundamental pollen character, shared by almost all pollen types
within the family, is the stratification of the pollen wall into a thick tectum
and foot layer and narrow or very narrow infratectum. Although there is some
variation, particularly in relation to endexinous thickenings, apertures, shape
and characteristics of the tectum, very few pollen grains diverge from this
basic wall stratification.
One of the most distinctive characters is the presence or absence of a
continuously thickened endexine in the mesocolpial region that is usually not
seen in the polar region. This feature is restricted to Pollen Types VII and
VIII which together account for 118 of the 398 species examined. In all other
pollen types the endexine is present and often noticeably thickened towards
and around the endoapertures but much thinner or even absent in the
mesocolpial region. In a few species the pollen has apertures surrounded by
a distinct costate 'ring' (Pollen Type V).
Comparisonof pollen types within systematicgroups

The distribution of the pollen types within Pennington's (1991) generic
arrangement is summarized in Table 1. Inevitably, because of the subtle but
significant variation in pollen characters, there may be continuous variation
within and between some subtypes.
Pollen Type I is common to tribes Mimusopeaeand Isonandreae.Within tribe
Mimusopeaeit is the predominant type in the largest genus, Mimusops (subtribe
Mimusopinae) although it is not recorded in the other genera of the subtribe.
In subtribe Manilkarinae it occurs in half the species of Manilkara and in
the monospecific genus Northia. It is not present elsewhere in this subtribe.
Polliniferous material for only five species in three genera was available from
the third subtribe ofMimusopeae, Glueminae.One of these species, Neolemonniera
clitandrifolia, has pollen of Type I. Madhuca, one of the two largest genera in
tribe Isonandreae,has Type I pollen in about two thirds of its species. In the

other large genus, Palaquium, it occurs in just under half the species. Of the
remaining five genera, Type I pollen has been recorded for one of the two
species of Diploknema which have been examined, and it is the only type
recorded for Payenaand Burckella.Outside these two tribes Pollen Type I is very
uncommon. It is found in one species of Sideroxylon,S. oxycantha(Sideroxyloideae),
and within the largest tribe, Chrysophylleae,in a few species in three genera
(Pichonia calomeris, Synsepalum cerasiferum, S. gabonensis, Xantolis assamica, X.
burmanicaand X. embellifolia). Of the four genera comprising the fifth, very
small tribe (Omphalocarpeae), the pollen of the monospecific Madagascar
genus, Tsebona macranthais of Type I.
Pollen Type II is most frequent in tribe Isonandreae.It occurs in the remain-
ing third of Madhuca species, in over 50% of Palaquium species and in four
of five species examined from Isonandra. Otherwise, Type II occurs very
sporadically throughout the three subtribes of tribe Mimusopeae, in four species
of Sideroxylon(Sideroxyleae)and in two species of Synsepalum(Chrysophylleae).
Pollen Type III has undistinguished tricolporate pollen grains, superficially
not readily recognizable as belonging to Sapotaceae. Nevertheless SEM
fractures and TEM thin sections (Fig. 15: E-G, L-M) show the wall strati-
fication to be typical of the family. The pollen morphology of one of the two
collections examined of Chrysophyllumrufumin most respects fits the description
of Type IIIC where both collections have been placed but the thickened poles
or 'polar caps' are curious. This striking increase in polar wall thickening
is caused by an unusually thick foot layer in combination with a narrow
endexine. The greatest concentration of species with Type III pollen are
in Manilkara (Mimusopeae:Manilkarinae), half the collections examined. How-
ever, it occurs sporadically throughout all the tribes with the exception of
Omphalocarpeae.
Pollen Type IV is recorded for only twenty one species that are somewhat
erratically distributed through all the tribes except Omphalocarpeae.It is more
distinctive than the previous pollen type and is found in all six collections of
Sarcospermaexamined. The widespread but sparse occurrence of this pollen
type throughout a large part of the family is particularly interesting since
Sarcospermarepresents one of the macromorphological extremes within the
family, and has at various times been removed from Sapotaceae.Lam (1925),
for example, considered that there were sufficient differences from Sapotaceae,
in particular locule number, some inflorescence features, the sub-opposite
leaves (alternate in Sapotaceae)and the auriculate petioles, to warrant separation
as a monogeneric family, the Sarcospermaceae (or Sarcospermataceae,Willis (1973)).
It is also interesting that in Xantolis, the only other genus sharing a similar
geographic distribution in India and the Far East, two of the six species
examined have pollen of this type.
Pollen Type V may be recognised by the unusually thick walls in pro-
portion to pollen grain size, except in subtype VD which is very thin-walled.
Furthermore the grains are most frequently subspheroidal, spheroidal or
prolate-spheroidal. The endoapertures, particularly in subtype VA, are often
surrounded by pronounced costae that are rarely recorded elsewhere in the
family. Most examples of this subtype are found in sections of Pouteria;
Oligotheca,Rivicoa, Antholucumaand Pouteriaand also in the genus Pichonia. The
morphology of the tectum of Pouteria campechiana (Fig. 21: J - K) is very
distinctive, although in all other respects the pollen is typical for subtype VA.

Subtype VB is limited to the extraordinary pollen of Pouteriakeyensis(Fig. 21:

L - M) with its massively thick ectexine and extremely narrow infratectum.
Pollen of Tridesmostemonhas been included in Pollen Type V (subtype VD)
although very thin-walled because it does not fit readily elsewhere and it
shows many of the characteristic features, especially its shape and costate
endoapertures. Subtype VE has only been recorded for Pycnandra (all four
species examined in the present study) although one collection of P. controversa
has Pollen Type VIIIE. With the exception of Tridesmostemon,the only other
record of Type V outside tribe Chrysophylleaeis in Autranella congolensis
(Mimusopeae).
Pollen Type VI is restricted with few exceptions to the genus Sideroxylon.
Within tribe Sideroxyleae, it is the only pollen type represented by a large
number of species. The sub-hemispherical poles are particularly distinctive,
giving many of the grains a 'lozenge-shaped' appearance (Fig. 24: A-E, F).
Outside the Sideroxyleaesuch grains occur in only six species from five genera
of the Mimusopeae and Chrysophylleae.The striate tectum is otherwise only
recorded in Pollen Type IX, where the grains can readily be distinguished by
other characters.
Pollen Type VII is the predominant pollen type in tribe Chrysophylleae,in
particular within the genera Chrysophyllum,Micropholis and Pouteria, where it
occurs in all sections of the genus except Antholucuma. With the exception of
Type XI it is the only pollen type in which most species have a protrudent
or semi-protrudent tectum (only about one third of the specimens examined
for Type VIII have this characteristic). Furthermore, apart from Type VIII,
the continuous endexinous thickening is found most frequently in association
with Type VII. It is the only pollen type where there is usually a clear dif-
ference in the appearance of the tectum between the apo- and mesocolpial
regions. It is predominant among the Neotropical species of the family but
many species, particularly of Pouteria, occurring mainly in Malesia, the Pacific
Islands, the Philippines, New Caledonia and Australia, also share it. Similar
pollen is also found in the two species of Delpydora(subtype VIIC), from West
Africa, although in this genus the exine is distinctly and uniquely corrugate
(Fig. 30: A-E).
Pollen Type VIII shares certain similarities with the pollen of a number of
species included in Type VI. However the presence of a continuous endexinnus
thickening and the absence of a striate tectum (except in Pradosia cuatrecasasii)
found
supports the circumscription of a separate pollen type. Such pollen is
mainly in genera of tribe and
Chrysophylleae, it is the predominant pollen type
in Niemeyera, Pradosia and Leptostylis but is rare in Chrysophyllumand Pouteria.
It also features prominently amongst the few specimens examined in tribe
Omphalocarpeae.
Although Types I- VIII account for most sapotaceous pollen, there are
some pollen types or subtypes that occur in only one genus (excluding mono-
specific genera) or section of a genus. Type VE is specific to Pycnandra, and
Type VIIC to Delpydora. However, the most distinctive examples are in the
remaining four pollen types.
Pollen Type IX comprises small subprolate or prolate-spheroidal grains
with a distinctly striate tectum, found only in Pouteria section Oxythece.They
can be readily distinguished from the striate grains of Pollen Type VI by size
and shape.

Pollen Type X is recorded for six of the seven collections of Elaeoluma
examined in the present study but has not been encountered elsewhere in the
family.
Pollen Type XI is distinguished by the presence of spinules, and subtypes
XIA & XIC, in particular, are congruent with currently recognized groups
(except Pouterialongifolia). Subtype XIA is particularly distinctive, and occurs
only in Pouteria longifolia and Sarcaulus. Subtype XIB has only been recorded
in two species of Pouteria, P. cladantha and P. vernicosa, both in section Fran-
chetella. This subtype is the only example outside Types I and III where a
granular tectum is recorded, but otherwise there are no features to suggest
an affinity with those pollen types. Subtype XIC has only been recorded for
the two species of Chromolucumaexamined.
Pollen Type XII includes the only spiny pollen grains found throughout the
entire family. Such grains are found in Diplo6n cuspidatum(subtype XIIA), and
in a somewhat different form in Micropholis retusa(subtype XIIB). The latter
is particularly anomalous, since it occurs in a genus which has otherwise been
found to have pollen exclusively of Type VII. There is no macromorphological
evidence to suggest that this species is not correctly placed in Micropholis.
For the present at least, the origins of the tew anomalous pollen morphologies
encountered remain obscure. Interestingly, C. marginatumsubsp. marginatum
is placed in the same section of Chrysophyllumas C. rufum, but in C. marginatum
subsp. marginatumthe thickening of the polar wall is caused by an increase in
the length of the columellae, rather than by additional thickening of the foot
layer.
The more or less porate pollen of Englerophytumstelechantha,could be con-
sidered to represent the extreme of aperture reduction in the family; the colpi
have virtually 'disappeared'.
Pollen morphologyas a taxonomiccharacter

Within Sapotaceae none of the five tribes recognized by Pennington (1991)
can be recognized solely by pollen morphology. However, there are definite
palynological trends that correlate quite well with Pennington's tribal arrange-
ment (Table 1). Types I, II and III predominate in tribes Mimusopeae and
Isonandreaebut are rarely found in Chrysophylleae(except in Synsepalum and
Xantolis) or Omphalocarpeae.Types V, VII and VIII are very common in
the tribes Chrysophylleaeand Omphalocarpeaebut absent from Mimusopeae and
Isonandreae. Type VI pollen is rarely found outside Sideroxyleae.Within the
Sideroxyleaethere are a significant number of species with pollen types charac-
teristic of Mimusopeae, Isonandreae and Chrysophylleae.Pollen types in some
instances clearly add support to generic groups; particularly within large
genera such as Pouteria where sections Antholucuma and Oxytheceare readily
defined palynologically. Occasionally the pollen of a second collection of a
species is sufficiently different to be placed in another pollen subtype; for
example, one collection of Elaeoluma nuda has pollen in subtype VC while the
other 2 collections of this species and the remaining species examined are in
Pollen Type X. This demonstrates the need for caution in assigning pollen
grains to species or, in a number of instances, to genera. More importantly
this underlines the need to examine at least two collections of a species wher-
ever practicable before attempting to make decisions regarding taxonomic
affinities. The variable nature of the pollen morphology even within quite

small taxonomic units is evident in such genera as Englerophytum,Niemeyera,

Pradosia and Synsepalum. In contrast the pollen of genera such as Elaeoluma,
Mimusops, Payena, Pycnandra, Sarcaulus and Sarcospermais fairly homogeneous.
The degree of overlap and the number of exceptions in pollen characters
between the major groups is of about the same order of magnitude as that in
macromorphological characters, and supports the view that tribal rather than
subfamilial rank is appropriate for the primary divisions of the family.
Pollen morphologicalconsiderations
The widely accepted functions of the pollen wall are those of protecting the
male gametophyte against excessive dehydration, transport of recognition
substances involved in pollen-stigma interaction (within or on the surface of
the exine) and subsequent germination. Ideally the pollen wall provides a
structure with sufficient elasticity to withstand changes in volume, but also
forming a more or less impenetrable barrier to excessive water loss. The shape
of some pollen types is not greatly altered by the state of hydration. Pollen
grains with longer colpi and without a continuous endexine in the mesocolpial
region (e.g. Pollen Type III 'Manilkara' (Fig. 16: B, C & E)) are susceptible
to infolding in the non-hydrated state or, as a result of acetolysis may collapse.
Many workers, including Heslop-Harrison (1979),-Muller (1979), Rowley
(1981) and Payne (1981) have reviewed the possible structure-function
relationships of the pollen wall, and the possible reasons for the seemingly
endless modifications which exist to the basic structure. Frequently the wall
has been viewed from a structural point of view with the tectum, foot layer
and infratectum being likened to beams and columns (Skvarla, Raven &
Praglowski (1976), Bolick (1981)). However, with regard to the modification
of the infratectum, the pollen of the Sapotaceaedoes not appear to conform to
this basic arrangement. Certainly in Pollen Type VII rigidity is achieved by
the modification of the endexine rather than the ectexine; instead of rigid
beams the tectum and foot layer of the ectexine have the appearance of two
flexible curves enclosing a narrow band of beads or spacers. Crane (1986)
comments that the densely granular, alveolar or irregularly columellate infra-
tectum is relatively well-adapted under normal conditions to counterbalance
stress by further compaction. Examination of acetolysed pollen with light
microscopy is particularly revealing, as 'lines of collapse' are often very
obvious. In those grains with continuous endexinous thickening, the fold or
collapse lines occur above and below but parallel to the equatorial axis (Fig.
29: B-D), whereas in grains with endexinous thickening more or less
restricted to the apertural area these 'lines' frequently tend to occur parallel
to the polar axis (Fig. 16: B-C).
The occurrence of a granular rather than a columellate pollen wall is well-
documented in gymnosperms (GullvAg 1966; Van Campo & Lugardon 1973;
Kurmann 1990). The granular wall or infratectum is also present in a wide
variety of angiosperm families, both primitive and more highly evolved,
including Annonaceae (Le Thomas 1980; Le Thomas & Lugardon 1976),
Magnoliaceae (Praglowski 1974), Leguminosae (Ferguson & Skvarla 1983),
Juglandaceae (Stone & Broome 1975), Sonneratiaceae(Muller 1978), Betulaceae
(Crane 1986) and Acanthaceae (Furness 1990). More remarkable than the
presence of a granular infratectum in Sapotaceaepollen is the extreme narrow-
ness of this stratum coupled with the exceptionally thick foot layer and

21
18
3 - 4-colporate
-0 4 - 5-cotporate
a) 15
c
x
Q)
c
u, 12
a)
E
a)
a-9
0
a)
-
P"
E
z
3
2n M f N
LM c-1
L LM 0 LLF)C
C'4rage
N 0 60 0-1 6 6 -1 6
LMo .
averagepolar length(gm)
FIG. 2a. Histogram to show relative average polar length range and distribution between 3-4
and 4-5-colporategrains.
tectum, none of which appear to provide more than the most minimal storage
or exit channels for the accommodation of tapetally derived recognition
substances. Modifications to the tectum observed in the apocolpium and
mesocolpium of the various pollen types apparently provide alternative
adaptations to storage and exit functions. The usually smooth apocolpium
with few perforations encountered in Type VII is counterbalanced by a
rugulate mesocolpium. In some other Pollen Types, in particular I, II & X
(Figs 10: A-B; 12: A; 38: M-P), the tectum arrangement in Type VII is
paralleled by an increase in the frequency and density of the apocolpial
perforations combined with a sparsely perforate mesocolpium.
The protrudent tectum, narrow endoaperture and reduced colpus associated
with Type VII suggest a very specialised protection of the gametophyte and
modification for the exit of the pollen tube. The possible advantages of the
distribution of the endexine in this pollen type or alternatively the endexine
distribution in some of the other pollen types where it is more restricted have
been discussed in Harley (1986b).
In common with the pollen of numerous species of plants, the endexine in
the polar regions of a grain may be very thin. In Sapotaceae,with few excep-
tions, the less frequent condition of endexine absent in the polar region is
observed in the pollen of most species.
The occurrence of 3-4 or 4- 5-colporate pollen grains within a single sample
is one of the characteristic features of the family and is not a particularly

40
3-4-colporate
30 [ 4- 5-colporate
-0
x
a)
rC
0)
E 20
shape ratio(P/R
FIG. 2b. Histogram to show relative shape class ratios and distribution between 3-4 and
4- 5-colporate grains.
useful character at generic level; for example, in the monospecific genus

Diploon (subtype XIIA), of which two collections were examined, one had 3-4
and the other 4-5-colporate pollen. Micropholis and Sarcaulus are the only
genera which have a consistent number of apertures (three). Aperture number
has been useful, however, in some sections of the larger genera such as
Pouteria, for example; sections Antholucumaand Pouteria. 3- 4-colporate pollen
grains predominate in Types VII, IX, X & XI, while 4-5-colporate grains
predominate in Types I, II, IV & V. Within a single sample pollen grains
with a greater number of apertures are no larger than those with a lesser
number of apertures. The possibility that 4-5-colporate grains might be
larger on average than 3-4-colporate grains was examined. 62 species had
both 3 and 4-colporate pollen whilst 110 had both 4 and 5-colporate pollen.
The 3- 4-colporate grains show a less dispersed size range. The most frequent
average polar length recorded for 3-4-colporate grains is about 10 ~m less
than that recorded for 4-5-colporate grains (Fig. 2a). An analysis of relative
occurrence and distribution of 3-4 and 4- 5-colporate pollen throughout the
shape classes reveals that 3- 4-colporate grains are similarly distributed in
both prolate-spheroidal to subprolate and prolate classes. 4-5-colporate
grains, on the other hand, are predominantly prolate-spheroidal and sub-
prolate (Fig. 2b). Generally the ratio of one aperture number to the other
within a given sample is in the region of 0-25-0-30 to 0-65-0-75. This
suggests that it could be related to the relative position of the four daughter

cells in the developing tetrad. The apertures are arranged according to the
laws of equidistance as defined by Tammes (1930):-'In an equatorial plane
with equal distances between them and an equivalent position with respect to
the poles'. Tammes (1930) cites a number of genera in different families
where a variable number of apertures exists between grains within a single
sample. He links it with polyploidy (of individuals rather than a species).
Tammes also discusses the possibility that it might be associated with an
unequal distribution of chromosomes, resulting in abnormal meiotic division
with a deviating number of daughter cells. This situation however, usually
results in cells of differing size, frequently infertile. In the Sapotaceaethis does
not appear to be the case; the pollen grains within a given sample are not only
similar to each other in size but have every outward appearance of normality.
A comparable situation in relation to variable aperture number has also been
observed in the pollen grains of the Meliaceae (Pennington & Styles, 1975) and
those of Alnus (Wodehouse, 1935).
Sowunmi (1978) summarizes and discusses some of the published data
relating to polymorphism in pollen, suggesting that whatever the differing
causative factors they almost certainly operate at meiosis.
Neither pollen development nor viability studies have been undertaken in
the present work and knowledge of pollen development or viability within the
Sapotaceaeis very limited. In relation to viability, Bhatnagar & Gupta (1970)
remark, not only in relation to Mimusops elengi but also more generally within
the family that, 'Although the plants flower profusely, seed set is relatively
low'. Bai & Subba Reddi (1980) and Kuruvilla (1989) record high pollen
output for Madhuca indica but nevertheless, Kuruvilla (1989) describes seed set
for the species as low. Studies with a much wider range of living material are
obviously needed.
Evolutionary trends
Two major trends are observed in the pollen morphology of the Sapotaceae,
of which the end results are best illustrated by Pollen Type VII, with a con-
tinuous equatorial endexinous thickening, reduced colpi and differentiated
tectum ornamentation and Pollen Types I and II where the endexinous
thickening is restricted to the area surrounding the endoapertures and
the tectum ornamentation shows little differentiation between meso- and
apocolpium.
The widely held view that the triporate condition is derived from the
tricolpate or tricolporate condition (e.g. Doyle 1969; Kuprianova 1969; Van
Campo 1976; Punt 1976) by a staged reduction in colpal length, suggests that
the frequently very reduced or even vestigial colpi in Pollen Types VII and
XI are more highly evolved than the longer colpi in most of the other pollen
types. The continuously thickened band of endexine could then be interpreted
as an adaptation of the more localised endexine, to protect the exine against
excessive harmomegathic stress (Harley, 1986b). There are considerably more
species with this pollen type in the Chrysophylleaethan in the Mimusopeae; the
number of species in the Sideroxyleaewith this type is apparently intermediate.
An increase in aperture number is also generally considered an advanced
feature although other factors, already discussed, may also be involved.
Spheroidal or oblate pollen grains are similarly considered more advanced
than prolate grains, particularly since they are frequently associated with

reduced apertures or the porate condition. The grains in Pollen Types IV and
V, are frequently prolate-spheroidal or spheroidal, but usually have a well-
defined colpus.
The granular ornamentation of the pollen in many species of Mimusops,
Madhuca, Manilkara, Payenaand Palaquiumspecies, although often anastomosed,
giving a slightly rugulate appearance never occurs as fully rugulate in these
groups. Pollen Types VI, VII & VIII, both with either striate or rugulate
pollen grains, very rarely display the granular or anastomosed granular
tectum type. If striate ornamentation evolved from anastomosed granulae it
is curious that intermediate examples are almost unknown. This suggests that
the granular and non-granular exines might represent differing evolutionary
trends. Possibly the granular or anastomosed granular exine ornamentation
is less derived in this family. Exceptionally, within the genus Pouteria, a small
group of species produce pollen with a granular tectum but with/no other
features suggestive of Pollen Types I or III.
The direction of most pollen evolutionary trends in the Sapotaceaeis equivocal
(Fig. 3). In general established evolutionary trends in pollen morphology,
linked with the most widely accepted evolutionary trends of other characters,
suggest that the tribes Chrysophylleaeand Omphalocarpeaeare more advanced
than tribes Mimusopeaeand Isonandreae.Many of the genera which show more
specialized macromorphological features are also those that have apparently
more specialized pollen. It is reasonable to assume that Pollen Type VII is
probably evolutionarily more advanced than the pollen of Types I and II. The
features described for Pollen Type VI might be considered as intermediate.
Differing trends may occur within the same genus, for example Pouteria,
although having a high percentage of species with Type VII pollen also has
many species with pollen which appears to have a less specialised morphology.
Plant characters do not evolve or adapt simultaneously with each other and
reticulate or mosaic patterns of similarity or differentiation, (heterobathmy
sensu Takhtajan, (1959)), are frequently apparent between species and alter-
native possibilities should not be ignored. Perhaps a simpler explanation, that
two parallel evolutionary lines are evident, might be more acceptable, par-
ticularly in view of the lesser number of apertures in the otherwise apparently
more specialized pollen type. This has so far been more or less supported by
much of the fossil evidence, summarized at the end of this paper (& Fig. 4),
which records both Mimusops and Pouteria-like pollen from the lower Eocene.
The affinities of the few records from the European Paleocene are less clear.
Longcolpi ) Short colpi Vestigial colpi

-- Prolate - Subprolate - Prolate-spheroidal Spheroidal---
--Ornamentation undifferentiated Ornamentationdifferentiated --

Infratectum columellate -Infratectum granulate
- Discontinuous equat. endex. thick.--Continuous equat. endex. thick.---~
-•- Tectumgranular - Anastomosedgranular- Striate - Rugulate-----

3 Apertures 4 Apertures 5 Apertures - --
-'--
Fi(;. 3. Summary of morphological characters from the pollen of the Sapotaceae with arrows
indicating possible evolutionary direction. (Arrows point in both directions where the direction
of a character trend is uncertain.)

Pollination
Information on pollination within the family is sparse, although it is not
unreasonable to assume from the overall morphology and habitat of the
majority of the species that a fairly limited range of vectors operates. It is quite
probable that many species are not pollinator-specific and can take advantage
of more than one vector type. Straka, Simon & Cerceau-Larrival (1967)
suggest 'entomophilie?, chiropterophilie?' for the Madagascar genera which
they examined. Ayensu (1974), considers that fruit bats in West Africa may
play a partial role, at least, in the pollination of Butyrospermumparkii whilst
foraging for the fleshy fruits. Whitmore (1978), describes the flowers as 'often
nocturnal and bat-pollinated'. The majority of Sapotaceaeare canopy trees of
lower montane primary tropical rainforest, particularly in South America. In
the Old World a few are understory forest trees, occasionally of secondary
forest. Pennington is of the opinion (pers. comm.) from field observations that
the floral presentation and the strongly scented flowers of many species,
coupled with their canopy position in the forest ecosystem, strongly indicates
insect or bat pollination; whether any of the evening or night flowering species
also attract moths is uncertain. Faegri and van der Pijl (1979) describe
and discuss bat blossoms and pollination in Madhuca species and Dumoria
(= Tieghemella), which, 'offer a very sweet and easily detached corolla.'
Pollinators include both fruit- and nectar-eating bats. Crepuscular and night-
flowering bat-adapted flowers in the tropics are typically strongly scented and
whitish or creamy in colour. In Sapotaceaethe flowers are borne in cauliflorous
inflorescences. Because spiny pollen is frequently considered to be an adap-
tation for entomophily, perhaps an insect vector may be at least partly
responsible for the pollination of Diplo6n, Sarcaulus and Micropholis retusa.
Subba Reddi (1976) discusses the possibility of wind pollination in Madhuca
indica and Bai & Subba Reddi (1980) suggest a wind-orientated pollination
syndrome for both Madhuca indica and Mimusops elengi. Pollination studies of
Madhuca indica by Kuruvilla (1989) record predation of the fleshy ripe corolla
by a variety of animals and birds and of the pollen by honey bees. The
study concludes however, that the species is anemophilous. Wind-pollinated
plants tend to produce greater quantities of pollen than those that are insect-
pollinated. Bai & Subba Reddi (1980) record a fairly high pollen output per
anther from both Madhuca indica and Mimusops elengi, and Kuruvilla (1989)
confirms the high pollen output from Madhuca indica. Pollen productivity for
other members of the family does not appear to have been recorded.
Fresh pollen has not been examined in the present study but morphological
observations suggest that it might be dry rather than sticky. Muller (1979)
points out that the granular infratectum does not usually have a well-developed
reservoir function. He cites examples from wind-pollinated Amentiferaewhere
the structure is 'correlated with a rather uniformly thin exine, lack of tryphine
causing a dry surface, and an absence of any sculptural development of the
tectum except small spinules.' The exine in pollen of the Sapotaceaeis not
usually thin but it certainly does not provide much facility for the retention
of tryphine in the tectum. Tryphine is responsible for the sticky surface
coating (pollenkitt) on pollen of some taxa, which may act as an adhesive in
insect pollination.
Further, careful field observations are needed to improve our knowledge
and understanding of the pollination strategies operating within the Sapotaceae.

Fossil pollen
A selective summary of recorded fossil sapotaceous pollen grains within the
literature suggests both Laurasian and Gondwanan origins for the family.
Raven & Axelrod (1974) discuss the possible origins of the Sapotaceae,postulating
a west Gondwanan origin, 'almost certainly before the close of the Cretaceous'.
Muller (1981) summarizes the state of knowledge of Sapotaceaefossil pollen up
until 1980. The earliest record which he cites is Sapotaceoidaepollenites robustus
Muller (1968), from the Senonian (upper Cretaceous) and Paleocene sediments
of north-west Borneo but the present author is not convinced that this record
is sapotaceous. The walls are very thick (up to 4? m) in relation to grain size
(32-(48)-64 ttm) and have a faint columellate pattern. Muller (1968) com-
ments 'this is a rather variable type of pollen but it is always characterized
by the heavy wall, the exine structure may be corroded easily however.'
Kedves (1967) summarizes, diagramatically, the distribution of Sapotaceae-like
pollen in the Palaeocene and Eocene of France, Germany and Hungary. Such
grains are rarely recorded in the Paleocene of these three countries and
although in the Eocene sapotaceous grains are somewhat more frequent they
are very dispersed.
Fossil pollen records for the family are sporadic but widespread, suggesting
a fairly cosmopolitan distribution extending, until the cooling events during
the Miocene (see Fig. 4), well into northern Europe; for example, Tricolporo-
pollenites glaber and T. globus from the Bakony Mountains, Hungary (Deik,
1960). Both these species have a majority of features indicative of sapotaceous
pollen but the pointed (T. glaber) or square-ended (T. globus) extremities of the
broadly lalongate endoapertures are unusual. These grains may be related to
Sarcospermabut their affinity might also be with the Burseraceae.Macko (1957)
records Chrysophyllum-likepollen grains from the lower Miocene in the Valley
of Klodnica (Poland). Tetracolporopollenites manifestus R. Pot. subsp. contractus
Pflug, from the middle Eocene, upper Oligocene to lower Miocene of Germany
(Thomson & Pflug, 1953); bears a close resemblance to Pollen Type VIA or
VID. Thomson & Pflug (1953) illustrate more than 20 examples of the fossil
species Tetracolporopollenites obscurusPflug. Amongst the 20 examples there are
very noticeable differences, particularly in endoaperture size relative to pollen
grain size. Some of these grains are clearly sapotaceous, (cf. Pollen Types I
or II, sens. lat.), while in other grains the endoapertures are too broad in pro-
portion to grain size to be matched to any recent sapotaceous genus. T.
sapotoidesPflug & Thoms. (Thomson & Pflug, 1953) is comparable to Pollen
Types I- III of the present study. Also from Germany (Potonie, Thomson &
Thiergart, 1950), Sapotaceoidaepollenites micromanifestusThoms. The illustrations
of this species are not very clear; it may well be sapotaceous but the endo-
apertures seem to be very wide and narrow, which is suggestive of the pollen
of some genera in the Burseraceae,e.g. Canarium L., CrepidospermumHook. f.
and TetragastrisGaertn. (Harley, unpub.). Gruas-Cavagnetto (1976) in her
studies of the Paleogene ( = Eocene + Oligocene) of southern England records
a number of different sapotaceous pollen types including: Tetracolporopollenites
halimbaenseKDS (cf. Pollen Type VIA & IIA), T. kirchheimeri(Reiss.) Thoms.
& Pflug (not Madhuca as indicated; the very short colpi, costate endoaperture,
protrudent tectum and fairly thick pollen walls suggest Pollen Type VA), T.
megadoliumR. Pot. (could be sapotaceous, maybe Pollen Type VI, but not
Mimusops, as she suggests, since it is prolate), T. obscurus(this may not be

Isonandra but possibly a less frequent sapotaceous type; there are some simi-
larities to Pollen Type VI but the perforate tectum is unusually coarse and
endoapertures very large), T. occultusPflug & Thoms. (probably Mimusops or
Palaquium). Harley, Kurmann & Ferguson (1991) record the occurrence of
Sapotaceae-like pollen grains from the middle Eocene, Isle of Wight, and
describe, from both light and transmission electron microscopy, a grain
resembling that of recent Tieghemellaheckelii and another grain with a very
similar appearance to either Pollen Type I or II.
In Africa, fossil records of Sapotaceaeinclude Psilastephanocolporitesmalacan-
thoidesJan du Chine et al. (1978), from the upper Eocene of southeast Nigeria,
which they compare with Pouteria( = Malacantha) alnifolia. Psilastephanocolporites
perforatus Salard-Cheboldaeff (1978), from the lower Miocene coastal sedi-
ments of Cameroon is compared to Butyrospermum( = Vitellaria)and Manilkara
but, as the epithet suggests, the tectum is perforate. Perforations in the tectum
of sapotaceous pollen are not often clearly resolved with the light microscope,
except (present paper) Pollen Types ID, IIB, XA (Fig. 39: M-P) and XB.
Possibly the grains represent one of the less frequent subtypes in either Pollen
Type I or II sens. lat. of the present arrangement. Similar comments apply
to the other fossil pollen type P. punctatus, described in the same paper. Sah
(1967) describes four new species of Sapotaceae-like fossil pollen from the
Neogene (= Miocene + Pliocene) of Burundi; Sapotaceoidaepollenites communis,
reminiscent of Pollen Type VIA; S. africana, which could be a Pollen Type
VIF, and S. obscurusand S. parvus, whose affinities are less certain although
they could be sapotaceous.
In India Ramanujam (1966) describes Sapotaceoidaepollenitesneyvelii (S.
arcotensein the plate legends), from the Miocene lignite of Madras. There are
records of a Pouteria-like pollen, lugopollis sp. Venkatachala & Rawat, from the
lower Eocene of Cambay but Rawat, Mukherjee & Venkatachala, (1977),
suggest a similarity with Sonneratiaceae.Records of Sapotaceoidaepollenites
oblongus
Venkatachala & Rawat, S. oblongatusVenkatachala & Rawat and Sapotaceoidae-
pollenites sp. Pot. Thoms. & Thierg. (Venkatachala & Rawat, 1972) are not
comparable with pollen of any recent genera seen by the present author.
However, IugopollistetraporitesVenkatachala & Rawat from Kauvery, Krishna-
Godvari and Cambay, (Venkatachala & Rawat, 1972, Venkatachala, 1974
and Rawat, Mukherjee & Venkatachala, 1977), which these authors compare
to Sonneratia alba J. E. Smith and which Muller (1978) considered to be
sapotaceous, is obviously Pouteria-like (cf. Pollen Type VIIA).
The lower Eocene of Australia has yielded Sapotaceoidaepollenitesrotundus
Harris (Harris, 1972 and Stover & Partridge, 1973), which Harris (1972)
compares with Tricolporopolleniteslatizonatus McIntyre from the middle to
upper Eocene of the New Zealand Tertiary (McIntyre, 1965, (wrongly cited
in Harris (1972) as McIntyre (1968)). McIntyre (1965) likens this pollen type
to Pouteria( = Planchonella)novo-zeylandica.The species certainly bears a striking
resemblance to some sapotaceous pollen grains although it lacks the continuous
endexinous thickening and (usually) narrower endoaperture associated with
most species of Pouteria. It is more likely to be either Pollen Type I or II.
In the U.S.A., records of sapotaceous pollen exist from the lower Eocene
(Claiborne Flora) (Fairchild & Elsik, 1969), (cf. Pollen Types IA-C or IIA
and Type VIIA), and upper Eocene, Frederiksen (1980), Chrysophyllum
brevisulcatumFrederiksen (either a Chrysophyllumor a Pouteria, Pollen Type

OD-D0-0
"
D oo Z
S23 3 (c)
.0 M (D
QUATERNARY:
QUATERNARY ,0 -
" .." -"C
TERTIARY - 0
: "
BOUNDARY
'
N
upper
03
I N- -,%(D' :3•-r
E
Miocene
early .D
upper
,
E-ocene - -
."
aleocene
upper ??- .:.: ~ o LB
j~
o
Z
,.
early o? 0
.
early 00
0* 0
*,..
TERTIARY0
TERTIARY I? I?
gq t C
CRETCEOU
BOUI`,JDARB
.
c D A
Key. c
ialeocene . . .
?'
.
'
TERTIARY
• •
CRETACEOUS ?
BOUNDARY A B C D E
Key..........
FC(;.4. A summary of the fossil records for the Sapotaceae, discussed in the text. The key relates
to accepted Sapotaceous records, ?? = probably not Sapotaceae. Geological time scale based on
Van Eysinga (1975).
Key: A = Pollen Types I, II or X
B = Pollen Type VI
C = Pollen Type VII
D = more than one sapotaceous pollen type recorded
E = Sapotaceae pollen, type not known.
Note the changed emphasis in the regional fossil localities after early-mid Miocene cooling.

VIIA), of southeastern U.S.A. Taylor (1989) has found Mimusops-like pollen
brevis Frederiksen, also from the Claiborne for-
grains, Tetracolporopollenites
mation. This is an important paper since the grains are described from both
light and scanning electron microscopy. A finely anastomosed-granular surface
is resolved. The endoapertures in proportion to grain size are surprisingly
large for sapotaceous pollen, reminiscent of the Madagascar genus Tsebona
macrantha. The fossil grains are, however, very much smaller. The Brandon
lignite (upper Oligocene) of Vermont has yielded several Mimusops- or
Manilkara-like pollens (Traverse, 1955). The four pollens which are ascribed
to Manilkara species match Pollen Type IA, while the two assigned to Mimusops,
M. elengi and M. mirabilis, are more difficult to reconcile since, although the
pollen has a number of sapotaceous features, the verrucate exine described for
the pollen of both species has not been encountered in the present study. The
work of Traverse (1955), is particularly interesting in relation to Sapotaceae:-
'The discovery of mimusopoid pollen in the deposit-it is quite abundant-
was perhaps the greatest single contribution of the pollen studies to the study
of the whole problem of the deposit. As a result of this identification, the most
abundant form of fruit in the deposit was also found to belong to the Mimusops
complex, and one of the unknown wood types was placed in the same group.'
Meso-American records include Chrysophyllum-likepollen and also, (assigned
to Protium Burm. f. (Burseraceae)),decidedly Pouteria-like pollen (Pollen Type
VIIA) from the Eocene of Panama (Graham, 1985 (Figs 42-44)), and
-
Chrysophyllumor a small Pouteria (18 9 x 13 -5 m) (Pollen Type VIIA or
VIIB) from the Oligocene of Puerto Rico (Graham & Jarzen, 1969).
In South America Lorente (1986) describes two species of Psilatricolporites
Van der Hammen, from the lower and middle Miocene of Venezuela. P.
maculosus Regali, Uesugui & Santos is reported as resembling Chrysophyllum
argenteumwhile PsilatricolporitespachydermatusLorente may be attributed to
Omphalocarpumon basis of size and endoaperture shape. Its general appear-
ance is however, more reminiscent of Burseraceaepollen.
Note added in proof: Xantolis racemosa (Dub.) Royen = Sideroxylon racemosum
(Dub.) Lecomte (p. 393, 491).
ACKNOWLEDGEMENTS
I would like to record my thanks to Dr Terry Pennington who invited me,

a very long while ago, to examine the pollen of the Sapotaceaeand to Dr Keith
Ferguson who has never failed in his encouragement and enthusiasm to see
this work completed. To them both I am also grateful for some very inter-
esting and stimulating discussions. To Dr Peter Crane (Field Museum of
Natural History, Chicago), I am indebted for his painstaking care in reviewing
the manuscript. His comments and criticisms have been invaluable.
APPENDIX
Summary of pollen morphologicalcharacteristicsof otherfamilies that are relatedto, or

that have some macromorphologicalor pollen characteristicscomparableto those of,
Sapotaceae.
Pollen of Sapotaceae has a number of distinctive characteristics; never-
theless, there are some tropical or subtropical families which have pollen that
has been or might at times be confused with that of Sapotaceae.Harley (1986a)

stated that; 'the pollen [of the Sapotaceae] cannot easily be confused with
that of the Ebenaceae,Myrsinaceae, Styracaceaeor Symplocaceae(families allied or
associated with the Sapotaceae)using light or electron microscopy.' This com-
ment was based on published and unpublished data, personal observations
with light microscopy and a small amount of electron microscopy. However,
it is difficult to summarize, using a single suite of characters, the various
characteristics which enable the identification of the pollen for each of these
families, except where the pollen for a family is stenopalynous. Some guide-
lines are offered below.
In Ebenaceae, Myrsinaceae, Styracaceae,and Symplocaceaenone of the grains
have a continuously thickened band of endexine such as that recorded in the
equatorial region of the grains included in Pollen Types VII and VIII of the
present paper. Absence of an equatorial endexinous thickening is also true for
Burseraceaeand Meliaceae, within the Sapindales, included in the present dis-
cussion because, although not closely related to Sapotaceae,their pollen shows
certain similarities which may cause confusion. Ebenaceaeand Styracaceae,both
included in the Ebenales, along with the Sapotaceae,were described by Erdtman
(1952) as having pollen similar to that of the Sapotaceae.On the other hand,
some closely related families have pollen that is quite distinct from that of
Sapotaceae. Symplocaceae,for example, has very distinct pollen but at various
times has been placed within the Ebenales or the Myrsinaceae.
Straka, Simon & Cerceau-Larrival (1967) describe and illustrate the pollen
of eight genera of the Myrsinaceaefrom Madagascar; Maesa Forssk., Monoporus
A.DC., Afrardisia Mez, OncostemumA. Juss., Badula A. Juss., Embelia Burm.
f., Rapanea Aubl. and Ardisia Sw.; Barry & Perez de Paz (1979) describe
pollen of a further two genera, PleiomerisA.DC. and HeberdeniaBanks, from
Macronesia. The most distinctive feature of Myrsinaceaepollen is its small size,
rarely in excess of 27 Am and frequently as little as 10 -0 Am polar length.
"-0
Furthermore, with the exceptions of Myrsine L. and Rapanea which appear to
be 4-5-colpate, the pollen is consistently tricolporate. Generally speaking the
endoapertures are very large in comparison to overall pollen grain size. Colpi
are usually long, at least three quarters the polar length. The pollen shape is
spheroidal, prolate-spheroidal or, occasionally, subprolate. The tectum appears
to be either scabrate, coarsely perforate or finely reticulate. There are a few
-
genera within the Sapotaceaethat have species with pollen grains less than 27 0
Am and, exceptionally, as small as 16 -0 Am. In none of these, however, is the
sum of all pollen characters sufficiently similar to be confused with pollen
grains from representatives of the Myrsinaceae, particularly in view of clear
differences in endoaperture size and tectum ornamentation.
For Ebenaceae,Ayala-Nieto & Ludlow-Wiechers (1983) describe four species
of Diospyros L. from Veracruz and Lieux (1982) describes another from south-
eastern U.S.A.. The pollen grains are consistently tricolporate, the colpi
usually being very long, often almost meeting at the poles; in some species
a distinct constriction is visible in the endoapertural region of the colpus.
There is frequently a distinct colpal margo which, in partially dehydrated
grains, has a ridge-like appearance, apparently due to a thickening of the
underlying endexine. The mid-mesocolpial area usually shows a slight variation
in tectal ornamentation. Similar characters are observed for the pollen of
Euclea Murr., the only other genus. Although the pollen of some species of the
Sapotaceaemay be tricolporate and lack a continuously thickened endexine in

the equatorial region, none of the other characters described for ebenaceous
pollen are recorded within the Sapotaceae.
The Styracaceaeis a fairly stenopalynous family that is palynologically quite
distinct from the Sapotaceae. Its most characteristic pollen features are con-
sistently tricolporate apertures, usually with a central constriction in the
colpus, and very large (in proportion to grain size) broadly lalongate endo-
apertures. The colpi are long and in many species almost meet at the poles.
In some cases, particularly where the pollen grains are partially collapsed, this
feature has caused them to be erroneously described, from light microscopy,
as syncolpate (Graue-Wiechers & Ludlow-Wiechers, 1984). Grains are usually
oblate-spheroidal, spheroidal or prolate-spheroidal in shape and in equatorial
outline the grains may be rounded or angular at the equator; the latter gives
the grains a squat, diamond-shaped appearance. In polar view the outline is
blunt-triangular. All these features have been well illustrated by various
authors, in particular for Styrax L.; Guayana Highlands, Maguire & Huang
(1978) and Veracruz, Graue-Wiechers & Ludlow-Wiechers (1984). Yuan-
Hui & Cheng-Hong (1985), provide a wider survey of genera in which they
include a number of scanning electron micrographs to illustrate fine detail of
tectum surface. These surfaces are generally dissimilar to those recorded for
the Sapotaceae. Exceptionally, the pollen of AfrostyraxPerkins & Gilg (Baas,
1972), has 3-porate grains reminiscent of those of Symplocaceae.
Lissocarpa Benth. (Lissocarpaceae), a genus closely allied to Ebenaceae and
Styracaceae,also has pollen grains very similar to those of Symplocaceae(White,
1981).
The pollen morphology of SymplocosJacq. (Symplocaceae)has been described
and/or illustrated by various authors, notably Van der Meijden (1970),
Maguire & Huang (1978), Barth (1979 & 1982) and Nagamasu (1989). The
pollen is very distinctive. The grains are oblate and either triporate or, more
frequently, tricolporate. The endoapertures are medium to large, in pro-
portion to overall grain size, and broadly lalongate or circular. The endexine
is thickened to form a distinct annulus around the endoaperture. The ecto-
aperture is a very reduced colpus that may be absent or extremely difficult
to distinguish in light microscopy. The tectum ornamentation is very variable
between species, ranging from psilate through irregular reticulate, verrucate
to echinate. There is no resemblance to the pollen of the Sapotaceae.Gupta
& Sharma (1977) describe and illustrate the pollen of a species named as
Palaquium ellipticum, but it bears no resemblance to the pollen of this species,
nor indeed to the pollen of any other species of Palaquium that has been
examined in the present study. It has much more in common with Symplocos,
and it would appear that their material may well have been misidentified.
Published pollen morphological data for the Burseraceaeis sparse and has
been summarized in Forman et al. (1989). There are a number of features
which help to distinguish pollen of the Burseraceae from the Sapotaceae.
Burseraceous pollen is almost invariably tricolporate and the apertures tend
to be one of two basic types. The less common aperture type in the family has
a circular endoaperture with more or less circular costae and a short colpus.
It is particularly associated with spheroidal or prolate-spheroidal grains from
genera such as Aucoumea Pierre, BurseraJacq. and Triomma Hook. f.. The
other aperture type usually has a narrowly lalongate endoaperture, which is
frequently very wide in the equatorial direction and costae that are located on

the equatorial margins. The colpi are long, three quarters to five sixths of the
total polar length. This aperture type is associated with subprolate or prolate
pollen grains and common to a great many genera in the Burseraceaeincluding
Canarium (Rumph.) L., CrepidospermumHook. f., Dacryodes Vahl., Protium
Burm. f., Tetragastris Gaertn. and Trattinickia Willd. In a few genera the
aperture is similar to that just described but the endoaperture is smaller
and the colpus has a central constriction, as in Boswellia Roxb. ex Colebr.
Differentiation in apocolpial and mesocolpial tectum ornamentation is not
recorded for pollen of this family.
Pennington & Styles (1975) describe the pollen of a number of genera of
the Meliaceae and most grains of this family may be distinguished from those
of the Sapotaceaeon one or a combination of characteristics. Columellae are
clearly visible in light microscopy in Aphanomixis Blume. The pollen of
ChisochetonBlume and Turraea L. share similar oblate pollen, which is 3-
colporate in Turraeabut frequently 4-colporate in Chisocheton.The endoaperture
is surrounded by a thickened costate ring and the colpus is very reduced. This
pollen has more in common with that of Symplocaceaethan Sapotaceae.In many
genera the grains have a form of lalongate endoaperture that does not occur
in the Sapotaceae.The extremities of the endoapertures are either pointed, as
opposed to blunt or round ended in the Sapotaceae,or, appear to split, some-
times in more than one direction, or they may be diffuse and difficult to
define. Genera with this endoaperture type include: Cipadessa Blume, Melia
L., Sandoricum Cav., Synoum A. Juss. and Trichilia P. Br. Other features
not commonly found in the pollen of the Sapotaceaeare broad colpi, central
colpal constrictions or a coarsely perforate to finely reticulate tectum. Aglaia
roxburghianaMiq. and A. sapindina Harms, have pollen with notable simi-
larities to SapotaceaePollen Type VII, but can be distinguished by their small
size, endoapertures which are very large in proportion to grain size and dis-
continuous endexine in the equatorial region. Pollen of some of the species
of the genus Amoora Roxb., for example A. rubizgnosaHiern, with tricolporate
apertures, may be confused with pollen of Sapotaceaealthough in fact the large
narrow lalongate endoaperture with costate thickenings on the equatorially
orientated margins are more similar to Burseraceae.The greatest confusion
between the pollen of Sapotaceaeand Meliaceaelies with those meliaceous genera
that have spheroidal or prolate-spheroidal, 4- 5-colporate pollen grains with
broad or circular endoapertures and colpi which are two thirds to three
quarters the polar length, for example; CabraleaA. Juss., Carapa Aubl. and
Ruagea Karst. These are notably similar to some of the subtypes within Pollen
Types IV and V of the Sapotaceae.DysoxylumBlume and Munronia Wight, share
many of these features but have a finely reticulate tectum to distinguish them.
Vavaea Benth., which also is comparable can be separated by its very thick
costae. Unfortunately there have been few electron microscope observations
of surface ornamentation of the tectum or of wall stratification, for any of
these families. Because these features are distinctive in sapotaceous pollen
such data may be extremely helpful in separating pollen of Sapotaceaefrom
similar forms in Burseraceaeand Meliaceae.

424 KEW BULLETIN
VOL.46 (3)
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& Rawat, M. S. (1972). Palynology of the Tertiary sediments in the
Cauvery basin. I. Palaeogene - Eocene palynoflora from the sub-surface.
In: Proc. Seminar on Palaeopalynology and Indian Stratigraphy (1971).
Univ. of Calcutta: 292-335, pls 1-5.
White, F. (1981). Lissocarpaceae. In, Bassett Maguire & collaborators: The
botany of the Guyana Highland, Part XI. Memoirs of the New York
Botanical Garden 32: 329-330.
Whitmore, T. C. (1978). Sapotaceae. In: V. H. Heywood (Ed.) Flowering
Plants of the World 132 -133. Oxford University Press. Oxford, London,
Melbourne.
Willis, J. C. (1973). A Dictionary of the Flowering Plants and Ferns. Eighth
edition, revised by H. K. Airy-Shaw. Cambridge University Press.
Wodehouse, R. P. (1935). Pollen grains: Their structure, identification and
significance in science and medicine. McGraw Hill Book Company Inc.,
New York & London.
Yuan-Hui, L. & Cheng-Hong, Y. (1985). Pollen morphology of Styracaceae
and its taxonomic significance. Acta Phytotaxonomica Sinica 23: 81-90.

Figs 5 - 43
[Light micrographs (LM): scale bars = 10 m. Scanning electron micro-
graphs (SEM): whole grains, scale bars = 10 /m; surface close-ups and
wall fractures, scale bars = 1 tm unless otherwise indicated. Transmission
electron micrographs (TEM): whole sections, scale bars = 10 /m, wall
sections, scale bars = 1 tm unless otherwise indicated].
Note-light micrograph descriptions: 'different grain' describes another
grain photographed from same sample, not one with a differing mor-
phology, although there may be very slight, insignificant differences.

,?4
Y! 44'Ift
2**%
Oro
1Cl
JI %
.
FIG. 5. [Subtype IA (part)]. A-C Mimusops obovata, Codd 9705: A whole grain, equatorial view
(SEM); B & C whole grains, equatorial views (LM) B high focus; C mid-focus. D & E Xantolis
burmanica, Lace 3189: D whole grain, equatorial view, mid-focus (LM); E whole grain, equatorial
view (SEM). F Madhuca aristulata, Scortechini 1984: close-up of mesocolpium (SEM). G Payena
lanceolata, Aniff 15541: close-up of mesocolpium (SEM). H Madhuca mindanaensis, Saikeh SAN
72241: close-up of mesocolpium (SEM). J Palaquium walsuraefolium, Jacobs 5076: close-up of
mesocolpium (SEM). K Manilkara cuneifolia, Gossweiler 8649: polar wall section (TEM). L Burckella
banikiensis, BSIP 5876: polar wall (TEM).

__ B W C'
I-
40;?_
L ipm 1IJ
FIG. 6. [Subtype IA (remainder)]. A Madhuca sericea, Boschproefstation92: whole grain, equatorial

view, high focus (LM). B & C Payena lucida, Parkinson 5299: whole grains, equatorial views, B
high focus; C different grain, mid-focus (LM). D-F, L & M Mimusops angel, Glover & Gilliland
912: D-F whole grains, equatorial view, D high focus; E mid-focus; F low focus; L equatorial
mesocolpial wall section (TEM); M polar wall section (TEM). G Northia seychellana,Jeffrey 1171:
whole grain, polar outline, low focus (LM). H & J Mimusops caffra, Barbosa 2633: whole grains,
equatorial view, (LM) H mid-focus; J low focus. K Madhuca spectabilis, Kostermans 5142: whole
grain, equatorial view, high focus.

?4 C~jor t
jawr
44" 41
C4
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J...
11
FIG. 7. [Subtype IB (part) & IE (part)]. A-D (Subtype IB), Payena endertii, Chai SAN 29390:
A whole grain, equatorial view (SEM); B whole grain, equatorial view, mid-focus (LM); C whole
grain, polar plane (TEM); D close-up of mesocolpium (SEM). (Subtype IE), E Manilkara littoralis,
Parkinson 66: whole grain, equatorial view (SEM). F & G (Subtype IB), F Palaquium sp. SAN
34407: close-up of mesocolpium (SEM). G. P. ottolandzeri, Koorders 10158p: close-up of
mesocolpium (SEM).

?
;ALr,?
-tA
A
i DO* . '
w pr - I*-f 1 I
C 4
1 0 I
F 1pm
FIG. 8. [Subtypes IB (remainder), IC, ID & IE]. A, B & E (Subtype IC), Tsebona macrantha,
Capuron SF 27675: A & B whole grains, equatorial view (LM), A high focus; B different grain,
high focus; E polar wall section (TEM). C & D (Subtype IE), C Manilkara littoralis, Parkinson 66:
whole grain, equatorial view (LM), low focus; D Madhuca motleyana, Sinclair 40328: whole grain,
equatorial view (LM), mid-focus. F (Subtype IB) Payena endertii, Chai SAN 29390: polar wall
section (TEM).

Ii
I
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LII
d
"r t
i-
41
FIG. 9. [Subtypes IF & IG]. A-G (Subtype IF), A Burckellafjiiensis, Howard H117: whole grain,
equatorial view (SEM). B & C Madhuca burckiana,Jacobs 5268: B whole grain, polar plane (TEM);
C polar wall section (TEM). D Manilkara trilora, Irwin 5042: close-up of mesocolpium (SEM).
E Burckella parviflora, Damaru 145: close-up of mesocolpium (SEM). F & G Manilkara pubicarpa,
FDBG 5860: whole grains, equatorial view (LM), F low focus; G mid-focus. H- K (Subtype IG),
H Palaquium obovatum, Mainigay 985: whole grain, equatorial view, mid-focus (LM). J & K
Madhuca neriifolia, Pennington 100: whole grain, equatorial view (SEM); K close-up of
mesocolpium (SEM).

4
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cra
,? soleeL' 4Le
c 40
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4=6 It
.IL tI
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FIG. 10. [Subtypes ID & IIA (part)]. A-F (Subtype ID), A Madhuca laurifolia, Curtis 2254:
whole grain, equatorial view (SEM). B Mimusops antongilensis, Capuron 22847SF: whole grain,
equatorial view (SEM). C-F Mimusops marginata, Strey & Moll 3679: C & D whole grains,
equatorial view (LM), C low focus; D high focus; E close-up of apocolpium (SEM); F close-up
of mesocolpium (SEM). G-L (Subtype IIA), Palaquium philippense, Loher 6564: G whole grain,
equatorial view (SEM); H whole grain, polar plane (TEM); J whole grain, equatorial plane
(TEM): K polar wall section (TEM); L close-up of mesocolpium (SEM).

41
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"g"
r
...
EAlp F
i!
Fic. 11. [Subtype IIA (remainder)]. A Palaquium rufolanigerum, Bujang S32502: whole grain,
equatorial view (SEM). B Madhuca sp., Pennington et al. 10250: close-up of mesocolpium (SEM).
C & D M. tubulosa, WhitmoreFRI 8816: C whole grain, polar outline, high focus (LM); D close-up
of mesocolpium (SEM). E M. orientalis, Brass 7747: whole grain, equatorial view, mid-focus
(LM). Palaquium gutta, 7' & P 362: whole grain, equatorial view, high focus (LM). G Labour-
donnaisia revoluta, Lorence 1602: close-up of mesocolpium (SEM). Palaquium leiocarpum, Paie S.
26046: close-up of mesocolpium (SEM).

It'I
%b
i?
5w ."A
G X, ??
Pra
FIG. 12. [Subtypes IIB & IIC]. A-E (Subtype IIB), A & B Palaquium cochlearifoliumAshton
BRUN 978: A whole grain, equatorial view (SEM); B close-up of apocolpium (SEM). C-E
Madhuca coriacea,Haviland 2118: C whole grain equatorial view (SEM); D close-up of mesocolpium
(SEM); E close-up of apocolpium (SEM). F - L (Subtype IIC), F Madhuca pallida, de Wilde & de
Wilde Duyfjies 14893: close-up of mesocolpium (SEM). G, K & L Palaquium lobbianum, Buwalda
4963: G whole grain, equatorial view, high focus (LM); K whole grain, equatorial view (SEM);
L close-up of mesocolpium (SEM). H P. neo-ebudicum, Bernardi 13186: whole grain, equatorial
view (SEM). J P. formosanum, Wilson 10999: whole grain, equatorial view, low focus (LM).

wAf
'*4W
!
.,
FIc. 13. [Subtype IID]. A-C Isonandra stocksii, RamamoorthyHFP 1550: whole grain, equatorial
view (SEM); B & C whole grains, equatorial view (LM), B high focus; C different grain, low
focus. D & H Isonandra villosa, Herb. Wight. 1733: D whole grain, equatorial view (SEM); H close-
up of mesocolpium (SEM). E & F Sideroxylonmascatense, Harris 16356: whole grains, equatorial
view (LM), E high focus; F different grain, mid-focus. G Sideroxylonwightianum, Champion s.n.:
close-up of mesocolpium (SEM). J-L Aulandra longifolia, J & M. S Clemens 21531: J whole grain,
equatorial view (SEM); K whole grain, equatorial view, low focus (LM); L close-up of
mesocolpium (SEM).

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Fic. 14. [Subtypes IIIA & IIIB (part)]. A-L (Subtype IIIA), A & B, D-F Manilkara bidentata
subsp. surinamensis, Ducke 1523: A whole grain, equatorial view (SEM); B close-up of mid-
apertural area (SEM); D- F whole grains, equatorial view (LM), D mid-focus; E low focus; F
different grain, low focus. C M. discolor, Chase 4664: whole grain, equatorial view (SEM). G &
H M. mochisia, Harris & Mwasumbi 3646: whole grains, equatorial view, 2 levels of low focus
(LM). J & K M. excelsa, Ducke 17626: J whole grain, equatorial view (SEM); K close-up of mid-
apertural area (SEM). L M. multifida, Santos 1094: whole grain, equatorial view, mid-focus (LM).
M-P (Subtype IIIB), Neolemonnieraogouensis, Thollen 146: M & N whole grains, equatorial view
(LM), M mid-focus; N different grain, low focus; P whole grain, equatorial view (SEM).

?
jip
I
5 "
C
C Ih~l~eman
FIG. 15. [Subtype IIIB (remainder)]. A-D Manilkara zapota, Lundell & Contreras19135: A whole
grain, equatorial view (SEM); B whole grain, equatorial view, low focus (LM); C whole grain,
polar view, mid-focus (LM ) [NB. cytoplasmic material has not cleared in B & C]. D-G
Mimusops schimperi, Burger 514: D whole grain, equatorial view (SEM); E whole grain, polar plane
(TEM); F whole grain, equatorial plane (TEM); G polar wall section (TEM). H & L Inhambanella
guereensis, Aubriville 4147: H whole grain, equatorial view (SEM): L polar wall section (TEM).
J & K Englerophytumletestui, Le Testu 8806: whole grains, equatorial view (LM), J mid-focus; K
low focus. M Madhuca curtisii, Ridley s.n.: polar wall section.

h,,,,r
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I
iJ( "" • I 'I
- ?
F,..
J~o "
.wK ,
FIG. 16. [Subtypes IIIC & IIID]. A-M (Subtype IIIC), A-C Chrysophyllum imperiale, cult.
Ireland s.n.: A whole grain, equatorial view (SEM); B & C whole grains, equatorial view (LM),
B mid-focus; C different grain, low focus. D Synsepalum passargei, Fanshawe 1925: whole grain,
equatorial view (SEM). E-G Neohemsleya usambarensis, Polhill et al. 4990: E whole grain,
equatorial view (grain rather collapsed) (SEM); F & G whole grains, equatorial view (LM), F
high focus; G low focus. H Nesoluma polynesica, Degener & Degener 30120: whole grain, equatorial
view, low focus (LM). J & K Pouteria ramiflora, Froes 1841: whole grains, equatorial view (LM),
J mid-focus; K low focus. L Pradosia beardii, Ayliffe 14407: whole grain, equatorial view, high focus
(LM). M Chrysophyllumrufum, Martius 521: whole grain, equatorial view, low focus (LM). N-R

(Subtype IIID), N & Q Pouteria reticulata subsp. reticulata, Prance et al. 8142: N whole grain,
equatorial view (SEM); Q close-up of mesocolpiurri (SEM). P & R Pradosia schomburgkianasubsp.
schomburgkiana,Prance & Ramos 23128: P whole grain, equatorial view (SEM); R close-up towards
apocolpium (SEM).
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00
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Fic;. 17. [Subtypes IVA & IVB (part)] A-N (Subtype IVA), A & B Sarcospermalaurina Ying Hu
12355A: A whole grain, equatorial view (SEM); B close-up of mesocolpium (SEM). C S. griffithii,
Thomson s.n.: whole grain, equatorial view (SEM). D-G Pradosia brevipes, Toledo & Gehrt s.n.: D
group of 3 grains at varying angles (SEM); E whole grain, equatorial view, low focus (LM); F
whole grain, polar view, mid-focus (LM); G close-up, mid apertural area (SEM). H & J
Sarcospermakachinense, Stainton 6725: whole grains, equatorial view (LM), H mid-focus, J different
grain, mid-focus. K & L S. arboreum,Henry 12837A: whole grains, equatorial view (LM), K high
focus; L different grain, low focus. M & N Pouteria rigida subsp. tomentosa,Maguire 32783: M whole
grain, equatorial view, mid-focus (LM); N whole grain, polar view, mid-focus (LM), [note the
planaperturate amb of N & F]. P (Subtype IVB), Xantolis siamensis, Kerr 10124: whole grain,
equatorial view (SEM).

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or,
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mwf
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FIG. 18. [Subtypes IVB (remainder( & IVC]. A & B (Subtype IVB), Xantolis racemosa, Petelot
871: A whole grain, equatorial view, mid-focus (LM); B close-up of mid-apertural area (SEM).
C - P (Subtype IVC), C - H Pouteria opposita, Ducke 308: C whole grain, equatorial view (SEM);
D close-up of mesocolpium (SEM); E section of wall towards and at the pole (TEM); F whole
grain, equatorial plan (TEM); G whole grain, polar plane (TEM); H whole grain, equatorial
view, high focus (LM). J Synsepalum msolo, Eggeling 6303: whole grain, equatorial view (SEM).
K - M Sideroxylonmicrophyllum, Decary s.n.: K whole grain, equatorial view, high focus (LM); L
whole grain, equatorial view (SEM); M close-up of mesocolpium (SEM). N Sarcospermalaurina,
Tsang 30726: whole grain, equatorial view, mid-focus (LM). P Synsepalumsubverticillatum, Greenway
& Rawlins 9431: whole grain, equatorial view, low focus (LM).

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-I
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41 h
L-dab-
Fic. 19. [Subtype IVD]. A- D & L Diploknema butyraceoides,Strachey & Winterbottom1: A whole
grain, equatorial view (SEM); B close-up of mesocolpium (SEM); C & D whole grains, equatorial
view, C low focus (LM); D mid-focus (LM); L mesocolpial wall section (TEM). E -J Sideroxylon
capiri subsp. capiri, Hinton 3833: E & F whole grains, equatorial view (LM), E high focus; F low
focus (LM); G 2 whole grains, polar planes (TEM); H polar wall section (TEM); J whole grain,
equatorial view (SEM). K Labourdonnaisia glauca, Vaughan 10516: whole grain, equatorial view
(SEM). M Manilkara bidentata subsp. bidentata, Broadway 6973: polar wall section (TEM).

IF'
W'l orl
v,
G ____i ? ?
.ool'
She-
FIG. 20 [Subtypes IVE & VA (part)]. A-J (Subtype IVE), A-D Tieghemellaheckelii, Leeuwenberg
2752: A whole grain, equatorial view (SEM); B polar wall section (TEM); C & D whole grains,
equatorial view (LM), C mid-focus; D low focus. E -J Palaquium quercifolium, Kostermans 11169:
E & F whole grains, polar view (LM), E mid-focus, 5-colporate; F mid-focus, 4-colporate; G-J
whole grains, equatorial view (LM), G mid-focus; H mid-low focus; J low focus. K- M (Subtype
VA), Pichonia lauterbachiana, Brass 22061: K whole grain, equatorial view (SEM); L & M whole
grains, polar view (LM), L mid-focus, 5-colporate; M mid-focus, 4-colporate.

wl W-F
jm! 41V
4Ni
Now ~ ~
FIG. 21 [Subtypes VA (remainder) & VB]. A-K (Subtype VA), A Pichonia lauterbachiana,Brass
21910: whole grain, equatorial view, mid-focus (LM). B P. solomonensis, Mauriasi et al. BSIP
17882: close-up of mesocolpium (SEM). C-F Pouteria laurifolia, C. T. White 12863: C whole
grain, equatorial view (SEM); D whole grain, equatorial view, low focus (LM); E & F whole
grains, polar view (LM), E mid-focus, 3-colporate; F mid-focus, 4-colporate. G & H Autranella
congolensis, Louis 3521: G whole grain, equatorial view, mid-focus (LM); H whole grain,
equatorial view (SEM). J & K Pouteria campechiana, Lundell & Contreras 19241: J whole grain,
equatorial view (SEM); K close-up of mesocolpium (SEM). L & M (Subtype VB), P. keyensis,
Koster BW4442: L whole grain, equatorial view (SEM); M whole grain, equatorial view, high
focus (LM).

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C
D~I
*!
!i
M*
FIGc. 22. [SubtypesVC & VE (part)]. A-J (Subtype VC), A & B Pichonia deplanchei, Deplanche
442: A whole grain, equatorial view (SEM); B whole grain, equatorial view, high-mid-focus
(LM). C & D P. deplanchei, McKee 13552: whole grains, equatorial view (LM), C mid-low focus;
D mid-focus. E & F Elaeoluma nuda, Philcox et al. 3257: E whole grain, equatorial view (SEM);
F equatorial view, mid-low focus (LM). G-J Pouteria robusta, Robyns 3324: G-H whole grains,
equatorial view (LM), G high focus; H mid-focus; J close-up, mid-apertural area (SEM). K - M
(Subtype VE), Pycnandrakaalainsis, Pennington &McPherson 10296: K whole grain, equatorial view
(SEM); L whole grain, equatorial view, mid-focus (LM); M whole grain, polar view, mid-focus
(LM).

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k.I
"
ly
-
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FIG. 23. [Subtypes VE (remainder) & VD]. A-H (Subtype VE), A-F Pycnandra aff. vieillardii,
Pennington & McPherson s.n.: A fractured grain to show endoaperture and inner surface (SEM);
B whole grain, equatorial view, high focus (LM); C whole grain, polar view, mid-focus (LM);
D fractured polar wall (SEM); E longitudinal fracture in endoapertural region (SEM); F close-up
of surface in mesocolpial area, near to aperture (SEM). G & H Pycnandra benthamii, McKee 5187:
G whole grain, equatorial view (SEM); H whole grain, equatorial view, mid-focus (LM). J &
K (Subtype VD), Tridesmostemonomphalocarpoides,Brenan & Onochie9434: J whole grain, collapsed,
equatorial view, mid-low focus (LM); K whole grain, collapsed, polar view, mid-focus (LM).

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4N
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Aw m
FiG. 24. [Subtypes VIA & VIB]. A-G & L (Subtype VIA), A-D Sideroxyloncelastrinum, Curtiss
1765: A whole grain, equatorial view (SEM); B & C whole grains, equatorial view (LM), B low
focus; C mid-focus; D close-up of mesocolpium (SEM). E. S. ibbarae, Lundell & Contreras 19767:
whole grain, equatorial view, low focus (LM). F & G S. puberulum, Bernardi 14781: F whole grain,
equatorial view, low focus (LM); G close-up of mesocolpium (SEM). L S. eriocarpum, Conzatti
1586: close-up of mesocolpium (SEM). H- K, M -Q(Subtype VIB), H - K S. confertum, Wright
s.n.: H whole grain, equatorial view (SEM); J whole grain, equatorial view, low focus (LM); K
close-up of mesocolpium (SEM). M & N S. jubilla, Ekman 15659: whole grains, equatorial view
(LM), M mid-focus; N low focus. P & Q Pouteria dictyoneurasubsp. dictyoneura, Wright 1329: P
whole grain, equatorial view (SEM); Q close-up of mesocolpium (SEM).

AO''
\aAt
FIG. 25. [Subtypes VIC & VID (part)]. A-K (Subtype VIC), A-D Sideroxylon occidentale,
Johnston 3904: A whole grain, equatorial view (SEM); B close-up of mid-apertural area (SEM);
C & D whole grains, equatorial view (LM), C mid-low focus; D low focus. E & F Pouteria
splendens, King s.n.: E close-up of mesocolpium and mid-apertural area (SEM); F whole grain,
equatorial view, high focus (LM). G & H Sideroxylonfloribundum subsp. floribundum, March 1562:
whole grains, equatorial view (LM), G mid-focus; H different grain, mid-focus. J & K Argania
spinosa, Gattefosse 810: J whole grain, equatorial view, high focus (LM); K close-up of
mesocolpium (SEM). L - P (Subtype VID), L & M Vitellariopsiskirkii, Paulo 117: L whole grain,
equatorial view (SEM); M close-up of mesocolpium (SEM). N & P Vitellaria paradoxa, Oldeman
911: whole grains, equatorial view (LM), N mid-focus; P different grain, low focus.

,i
F G
L 2
u pm
F iGPS~~
FIG. 26. [Subtypes VID (remainder) & VIE]. A & B (Subtype VID), Sideroxylonfoetidissimum
subsp. foetidissimum, Curtiss 1759: A whole grain, equatorial view, mid-focus (LM); B close-up of
mesocolpium (SEM). C-M (Subtype VIE), C S. rotundifolium, Harris 11040: whole grain,
equatorial view, low focus (LM). D-G S. cubense, Ekman 1875: D close-up of mid-apertural area
(SEM); E whole grain, equatorial view (SEM); F & G whole grains, equatorial view (LM), F
mid-focus; G different grain, low focus. H S. montanum, Purdie s.n.: whole grain, equatorial view,
low focus (LM). J S. portoricensesubsp. minutiflorum, Hinton et al. 10416: whole grain, equatorial
view, low focus (LM). K & L Viterllaria paradoxa, Coull 1: K close-up of mid-apertural area
(SEM); L whole grain, equatorial view, low focus (LM). M Sideroxylongerrardianum, Barron 1574:
close-up of mid-apertural area (SEM).

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161
Fic. 27. [Subtypes VIF & VIIA (part)]. A-G (Subtype VIF), A-C Capurodendronrubrocostatum,
Humbert & Perrier2349: A close-up of mid-apertural area (SEM); B whole grain, equatorial view
(SEM); C whole grain, equatorial view, mid-focus (LM). D-F C. bakeri, Scott Elliot 2969: D
transverse fracture, mesocolpial wall (SEM); E & F whole grains, equatorial view (LM), E mid-
focus; F low focus. G Sideroxylonfloribundumsubsp. belizense, Schipp 1269: whole grain, equatorial
view, mid-low focus (LM). H-K (Subtype VIIA), H Pouteria caimito, FDBG 3652: whole grain,
equatorial view (SEM). J & K P. brownlessiana, K.]J. White s.n.: close-up of mesocolpium (SEM).

II
E1 o, l
,G.
A A6
pA
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W.e
M
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AND
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FIG. 28. [Subtype VIIA (remainder)]. A-D Micropholis mensalis, Maguire 24452: A whole grain,
polar plane (TEM); B - C whole grains, equatorial view (LM); B high focus; C different grain,
mid-focus; D same grain as B, low focus. E - G Aubregrinia taiensis, Enti FH 6871: E whole grain,
equatorial view, mid-focus (LM); F & G whole grains, polar view, mid-foci (LM). H & J Pouteria
sp. McKee 14204: H whole grain, equatorial view, low focus (LM); J whole grain, equatorial view
(SEM). K & L P. maclayana, Dennis et al. BSIP 8503: K close-up of apertural area (SEM); L whole
grain, equatorial view (SEM). M P. decussata, Ducke 17607: whole grain, equatorial view, mid-
focus (LM). N & P P. rubicunda, McKee 7771: N close-up of mesocolpium (SEM); P close-up of
apocolpium (SEM).

AVO
IL
FGc. 29. [Subtype VIIB]. A Ecclinusa lancifolia, Spruce 1949: whole grain, equatorial view
(SEM). B-D Chrysophyllumwelwitschii, Bos 1860: whole grains, equatorial view (LM), B high
focus; C mid-focus; D low-focus. E-H C. cuneifolium, Granville T.1087: E section through
whole grain, polar plane (TEM); F- H whole grains equatorial view (LM), F high focus, G mid-
focus; H low focus. J - N C. sanguinolentumsubsp. balata, Schultes & Lopez 9980: J - L whole grains,
equatorial view (LM), J high focus; K mid-focus; L low focus; M close-up of mesocolpium
(SEM); N close-up of apocolpium (SEM).

456 KEWBULLETIN
VOL.46 (3)
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Fa;. 30. [Subtypes VIIC & VIID (part)]. A-F (Subtype VIIC), A-D Delpydora gracilis,
Baldwin 6715: A whole grain, equatorial view (SEM); B whole grain, equatorial view, mid-focus
(LM); C close-up of apertural area (SEM); D detail of 'merged' appearance of endoapertures in
a 5-colporate grain (LM). E & F D macrophylla, Le Testu s.n.: E whole grain, equatorial view
(SEM); F fracture showing endoaperture (SEM). G-K Sideroxylon sp., Bouton s.n.: G whole
grain, equatorial view (SEM); H section through whole grain, polar plane (TEM); J section
through whole grain, equatorial plane (TEM); K close-up of apertural area (SEM).

,i??
?i,~
FIG. 31. [Subtypes VIID (remainder) & VIIE (part)]. A-C (Subtype VIID), Sideroxylon sp.,
Bouton s.n.: whole grains, equatorial view (LM), A high focus; B mid-focus; C different grain,
low focus. D - M (Subtype VIIE), Pradosia subverticillata, Ducke 812: whole grain, equatorial view
(SEM). E - F Pouteriatorta, Sandwith 347: E whole grain, equatorial view, low focus (LM); F whole
grain, equatorial view (SEM). G Pouteria pohlmannia, L. S. Smith 12486: whole grain, equatorial
view, mid-focus (LM). H P. cladantha, Clarke &Maquirino s.n.: whole grain, equatorial view, mid-
focus (LM). J & K P. obovoidea, Gafui et al. BSIP 1250: whole grains, equatorial view (LM), J
mid-focus; K low focus. P. procera, Martius 398: close-up of apertural area (SEM). M P. baillonii,
Green 1712: close-up of apertural area (SEM).

458 VOL. 46 (3)
KEWBULLETIN
D•"N"FM"
Itt
Mo o K ......... . ..D
Fic. 32. [Subtypes VIIE (remainder) & VIIF (part)]. A-G (Subtype VIIE), A-B Pouteria
australis, Mueller s.n.: A section through whole grain, polar plane (TEM); B longitudinal section
through endoapertural region (TEM). C P. sp. (Planchonella glabra) Ridley, 15770: close-up of
mesocolpium (SEM). D & E Chrysophyllumboivinianum, SF 2521: whole grains, equatorial view
(LM), D mid-focus; E low focus. F & G Pouteria toricellensis, Hertz & Katik NGF 42938: whole
grains, equatorial view (LM), F high focus; G mid-focus. H - L (Subtype VIIF), H Chrysophyllum
ovale, Klug 2324: whole grain, equatorial view (SEM). J C. sparsiflorum, Delascio & Liesner 7075:
whole grain, equatorial view (SEM). K & L C. lucentifolium subsp. pachycarpum, Silva & Souza
2555: whole grains, equatorial view (LM), K mid-focus, L different grain, low focus.

N P.
Fic. 33. [VIIF (remainder) & VIIG (part)]. A-L (Subtype VIIF), A-C Sideroxylon saxorum,
Decarv 8429: A whole grain, equatorial view (SEM); B & C whole grains, equatorial view (LM),
B high focus; C different grain, mid-focus. D & E S. salicifolium, March 1609: whole grains,
equatorial view (LM), D low focus; E different grain, mid-focus. F & G Micropholis gardneriana,
Gardner 3310: whole grains, equatorial view (LM), F high focus; G low focus. H-K Pouteria
obovata, Procter 3950: H section through whole grain, polar plane (TEM); J & K whole grains,
equatorial view (LM), J high focus; K mid-low focus. L. P. linggensis, VersteeghBW 3927: whole
grain, equatorial view, high-mid-focus (LM). M-P (Subtype VIIG), M P. gardneri, Hassler
7514: whole grain, equatorial view (SEM). N & P Chrysophyllum aff. flexuosum, Pinheiro 2178:
whole grains, equatorial view (LM), N high focus; P different grain, low focus.

460 KEW BULLETIN VOL. 46 (3)
~t... •"
__ ?
GP
K * 1
L _M
Flc. 34. [Subtype VIIG (remainder)]. A. Chrysophyllumviridifolium, Gardner 938: whole grain,
equatorial view (SEM). B & C Pouteria unmackiana, Hyland 9015: whole grains, equatorial view
(LM), B mid-focus; C low focus. D P. sarcospermoides,Carr 12933: high-mid-focus (LM). E-H
Breviea leptosperma,Tisserant 1429: E & F whole grains, polar view (LM), E mid-focus, 4-colporate;
F mid-focus, 3-colporate; G whole grain, equatorial view, mid-focus (LM); H section through
whole grain, polar plane (fEM). J Pradosia ptychandra,Mori & Veyret8982: whole grain, equatorial
view (SEM). L Pouteria multinervis, Clemens 1971: whole grain, equatorial view, mid-focus (LM).
M P thyrsoidea, Walker & White 84A: whole grain, equatorial view, high-mid-focus (LM).

'
E FD
J K
Mr
Fic. 35. [Subtypes VIIIA & VIIIB]. A-K (Subtype VIIIA), Pouteria alnifolia, Keay FHI 21076:
whole grain, equatorial view (SEM). B Englerophytummagalismontanum, Kennedy 2313: whole grain,
equatorial view (SEM). C & D Leptostylis petiolata, McKee 13084: C whole grain, equatorial view
(SEM); D whole grain, equatorial view, mid-focus (LM). E & F Niemeyera balansae, McKee 9831:
whole grains, equatorial view (LM), E high-mid-focus, F mid-focus. G & H Chrysophyllum
lanatum, Cuatrecasas 19187: whole grains, equatorial view (LM), G high focus; H low focus. J &
K Pradosia atroviolacea, Ducke 1800: whole grains, equatorial view (LM), J low focus; K different
grain, low focus. L-N (Subtype VIIIB), L & M Omphalocarpumbequaertii, Mildbraed 9021: L
whole grain equatorial view (SEM); M whole grain, equatorial view, high focus (LM). N Pouteria
sericea, Parker 509: close-up of mesocolpium towards aperture (SEM).

E7
1
,IiJt
Gt
Jb
K1
FIG. 36. [Subtypes VIIIC, VIIID & VIIIE (part)]. A-D (Subtype VIIIC), Sideroxylon
reclinatum, Fredholm 5832: A whole grain, equatorial view (SEM); B & C whole grains, equatorial
view (LM), B low focus; C different grain, high focus. E -J (Subtype VIIID), E S. portoricense
subsp. portoricense, Harris 5388: whole grain, equatorial view, mid-focus (LM). F-J Pradosia
cuatrecasasii, Cuatrecasas 13988: F whole grain, equatorial view (SEM); G half-section through
grain, polar plane (TEM); H & J whole grains, equatorial view (LM), H mid-focus; J different
grain, mid-focus. K-M (Subtype VIIIE), K & L Omphalocarpumelatum, FHI 54070: K close-up
of mesocolpium (SEM); L whole grain, equatorial view (SEM). M Pycnandra controversa,McKee
5100: whole grain, equatorial view (SEM).

F__G
J K
FIG. 37. [Subtypes VIIIE (remainder) & VIIIF]. A-D (Subtype VIIIE), A & B Pycnandra
controversa, McKee 5100: A whole grain, equatorial view, mid-focus (LM); B detail of
endoaperture, equatorial view, low focus (LM). C & D Omphalocarpummayumbense, Gossweiler
7830: whole grains, equatorial view (LM), C mid-focus; D different grain, mid-focus. E-K
(Subtype VIIIF), E-G Pouteria sp. (Aningeria pseudo-racemosa), Faulkner 729: E whole grain,
equatorial view (SEM); F & G whole grains, equatorial view (LM), F mid-focus; G different
grain, high focus. H - K Leptostylisfilipes, Webster& Hildreth 14665: H close-up of mesocolpium
(SEM); J & K whole grains, equatorial view (LM), J mid-focus; K different grain, high focus.

I~Y~e~I D
4r e,
;141
FIG. 38. [Pollen Type IX & Subtype XA (part)]. A-L (Pollen Type IX), A-C Pouteria
pallida, Ernst 1798: A close-up of mesocolpium (SEM); B whole grain, equatorial view (SEM);
C whole grain, equatorial view, low focus (LM). D & E P. cuspidatasubsp. cuspidata,Pranceet al.
5541: D whole grain, equatorial view (SEM); E close-up of mesocolpium (SEM). F-H P.
scrobiculata,
Steyermarket al. 92334: F close-up of mesocolpium (SEM); G whole grain, equatorial
view (SEM); H whole grain, equatorial view, low focus (LM). J & K P. cuspidatasubsp. dura,
Maguireet al. 56468:whole grains, equatorial view (LM), J 5-colporate, low focus; K different
grain, 4-colporate, mid-focus. L P. gabrielensis,
Maguireet al. 41874: whole grain, equatorial view,
low focus (LM). M-P (Subtype XA), M Elaeolumaglabrescens,Spruce2029: whole grain,
equatorial view (SEM). N E. nuda,Maguire24596: whole grain, equatorial view (SEM). P. E.
schomburgkiana, Maguire& Fanshawe32176: whole grain, equatorial view (SEM).

dFI
K
_ _ , E
-04O --0
*1
M '• N P p L
c? .
A-
Q -.
RL *
.
-.
Zoe
"IMI
FIc. 39. [Subtypes XA (remainder) & XB]. A-E (Subtype XB), Elaeoluma schomburgkiana,
Spruce 1836: A close-up of mesocolpium (SEM); B polar wall fracture (SEM); C - E whole grains,
equatorial view (LM), C high focus; D mid-focus; E low focus. F-S (Subtype XA), F, M-P
E. nuda, Maguire et al. 42219: F close-up of mesocolpium (SEM); M-P whole grains, equatorial
view (LM), M low focus; N mid-focus; P high focus. G & H, Q- S E. glabrescens, Spruce 2029:
G close-up of mesocolpium (SEM); H polar wall fracture (SEM); Q- S whole grains, equatorial
view (LM), Q high focus; R high-mid-focus; S low focus. J-L E. schomburgkiana, Maguire &
Fanshaiwe32176: whole grains, equatorial view (LM), J mid-focus; K mid -low focus; L low focus.

-44f
17,
ON
2.r,
.
vt 94
Itnr-
Ali
FIc. 40. [Subtypes XIA & XIB]. A-L, W-Y (Subtype XIA), A-F Sarcaulus brasiliensis subsp.
brasiliensis, Mori &Bolten 8385: A whole grain, equatorial view (SEM); B close-up of mesocolpium
(SEM); C-E whole grains, equatorial view (LM), C mid-focus; D high focus; E low focus; F
whole grain, polar view, high focus (LM). G & H S. wurdackii, Wurdack 2494: G close-up of
apertural area (SEM); H section through whole grain, polar plane (TEM). J-L S. oblatus,
Pennington & Tenorio 10731: J close-up towards apocolpium (SEM); K whole grain, equatorial
view, mid-focus (LM); L whole grain, polar view, low focus (LM) [NB cytoplasmic material has
not cleared in K or L]. W-Y Pouteria longifolia, Beck 8547: W & X whole grains, equatorial view
(LM), W high-mid focus; X mid-low focus; Y whole grain, polar view, mid-focus (LM). M-V
(Subtype XIB), M-P P. vernicosa, Krukoff 8192: M whole grain, equatorial view (SEM); N &
P whole grains, equatorial view (LM), N low focus; P mid-focus. Q- S P. aff. vernicosa, Pennington
22773: Q whole grain, equatorial view, mid-focus (LM); R whole grain, polar view, mid-focus
(LM); S section through whole grain, polar plane (TEM). T-V P. cladantha, Cid et al. 1945: T
close-up of mesocolpium (SEM); U & V whole grains, equatorial view (LM), U high focus: V
mid-focus.

C D
40,V1
hg~-Y?~- ?~~~-1
- r~I 'i
1C1
rl ~ t
,r~
~~I
K00\1
ar,
FIG. 41. [Subtypes XIC, XIIA (part) & XIIB (part)]. A-F (Subtype XIC), A-E Chromolucuma
rubriflora, Ducke 22235: A whole grain, equatorial view (SEM); B section through whole grain,
polar plane (TEM); C & D whole grains, equatorial view (LM), C mid-focus; D high focus; E
section through polar wall (TEM). F C. baehniana, FDBG 3436: close-up of apertural area (SEM).
G- L (Subtype XIIA), Diploon cuspidatum, G, H & K, Handro 956: G whole grain, equatorial view
(SEM); H close-up towards apocolpium (SEM); K section through whole grain, polar plane
(TEM); J & L Wurdack & Monachino 39595: J whole grain, equatorial view (SEM); L close-up
of apertural area (SEM). M (Subtype XIIB), Micropholis retusa, Spruce 2735: close-up of apertural
area (SEM).

D 1 pm
PB G
J K
2Pm
f_ _ _ 0 ___
oAA
Fic. 42. [Subtypes XIIA (remainder), XIIB (remainder) & Anomalous Pollen Type II]. A-D
(Subtype XIIA), Diploan cuspidatum, Wurdack & Monachino 39595: A- C whole grains, equatorial
view (LM), A high focus; B mid-focus; C low focus; D section through spine and polar wall
(TEM). E-J (Subtype XIIB), Micropholis retusa, Spruce 2735: E section through spine and wall
(TEM); F-H whole grains, equatorial view (LM), F high focus; G mid-focus, H low focus; J
section through apertural area (TEM). K - Q Englerophytumstelechantha,Leeuwenberg5544: K & L
whole grains, equatorial view (LM), K mid-focus; L different grain, mid-focus, M whole grain,
polar view, low focus (LM) [NB cytoplasmic material present in K, L & M]; N whole grain,
equatorial view (SEM); P whole grain, equatorial plane (TEM) [cytoplasmic material present];
Q half section through grain, polar plane (TEM).

tA
" ~do
_
~ ~•
t,. i,
.
...
r 1I~MEMOt
hlL~
Fic. 43. [Anomalous Pollen Type I]. A-H [Anomalous Subtype IA]. Chrysophyllummarginatum
subsp. marginatum, A Regnell s.n.: whole grain, equatorial view (SEM). B-E Morong 701: B
fractured grain, equatorial view (SEM); C 2 whole grains, equatorial plane (left) and polar plane
(right) (TEM); D & E whole grains, equatorial view (LM), D high focus; E mid-focus. F & G
Reitz & Klein 8070: F whole grain, equatorial view (SEM); G close-up of mesocolpium (SEM).
H Heringer el al. 3446: whole grain, equatorial view, mid-focus (LM). J-L [Anomalous Subtype
IB] . C. inornatum, Reitz & Klein 2582: J close-up of mesocolpium towards apertural area (SEM);
K & L whole grains, equatorial view (LM), K mid-focus; L low focus.

470 VOL.46 (3)
KEWBULLETIN
TABLE1. Distribution of pollen types within Pennington's (1991)
taxonomic arrangement. [Numbers = no. of species examined,
AN = anomalous pollen]
POLLEN TYPES
MIMUSOPEAE I II III IV V VI VII VIII IX X XI XII AN.
MIMUSOPINAE
1. Mimusops 13 2
2. Vitellariopsis 1
3. Autranella 1
4. Tieghemella 1 1
5. Baillonella None examined
6. Vitellaria 1
MANILKARINAE
7. Manilkara 13 2 15 1
8. Labramia None examined
9. Faucherea 1
10. Northia 1
11. Labourdonnaisia 2 1
12. Letestua 1
GLUEMINAE
13. Inhambanella 1 1
14. Neolemonniera 1 1
15. Lecomteodoxa None examined
16. Gluema None examined
17. Eberhardtia None examined
ISONANDREAE
18. Palaquium 13 20 1 1
19. Aulandra 1
20. Isonandra 4 1
21. Madhuca 23 10 1
22. Payena 8
23. Burckella 5
24. Diploknema 1 1 1
SIDEROXYLEAE
25. Sideroxylon 1 4 2 2 21 5 2
26. Neohemsleya 1
27. Nesoluma 1
28. Argania 1
29. Sarcosperma 5
30. Diplon 1
CHRYSOPHYLLEAE
31. Pouteria
Sect. Oligotheca 2 35 3
Sect. Pierrisideroxylon 4
Sect. Gayella 1 1 3
Sect. Rivicoa 1 3 1 2
Sect. Aneulucuma 3
Sect. Antholucuma 4
Sect. Pouteria 1 5
Sect. Oxythece 2 3 4
Sect. Franchetella 3 1 11 2
32. Aubregrinia 1
33. Breviea 1
34. Micropholis
Sect. Micropholis 7
Sect. Exsertistamen 1
35. Chromolucuma 2
36. Chrysophyllum
Sect. Aneuchrysophyllum 1 1 11
Sect. Donella 5
Sect. Ragala 1

TABLE 1. cont... I II III IV V VI VII VIII IX X XI XII AN.
Sect. Prieurella 3
Sect. Chrysophyllum 2 5 2
Sect. Villocuspis 3
37. Ecclinusa 3
38. Delpydora 2
39. Pichonia 1 3
40. Sarcaulus 3
41. Elaeoluma 1 4
42. Niemeyera 1 1 5
43. Pradosia 2 1 2 4
44. Leptostylis 1 2
45. Pycnandra 5 1
46. Synsepalum 2 2 4 2
47. Englerophytum 1 1 2 1
48. Xantolis 3 1 2
49. Capurodendron 2
OMPHALOCARPEAE
50. Tsebona 1
51. Magodendron 1
52. Omphalocarpum 4
53. Tridesmostemon 1
Note to Table 1: 15 species have pollen assigned to 2 or, in one case 3, pollen types. This explains
the difference between the number of species shown in the table (414) and the number of species
examined (398).
Caption for Table 2:
Explanation of table columns:
1. Polar length, average and ranges.
2. Equatorial width, average and ranges.
3. Shape class, P/E (Erdtman, 1969): *88 - 99 = oblate-spheroidal, = spheroidal,
1P01 - 13 = prolate-spheroidal, 1- 14 - I - 32 = subprolate and 1 -33 -1"0099 = prolate.
4. Aperture1" number and type: P = porate, V = colpi vestigial, 0-25-0 1" =
85 approximate
length of colpi in proportion to polar length.
5. Apocolpial wall thickness.
6. Mesocolpial wall thickness.
7. Endexinous thickening continuous in equatorial zone, (+) or (-). NB This column is
included since a continuous endexinous thickening is not necessarily indicated when the
mesocolpial wall is thicker than the polar wall (columns 5 & 6), this can be the result of a
thicker ectexine.
8. Composition of apocolpial infratectum (only scored for species where the wall has been
observed from SEM fractures or TEM thin sections): VN = very narrow, SF = structures
few, SC = short columellae, LC = long columellae, VLC = very long columellae
(extremely rare), GC = granular columellate, G = granular and DG = dense granular.
9. Endoaperture shape: NL = narrow lalongate (shorter diameter less than half the longer
diameter), BL = broad lalongate (shorter diameter more than half the longer diameter),
EXBL = extreme broad lalongate, C = circular (or almost) and BLO = broad lolongate
(very rare).
10. Tectum protrudent (+), slight (SL) or not (-); COS costate.
11. Tectum characteristics: BRE = broken reticulate, CG = coarsely granular, CLRR =
coarse, -low-relief rugulate, COR = corrugate-rugulate, CPER = coarsely perforate,
CRE = coarsely reticulate, CRU = coarsely rugulate, CS = coarse slits, CSR = coarsely
striate-rugulate, CST = coarsely striate, FAG = fine, anastomosed granular, FG finely
granular, regular or irregular, FLRR = fine, low-relief rugulate, FOS = fossulate-rugulate,
FOV = foveolate, FRE = finely reticulate, FRU = indistinctly or finely rugulate, FSR =
finely striate-rugulate, FST = finely striate, INS = insulate, LRR = low-relief rugulate,
MF = microfossulae, PER = distinctly or coarsely perforate, PS = psilate, RID = ridges,
RU = rugulate, SCA = scabrate, SPL = spinules, SPN = spines, SPS = subpsilate,
SR = striate-rugulate, ST = striate and VFSR = very finely striate-rugulate.
12. Differentiation in patterning between apocolpium and mesocolpium, ( + ), slight (SL), not
(-) or OCC = occasionally, (some grains in the sample).
13. Pollen type.
14. Pollen type in Flora Neotropica monograph, see Harley (1990a).
* adjacent to country of origin indicates that pollen is illustrated.
NB Blank space indicates that data for character has not been recorded.

TABLE2. Pollen morphological data for genera and species examined. Systematic arrangem
1. 2. 3. 4. 5. 6. 7.
Tribe MIMUSOPEAE Hartog

Subtribe MIMUSOPINAE Aubr6v. [Mimusopineae]
1. Mimusops L.
M. aedificatoriaChambo 10 Kenya (24-)29-2(-32) 4-5 -
M. angel Glover & Gilliland (32-)35"6(-38) 1"22 0"75 2"0 4"0-4"5
912 Somalia * 1 07 4 -
M. antongilensisCapuron 22847 SF Madagascar * (36-)33"4(-42) (29-)31"1(-34)
(43-)47 1(-50) 1 07 4-5 0"75
0-75 3"0 5"0 -
Barbosa 2633 Mozambique * (42-)44
(47-)50"4(-55)
7(-49) (34-)36 2(-39) 1 23 4 0 75 2-0
2"0 3"0-4"0 -
M. caffra
4-5 -
M. dispar Pentz 1 S. Africa (25-)26 7(-28) 4-0-5"0
Bernardi 14348 Sri Lanka (33-)35'0(-36) (28-)28 7(-30) 1"31 3-4 0"75 1"5 3"0-4"0 -
M. elengi
M. fasciculata Westerhuis BW 5444 Pap.N.Guinea (32-)34'3(-35) (32-)35 2(-38) 14 3-4
11"19 0"85 2"0 5"0 -
Welwitsch 48136 Angola (38-)40"2(-43) (30-)34 5(-41) 1 12 4 0"65 5-0
3"0-4"0 4"0-5"0 -
frondosa
M.
M. fruticosa Greenway 9645 Kenya (34-)38"6(-42) 1 12 4-5 0"65 2"0-2"5 -
Lawton 1831 (37-)38"9(-40) 1 22 4 0"75 1"5-2-0 5"0 -
M. kummel Nigeria (42-)43"8(-46)
(39-)41 7(-44)
S. Africa * (38-)40 9(-43) (32-)34"1(-37) 1 47 4-5 0"65 2-0
2"0 4"0-5"0 -
M. marginata Strey & Moll 3679
Codd 9705 S. Africa * (51-)51 9(-53) (25-)27"8(-29) 1 15 4-5 0"75 4"0-5"0 -
M. obovata
Semsei 1022 Tanzania (46-)47 4(-50) (44-)45"0(-46) 1 18 4 0"75 2"5 4"0 -
M. riparia
* (40-)42 2(-44) 1 08 4-5 00"75
65 2"0-2"5 0-4 0
1-5-2 0 34"0-5"0 -
M. schimperi Burger 514 Ethiopia (37-)40"1(-42)
(38-)38 8(-40)
4 -
M. seychellarumJeffrey 568 Seychelles (33-)35 3(-37)
(25-)28"0(-32) 1"16 0"85 2"0 5"0
2. Vitellariopsis Baill.
V kirkii Paulo 117 Tanzania * 4-5 2-0 3-0 -
1-48
(31-)36"1(-39) (23-)24"4(-28) 0"75
3. Autranella Chev.
A. congolensis Louis 4048 Zaire 4 -
A. congolensis Louis 3521 Zaire * (48-)49"8(-51) (42-)43"6(-45) 1 00
1"14 4 0 65
0"50 2 0
2"0-3"0 3 0-5 0 -
3-0-4"0
(59-)60 8(-65) (59-)60 8(-65)
4. Tieghemella Pierre
Michelson 892 Zaire (47-)48 6(-50) (38-)38 8(- 40) 1 25 4- 5 0 65 1-0 4 5- 50 -
T. africana 0 75 1 5-2 0 3 0 -
Leeuwenberg 2752 Ivory Coast * (35-)36 7(-38) (28-)24 4(-34) 1-22 5
T. heckelii
6. Vitellaria Gaertn. f.
V. paradoxa Oldeman 911 Ivory Coast * 4-5 0-85 2-0-2-5 +
Coull 1 Ghana * (39-)42"2(-48) (28-)30"9(-39) 1-33
1"36 4 2-0
1"0-1"5 +
V. paradoxa
(41-)44"1(-47) (29-)33"1(-36) 0'85 3"0

TABLE 2. Continued.
1. 2. 3. 4. 5. 6. 7.
Subtribe MANILKARINAE Aubrev. [Manilkarineae]

7. Manilkara Adans.
M. bidentata subsp. bidentata
Broadway 6973 Trinidad 38-0 38-0 1 00 4-5 0 85 1 5 15 -
M. bidentata subsp. surinamensis
Ducke 2501 Brazil (40-)41"2(-43) (32-)33"2(-34) 1"24 4-5 0"85 2"0 20 -
M. bidentata subsp. surinamensis
Ducke 1523 Brazil * (40-)42"0(-45) (31-)32"4(-33) 1"30 4 0"85 3"0 30 -
M. butugi Meyer 8076 Sudan (35-)37"9(-41) (28-)30 1(-32) 1 26 4-5 0"75 1 5 30 -
M. concolor Gomes & Sousa 3799 Mozambique (31-)33"7(-36) (24-)26 6(-28) 1 27 4-5 0"85 1 5 1-5 -
M. cuneifolia Gossweiler 8649 Angola * (38-)42"4(-45) (23-)28 2(-36) 1 50 4-5 0"85 1 5 15 -
M. dawei Gillman 390 Uganda (53-)55"6(-60) (39-)43 6(-49) 1 27 4-5 0-75 1 0-1"5 2"0-2"5 -
M. discolor Williams 663 Kenya (28-)30"1(-33) (29-)30 5(-33) 99 4-6 0 75 2 0 2"0 -
M. discolor Chase 4559 Zimbabwe (24-)24"7(-25) (22-)22 0(-22) 1 12 4-5 0 75 1 0 1.0 -
M. discolor Chase 4664 Zimbabwe * (32-)34"0(-35) (32-)33 0(-34) 1 03 4-6 0 85 10 1.0 -
M. excelsa Ducke 17626 Brazil * (43-)44"0(-46) (36-)38 7(-41) 1 14 5 0 75 2 0 2"0 -
M. hexandra Gamble 27116 India (30-)33"9(-35) (23-)24"3(-25) 1 39 4-5 0 85 2 0 2"0 -
M. hoshinoi Kanehira 2139 Caroline Is. (43-)45-0(-48) (39-)40"0(-41) 1 12 4 0 75 2 0-2"5 2"0-2"5 -
M. huberi Guedes 175 Brazil (35-)36-5(-38) (28-)28"2(-29) 1 29 4 0 85 2 0 2"0 -
M. jaimiqui subsp. emarginata
Simpson 312 U.S.A. (46-)47-8(-49) (41-)43-5(-45) 1"10 4 0"75 2"0 2"0 -
M. jaimiqui subsp. haitensis
Ekman 15184 Domin. Repub. (35--)37-3(-40) (28-)32"2(-35) 1"16 4 0"65 2"0 2"0 -
M. letestui Le Testu 9327 Gabon (47-)49"9(-55) (40-)43"5(-46) 1-15 4-5 0"75 2"0 3-0 -
M. littoralis Parkinson 66 Andaman Is. * (58-)62"3(-65) (50-)56"7(-65) 1-10 4-5 0-75 1"0-1"5 3"0-4-0 -
M. maxima Pinheiro 1848 Brazil 45"0 40"0 1 12 4-5 0"65 2"0 2"0 -
M. mochisia Harris & Mwasumbi
3646 Tanzania * 1 29 4-5 0 85 1 -
M. Santos 1094 Brazil * (25-)27"3(-28) (20-)21"1(-22) 1 15 4 0 75 2 0-1"5
5 1"0-1"5 -
multifida (38-)39"0(-40) (33-)34"0(-35) 2"5
M. multinervis Enti FH 6827 Ghana (27-)28"3(-30) (20-)21"1(-22) 1 34 5-6 0 85 2 0 2"5 -
M. obovata Lowe 3611 Nigeria (38-)40- 1(-44) (30-)30"6(-33) 1 31 4 0 85 1 5-2"0 1"5-2"0 -
M. pubicarpa FDBG 5860 Guyana (39-)42"4(-45) (34-)37"0(-40) 1 14 4-5 0 85 2 0 2"0 -
M. roxburghii Herb. Wight. s.n. India (50-)54"1(-55) (35-)37"6(-40) 1 44 4-6 0 75 2 0 2"0-2"5 -
M. samoensis Whitmee 226 Samoa (55-)57"2(-61) (49-)52"2(-55) 1 09 4 0 65 2 0-2"5 2"0-2"5 -
M. staminodella Lundell & Contreras
20833 Guatemala (43-)45"0(-49) (43-)44"7(-49) 1-01 4-5 0"65 2"5 2"5 -

TABILE 2. Continued.
1. 2. 3. 4. 5. 6. 7.
M. subsericea Miers 3677 Brazil 4-5 0-75 2-5 -

M. sulcata Verdcourt 5298 (44-)45"4(-48)
31 0 (39-)42"2(-45)
27-0 1"07
1 15 3-4 2"5
1 5 1 5 -
Kenya
M. Irwin 5042 Brazil * 4-5 0"85 2-0 -
triflora
M. valenzuelanaEkman 8347 Haiti (43-)45-2(-48) (38-)39"4(-40) 1"15 4 0"75 2"0 -
M. vitiensis A. C. Smith 1461 (31-)42"3(-44) (35-)37"0(-40) 1"14 4-5 0"75 3"0 3"0 -
Fiji
M. Lundell & Contreras 19135 (39-)41"4(-45) (32-)35"2(-38) 1"18 0"75 3"0 4"0
zapota
19135 Guatemala * 4 -
(49-)51"4(-58) (47-)49"8(-55) 1"03 0"75 1"5 1"5
9. Faucherea Lec.
F laciniata Thouvenot 77 Madagascar 1 13 4-5 0-75 -
(48-)52"3(-56) (43-)46"2(-50) 1"5 1"5
10. Northia Hook. f.
N. seychellana Jeffrey 1171 * 1 22 4-5 0 65 2 2 5-3-0 -
Seychelles (36-)40 4(-44) (29-)33 0(-39)
0-3"0
11. Labourdonnaisia Boj.
L. calophylloidesBernardi 14717 Mauritius 4-5 -
L. glauca Vaughan 10516 Mauritius * (47-)51"9(-57)
48-0 (43-)48"9(-54) 1"06 4-5 0"75
0-75 1"5-2"0 3"0-5-0
1-0 -
L. revoluta Lorence 1602 Mauritius * (40-)44"3(-48) 1"08 4-5 0-75 -
(41-)44"9(-47) (38-)39"9(-42) 1"12 1"0-1-5 3"0-4"0
12. Letestua Lec.
L. floribunda Le Testu 1890 Zaire (39-)42-4(-44) 4-5 2-5-3-0 -
(27-)30"7(-33) 1"38 0"75 1"5-2"0
Subtribe GLUEMINAE Baehni
13. Inhambanella Engl. ex Dub.
ILguereensis Aubrbville 4147 Ivory Coast * 1 09 4 -
L henriquesii Goldsmith 176/62 Zimbabwe (36-)40"0(-42) (36-)36"6(-39) 4 0"75 2"0 3"0 -
(36-)41"3(-45) (31-)33"4(-36) 1"24 0"75 2"0-2-5 3"0
14. Neolemonniera Heine
N. clitandrifoliaBimuyo 45457 Nigeria 4-5 -
N. ogouensis Thollen 146 Gabon * (34-)35"7(-37) (29-)29"8(-31)
(41-)44-5(-46)
1"19
1-14 4-5 0"75 2"0-2"5 3"0-3"5 -
(47-)50"7(-54) 0"75 2"0-3"0 2"5-3"0
Tribe ISONANDREAE Hartog
18. Palaquium Blanco
P. burckii Beguin 262 Sumatra 1-10 4 -
P. calophyllum Chai S. 18487 Sarawak (62-)67"7(-74) (55-)61"2(-69) 4 0"75 3"0-4-0 3"0-4"0 -
0-75 1-5
P. cochlearifoliumAshtonBRUN 978 Brunei * (47-)50"0(-53) (36-)40'7(-44) 1"23 4-5 0-65 3"5-4"0 -
(53-)64"0(-70) (50-)57"7(-62) 1"11 4"0 5"0

TABLE2. Continued.
1. 2. 3. 4. 5. 6. 7.
P. dasyphyllum Sinangul SAN 56204 Sabah (42-)44 2(-46) 4 0 75 15 2-0 -

(33-)34"8(-37) 1"27 3 0 85 2 5 3-0 -
P. ellipticum Gamble 1514 India (40-)43 5(-46)
A. C. Smith 7616 (34-)36"1(-38) 1"20 4 0 65 2 5 3-0 -
P. fijiense Fiji (39-)43 5(-46)
P. formosanum Wilson 10999 Taiwan * (34-)42 6(-48) (30-)34"2(-40) 1"25
1 19 4-5 0 75 2 0 3-0 -
P. galactoxylumWomersley s.n. Pap.N.Guinea (46-)51 5(-55) (33-)35"8(-42) 1 24 5 0 75 1 5-2-0 4-0 -
P. grande Waas 432 Sri Lanka (39-)41"6(-45) 1 17 4 0 65 2 0 2-0 -
(63-)68 2(-70)
T. & P. 362 * (54-)58"0(-60) 4 6-0 -
P. gutta Malay Pens. (39-)41 9(-45) 1 16 0 75 4-0
P. hexandrum Ridley s.n. (47-)48"8(-51) 1 33 4 0 -
Malay Pens. (30-)32 2(-34) 85 2-0
(23-)24"2(-27) 1 23 3 0 75 2"0 3-0 -
P. 'hornei Berry 264 Fiji (40-)43 3(-46)
P. leiocarpum Paie S.26046 Sarawak * (44-)50 3(-53) (32-)35"0(-40) 1 25 4 0 75 2"0 2-5-3-0 -
P. lobbianum Buwalda 4963 Moluccas * (51-)53 5(-57) (37-)40"0(-43) 1 10 4-5 0 75 2"0-3"0 2-5-4-0 -
P. microphyllumOgata KEP 110385 Malay Pens. (45-)48"5(-53) 1 11 3-4 2"0 2-0-2-5 -
(44-)53 4(-58)
P. multiflorum Haviland 1890 Sarawak (38-)47"9(-50) 1 47 3-4 0"75
0-75 1"0-1-5 2-5-4-0 -
(32-)36 1(-38)
P. neo-ebudicumBernardi 13186 Vanuatu * (50-)52-6(-56) (23-)24"5(-26) 1 06 4-5 1"5 3-0 -
P. obovatum Maingay 985 Malay Pens. * (60-)66 1(-70) (44-)49"3(-55) 1 00 5 0"65 2"0 6-0 -
(60-)66 1(-70)
P. ottolandieri Koorders 10158p Java * (38-)42-7(-49) 1 40 4 0"65 5"0 3-5-4-0 -
P. philippense Loher 6564 Philippines * (50-)59"9(-62) 1 18 4-5 00"85
75 2"5-3-0 3-0 -
P. polyanthum Wallich 4166 India (50-)52"4(-55) (42-)44"3(-46) 1 08 3-4 0 75 2"0 4-0-5-0 -
P. pseudorostratum (42-)49"2(-54) (39-)45"3(-50) 3"0-4"0
Zehnder 9433 Sarawak (48-)52 4(-56) (35-)39 5(-44) 1 32 4 0-65 1 5-2 0 4-0-5-0 -
P. quercEfoliumKostermans 11169 Java * (38-)40-5(-44) 4-5 0-75 2-0-3-0 -
P. ridleyi Endert 579 Sumatra (35-)36"6(-38) 1"11 4 2"0-3"0 2-5-3-0 -
P. rioense J. & M. S. Clemens (29-)33"9(-38) (23-)25"8(-29) 1"31 0"75 2"0
40446 Sabah 3-4 0-65 3-0-3-5 -
P. rivulare Chai S.37358 Sarawak (48-)51"4(-57) (40-)43"5(-47) 1"18 4 0-75 1"0-1"5
1-5-2-0 2-5 -
(44-)48 7(-53)
P. rostratum Cockburn FRI 7710 Malay Pens. (50-)58-4(-64) (38-)41"8(-47) 1"16
1 11 4-5 0-65 6-0 -
Bujang S.32502
P. rufolanigerum Sarawak * (44-)46-0(-'49) (44-)52"5(-60) 4 3"0-4"0 3-0 -
P. sericeum Murthy & Ashton (40-)43"0(-46) 1"06 0"65 2"0
23343 Sarawak 1-34 4 5-0 -
P. stipulare Richards 2605 Sarawak (44-)47"1(-50) (33-)35"1(-38)
(45-)48-6(-55) 1 16 4 0"75
0-85 4"0 3-0-4-0 -
P. walsuraefoliumJacobs 5076 Sarawak * (53-)56"6(-61) 4 0-85 2"0-3"0 -
Sri Lanka (34-)39"1(-43) (31-)35"5(-41) 1"10 4 0-75 1"5 1"5-2-0
3-0-5-0 -
P. sp. Jayasuriya 1920
P. sp. Soepadmo & Mahmud 1115 (56-)58"4(-62) (48-)50"9(-56) 1"14 2"5-3-0
1115 Malay Pens. 4-5 0 50 1 0-1 5 3-0-4-0- -
P. sp. Streimann & Helean NGF 18338 (60-)63"2(-70) (54-)56"2(-60) 1"12
NGF 18338 Pap.N.Guinea 3 3-0-3-5 -
(55-)59"0(-65) (45-)48"1(-52) 1"22 0"65 2-0-3"0

TABLE 2. Continued.
1. 2. 3. 4. 5. 6. 7.
P. sp. SAN 34407 Sabah * 4 0-75 3-5-4-0 -

(69-)71"4(-74) (59-)60.8(-62) 1.17 3"5-4-0
19. Aulandra Lam
A. longifolia J. & M. S. Clemens
21531 Borneo * (47-)51 7(-56) (33-)35 8(-39) 1 44 3-4 0 85 1 5-2 0 2 5 -
20. Isonandra Wight
I. lanceolata Waas 1106 Sri Lanka 4-5 -
(47-)48"1(-51) (36-)37"7(-41) 1"27 0"75 1"5-2"0 2"5-3"0 -
I. perrottetiana Herb. Wight. 1734 India 4-5
I. stocksii HFP (40-)40"8(-42) (30-)31"4(-34) 1"29 0"85 2"0 3"5
Ramamoorthy
1550 India * 4 2-5 -
IL villosa Herb. Wight. 1733 India * (45-)47"2(-49) (34-)36"0(-39) 1"31 4-5 0"85 2"5 -
I. zeylanica & Cramer (42-)43"2(-44) (34-)34"4(-36) 1"25 0"75 1"0 2"5
Jayasuriya
777 Sri Lanka 4 0-65 -
(55-)57"8(-60) (34-)34"5(-35) 1"66 0"5 3"0
21. Madhuca Hamilton ex Gmel.
M. aristulata Scortechini 1984 Malay Pens. * 3-4 2 0 -
M. betis Merrill 2066 (45-)46"4(-48) (36-)38"2(-31) 1"21 4-5 0"75 2 2"0 -
Philippines
M. bourdillonii Bourdillon 386 India (52-)54"0(-55) (45-)47"2(-49) 1"14
1 08 3-4 00"75
85 2 0-3"5
5 3"5 -
M. burckiana Jacobs 5268 Sarawak * (44-)46"5(-49) (40-)43"2(-46) 4 0 75 2 2"5 -
M. coriacea Haviland 2118 Sarawak (43-)46"2(-49)
60-0 (38-)40"7(-45) 1"13 5-3"0 2"5-3"0 -
55-0 1-09 4 0 65 1 5-2"0 1 5-2 0
M. costulata Gibot 35876 Sabah 1-23 4 0 75 2 0 -
M. curtisii Jacobs 5011 Sarawak (45-)45"5(-46) (35-)37"0(-39) 4-5 0 50 3 2"0 -
(58-)59-5(-64)
M. curtisii Ridley s.n. Malay Pens. * (57-)60"1(-63) 1"01 4-5 0 65 3 0-5"0
5-4-0
3"0-5"0 -
M. decipiens Ridley 11371 (52-)55"8(-60) (49-)55"8(-59) 1"00 3-4 0 75 3-0 3-5-4"0 -
M. elmeri Binson & Bongsu (53-)55"0(-59) (46-)47"6(-1U) 1"15 3"0
63018 Sabah 1 22 4 2-5-3-0 -
M. kingiana Mujin SAN 37829 Sabah (50-)52"0(-58) (40-)42"6(-45) 1 12 4-5 0"75
0-65 2"5-3"0 -
M. korthalsii Gibot 35964 Sabah (49-)52"4(-55) (43-)46"9(-50) 1 06 4-5 0-75 2"5 4"0 -
(46-)47"4(-50) (43-)44"8(-46) 2-0-2"5 2-0-2"5
M. laurifolia Curtis 2254 Malay Pens. * (43-)46 1(-60) 1 13 4 -
M. leucodermis Ledermann 9733 (37-)40"9(-46) 1 15 4 0"75 1"5-3-0 1"5-3-0 -
Pap.N.Guinea (55-)49 7(-58) (46-)48.0(-51)
M. longifolia Ritchie 1830 India 1 12 4-5 0"65 1"5-2"5 1"5-2"5 -
(44-)48 0(-50)
M. malaccensis Nadiman s.n. (39-)42"9(-45) 1 21 4 0"75 3"0-3"5 3"0-3"5 -
Singapore (41-)45 5(-49)
M. microphylla Walter s.n. Sri Lanka (35-)37"6(-41) 1 12 4 0"75 2-5-3"0 2"5-3"0 -
(50-)54 0(-58) 3-0-4-0
M. mindanaensis Saikeh SAN 72241 Sabah * (38-)39 7(-41) (45-)48"0(-53) 1 13 4 0"75
0-75 3"0-4"0 -
1-0-2-0
M. motleyana Sinclair 40328 * (32-)35"1(-38) 21"0-2-0 -
Singapore (70-)75 5(-82) 1-05 4-5 5
M. neriifolia Herb. Hook. (Penn. (70-)71"5(-76) 0"65 4"0
India * 4-5 -
100)
(47-)47"0(-47) (45-)46"3(-48) 1"01 0"65 2"0-2"5 2"0-25

TABLE2. Continued.
1. 2. 3. 4. 5. 6. 7.
M. obovatifolia Cenabre 29199 Philippines 4-5 0-75 2-5-3-0 -

(79-)84"7(-93) (70-)75"3(-82) 1"12 2-5-3"0 -
M. orientalis Brass 7747 Pap.N.Guinea* 4
(50-)54-2(-59) 1-14
M. pallida de Wilde & de Wilde (59-)61"7(-65) 0"65 2"5-3"0 4"0
Duyfjies 14893 Sumatra * 5 40 -
M. palustris Corner 21330 Malay Pens. (43-)46"5(-49) (43-)45"5(-47) 11"02
11 4-5 0"65 3"0 -
M. pierrei Haviland & Hose (50-)53"2(-57) (45-)47"8(-50) 0"65 1"5 1"5
3481 Sarawak 4-5 2-5 2-5 -
M. pierrei Kerr 4352 Thailand (75-)78"5(-82) (70-)73"7(-78) 1"06 4 0"65
0-75 2-0-3-0 -
(36-)38-5(-42)
M. pubicalyx Otik 4775 Sabah (44-)46"2(-50) 1"20
1 14 4-5 3"0-4"0 -
(43-)45-5(-48) (39-)40-0(-41)
* (40-)42-0(-46) 4 0"65 1"5 2"0 -
M. sericea Boschproefstation 92 Sumatra
M. spectabilis Kostermans 5142 Borneo * (33-)35"6(-43) 1"18
1-06 4 0"75
0 ,75 2"5 3"0 -
M. stipulacea Winit 1295 Thailand (43-)46"8(-50) (39-)44"0(-47) 4 2"5 3"0 -
M. subquincuncialis
(41-)42"4(-45) (33-)34"0(-35) 1"25 0"65 2"0 2"5
Tsang 30271 Indo-China 33 0 27 0 1 22 0 85
M. tubulosa Whitmore FRI 8816 Malay Pens. * (47-)48-5(-50) (43-)44-5(-46) 1-09 4 -
M. sp. Kostermans 10052 Borneo 4-5 0"75 2"5 2"5 -
(45-)51"2(-55) (45-)49"0(-53) 1"04 0"65 2"0 2"0
M. sp. Pennington et al.
10250 Malay Pens. * 4 3-0 -
(45-)46"6(-50) (39-)39"6(-41) 1"18 0"75 2"5
M. sp. Pennington et al.
10234 Malay Pens. (31-)33-8(-35) 1-14 4-5 0-75 -
(36-)38"5(-42) 2"5 3"0
22. Payena A.DC.
P. acuminata Whitmore FRI 8503 Malay Pens. (30-)33 5(-36) 1-24 3-4 0-75 2 3 -
P. dasyphylla Yeop FD 3650 (39-)41"5(-44) 55-3"0 3 00-3"5 -
Malay Pens. (56-)58 0(-60) (51-)53 0(-55) 1 09 4-5 0 85 2
P. endertii Chai SAN 29390 Borneo * (61-)66 8(-74) 1-07 4 0-75 3 5 0 -
P. lanceolata Aniff 15541 Malay Pens. * (69-)72"0(-79) (39-)41 6(-44) 4 2 00-4"0 3 0 -
P. lucida Parkinson 5299 Burma * (46-)47"6(-50) (38-)40 3(-42)
1"14 4-5 0"75 3 5 5 0 -
(48-)50"2(-52) 1"24 0"75 -
P. maingayi Ogata KEP 105178 Malay Pens. (52-)54 1(-55) 3-4 4 0-4-5 4
P. microphylla NIFS 28096 Borneo (55-)56"2(-57) 1"03 4 0"75 4 0 0-4"5 -
P. obscura (50-)52"0(-55) (48-)49"3(-51) 1"05 4 0"75 4"0 -
Ridley 9203 Singapore 2
(65-)66"7(-69) (55-)55"7(-57) 1"19 0"75 0-3"0 3"0-4"0
23. Burckella Pierre
B. banikiensis BSIP 5876 Solomon Is. * 4-5 0-75 -
B. fijiensis Howard H117 Fiji * (51-)54"4(-56) (46-)47"4(-50) 1"15 4-5 0-75 1-5-2"0 4"0 -
B. macropoda Carr 15783 (58-)60"7(-64) (52-)55"9(-60) 1"08 4-5 0-75 3"0 3"5-4"0 -
Pap.N.Guinea (45-)48-8(-55)
B. obovata Waterhouse 870 New Britain (42-)45"1(-47) 1"08 4-5 3"0 4-0-5"0
3-5 -
0-75
* (60-)61"4(-63) (54-)55"6(-57) 1"10 2"0 -
B. parvifolia Damaru 145 Fiji 1-06 4-6
(45-)49"8(-57) (42-)46"9(-55) 0"75 2"0-2"5 4"0-5-0

TABLE2. Continued.
1. 2. 3. 4. 5. 6. 7.
24. Diploknema Pierre

D. butyraceoides Strachey &
Winterbottom 1 India * (44-)48-4(-53) 4 0-75 -
India (37-)41"8(-47) 1"16 4 3"0 5"0 -
Kanjilal 3137
D. butyraceoides (45-)48-6(-54)
D. krabiensis Hansen & Smitinand (51-)55"7(-61) 1"14 0"75 1"5 3"0
12352 Thailand 1 18 4 -
(41-)43"0(-46) (33-)36"4(-39) 0"75 3"0 3"0
Tribe SIDEROXYLEAE Radlk.
25. Sideroxylon L.
S. americanum Wilson 8349 Bahamas 3-4 -
S. borbonicum Friedman 1280 Mascarene Is. (32-)36"2(-40) (25-)27"2(-29) 1"33 3-4 0"75
0-75 1"5 2"5 -
S. boutonianumGueho 18442 Mauritius (31-)35"9(-39) (25-)27'1(-29)
(25-)26.4(-28) 1"32 3-4 0-35 1"5 3"0-3-5 -
S. capiri subsp. capiri (36-)38"4(-41) 1"45 1"0 3"0
Hinton 3833 Mexico * (55-)56 8(-59) 4-5 2 5 3 0 -
S. celastrinum Curtiss 1765 U.S.A. * (32-)34 2(-36) (47-)47"4(-49) 1"20 4-5 0"75
0-75 15 15 -
S. celastrinum Hayes 451 Guatemala (35-)35 2(-36) (22-)23"6(-25)
(24-)24 7(-26) 1"45
1 42 4 0 85 15 15 -
S. confertum Wright s.n. Cuba * (37-)37 7(-39) 3 10 15 -
S. contrerasii Contreras 10721 Guatemala (27-)28 4(-30) (25-)27"0(-29) 1"40
1-53 4 0"75 10 2 0 -
S. cubense Ekman 1875 Haiti * (38-)38 9(-41) (18-)18"5(-19) 1-18 4 0"35
0-65 2 0 2 5 -
S. discolor Smith & Lavranos (31-)32'7(-37)
428 Socotra 4 -
S. eriocarpum Conzatti 1586 Mexico * (40-)43"7(-49) (39-)42"8(-49) 1"02 4 0"75 1"0-1"5 3"0 -
S. floribundum subsp. belizense (30-)32"0(-34) (22-)23"6(-26) 1"36 0"75 1"5 1"5
Belize * 3 -
Schipp 1269 1-24
S. floribundum subsp. floribundum (35-)36"6(-40) (27-)29"6(-36) 0"75 3"0 3"5
March 1562 Jamaica * (37-)37-8(-40) 4 -
S. foetidissimum subsp. foetidissimum (30-)32"5(-36) 1"16 0"65 3"0 4"0
Curtiss 1759 U.S.A. * 4-5 -
(51-)61"4(-66) (40-)46"3(-52) 1"33 4 0"75
0-75 2"0 3"0 -
S. galeatum Jauffret s.n. Rodriguez I.
S. gerrardianumBarron 1574 Madagascar * (48-)50"8(-55) (42-)44"6(-49) 1"14 4 1"5-2"0 2"0-3"0 -
S. ibarrae Lundell & Contreras (47-)49-5(-57) (34-)36"8(-41) 1"34 0"75 4"0 4"0-5-0
19767 Guatemala * 3-4 -
(32-)33"5(-35) (21-)23"0(-25) 1"45 4 0"75 1"0 3"0 -
S. inerme Magogo & Innes 404 Tanzania (22-)23 1(-25)
S. jubilla Ekman 15659 Cuba * (28-)30,2(-32) 1"31 3 0"65 1"0 3"0 -
S. leucophyllumMoran 13056 Mexico (22-)23"7(-25)
(30-)32 1(-34) (15-)16"4(-18) 1"44
1-38 4 0"65 2"0
10 3"0 -
(21-)23"3(-25) 0"75 2"0

T•ABI.E 2. Continued.
1. 2. 3. 4. 5. 6. 7.
S. lycioides Gattinger s.n. U.S.A. 4 -

S. mascatense Harris 16356 Afghanistan * (27-)29"6(-33) (18-)20"0(-22) 1"48 4-5 0"75 1"0 1"5 -
S. mermulano Perez s.n. Canary Is. (25-)27"5(-30) (19-)20"3(-21) 1"34
1 18 3 0"85 1"0-1"5 2"0 -
S. microphyllumDecary s.n. Madagascar * (31-)35"3(-38) (28-)30"0(-33) 3 0"75 1"5 3"0 -
S. montanum Purdie s.n. Jamaica * (33-)34"5(-36) (28-)28"7(-31) 1"20 4-5 0"75 2"0-3"5 2"0-3"5 -
S. obtusifolium subsp. buxifolium (41-)43"4(-46) (32-)34"3(-37) 1"26 0"75 2"5 3"5
Simmonds 14680 Trinidad 4-5 -
S. occidentale Johnston 3904 Mexico * (29-)31"5(-33) (24-)25"7(-27)
(25-)26 8(-28) 11"23
20 4 00"75
75 1"0 31"5
0 -
S. oxycanthum Moone 8535 Ethiopia (29-)32"3(-36) 3-4 2"0 -
S. portoricense subsp. minutiflorum (33-)35"3(-38) (29-)31"4(-33) 1"12 0"65 1"0 1"0-1"5
Hinton et al. 10416 Mexico * 4 -
S. portoricense subsp. portoricense (36-)38'6(-42) (29-)31"3(-34) 1"23 0"75 2"0 3"0
Harris 5388 Jamaica * 4 0-75 -
S. portoricense subsp. portoricense (36-)39"4(-43) (27-)29"0(-34) 1"35 1"5 3"5
Ekman 8737 Haiti 1 14 4 -
S. puberulum Bernardi 14781 Mauritius * (48-)50"8(-55) (42-)44"6(-49)
(26-)27 5(-29) 1 58 4 0"75 -
S. reclinatum Fredholm 5832 U.S.A. * (42-)43"5(-45) (18-)19 1(-21) 1 26 4 0"65 1"0
1-5 2"0-3"0 -
S. rotundifoliumHarris 11040 Jamaica * (21-)24"0(-26) (20-)23 9(-25) 1 30 4 0"65 2"0 -
S. salicifolium March 1609 Jamaica * (28-)31"1(-33) (15-)15 8(-17) 1 69 3 0"75 1"0 1"5 -
S. saxorum Decary 8429 Madagascar * (24-)26"8(-29) (17-)19 2(-21) 1 41 3 0"75 1"0 2"0 -
S. tenax Curtiss 5678 U.S.A. (26-)27"1(-28) (20-)20 5(-21) 1 27 4 0"75 1"0 1"5-2"0 -
S. wightianum Champion s.n. Hong Kong * (25-)26"0(-27) 1 57 4 0"75 0"5 2"0 -
S. sp. Bouton s.n. Mauritius * (31-)34"6(-37) (20-)22"0(-25) 1 37 4 0"75 1"0
3-0 2"0 +
(43-)45"8(-50) (29-)33"5(-38) 0"35 3"5-4"0
26. Neohemsleya Pennington gen.nov.ined.
N. usambarensisPolhill et al. 4990 Tanzania * 3 -
(22-)22"9(-24) (17-)18"7(-20) 1"22 0"75 2"0 2-5-3"0
27. Nesoluma Baill.
N. polynesica Degener & Degener
30120 Hawaii * 3-4 -
0-65
(30-)35"5(-37) (24-)25"3(-26) 1"40 1"0 2"0
28. Argania Roem. & Schult.
A. spinosa Gattefosse 810 Morocco * 4-5 0-75 -
(29-)30'5(-32) (23-)25"0(-27) 1"22 1"5-2"0 1"5-2"0
29. Sarcosperma Hook. f.
S. arboreum Henry 12837A China * 4-5 2-0 -
S. griffithii Thomson s.n. India * (29-)32"4(-35) (27-)30"3(-33) 1"07
1 12 4 0"65 2-0 2"0
2-0 -
(26-)28"4(-30) (24-)25"4(-28) 0"75

TABLE2. Continued.
1. 2. 3. 4. 5. 6. 7.
S. kachinense Stainton 6725 India * 4 -

S. laurina Ying Hu 12355A Hong Kong * (34-)37"9(-40) (33-)36"7(-39) 1"03
1-18 4 0"75
0-75 1"5 2"0
2-0 -
S. laurina Tsang 30726 Indo-China (25-)26"4(-28) (20-)22"4(-24) 1-10 4-5 2"0
1-5 -
S. paniculatum NG FRI 1379 Malay Pens. (26-)27"2(-29) (23-)24"7(-26)
(21-)23 1(-25) 4 0"75 2"0
2-0 -
(23-)24"6(-27) 1"06 0"75 2"0
30. Diplo6n Cronq.
D. cuspidatum Handro 956 Brazil * 3-4 2-0 -
D. cuspidatum Wurdack & (24-)25"7(-28) (23-)23"5(-24) 1"09 0"75 2"0
Monachino 39595 Venezuela * (23-)24-0(-25) 4-5 -
(23-)24"0(-26) 1"00 0"65 1"5 1"5
Tribe CHRYSOPHYLLEAE Hartog

31. Pouteria Aubl.
Sect. 1. Oligotheca(A.DC.) Baehni

P. amieuana Aubr6ville & Heine
255 N. Caledonia (44-)46 0(-48) 1 23 4
P. australis Mueller s.n. Australia * (42-)44 2(-48) (34-)37"5(-39)
(32-)33 1(-35) 1 33 4 1-0 +
P. baillonii Green 1712 N. Caledonia * (52-)55 5(-59) (36-)41 7(-44) 1 33 4 0"65 3"0-4"0 +
P. balanseana McKee 14405 N. Caledonia (37-)39 0(-42) (31-)33 4(-36) 1 17 4 0"65 2-5-4"0 5"0-7"0 -
P. baueri McKee 14270 N. Caledonia (42-)45 3(-52) (29-)31 0(-33) 1 46 4 0"75 11"0-1-5 1"0-1"5 +
P. brownlessianaK. J. White s.n. Australia * (47-)50 6(-55) (40-)42 2(-45) 1 20 4 0"65 0-1"5 4"0
1 0-1-5 +
P. cinerea McKee 9934 N. Caledonia (46-)49 1(-54) (39-)42 0(-46) 1 17 4 0"65 1 0-1-5 4"0-5"0 +
P. cotinifolia Story & Yapp 97 Australia (45-)48 3(-51) (35-)37 9(-40) 1 27 4 0"65 2 3-0-5"0 +
P. firma Kerr 15958 Thailand (35-)36 6(-38) 1 27 4 0"50 2 00-3"0 4"5-5"0 +
P. firma Anta 232 Bangka I. (35-)36 0(-37) (27-)28"8(-31) 1 30 4 0"75 1 3"0-4"0 +
P. grandifolia Wallich 4155 India (37-)40 0(-44) (25-)27"6(-29) 1 19 4 0"75 2 05-2"0 3"5 +
P. keyensis Koster BW 4442 Irian Jaya * (40-)42 5(-45) (31-)33-7(-37) 1 07 4 0"50 3 5-4-0 4"0 -
P. laurifolia C. T. White 12863 Australia * (30-)33 8(-40) (37-)39"7(-43) 99 3-4 0"75 2 5 3"5-4"0 -
P. linggensis Versteegh BW 3927 IrianJaya * (27-)29 0(-31) (30-)34"1(-39) 1 32 3 0"75 10 2"5-3"0 +
P. maclayana Dennis et al. BSIP (19-)22"0(-23) 0"75 2"5
8503 Solomon Is. * 4 V +
P. macrantha Vidal 439 Philippines (40-)42"5(-44) (31-)31"7(-32) 11"34
33 3-4 2"0 4"0 +
P. malaccensis Ridley 6130 Singapore (50-)52'7(-55) (36-)39"5(-42) 1 16 4-5 0"65 2"0 4"0 +
P. myrsinoides London Exhib. 1862 Australia (43-)44'8(-47)
(50-)50-8(-55) (36-)38"6(-40) 4 0"65 2"0 4"0 +
P. nitida Backer 11006 Java (37-)38"3(-42) 1"32
1-51 4 0"50 2"0 5"0 +
(32-)34"2(-37) (21-)22"6(-25) 0"65 1"5 3"0-4"0

2. Continued.
ITABLE.
2. 3. 4. 5. 6. 7.
P. obovata Procter 3950 * 3 -

Seychelles (28-)29"6(-31) (19-)19"7(-21) 1"50 0"85 1"5-2"0 1"5-2-0
P. obovata Burkill 2611 Singapore (30-)32"4(-34) (23-)24"9(-27) 1"30 3 0"75 2"0 3"0 -
P. obovoidea Gafui et al. BSIP
1250 Solomon Is. * (31-)33 7(-36) (22-)24"4(-28) 1"38 4 0 75 2"0-2"5 3"0 -
P. oppositifolia Ducke 1760 Brazil (31-)32 9(-34) (23-)24"5(-26) 1"34 3 0 65 2 0 3"0 -
P. pinifolia McKee 14559 N. Caledonia (73-)78 7(-90) (64-)72 4(-82) 1 09 5 0 50 1 0-2 0 4 0 +
P. pohlmannia L. S. Smith 12486 N. Australia * (31-)33 0(-35) (21-)23-9(-27) 1"38 4 0 65 1 5-2"0 3"0-4-0 +
P. rubicunda McKee 7771 N. Caledonia * (55-)58 5(-61) (38-)40 1(-42) 1"46 4 0 50 1 5 4-0-5"0 +
P. sandwicensis Rock 8064 Hawaiian Is. (31-)33 4(-36) (23-)24 5(-26) 1 36 3-4 0 65 2 0 2"5-3"0 -
P. sarcospermoides
Carr 12933 Pap.N.Guinea* (52-)54"1(-55) (44-)45 6(-49) 1 19 4-5 0 65 1 0-2"0 3"5-5"0 +
P. sebertii McKee 15416 N. Caledonia (42-)44"5(-46) (35-)38 0(-40) 1 17 4 0 65 1 5-2"0 3-0-4"0 +
P. sericea Parker 509 Australia * 1 16 4 0 50 2 0-3"0 +
(41-)43"6(-47) (36-)37 6(-40) 5'0
P. sericea Brown 2823 Australia (32-)36"4(-40) (24-)25 2(-27) 1 44 4 0 85
P. singuliflora Webb & Tracy (51-)54"4(-56) (42-)44 7(-46) 1 22 4-5 0 65 2"0 5"0 +
10800 Australia
P. toricellensis Hertz & Katik NGF
42938 Pap.N.Guinea* (38-)40"7(-43) (26-)28"4(-30) 1"43 3-4 V 1"5-2"0 3"0-4"0 +
P. unmackiana *
Hyland 9015 Australia (44-)46"2(-49) (35-)39"4(-42) 1"17 4 0"65 2"0-3"0 4"0 +
P. velutina Elmer 13130 Philippines (34-)35-1(-36) (26-)26"6(-28) 1"32 3-4 0"85 1"5-2"0 2"5-3"0 +
P. wakere McKee 15420 N. Caledonia (35-)37-3(-40) (31-)31 3(-32) 1"19 4 0"65 2"0-2"5 5"0 +
P. sp. McKee 14204 N. Caledonia * (38-)39"4(-41) (26-)28-2(-30) 1"40 3-4 0"65 1"5-2"0 3"0-3"5 +
P. sp. (Planchonella glabra)
*
Ridley 15770 Malay Pens. (44-)46"6(-50) (30-)33"3(-36) 1"40 4 0"65 2-0 4"0-5"0 +
P. sp. FD 1034 Fiji (29-)31 3(-35) (21-)23"4(-25) 1"33 4 0"65 1"5 4"0 +
P. sessilis Vodonaivala s.n. Fiji (30-)31-6(-35) (19-)20"1(-23) 1-57 3 0"75 1"0-1-5 3"0 +
P. sp. (Planchonella thiensis)
Pennington &
McPherson 10282 N. Caledonia (39-)42"6(-44) (28-)29"8(-32) 1"43 4 0"50 2"0 4"0-5"0 +
Sect. 2 Pierrisideroxylon
Pennington comb.nov.ined.
P. hochreutineriCarr 13023 Pap.N.Guinea (59-)60"6(-63) (44-)46"0(-50) 1 32 4 0"65 2"0 5.0 +
P. kaernbachiana Carr 5420 Pap.N.Guinea (45-)49"3(-53) (35-)37"0(-41) 1 33 4 0"65 2"0 5-0 +
P. multinervis Clemens 1971 Pap.N.Guinea* (38-)39"7(-42) (27-)29"1(-32) 1"36 4-5 0"65 2"0 4"0 +
P. thyrsoidea Walter & White 84A Solomon Is. * (47-)49 5(- 53) (38-)40 2(-43) 1 23 4 0 75 1 5- 20 1 5- 2 0 -
P. vrieseana Vogel 4424 Moluccas (40-)43 0(-45) (29-)30 8(-32) 1 39 3-4 0 65 1 5-2 0 2 5-3 5 +

TABLE 2. Continued.
1. 2. 3. 4. 5. 6. 7. 8
Sect 3. Gayella(Pierre) Pennington

P. decussata Ducke 17607 Brazil * 4 0.65 +
P. eugeniifolia Cowan & Soderstrom 2198A (34-)36"2(-39) (24-)25"0(-26) 1"46 1"5 3"0
2198A Guyana 3 0-75 3-0 +
1-44 3 0-65 +
P. eugeniifolia Silva & Bahia 3093 Brazil (33-)35"8(-39) (23-)25"8(-28) 1"38 1"0
Beck 8547 Bolivia * 3 V 1.0
2"0 10
4"0
P. longifolia (33-)34"8(-37) (23-)24"2(-26) -
P. nemorosa Steinbach 6666 Bolivia (20-)21"0(-23)
(28-)30-0(-32) (18-)19"1(-20) 1"13 3 +
Reed s.n. Chile (19-)20"2(-21) 1-22
1"48 3 0"85 1"5 2"5 +
P. splendens
Chile 4-5 2-5 + V
P. splendens Capt. King s.n. (43-)43"7(-45) (35-)35"7(-38) 0"75 2"0 3"0
4-0
(38-)42"5(-46) (36-)40"2(-43) 1"06 0"75
Sect 4. Rivicoa (A.DC.) Baehni
Keay FHI 21076 Nigeria * 4 25
P. alnifolia -
P. altissima Aubreville 86 Ivory Coast (40-)43"5(-41) (34-)36"4(-41) 1"19 4 0"75 2"0 +
P. campechiana Lundell & Contreras (31-)32"7(-36) (23-)24"5(-26) 1"33 0"75 1"5-2"0 3"0
Guatemala * 4-5 - V
-19241 0-50 5-0
Mexico 1"04 4 - V
P. campechiana Hinton 5391 57"0 (54-)54"5(-55) 3"0
P. macrophylla Baker 67 Brazil (55-)55"2(-56) (53-)54"5(-55) 1"01 4-5 0"65 4"0 5"0
Robyns 3324 Zaire * (56-)63"0(--71) 1"04 4 2-5-3-5 4"0 -
P. robusta (55-)60"3(-65) 0"65 3"0
Brazil (37-)39"9(-42) (33-)34"3(-35) 1"16 4-5
1-29 V
0"175 2"5-4-5 +
P. speciosa Lucke 22264 + D
P. speciosa Archer 7718 Brazil (67-)71"2(-75) 4-5 V 2"0 6"0
(86-)92"2(-97)
P. sp. (Aningeria pseudo-racemosa) (64-)71"7(-74) (56-)61"7(-65) 1"16 4"0 6"0
Faulkner 729 Tanzania * 4-5 +
(49-)52"9(-56) (36-)39"3(-41) 1"35 0"75 2"5-3"5 5"0-6-0
Sect. 5. Aneulucuma(Radlk.) Pennington
P. ephedrantha Krukoff 5422 Brazil 3 2-0 5-0 +
Martius 398 Brazil * 3 0-50 + S
P. procera (18-)19"7(-21) 1"67
Hinton 4597 Mexico (32-)33"0(-35) (23-)25"0(-26) 1"46 4 0"50 1"0 5-0
3"0 + G
P. sapota (35-)36"6(-38)
(37-)44"1(-48) (28-)33"8(-36) 1"30 0"50 2"0
Sect. 6. Antholucuma
(A.DC.) Eyma
P. domingensis subsp. cuprea
Ekman H4695 Haiti 4-5 0-50 6-0 - V
Brazil (60-)63"7(-65) (66-)67"2(-69) 0"95 4-5 2"5 5-0 -
P. grandiflora Glaziou 8227
(63-)64"0(-65) (60-)62"8(-66) 1"02 0"65 4"0 -
P. multiflora Hart 6961 Trinidad 4-5 4-0 5-0 V
P. venosa subsp. amazonica (45-)47"1(-50) (46-)48"0(-50) 0"98 0"50
FDBG 2397 Guyana 4-5 9P50 4-0 + V
(60-)64"5(-68) (60-)62"7(-65) 1"03 2"0

"TABLE2. Continued.
1. 2. 3. 4. 5. 6. 7.
Sect. 7. Pouteria
P. caimito FDBG 3652 Guyana * 3 +
P. caimito FDBG 2320 Guyana (41-)43"4(-48) (25-)28"8(-32) 1"51 4 0"50 2"0
2-0 5"0 +
P. gardneriana Rambo 37978 Brazil (50-)52"0(-55) (36-)38"0(-39) 1"37 4 0"50 3-0 5"0 +
P. glomeratasubsp. glonmcrata (43-)44"5(-46) (40-)41"6(-44) 1"06 0"65 4"0
Gaumer s.n. Mexico 4 1-0 1-5 +
P. gomphiaefoliaFroes 191 Brazil (35-)36"8(-40) (31-)32"8(-35) 1"12 3 0"50 2-0 +
P. guianensis Sandwith 550 Guyana (42-)44"2(-47) (34-)35"2(-37) 1"25 3 0"65 3-0 4"0 +
P. guianensis Hart 5982 Trinidad (48-)51"0(-54) (30-)33"5(-37) 1"52 3 0"65 2-0 4"0
4-0 +
P. torta Sandwith 347 Guyana * (62-)62"5(-63) (50-)50"5(-52) 1"24 3 0"50 +
P. tortasubsp. glabra (56-)60"4(-63) (44-)46"6(-48) 1"29 0"65 1"0 3"0-4-0
Schunke 4373 Peru 3 0-75 +
P. tortasubsp. torta (50-)53"1(-56) (35-)37"7(-40) 1"40 2"0 4"0
Hatschbach 1399 Brazil (46-)48 4(-51) 3 +
(34-)36"0(-39) 1"34 0"50 2"0
Sect. 8. Oxythece(Miq.) Eyma
P. ambelaniifoliaTillett & Tillett 45518
Guyana 1-32 3-4 +
P. cuspidatasubsp. cuspidata (24-)24"9(-25) (18-)18"8(-20) 0"75 1"0 2"0
Jenman 924 Guyana 3-4 0-65 15 -
P. cuspidatasubsp. cuspidata (19-)20"7(-22) (14-)14"7(-15) 1"40 1"5
Prance et al. 5541 Brazil * 4-5 1-5 +
P. cuspidatasubsp. dura (19-)19"6(-20) (15-)15"9(-17) 1"23 0"75 2"5
Sandwith 328 Guyana (17-)18-3(-19) 4 15 -
P. cuspidatasubsp. dura (20-)23"0(-25) 1"25 0"65 1"5
Maguire et al. 56468 Brazil * 4 10 -
P. cuspidatasubsp. robusta (20-)21"7(-23) (17-)17"8(-19) 1"21 0'85 1"0
BW 6613 Surinam 4 0-65 2-0 +
P. cuspidatasubsp. robusta (18-)18"9(-20) (14-)14"4(-15) 1"31 2"5
Schunke 2146 Peru 3-4 0-75 1-5 +
P. gabrielensis Maguire et al. 41874 Venezuela * (21-)23"3(-26) (20-)21"2(-23) 1"09
1-17 3-4 1"0 +
0-65
P. kaieteurensis Cowan & Soderstrom (21-)21"7(-22) (17-)18"4(-19) 1"0 1"5
1960 Guyana 3 0-75 1-5 +
P. opposita Ducke 308 Brazil * (21-)23"2(-25) (17-)18"5(-20) 1"25 3-4 2"0 +
(16-)17-5(-18)
P. pallida Ernst 1798 Dominica * (22-)23"1(-24) 1"32 3-4 0"75
0-65 1"0
2-0 1"5
2-5 +
(17-)18-7(-21)
P. rigidasubsp. tomentosa (23-)24"6(-26) 1"31
Maguire 32783 Venezuela * (23-)24-1(-25) (20-)21 9(-24) 1-10 4 0-65 2-0 3-0 + BL

TABLE 2. Continued.
1. 2. 3. 4. 5. 6. 7. 8.
P. scrobiculata Steyermark et al.

92334 Venezuela * 1-32 3-4 3-0 +
(21-)21"7(-23) (15-)16"4(-17) 0"65 2"5
Sect. 9 Franchetella
(Pierre) Eyma
P. bangii Ule 9693 Brazil 3 1-0 1-5 -
P. bangii BW 6719 Surinam (24-)25"0(-26) (18-)19"3(-21) 1"29 3-4 0"75 3-5-4-0 +
P. belizensis Lundell 2767 Guatemala (27-)29"2(-30) (17-)19"2(-21) 1"52 4 0"75 1"5-2"0 3-0 + VN
P. cladantha FDBG 2233 (27-)28"6(-30) (21-)21"2(-22) 1"35 3 0"65 1"0
Guyana
P. cladantha Cid et al. 1945 Brazil * (18-)18"7(-20) (13-)14"7(-17) 1"27 3 0"65 2%5 +
P. cladantha Clark & Maquirino (22-)23"4(-25) (18-)19"5(-21) 1"20 0"65 1"5
s.n. Venezuela * 3 +
P. coriacea Russell 12644 Trinidad (30-)33"6(-37) (19-)19"8(-21) 1"69 3 0"75 2"0 3"0 +
P. dictyoneura subsp. dictyoneura (24-)25"6(-27) (14-)15"2(-17) 1"68 0"75 1"0 3"0
Wright 1329 Cuba * 4 3-0 +
(31-)32"2(-34) 20"0 1"61 0"75 2"0
P. durlandii Steyermark et al. 123196
Venezuela 3-4 +
P. durlandii subsp. durlandii (26-)27"5(-29) (18-)19"7(-22) 1"39 0"50 1"0 3".
Lundell 16759 Guatemala 1-38 3-4 +
P. durlandii subsp. durlandii (25-)26"3(-28) (18-)19"0(-20) 0"65 1"0 2"5
Gentle 1809 Belize 1 29 4 1-5 +
P. Sandwith 573 (29-)30"9(-32) (22-)23"9(-25) 1 66 3 0"65 3"0 +
egregia Guyana
P. Melinon s.n. Fr.Guiana (27-)28"5(-29) (17-)17"2(-18) 1 64 4 0"75
0-65 1"5 3"0 +
engleri
P. Hassler 7514 (35-)37"8(-41)
* (26-)29 (21-)23"1(-24) 1 19 3 1"0 2"0 + GC
gardneri Paraguay 1(-32)
P. Prance&Silva59467 Brazil (22-)24'4(-26) 1 31 3 0"65
0-65 1"0 3"0 +
gardneri 2"0
P. Froes 1841 Brazil * (28-)29"7(-31) (21-)22"6(-24) 1 59 3 1"0 +
ramiflora (15-)18-0(-20)
P. ramflora Irwin & Soderstrom (25-)28"6(-30) 0"75 1"0 3"0
5892 Brazil 3 3-0 +
P. ramiflora Ratter & Ramos 444 Brazil (27-)28"1(-31)
(30-)31 5(-34) (23-)24"7(-29)
(24-)25 2(-26) 11"14
25 3 00"65
75 2"0
P. ramiflora Spruce 926 Brazil 1-45 3 2-0 +
P. reticulata subsp. reticulata (35-)35"2(-36) (23-)24"2(-26) 3"0
FDBG 2413 Guyana 1-38 3-4 15 -
P. reticulata subsp. reticulata (25-)26"8(-28) (19-)19"4(-20) 0"75 1"0
Prance et al. 8142 Brazil * 3 20 - GC
P. reticulata subsp. reticulata (22-)24'9(-27) (14-)17"3(-21) 1"44 0"75 1"0
Krukoff 6265 Brazil 3 0-75 1-0 3-0 +
(23-)24"3(-26) (18-)21"5(-23) 1"13

TABLE2. Continued.
1. 2. 3. 4. 5. 6. 7. 8.
P. retcutdatasubsp. reticulata
Schipp 1238 Belize 3-4 0-75 1-5 2-0 -
P. reticulatasubsp. reticulata (25-)27"1(-30) (21-)22"5(-25) 1"20
Gentle 1208 Belize 3-4 0-75 15 15 -
P. sagotiana Hostmann 1315 Surinam (23-)25"8(-28) (19-)19"8(-23) 1"30 3 0-75 0-5 1-5 + SC
P. ucuqui Schultes & Lopez (15-)16"0(-17) (11-)11"6(-12) 1"38
9560 Brazil 3 V 1.0 2-0 + GC
P. ucuqui Schultes & Lopez (26-)27"1(-29) (17-)18"0(-20) 1"50
9484 Brazil 3 1-5 3-0 +
P. vernicosa Krukoff 8192 Brazil * (24-)25"5(-27) (15-)16'1(-17) 1"58
1 12 3 0"65
0-65 1-5 15 -
P. sp. aff. vernicosa (18-)18"5(-20) (15-)16"5(-17)
Pennington 22773 Brazil * 3 1-5 15 - LC
(17-)18"2(-19) (14-)14"7(-16) 1"24 0"75
32. Aubregrinia Heine
A. taiensis Enti FH 6871 Ghana * 3-4 +
(36-)38"3(-41) (25-)28"5(-31) 1"34 0"65 1-0-1"5 3"0-4-0
33. Breviea Aubr. & Pellegr.
B. leptosperma Tisserant 1429 Zaire * 4 2-0 4-0 + VN
(35-)38"2(-40) (27-)30'1(-32) 1"27 0"65
34. Micropholis (Griseb.) Pierre
Sect. 1. Micropholis
M. compta Santos 1224 Brazil (31-)34-4(-37) (22-)22-6(-24) 3 1-5 3-0 + DG
M. gardneriana Gardner 3310 Brazil * (25-)28-5(-30) (18-)19-0(-20) 1"52 3 0"50 1-0 2-0 +
M. guyanensissubsp. guyanensis 1"50 0"50
Schulz 7296 Surinam (15-)18-7(-22) 1 45 3 0-50 1 0 2-0 +
M. melinoniana Matuda 4195 Mexico (25-)27"1(-31)
(34-)38-3(-41) (23-)26-7(-29) 1 43 3 0-50 2 0 3-0 +
M. mensalis Maguire 24452 Surinam * (35-)37-1(-39) 1 38 3 V 1 5 3-0 + SC/
M. resinifera Ducke 22259 Brazil (28-)29-9(-32) (25-)26"8(-29)
(19-)20-4(-21) 1 46 3 0-50 1 5 3-0 +
M. retusa Spruce 2735 Brazil * 1 15 3 V 1.0 1.0 - SC/
M. venulosa Harley et al. 10523 Brazil (24-)27"0(-33) (23-)23"4(-25)
(18-)20-1(-22) 1 52 3 0-50 1 0 2-5 +
Sect. 2. Exsertistamen (29-)30"7(-32)
Pennington
M. maguirei Maguire et al. 36776 Venezuela 3 0-50 1-5 4-0 +
(43-)44"3(-46) (32-)35"0(-38) 1"26
35. Chromolucuma Ducke
C. baehniana FDBG 3436 Guyana o * (32-)34"0(-35) 25-0 3 0-50 1-0 2-0 - SC
C. rubriflora Ducke 22235 Brazil * 1"36 3 V 2-0 3-0 - SC
(31-)-34"2(-37) (26-)28"2(-30) 1"21

TABLE2. Continued.
1. 2. 3. 4. 5. 6. 7.
36. Chrysophyllum L.
Sect. 1. Aneuchrysophyllum Engl.
C. albidum Oldeman 140 Ivory Coast 3 0-65 +
C. bangweolenseAstle 713 Zambia (29-)29"4(-30)
(30-)35 1(-38) (18-)19"2(-20) 1"53 3 11"5 3"0-3"5 +
C. beguei Enti FH 6714 Ghana (24-)28"0(-30) 1"25 3 0"50 2-5-3"0 +
C. (Gambeya boukokoensis) (29-)29"3(-30) (20-)20"6(-22) 1"42 0"65 2"0 3"0
Le Testu Gabon (31-)31 7(-34) 3 3-0 +
C. boivinianum Capuron SF 11403 Madagascar (19-)19"5(-20) 1"63 3 0"65 1"5 +
C. boivinianum SF 2521 Madagascar (30-)32"3(-35) (23-)24"2(-26) 1"33 3 0"65 2"0 3"0 +
C. eximium Ducke 24884 Brazil (35-)35"7(-36)
(23-)25 2(-28) (24-)24"5(-26)
(20-)21 0(-23) 11"47
20 3 00"65
65 11"0
5 22"5
0 -
C. giganteum Geerling & Bokdam
2319 Ivory Coast 4-5 +
C. gonocarpum Hassler 12382 Paraguay (39-)42"4(-44) (29-)34'7(-39)
(21-)23 1(-25) 1"22 3-4 0"50 2"0 3"0 +
(29-)31"6(-34) 1"37 0"65 1"0 3"0
C. gonocarpum Ibarrola 3563 Argentina 3-4 1-5 +
C. imperiale cult.Ireland s.n. cult. * (25-)28"3(-30) (18-)18"5(-20) 1"53 3 0"75 2"0 -
C. lanatum Cuatrecasas 19187 Colombia * (34-)34"8(-35) (24-)25"5(-28) 1"36 3-4 0"75 1"5 1"5 +
C. lucentifolium subsp. pachycarpum (41-)43"1(-45) (35-)36"2(-39) 1"19 0"65 2'0 4"0
Silva & Souza 2555 Brazil * 4 +
C. pomiferum FDBG 2641 Guyana (27-)28"6(-30)
(33-)34 0(-35) (20-)21"2(-23)
24-0 1"35 3 0"75
0 65 2"0 3"5
Ducke 15802
C. venezuelanense Brazil 1"42 3 1"0 2"5 +
Ducke
C. venezuelanense 602 Brazil (34-)37"5(-40) (23-)25"0(-27) 1"50 3 0"65 1"0 3"0 +
C. viride Klein 1718 Brazil (32-)32"5(-33) (24-)24"7(-25) 1"31 4-5 0"75 2"0 3"5 +
(31-)32"5(-33) (27-)28"0(-29) 1"16 0"65 1"0 2"5
Sect. 2. Donella(Pierre ex Baill.) Engl.
C. pruniforme Deighton 2718 Sierra Leone (27-)27 3(-28) 1 32 3
C. roxburghii Thwaites CP2689 Sri Lanka (29-)31 1(-33) (19-)20"7(-22) 1 31 3 0"50 1"0 3"0 +
C. viridifolium Gardner 938 Kenya * (29-)32 3(-34) (22-)23"7(-25) 1 13 3 0"50 1-5-2"0 3"0
30 -
C. welwitschii Leeuwenberg 2267 Ivory Coast (29-)30 1(-32) (25-)28"5(-30) 1 48 3 0"50 2"0
1-5
C. welwitschii Jordan 254 Sierra Leone (31-)32 7(-35) (18-)20-3(-24) 1 26 3 0"50 3'0 +
C. welwitschii Bos 1860 Liberia (40-)41 6(-48) (25-)26"0(-27) 1 70 3 0o50
0"50 1"0 2"5 +
C. sp. (Donella ogowensis) (23-)24"4(-28) 1"0 3"0-4"0
Le Testu 2213 Gabon 3 V +
(42-)45"2(-54) (22-)24"4(-31) 1"85 1"0-1"5 3"0-4"0
Sect. 3. Ragala (Pierre) Pennington
C. sanguinolentum subsp. balata
Klug 3699 Peru 3-4 0-50 15 20 -
(33-)33"5(-34) (22-)22"5(-23) 1"49

TABLE 2. Continued.
1. 2. 3. 4. 5. 6. 7. 8
C. sanguinolentum subsp. balata

Schultes & Lopez
9980 Colombia * 3 - C
(34-)35"4(-37) (20-)21"0(-22) 1"69 0"50 1"5 3"5
Sect. 4. Prieurella(Pierre) Pennington
C. amazonicum Schunke 2144 Peru 3 +
C. colombianumCuatrecasas 15219 Colombia (28-)28"8(-29) (16-)17"2(-18) 1"67 3 0"75
0-65 2"0 3"0 +
C. cuneifolium Granville T. 1087 Fr. Guiana * (29-)29"7(-30) (19-)20"0(-21) 1"49 3 1"5 3"0 + C
1-70
(33-)36"3(-39) (20-)21"4(-23) 0"50 1"0 4"0
Sect. 5. Chrysophyllum
C. argenteum subsp. argenteum
Jones s.n. Domin. Repub. 3 +
C. argenteum subsp. auratum (26-)27"3(-29) (17-)17"4(-18) 1"57 0"65 1"5 3"0
Schunke2142 Peru 3 + C
C. cainito Valeur 677 Domin. Repub. (30-)31"6(-35) (19-)20"4(-22) 1"55 3-4 0"75 1"5 3"0 + C
C. cainito Docters & (23-)25"6(-27) (14-)14"8(-17) 1"73 0"65 1"0 2"5
Leeuwenberg s.n. Java 3 +
(26-)27"7(-32) (16-)18"0(-20) 1"54 0"35
C. hirsutum Gentry & Dwyer
4800 Panama 3 +
C. inornatum Reitz & Klein 2582 Brazil * (30-)30"5(-31) (19-)20"0(-21) 1"53
1 68 3 0"65
0 85 31"0
0 3"0 + V
(33-)34"4(-36) (19-)20"5(-22) 1"0
C. marginatum subsp. marginatum
Malme 1394 Brazil 1 82 3 0 65 4 0 + V
(26-)26"6(-27) (14-)14"6(-17) 2"0
Heringer et al. 3446 Brazil * 1 58 3-4 0 75 5 0 + V
(28-)29"4(-30) (17-)18"6(-20) 3"0
Regnell s.n. Brazil * 1 72 4 0 65 5 0 + V
(28-)29"0(-30) (15-)16"9(-18) 3"0
Reitz & Klein 8070 Brazil * 1 38 3 0 75 2 5 + V
(21-)22"0(-23) (15-)16"0(-17) 2"0
Morong 701 Paraguay * 1 41 4 0 65 3 0 + V
(26-)26"3(-27) (18-)18"6(-20) 2"0

TABLE 2. Continued.
1. 2. 3. 4. 5. 6. 7. 8
C. mexicanum Pennington &

Sarukhin 9126 Mexico 3 +
C. mexicanum Contreras 2782 Guatemala (22-)23"4(-25) (13-)14"0(-15) 1"67
1-44 3 0"65 2"0 2"5 +
C. oliviforme Curtiss 1756 U.S.A. (22-)23"9(-25) (15-)16"5(-17) 3-4 0"65 1-0
1"0 2"0 +
C. ovale Klug2324 Peru * (29-)32"7(-34) (21-)22"5(-24) 1"45 3 0"75 1-5 2"0 +
C. rufum Glaziou 11146 Brazil (23-)23"5(-24) (13-)13"5(-14) 1"74 3
1-40 0"65 3"0
20 -
C. Martius 521 Brazil * (24-)24-9(-26)
(30-)32"0(-33) (22-)22"8(-23) 3 0"75 2"5 15 - G
rufum
(14-)15"1(-16) 1"65 0"85 1"5
Sect. 6. Villocuspis A.DC.
C. flexuosum Glaziou 17124 Brazil 3 +
C. aff.flexuosum Pinheiro 2178 Brazil * (34-)36"2(-40) (28-)29"5(-31) 1"23 3 V
0"65 2"0 3"0 + D
C. sparsiflorum Delascio & Liesner (38-)41"4(-45) (30-)33"4(-38) 1"24 2"0 3"0
7075 Venezuela 3 2-0 + G
C. splendens Santos 2714 Brazil (20-)21"3(-22) (16-)16"3(-17) 1"31
1-30 3 0"65
V 1"0 +
(30-)32"4(-35) (24-)25"0(-27) 2"0 3"0
37. Ecclinusa Mart.
E. guianensis FDBG 2070 Guyana 3 +
E. lancifolia Spruce 1949 Brazil * (38-)39"5(-41)
(49-)51-2(-54) (30-)31"3(-32) 1"26 3 0"50 1"0 4-0
3"0 +
E. Hoehne SP 28590 Brazil (33-)36"4(-40)
(31-)33 1(-35) 1"41 3 0"50 1"5 3-0 +
ramiflora
E. ramiflora Hoehne s.n. Brazil (41-)42"4(-47) 1"28 3 0"50 1"0 3-0 +
(30-)34"6(-37) (24-)26"5(-32) 1"30 0"65 1"5
38. Delpydora Pierre
D. gracilis Baldwin 6715 Liberia * (41-)42-4(-46) 4-5 + C
D. macrophylla Le Testu Gabon * (61-)64"3(-69) 1"52 4 0"50 2"0-2"5 5"0 +
(61-)68"0(-78) (42-)44"7(-48) 1"52 0"50 2"0-2"5 6"0-8"0
39. Pichonia Pierre
P. calomeris McKee 14681 N. Caledonia 3 40 -
P. deplanchei McKee 13552 N. Caledonia * (52-)55"8(-59) (43-)49"0(-57) 1"14 4 0"75
0-65 3"0 +
N. Caledonia * (35-)37"4(-42) 1"16 4 2"0-3"0 5-0-60
5"0-6"0 -
P. deplanchei Deplanche 442 (40-)43"4(-45)
(41-)43-5(-45) 1-18
P. Brass 22061
lauterbachiana Pap.N.Guinea (33-)36"7(-40) 4-5 9'65 3"0-4"0
P. Brass 21910
lauterbachiana Pap.N.Guinea* (36-)37"8(-40) (33-)35"4(-38) 1"06 4-5 0"65 3"0 5"0 -
P. Womersley 19233
lauterbachiana Pap.N.Guinea (45-)49"7(-53)
(43-)44-8(-47) (41-)46"5(-51) 1"07 4-5 0"75 2"0-2-5 3"0
P. solomonensis Mauriasi et al. BSIP (40-)42"0(-43) 1"06 2"0 3"0
17882 Solomon Is. * 4-5 0-65 -
(44-)45"3(-48) (42-)44"7(-46) 1"01 3"0 4"0-50
40. Sarcaulus Radlk.
S. brasiliensis subsp. brasiliensis
Mori & Bolten 8385 Surinam * 3 V 1.0 1-0 -
(18-)18"7(-21) (18-)18"4(-20) 1"02

TABLE 2. Continued.
1. 2. 3. 4. 5. 6. 7
S. oblatus Pennington &

Tenorio 10731 Ecuador * 3 V 1.5 1-5 -
S. wurdackii Wurdack 2494 Peru * 17"0
18-0 17"0 1"00 3 V 1.0 -
41. Elaeoluma Baill. 18"0 1"00 1"0
E. crispa Clarke & Maquirino
8229 Venezuela 3 0-75 3-0 +
E. glabrescens Spruce 2029 Brazil * (30-)30"3(-31) (20-)22"3(-25) 1"36
1 16 3-4 2"0 - S
E. nuda Philcox et al. 3257 Brazil * (31-)34"0(-36) (28-)29"4(-30) 4-5 0"65 + V
E. nuda Maguire et al. 42219 Venezuela * (35-)37"5(-39) (30-)30"8(-33) 1"21 4 0"65 2"0
2-0 4"0 +
E. nuda Maguire 24596 Surinam * (30-)31"6(-34) (25-)26"1(-27) 1"21 4 0"75 3"0
3-0 +
E. schomburgkiana (28-)29"8(-33) (21-)22"6(-25) 1"32 0"65 2"0
Spruce 1836 Brazil * 1-27 3 0-75 3-0 - S
E. schomburgkiana (27-)29"0(-30) (21-)22"8(-25) 3"0
Maguire & Fanshawe
32176 Guyana * 3-4 -
(21-)23"1(-25) (20-)20"7(-21) 1"11 0"75 2"0 2"0
42. Niemeyera F.v.Muell.
N. antiloga Moriarty 577 Australia 4 -
N. balansae McKee 9831 N. Caledonia * (30-)31"3(-33) (25-)27"3(-29) 1"15 4-5 00"75
65 32"0
0 2"5-3-0 -
N. chartacea Hyland 9167 Australia (34-)37"9(-40) (29-)31"9(-35) 1"19 4 0 75 1 0-1-5 3"0-4"0 -
N. chartacea Brass 2218 Australia (30-)31"0(-33) (22-)24"0(-25) 1"29 4 0 75 1 0 1"0-1"5 -
N. deplanchei McKee 13654 N. Caledonia (30-)32"7(-34) (26-)26"7(-28) 1"22 4 0 75 15 2"0-2"5 -
(20-)21-4(-23)
N. francei McKee 14518 N. Caledonia (27-)28"4(-30) 1"33
1 10 4 0 75 15 2"5-3"5
3-0 -
(33-)33-7(-36)
N. lissophylla McKee 14590 N. Caledonia (29-)30"5(-33) 3-4 0 65 1 0-1-5 +
N. sp. Pennington & (28-)30"6(-32) (24-)26"8(-28) 1"14 3"0
McPherson 10311 N. Caledonia 3 2-5 30 -
(38-)40"1(-42) (31-)35"1(-39) 1"14 0"65
43. Pradosia Liais
P. atroviolacea Ducke 1800 Brazil * 4 0-75 +
(21-)25 1(-28) 4-0
P. beardii Ayliffe 14407 Trinidad * (30-)32"3(-34) 1"29 3 1"5 20 -
(20-)23 2(-26)
P. brevipes Toledo & Gehrt s.n. Brazil * (27-)29"9(-30) 1"29 4-5 0"85 1"5 +
(23-)24 5(-26)
P. cuatrecasasii Cuatrecasas 13988 Colombia * (26-)27"5(-29) 11"12
13 3-4 0"50
0-65 2"0 4"0 + V
(29-)31 3(-33)
P. ptychandra SF 7240 Fr. Guiana (31-)35"3(-36) (17-)19 2(-20) 3-4 2"0 4"0 +
P. ptychandra Mori & Veyret 8982 Fr. Guiana * (21-)22"1(-23) (26-)28 6(-33) 1"15 3-4 0"50 1"5 3"0 +
P. schomburgkiana subsp. schomburgkiana (28-)30"0(-32) 1"05 0"50 1"0 2"0
Prance & Ramos
23128 Brazil * 3 20 -
(25-)28"0(-30) (20-)21"4(-23) 1"31 0"85 1"0

TABLE2. Continued.
1. 2. 3. 4. 5. 6. 7. 8
P. schomburgkiana subsp. schomburgkiana -

Stockdale 178 Brazil 1-38 3
Brazil * (33-)34-0(-35) 4 +
P. subverticillataDucke 812 (24-)24"6(-26) 0"65
0-65 2"5 3"0
(30-)31"1(-34) (20-)21"1(-23) 1"47 2"0 4"0
44. Leptostylis Benth.
L. filipes Webster & Hildreth
N. Caledonia * 1-28 4-5 -
14665 0-75
N. Caledonia 4-5 1-5-2-0 -
L. grandifolia McKee 14451 (49-)52"2(-56)
(53-)55-6(-58) (39-)40"6(-45) 1-11 2-0-2"5 4"0
McKee 13084 N. Caledonia * 4-5 -
L. petiolata (44-)50"1(-53) 0"75 1"0-1-5
(45-)48"5(-52) (37-)39"5(-41) 1"23 0"65 1"0-1"5 3"5-5"0
45. Pycnandra Benth.
3-4 -
P. benthamii McKee 5187 N. Caledonia *
+
P. controversa McKee 5100 N. Caledonia * (46-)50"3(-54) (43-)46"9(-51) 1"07 4 0"65
0-75 3"0 4"0-5-0
+
P. controversa McPherson 2885 N. Caledonia (42-)45"5(-49) (32-)33"3(-35) 1"37 4 2"5-3"0 4"0-6-0
P. kaalainsis Pennington & (46-)48"5(-51) (42-)44"9(-47) 1"08 0"65 3"0 4"5-5"0
McPherson 10296 N. Caledonia 3-4 --
*
P. vieillardii McPherson 2962 N. Caledonia (37-)39"3(-41) (35-)38"1(-41) 1"03 4 0"65 3"0-3"5 4"0
P. aff. vieillardii Pennington & (40-)42"0(-44) (35-)37"5(-40) 1"12 0"65 3"0 4"0-5"0
McPherson s.n. N. Caledonia 3-4 -
*
P. sp. Pennington & (44-)46"1(-49) (39-)42"4(-46) 1'09 0"75 3"0-4"0 5"0-6"0
3-4 -
McPherson 10283 N. Caledonia 0-65
(37-)39"2(-41) (35-)36"4(-38) 1"08 2"5 4"0
46. Synsepalum (A.DC.) Baill.
Brenan 9225 1 15 4 2 3 0 -
S. afzelii Nigeria
Lawler & Hall 304 Nigeria (31-)34 7(-37) 1 13 4-5 0 65 1 0-1-5 3 0 -
S. brevipes (34-)35"2(-36) (30-)30"5(-32) 0"65
0 75 1 0-2"5
0 2 0-2-5 -
1046 Tanzania 1 15 3-4
S. cerasiferum Semsei (30-)33 1(-35) (28-)30"8(-35)
0 75 1 5 1 5 -
Breteler 1056 Cameroon (28-)29 2(-31) 1 14 4-5
S. dulcificum (26-)28"7(-32)
Gabon 1 15 4 0 65 2 5 2 -
S. gabonense Le Testu 3680 (24-)25"7(-27)
0 85 1 5 1 5
1 17 3-4 -
S. glycydorum Latilo FHI 41350 Nigeria (38-)38"9(-40) 5-3"0
Eggeling 6303 Tanzania * (43-)44"8(-46) 1 15 4 0 75 1 2 -
S. msolo * (30-)31"5(-33) (26-)27"0(-28) 1 0 1 55-3"0
Fanshawe 1925 Zambia (27-)27 8(-29) 1 56 3 0 75 0-2"0 -
S. passargei (29-)30"6(-33) (25-)26"6(-28)
Louis 10180 Zaire (38-)42 1(-45) 4 0 75 2 0 2 5 -
S. sereti (17-)17"8(-18)
S. subverticillatum (34-)38"7(-44) 1"09
Greenway & Rawlins
9431 Kenya * (30-)33 6(-36) (25-)27 4(-30) 1 23 4-5 0 65 1 5-2 0 4 0-5 0 +

TABLE 2. Continued.
1. 2. 3. 4. 5. 6. 7.
47. Englerophytum Krause

E. letestui Le Testu 8806 Gabon * 4 3-0-45 -
E. magalismontanum (27-)29"5(-32) (22-)24"9(-27) 1"18 0"65 2"5
* -
Kennedy 2313 Nigeria (26-)27"7(-29) (21-)21"6(-23) 1"28 3-4 0"75 1"5-2-0 3"0-35
E. natalense Faulkner 1080 Tanzania (27-)28"2(-30) (21-)21"2(-22) 1"33 4 0"65 1"0-1-5 3"0 -
E. oblanceolatumAubrbville 4133 Ivory Coast (23-)23"6(-24) (19-)20"4(-22) 1"16 4-5 0"75 1"0-1-5 2"0-2"5 -
E. stelechantha Leeuwenberg 5544 Cameroon * 1"13 3 V 1"0 1"5-30 -
(21-)24"3(-28) (20-)21"5(-25)
48. Xantolis Rafin.
X. assamica Masters s.n. India (36-)38"8(-41) (30-)32-6(-35) 1 20 4 0 75 2"0 2 0 -
X. burmanica Lace 3189 Burma * 1 24 4 0 75 2 -
(32-)40"9(-44) (28-)33"0(-36) 2"5-3-0 5-3"0
X embeliifolia Chun & Tso 44618 China (47-)51"0(-54) (38-)44"2(-47) 1 15 4-5 0 75 2"0-2-5 2 0-2"5 -
X. racemosa Petelot 871 China * 1 28 4 0 75 1 5 -
X. siamensis Kerr 10124 Thailand * (27-)29"1(-32) (21-)22"8(-28) 1 35 4 0 75 1"5
2-0 2 0 -
(27-)30"6(-32) (20-)22"7(-24)
X. tomentosa Haines s.n. India (37-)38-4(-40) (27-)29-4(-33) 1 31 4-5 0 75 2"0-3-0 2 0-3-0 -
49. Capurodendron Aubrbv.
C. bakeri Scott Elliot 2969 Madagascar * 1 19 3 3-0 -
C. rubrocostatum Humbert & Perrier (35-)35"6(-37) (28-)30"0(-31) 0"75 3"0
2349 * 3 2-0 -
Madagascar (35-)36"2(-38) (27-)27"9(-30) 1"29 0"75 2"0
Tribe OMPHALOCARPEAE Dub.
50. Tsebona Cap.
T macrantha Capuron SF 27675 Madagascar * (68-)74-4(-77) (55-)63-0(-70) 1-18 4-5 -
0"75 3"0 5"0
51. Magodendron Vink.
M. venefici Hoogland 4625 Pap.N.Guinea (45-)49-5(-53) (36-)40"8(-45) 1 21 4 0-65 2"5 3"5-4"0 -
52. Omphalocarpum Pal. Beauv.
O. ahia Chipp 242 Ghana 4 -
* (53-)57"0(-60) (40-)44"0(-46) 1"29 0"65 1"5-2-0 6"0-7"0
-
O. bequaertii Mildbraed 9021 Cameroon (40-)43"3(-46) (36-)39-0(-42) 1"11 4 0"65 2"0-3-0 3-0-40
FHI 54070 * (53-)57-0(-60) -
O. elatum Nigeria
(40-)44"0(-46)
1 29 4
* 0"65 1"5-2-0 6"0-7"0 -
O. mayumbenseGossweiler 7830 Angola
(54-)60"3(-73) (43-)47"3(-59) 1"27
3-4
2"0-3-0 4"5-60
53. Tridesmostemon Engl.
T. omphalocarpoides
Brenan & Onochie
9434 Cameroon * 49 0 45 0 1 08 4 0 35 1 0 1 0 -


Harley 1991 Pollen Morphology Sapotaceae

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Harley 1991 Pollen Morphology Sapotaceae

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The Pollen Morphology of the Sapotaceae

Author(s): Madeline M. Harley

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Accepted for publication December 1990.

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MATERIALS AND METHODS

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Number of specimens examined: 472

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KEY TO POLLEN TYPES

1. Average polar length not exceeding 35

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DESCRIPTIONS OF POLLEN TYPES

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length. Endoapertures usually broad occasionally narrow. Wall thickness,

Key to subtypes of Pollen Type I

SubtypeIA. (Figs 5: A-L; 6: A-M)

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SubtypeIC. (Fig. 8: A-B, E)

Species included: Omphalocarpeae;Tsebona macranthaCap.

Subtype ID. (Fig. 10: A-F)

Subtype IE. (Figs 7: E; 8: C-D)

Species included: Mimusopeae: Manilkarinae; Manilkara littoralis Dub.

SubtypeIF. (Fig. 9: A-G)

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Isonandreae;PalaquiumpolyanthumEngl., Madhuca bourdilloniiLam, M. burckiana

SubtypeIG. (Fig. 9: A-G)

POLLEN TYPE II. 'PALAQUIUM'.

Number of specimens examined: 49

Key to subtypesof Pollen Type II

SubtypeIIA. (Figs 10: G-L; 11: A-H; 12: C-E)

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Species included: Mimusopeae:Manilkarinae;Manilkara discolor(Sond.) Hemsl.

SubtypeIIB. (Fig. 12: A-B)

Species included: Isonandreae;Palaquiumcochlearifolium

SubtypeHIC. (Fig. 12: F-L)

Species included: Isonandreae;Palaquium formosanum Hayata, P. lobbianum

Subtype IID. (Fig. 13: A- L)

Species included: Mimusopeae: Mimusopinae; Tieghemella africana Pierre.

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POLLEN TYPE III. 'MANILKARA'.

Number of specimens examined: 51

Pollen grains prolate-spheroidal, subprolate or prolate, infrequently

Key to subtypesof Pollen Type III

Subtype liIA. (Fig. 14: A- L)

SubtypeIIIB. (Figs 14: M-P; 15: A-M)

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Subtype IIIC. (Fig. 16: A- M)

Species included: Sideroxyleae;NeohemsleyausambarensisT. D. Penn., Nesoluma

Subtype IIID. (Fig. 16: N-R)

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Key to subtypes of Pollen Type IV

SubtypeIVA. (Fig. 17: A-N)

Subtype IVB. (Figs 17: P; 18: A-B)

Species included: Chrysophylleae;Xantolis racemosa(Dub.) Royen, X. siamensis

SubtypeIVC. (Fig. 18: C-P)

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Species included: Mimusopeae: Manilkarinae; Manilkara bidentata (A.DC.)

Subtype IVE. (Fig. 20: A-J)