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MES SMITHS0NIAN_INSTITUT|0N NOlinillSNI NVINOSHillMS S3 I a Va a ^ SMITHSONIAN _ INSTITl

JOAN W. NOWIGKE
and

JOHN J. SKVARLA

SMITHSONIAN CONTRIBUTIONS TO BOTANY • NUMBER 37

SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION

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Contributions to Knowledge in 1848 and continuing with the following active series:

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Smithsonian Contributions to Paleobiology
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Smithsonian Studies in History and Technology

In these series, the Institution publishes small papers and full-scale monographs
that report the research and collections of its various museums and bureaux or of
professional colleagues in the world cf science and scholarship. The publications are
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S. Dillon Ripley

Secretary

Smithsonian Institution

■T. .

'•1

SMITHSONIAN CONTRIBUTIONS TO BOTANY • NUMBER 37

Pollen Morphology and the
Relationship of the
Plumbaginaceae, Polygonaceae,
and Primulaceae to the
Order Gentrospermae

Joan W. Nowicke
and John J. Skvarla

ISSUED

f

: ; i

HUG 18

SMITHSONIAN INSTITUTION PRESS
City of Washington
1977

ABSTRACT

Nowicke, Joan W., and John J. Skvarla. Pollen Morphology and the Relation-
ship of the Plumbaginaceae, Polygonaceae, and Primnlaceae to the Order Centro-
spermae. Sinilhsonian Coniributions to Pwlany, number 37, 64 pages, 200 figures,
5 tables, 1977. — Three families, Plumbaginaceae, Polygonaceae, and Primnlaceae,
are considered to be related to or derived from the Order Centrospermae by vari-
ous authors. These three families have anthocyanin pigments in contrast to the
betalains found in all but two families in the Centrospermae. In addition, all
three are known to have starch-type sieve-tube plastids in contrast to the protein
type found in all centrospermous families. Examination of the pollen of 134
species by SEM, TEM, and light microscopy revealed great diversity, especially in
the Polygonaceae, but not tlie spinulose and tubuliferous/punctate ektexine,
which characterizes the vast majority of the centrospermous taxa. Recent evidence
argues against a close relationship of the Plumbaginaceae, Polygonaceae, and
Primidaceae with the Centrospermae, and the absence of any pollen types com-
mon to the three families further suggests that they are not closely related to each
other.

Official publication date is handstamped in a limited number of initial copies and is recorded
in the Institution’s annual report, Sinitlisotiian Year. Series cover design: Leaf clearing from the
katsura tree Cercidiphyllum japonicum Siebold and Zuccarini.

Library of Congress Cataloging in Publication Data
Nou'icke, Joan W.

Pollen morphology and the relationship of the Plumbaginaceae, Polygonaceae, and Primnlaceae
to the order Centrospermae.

(Sinith.sonian contributions to botany ; no. 37)

Bibliography: p.

1. Plumbaginaceae. 2. Polygonaceae. 3. Primnlaceae. 4. Centrospermae. 5. Palynotaxonomy.
6. Pollen morphology. 1. Skvarla, John J., joint author. 11. Title. III. Series: Smithsonian
Institution. Smithsonian contributions to botany ; no. 37.

QKL.S2747 no. 37 [QK495.A12] oSL.OSs [583’.672] 77-608070

Contents

Page

Introduction 1

Materials and Methods 3

Results of Pollen Analyses 4

Centrospermae 4

Plumbaginaceae 5

POLYGONACEAE 8

Primulaceae 11

Discussion and Conclusions 12

Literature Cited 15

Tables 16

Figures 23

iii

Pollen Morphology and the
Relationship of the
Plumbaginaceae, Polygonaceae,
and Primulaceae to the
Order Centrospermae

Joan W. Nowicke
and John J. Skvarla

Introduction

The Centrospermae (Caryophyllales) represents
one of the most controversial orders in the angio-
sperms. This group of at least 10 families, which in
the past has been cited as one of the few natural
orders based mostly on morphological characteris-
tics, has unique N-containing pigments, the beta-
lains, and a distinctive structure in the sieve-tube
plastids. Both Cronquist (1968) and Takhtajan
(1969) unite the betalain families, Aizoaceae, Ama-
ranthaceae, Basellaceae, Cactaceae, Chenopodiaceae,
Didiereaceae, Nyctaginaceae, Phytolaccaceae, and
Portulacaceae, as well as two anthocyanin families,
Caryophyllaceae and Molluginaceae, in the order
Caryophyllales (± Centrospermae). Both authors
include Halophytum ameghinoi Spegazzini (a beta-
lain taxon) and the Gyrostemonaceae in this order:
Cronquist (1968) treats Halophytum as a monotypic
genus in the Chenopodiaceae while Takhtajan
(1969) gives it family status; Cronquist regards the

Joan W. Nowicke, Department of Botany, National Museum
of Natural History, Smithsonian Institution, Washington,
D. C. 20560. John J. Skvarla, Department of Botany and Micro-
biology, University of Oklahoma, Norman, Oklahoma 75019.

Gyrostemonaceae as part of the Phytolaccaceae, and
Takhtajan separates it as a distinct family. Takh-
tajan recognizes the Tetragoniaceae as a separate
family while Cronquist includes it in the Aizoaceae.
Dysphania is treated as a member of the Chenopo-
diaceae by both authors; and finally Takhtajan also
places the Hectorellaceae and the Bataceae in this
order.

Thorne (1968) in a synopsis of angiosperm classi-
fication has placed the betalain families in a sub-
order, Chenopodiinae, one of two constituting the
order Chenopodiales. He recognized the Gyrostemo-
naceae and the Halophytaceae as distinct families
but treated the Molluginaceae as a subfamily in the
Aizoaceae, the Achatocarpaceae as a subfamily in
the Phytolaccaceae, and Dysphania as a member of
the Chenopodiaceae. The other suborder, Caryo-
phyllinae, consists of two families, the Caryophylla-
ceae and the Polygonaceae. The next order, Bati-
dales, consists of only the Batidaceae, but Thorne
apparently regards this family as somewhat related
to the Chenopodiales since both orders are united
under a superorder, Chenopodii florae.

Mabry, Taylor, and Turner (1963) and Behnke
and Turner (1971) have treated the betalain fami-

1

2

SMITHSONIAN CONTRIBUTIONS TO BOTANY

lies as a distinct group, separate from, yet closely
allied to, the two anthocyanin families, Caryo-
phyllaceae and Molluginaceae.

Evidence from ultrastriictural research on sieve-
tube plastids (Behnke and Turner, 1971; Behnke,
1976) and pollen morphology (Nowicke, 1975;
Skvarla and Nowicke, 1976) has reinforced the close
tie between the betalain families and the Caryo-
phyllaceae and the Molluginaceae. In an investiga-
tion of these plastids (colorless leucoplasts found in
the sieve-tubes) of the above-mentioned families
(Behnke and Turner, 1971) and the Magnoliidae
and Ranunculidae (Behnke, 1971), two main types
were found: plastids accumulating only starch and
designated as the S-type; and plastids accumulating
at least some protein, variously deposited, and desig-
nated as the P-type. The betalain families and the
Caryophyllaceae and the Molluginaceae all had the
P-type plastid in which proteinaceous fdaments
formed a peripheral ring usually around a crystal-
loid. Within the above-mentioned families, Behnke
(1976) was able to distinguish three groups based on
the crystalloid: globular, the most common; polyg-
onal, found in the Caryophyllaceae and in two
other genera, Limeiim (Molluginaceae) and Stegno-
sperma (Phytolaccaceae); and crystalloid-free, hav-
ing only the ring, found in two families, the
Amaranthaceae and the Chenopodiaceae.

Examination of the pollen of 177 species (No-
wicke, 1975) by light microscopy and SEM revealed
that 85% had a similar type of ektexine pattern or
sculpture, which was described as spinulose and the
tectum perforate, the openings described as tubules
or punctae. This was the predominant ektexine pat-
tern in every betalain family and in the Caryo-
phyllaceae and the Molluginaceae; in some families,
i.e., the Phytolaccaceae and the Molluginaceae, this
pattern was the only one found. Examination of
selected taxa by TEM (Skvarla and Nowicke, 1976)
revealed that the predominant pollen-type in the
betalain families and in the Caryophyllaceae has a
similar, sometimes identical, wall structure.

Pollen of the Achatocarpaceae, Bataceae, Gyro-
stemonaceae, and Theligonaceae was also examined,
but none of the species in these small families has
the spinulose and tubuliferous/punctate ektexine
described above. The Bataceae, Gyrostemonaceae,
and Theligonaceae all have the starch-type plastid
and lack the betalain pigments. Thus the evidence

from palynology, pigmentation, and plastid struc-
ture indicates that these three families are not
closely related to the Centrospermae. The relation-
ship of the Achatocarpaceae, a family unknown
with regard to pigmentation, to the Centrospermae
is more obscure — the pollen morphology does not,
in Nowicke’s opinion (1975) support a close tie, but
on the other hand the two genera that comprise the
family, Achatocarpus and Phaulothamnus, do have
the P-type of plastid (Behnke, 1976).

The Gyrostemonaceae have in fact been the sub-
ject of a multidisciplinary study (Goldblatt, et al.,
1976) in which all lines of evidence, including cyto-
logical, argue strongly against any relationship of
this family to the Centrospermae.

Although there is almost universal agreement on
treating the betalain families as a closely related
group, the precise definition of the order Centro-
spermae, i.e., to include or exclude the Caryo-
phyllaceae and Molluginaceae, may never be re-
solved to everyone’s satisfaction. Of more interest
in view of the accumulating data on the Centro-
spermae (sensu lato) is the question of the deriva-
tion of the families Plumbaginaceae, Polygonaceae,
and Primulaceae from this order.

Takhtajan (1969) considers the Polygonales, con-
taining only the Polygonaceae, as near the Caryo-
phyllales, especially the Portulacaceae and Basella-
ceae, and probably derived from the same stock as
the Caryophyllales (Centrospermae). His views on
the Plumbaginaceae are similar: nearest to Portula-
caceae and Basellaceae and probably derived from
the same stock as Caryophyllales. He does admit
that the pollen morphology of the Plumbaginaceae
is different — apparently from that of Portulacaceae
and Basellaceae.

According to Cronquist (1968:185), “both the
Polygonaceae and Plumbaginaceae are pretty clearly
related to the Caryophyllales. Both have a single,
basal, bitegmic, crassinucellate ovule in a com-
pound, unilocular ovary, and both have trinucleate
pollen. These characters are not known to occur in
combination outside the subclass Caryophyllidae.
The Polygonaceae are further bound to the Caryo-
phyllales by similarities in the pollen and by a more
or less transitional group of genera which are vari-
ously referred to the Caryophyllaceae or treated as
a separate family, Illecebraceae. The Plumbagina-

NUMBER 87

3

ceae are somewhat more isolated but may also be
derived from the Caryopliyllaceae.”

In one respect the system of Thorne (1968) paral-
lels those of Cronquist (1968) and Takhtajan (1969)
in that all three authors consider the Polygonaceae
as related to the Caryopliyllaceae. Although Thorne
treats the Primulales and Plumbaginales as adjacent
orders indicative of some relationship, they are far
removed from the Chenopodiales, at least in his
linear sequence.

Philipson (1975) in a review paper on evolution-
ary lines in the dicotyledons states that there is
general agreement to exclude the Primulaceae from
the subclass Caryophyllidae (sensu Cronquist, 1968).
“Almost as universal is the acceptance of the Poly-
gonaceae as at least peripheral to the group. The
position of the Plumbaginaceae remains more de-
batable” (Philipson, 1975:74).

Hutchinson (1959:117) regarded the Caryophyl-
lales (Elatinaceae, Molluginaceae, Caryopliyllaceae,
Ficoidaceae, and Portulacaceae) as “a prolific herba-
ceous group which has given rise to apetalous orders
such as Polygonales, Clienopodiales, and perigynous
petaliferous families as Lythrales, besides sympeta-
lous groups as Gentianales and Primulales.” The
Illecebraceae are given family status and included
as the only other family in the Polygonales. It is
noteworthy that Hutchinson included Herniaria,
Paronychia, and Scleranthus, as well as Illecebriim,
among the genera listed at the end of the family
description. He united the Primulaceae and Plum-
baginaceae as the order Primulales, closely related
to the Caryophyllaceae and perhaps the Saxifraga-
ceae. The Primulaceae, Polygonaceae, and Plumba-
ginaceae contain the anthocyanin pigments, and the
last two, at least, are known to have the starch-type
of plastid (Behnke and Turner, 1971) and appar-
ently the Primulaceae also has this type (Behnke,
pers. comm.).

In an effort to resolve the limits and relationships
of the Centrospermae, pollen of Plumbaginaceae,
Polygonaceae, and Primulaceae were examined by
light microscopy, SEM, and TEM. For purposes of
comparison and the reader’s convenience, we have
included a brief discussion and a SEM and TEM
of a species from several betalain families, and from
the Caryophyllaceae and Molluginaceae.

Acknowledgments. — In an extensive study like
this numerous persons have contributed in one way

or another, but Janice Bittner at the Smithsonian
Institution and William Chissoe at the University
of Oklahoma deserve special thanks for their fine
technical assistance and helpful suggestions. The
scanning electron photographs were prepared by
the Scanning Electron Microscope Laboratory at the
Smithsonian Institution. We also wish to thank
Aaron Goldberg and Stanwyn G. Shetler for their
critical review and Clara Ann Simmons for her
assistance in the preparation of the manuscript.
This research was supported in part by BMS grant
75-19846 to John J. Skvarla.

Materials and Methods

Pollen of 20 species in the Plumbaginaceae
(Table 3), 85 in the Polygonaceae (Table 4) and 29
in the Primulaceae (Table 5) was examined by light
microscopy, scanning electron microscopy (SEM),
and transmission electron microscopy (TEM). Also
examined and included are 38 additional taxa from
the centrospermous families (Table 2). Pollen sam-
ples were removed from herbarium material and
acetolyzed according to procedure outlined in Erdt-
man (1966). Samples for the SEM were vacuum-
coated with gold, and examined and photographed
with a Cambridge Stereoscan MK IIA microscope.
Samples for the TEM were incorporated into agar,
dehydrated through increased concentrations of
ethyl alcohol, and subsequently embedded in
araldite-epon resins (Skvarla, 1973). Some pollen
samples were stained in 0.125% OsO^ in 0.1 M so-
dium cacodylate buffer for two hours prior to agar
incorporation. Thin-sections were made with dia-
mond knives, collected on iincoated grids and
stained with uranyl acetate and then lead citrate.
Electron microscope observations were made with
a Philips model-200 transmission electron micro-
scope. Slides of all samples are deposited at the
Palynological Laboratory, Department of Botany,
Smithsonian Institution.

This study is a comparison of the pollen mor-
phology of three families, Plumbaginaceae, Poly-
gonaceae, and Primulaceae, with the pollen of the
Centrospermae, and detailed measurements and/or
descriptions are not given. The species examined
are listed in Tables 2-5; the names were taken from
the herbarium labels or the most recent annota-
tions. The identifications would have to be mis-

4

SMITHSONIAN CONTRIBUTIONS TO BOTANY

taken at the level of family to be significantly mis-
leading in this type of study, but all vouchers and
geographical locations are given in Tables 2-5.

Results of Pollen Analyses

Pollen analysis of the Plumbaginaceae, Polygona-
ceae, and Primulaceae revealed diverse forms within
each family, especially the Polygonaceae, but no dis-
tinctive pollen type(s) was common to all three
families. For this reason each family will be dis-
cussed separately following a review of the Centro-
spermae.

Centrospermae
Figures 1-28

The common pollen types in the Centrospermae
are illustrated in Figures 1-6: 3-colpate, pantopo-
rate, and pantocolpate, all with a spinulose and
tubuliferous/punctate ektexine. With the exception
of some taxa in the Caryophyllaceae which are
3-colporoidate, the apertures in the Centrospermae
pollen are simple. The most variable class of aper-
tures is that of pantoporate. Anredera scandens
(Figure 3) is usually 6-porate, with one pore on each
plane of the roughly cube-shaped grain. Gymnocar-
pos fruticosiim pollen (Figure 4), with large, sunken
pores in a geometric pattern, contrasts sharply
with that of Chenopodiiun ambrosioides (Figure 5),
which has numerous small pores.

Figures 7 through 12 illustrate at high magnifica-
tion (X 7500) the ektexine surface, spinulose, and
tubuliferous/punctate, which characterizes the vast
majority (85%) of the centrospermous taxa. Hama-
tocactus septispinus pollen (Figure 12) has the
largest perforations among the species examined
by SEM.

Figures 13 through 18 illustrate specialized or un-
usual forms, but at least one species, Cardionema
ramosissima (Figure 16), has the typical ektexine.
Two other taxa, PsilotricJnim amplum (Figure 13),
considered unusual because of the stellate aperture
plates, and Herniaria glabra (Figure 15), in which
the grains have a rare tetrahedral shape, also have
the common ektexine pattern but a modified ver-
sion of it — the spines and perforations are much
reduced.

The internal structure of the exine in various
centrospermous pollen types is illustrated in Fig-
ures 19-28. Characteristically in the centrospermous
taxa, the ektexine is well developed while the en-
dexine, with few exceptions (e.g., Mesembryanthe-
mum variabile, Figure 19; Herniaria glabra. Figure
24), is developed only in the region of the aperture.
Apertures with spine-shaped flecks of ektexine are
common in the Centrospermae and in this study are
illustrated by Opiintia (Figure 22), Limeiim (Fig-
ure 26), and Gymnocarpos (Figure 23). Representa-
tive pollen from nine families of the Centrospermae
were examined by TEM. Comparison of these data
with those of earlier TEM work in the Centro-
spermae (Skvarla and Nowicke, 1976) allows us to
expand our knowledge of this order, and a brief
summary for each family is presented below.

Aizoaceae. — Mesembryanthemiim variabile pol-
len (Eigure 19) is considerably different from that
of M. crystallinum (Skvarla and Nowicke, 1976, fig.
13) in possessing a thinner tectum and columellae,
as well as an extremely thick foot layer and a nar-
row but consistent endexine. Mesembryanthemiim,
however, is a large and unsatisfactory genus and
structural variation is not unexpected. The exine
stratification of M. variabile bears a striking simi-
larity to that of Boerhavia erecta of the Nyctagina-
ceae (Skvarla and Nowicke, 1976, fig. 30).

Amaranthaceae. — The pollen of this family has
been shown with SEM and TEM to be highly pleo-
morphic (Nowicke, 1975; Skvarla and Nowicke,
1976). The exine structure of Psilotrichum amplum
(Eigure 20) extends this pleomorphism by indicat-
ing a complete absence of endexine.

Baseli.aceae. — Some pollen in this family is
unique in having a cuboidal shape (Nowicke, 1975;
Skvarla and Nowicke, 1976). The structure of
Anredera scandens pollen (Figure 21), is quite dif-
ferent from that of Basella alba (Skvarla and
Nowicke, 1976, fig. 19) but similar to that of
Boussingaultia gracilis (Roland, 1968, pi. 8: fig. 4),
a more typical member of the family.

Cactaceae. — The structure of Opuntia lindhei-
rneri pollen (Figure 22) is consistent with other ex-
amples of the Cactaceae. Comparison of this micro-
graph with one of Alhiaudia (Skvarla and Nowicke,
1976, fig. 28) again underscores the close relation-
ship of the Cactaceae and Didieriaceae.

NUMBER 37

5

Caryophyllaceae. — The two taxa examined,
Gymnocarpos fruticosiim (Figure 23) and Herniaria
glabra (Figure 24), correspond in exine structure to
that previously noted for the family (Skvarla and
Nowicke, 1976, figs. 41-48). Additional comments
on H. glabra will be given later.

Chenopcdiaceae. — The exine structure of Che-
nopodium ambrosioides (Figure 25) agrees with
other examples in the family: thick tectum with
few spines and a thin foot layer.

Molluginaceae. — The exine structure of this
family has not been previously examined. Pollen of
Limeiim viscosttm (Figure 26) appears similar to the
pollen of Cometes siirattensis of the Caryophyl-
laceae (Skvarla and Nowicke, 1976, fig. 41), as well
as to Mesembryanthemiim crystallimim of the
Aizoaceae (Skvarla and Nowicke, 1976, fig. 13).
These relationships are of interest in the context
of the introductory remarks concerning the Mol-
luginaceae.

Nyctaginaceae. — Abronia angustifoUa (Figure
27) pollen morphology differs from that of previous
taxa examined by lacking an endexine. As indicated
earlier (Skvarla and Nowicke, 1976), however, the
Nyctaginaceae display a spectrum of exine mor-
phology.

Portulacaceae. — The exine morphology of
Naiocrene parvifolia (Figure 28) is similar to that
of other taxa examined in the family, most notably
Calyptridium umbellatum (Skvarla and Nowicke,
1976, fig. 38).

Several taxa whose placement or affinities are in
dispute were also examined. GeocarpoJi ininimiirn
(Figure 9), the single species of this monotypic
genus, has an ektexine pattern that is characteristic
of the order Centrospermae, but the presence of
anthocyanins would seem to restrict the family
placement to Caryophyllaceae or Molluginaceae.
Two more monotypic genera, Hectorella and
Lyallia, have been united as a separate family,
Hectorellaceae (Philipson and Skipworth, 1961), or
included in the Caryophyllaceae (Eckhardt, 1964).
Material of Hectorella was not available, and a
pollen sample of Lyallia kergiielensis yielded only
10-12 grains, all of which were 3-colpate with a
spinulose and sparsely punctate ektexine. As with
Geocarpon, the pollen morphology of Lyallia ap-
pears characteristic of the Centrospermae, but in-
conclusive with regard to status or family affinity

since these types are of wide occurrence in the
order. The evidence from plastic! structure is more
definitive and supports the treatment of Hectorella
and presumably also Lyallia as a separate family,
or at least argues for their exclusion from the
Caryophyllaceae. Hectorella has the P-type plastic!
but lacks the polygonal central crystalloid that char-
acterizes the Caryophyllaceae (Behnke, 1975).

The pollen types in Figures 15 and 16 are rep-
resentative of a small group of genera that have
been treated as a subtribe, the Illecebrinae, in the
Caryophyllaceae (Pax and Hoffman, 1934) or given
family status, the Illecebraceae (Hutchinson, 1959).
Pax and Hoffman regarded the subtribe as consist-
ing of four genera, Illecebrum, Haya, Cardionema,
and Chaetonychia, while Hutchinson’s concept of
the Illecebraceae includes several additional genera,
among them, Herniaria. Pax and Hoffman placed
this particular genus in the first subtribe, Parony-
chiinae, with the Illecebrinae as the second. Cron-
quist (1968:185) regards these genera as a transi-
tional gi'oup between the Caryophyllaceae and the
Polygonaceae. Figure 15 is a single grain of Her-
niaria glabra and is extremely similar to those of
Illecebrum verticillatxim in scanning electron mi-
croscopy and light microscopy. Herniaria glabra is
illustrated because the material of Illecebrum was
insufficient for examination by TEM. Figure 16 is
Cardionema ramosissima; material of Haya and
Chaetonychia was not available.

For a detailed discussion of pollen morphology
in the order Centrospermae the reader is advised to
consult Nowicke (1975) and Skvarla and Nowicke
(1976).

Plumbaginaceae
Fir.URE.s 29-81

Twenty species and one variety, representing
nine genera (Table 3), were examined by light
microscopy and SEM, and eleven of these species
were sectioned and examined by TEM. In all
taxa, the apertures appeared to be simple, mostly
3-zonocolpate, rarely 4- (or 5-)zonocolpate, and in
one collection of Ceratostigma ivillmottianum (Eig-
ure 59) the grains were pantocolpate. The ektexine
was either reticulate or verrucose, two sharply dis-
tinct morphologies with no intermediate forms.
These correspond to the two pollen types noted for

6

SMITHSONIAN CONTRIBUTIONS TO BOTANY

the family by Erdtman (1966:325); the Plumbago
type, which is verrucose, and the Armeria type,
which is reticulate. The Plumbago type (Figures
29-40, 59-64) has a well-defined foot layer, highly
irregular columellae and a continuous tectum. Ex-
tending from the latter is another set of columellae,
designated as verrucae, thicker than those below the
tectum, with the terminal sculpturing appearing as
fine bristles in TEM. The Armeria type (Figures
41-58) has a foot layer supporting straight, regular
columellae, which are fused distally into an incom-
plete tectum of the reticulate configuration.

Heterostyly, frequently associated with dimorphic
stigmas and dimorphic pollen grains, is a well-
documented phenomenon in the Plumbaginaceae
(Baker, 1948, 1953, 1966; Dulberger, 1975; Philipp,
1974). Although the collections at the U.S. National
Herbarium are too limited and valuable to permit
a study of heterostyly per se, and this is not our in-
tention, the survey of the Plumbaginaceae revealed
seemingly controversial results. Neither author
claims taxonomic expertise in the Plumbaginaceae,
but the collection sampled in each case was similar
to or within the range of variation of the remaining
collections of the particular species as identified.
Few specimens had sufficient numbers of open
flowers to permit a designation of short- or long-
styled with any degree of confidence, nor could we
follow Baker’s (1966) scheme of labeling the two
Armeria forms “A” or “B” since this too is based
on style length. However, the results in two species
with the Plumbago type reinforce each other, and
the pollen forms found in four species with the
Armeria type reinforce each other. The following
discussions apply only to the particular collections
listed in Table 3 for each species, and within each
major type the different grains are referred to as
forms.

What appears to be the two forms in the Plum-
bago type pollen can be illustrated by two collec-
tions of Ceratostigma griffithii from China (Figures
29-32) and two collections of Plumbago europaea
(Figures 35-38). In each species there is a form with
pointed verrucae (Figures 32, 38) and a form with
more rounded verrucae (Figures 30, 36). Whether
these two forms actually represent dimorphic pollen
and are associated with heterostyly remains to be
answered, but our results agree, for the most part,
with those of Erdtman (1970).

Baker (1948, 1966) regarded Ceratostigma and
Plumbago as having monomorphic pollen, but the
similarity of the two forms makes them difficult to
distinguish using only light microscopy. As a result
only those species examined also by SEM have been
designated as having pointed verrucae or rounded
verrucae in Table 3. Collections having the pointed
verrucae were found in Plumbago rosea (Figures 33,
34) and in Ceratostigma ivillmottianurn (Figures
59-62). A collection identified as Plumbago auri-
ciilata (Figures 39, 40) had the more rounded
verrucae.

The above reservation does not apply to the
Armeria type in which the two forms are strikingly
distinct even in light microscopy. The Armeria
forms can be illustrated by two collections each
of Armeria ynaritima (Figures 43-46), Limonium
vulgare (Figures 51, 52), and Statice sinuata (Figures
55-58) and one collection of Statice tenella (Figures
47-50) with several unattached branches, presum-
ably from different plants. In each of the above
species there is a prominently (or coarsely) reticu-
late form (Figures 45-48, 51, 55, 56) and a finely
reticulate form (Figures 43, 44, 49, 50, 52, 57, 58).
While each form exhibits subtle interspecific differ-
ences, i.e., the coarsely reticulate grain of Armeria
maritwia may be slightly different from the coarsely
reticulate grain in Statice sinuata, the classification
of grains as to coarsely or finely reticulate could
have been made without knowledge of the alternate
form. The remaining Armeria types illustrated by
SEM are Goniolinum collinum (Figures 53, 54) and
Limonium viciosoi (Figures 41, 42). In both species
the collection sampled had the coarsely reticulate
form.

TEM observations corroborate the dimorphism
shown by SEM. The verrucose type is represented
by Ceratostigma griffithii (Figure 69): the fine
bristle-like columellae heads correspond to the
rounded verrucae illustrated in Figure 30 by SEM;
ill Figure 70, the verrucae have thicker bristles, and
those with perfect longitudinal section illustrate the
large blunt protrusion of the pointed verrucae
shown in Figure 32 by SEM. In Armeria maritirna
(Figures 67, 68) and Statice sinuata (Figures 77-80),
representative of the reticulate morphology, TEM
reinforces the differences in diameter and length
of the columellae between the finely and coarsely
reticulate forms, and also the existence of a fine

NUMBER 37

7

network of sporopollinen around the muri of the
coarsely reticulate forms (Figure 67), which was not
readily apparent in the finely reticulate forms. Our
observations of A. maritima agree with the TEM
investigations of Erdtman and Dunbar (1966).
These workers designated the coarsely reticulate
exines as “A-line” and the more finely reticulate
exines as “B-line,” Direct correlations in exine struc-
ture between our TEMs (Figures 67, 68) and those
of Erdtman and Dunbar (1966, figs. 1, 2) are ob-
vious.

In all samples examined the endexine pollen wall
unit that characteristically occurs beneath the foot
layer in the majority of angiosperm pollen grains
is highly reduced in the Plumbaginaceae. We have
noted its presence only in the aperture region, and,
therefore, it is depicted in just a few of the trans-
mission electron micrographs included in this report
(Figures 65, 66, 68, 71, 73, 74, 80). Internal foramina
in the columellae and foot-layer (Figures 65-67, 75)
are also found in the Plumbaginaceae. This feature
has been described in the large family Compositae
by Skvarla and Larson (1965). These holes are of
sporadic distribution and are most commonly noted
in the Armeria pollen type.

In Ceratostigma willmottianum, a collection from
India had 3-colpate grains (Figures 61, 62), and a
collection from China had pantocolpate grains (Fig-
ures 59, 60). The pantocolpate grain has larger clus-
ters of verrucae, but at high magnification, X 7500
(Figures 60, 62), the tips of the verrucae in both
forms are very similar in the terminal structure.
These results do not correlate with those of Cera-
tostigma griffithii and Plumbago europaea, but di-
morphic pollen grains based on aperture structure
are not unknown (Kohler, 1976). Additional sam-
ples are needed to confirm the alternate form as
3-colpate.

As stated above, this study was not intended to
document or identify the species of Plumbaginaceae
with dimorphic pollen. We are aware that a de-
tailed investigation of each of the species listed in
Table 3 might well reveal that a majority of these
have dimorphic grains. Also, most pollen samples in
this family consist of a single flower with the five
anthers. This sampling technique limits damage to
the specimen, but does not yield large numbers of
grains.

If, especially in the Armeria types, 15 or 20 an-

thers from each collection had been examined, the
larger sample might have produced results more
compatible with those of previous workers. In the
case of Limonium vulgare^ examination of four col-
lections (Table 3) revealed three with the coarsely
reticulate form (Figure 51) and one with the finely
reticulate form (Figure 52). Our coarsely reticulate
form corresponds exactly with the “A” grain of
Baker (1966, fig. 1), but we did not find the form
which he illustrated as the alternate or “B” (1966,
fig. 2) and described as bearing a pattern of small
spines on the surface of the grain. In fact, the closest
similarity to his “B” grain is Plumbago rosea (Fig-
ures 33, 34). It seems unlikely that both major
pollen types, which are morphologically and struc-
turally very different, would be found in one species,
but our results do not exclude this possibility.

Another problematical taxon is Armeria maritima
var. sibirica (Turczaninow) Lawrence. This particu-
lar variety, sometimes treated as a subspecies or even
as a species, has been investigated by a number of
workers, with differing, but not necessarily mutually
exclusive, results. Both Baker (1966:355-356) and
Philipp (1974:41) regard “sibirica” as monomorphic,
having the papillate stigma and Type A pollen
(coarsely reticulate form). However, in a study of
collections made between longitudes 30°W and
60°E, Praglowski and Erdtman (1969) described six
pollen forms, and found as many as four in a single
anther. These include an “A” and “B” that cor-
respond to our coarsely and finely reticulate forms.
Of all the Plumbaginaceae examined in our study,
39 collections representing 20 species, only Armeria
maritima var. sibirica had both forms within a
single sample. The holdings of var. sibirica at
the United States National Herbarium are mostly
from Greenland and Canada, with only two from
Norway. Pollen was removed from a single inflores-
cence on each of six collections (Table 3) . The
SEMs, taken at a range of low magnifications, illus-
trate mixtures of the finely and coarsely reticulate
Armeria forms. Unfortunately this variety was ex-
amined late in the study and the micrographs could
not be included here. However, two observations
seem worthy of note: the coarsely reticulate form
appears to be predominant, and, secondly, the dis-
tinction between the two forms is not as striking
as in the other dimorphic Armeria taxa. Philipp
(1974, fig. 1) illustrates grains from dimorphic

8

SMITHSONIAN CONTRIBUTIONS TO BOTANY

Danish Armeria maritima (1974, fig. 1a, b) which
agree very well with our Figures 43-46, but more
importantly the two SEM’s from monomorphic
Greenland plants (1974, fig. 1c,d) also show a pos-
sible dilution of the distinction: the mesh of what
could be the finely reticulate form (fig. Ic) is larger,
and the mesh of what could be the coarsely reticu-
late form (fig. Id) is slightly smaller.

Philipp (1974:49) also cites the existence of large
and small pollen grains in the hybrids between
the dimorphic subspecies, A. maritima ssp. mari-
tima X ssp. elongata. The proportion of the grains
varies widely; in one cyme the percentage of large
grains in open flowers ranged from 45 to 90. In one
plant the maximum variation between the cymes
was from 24% to 90% large grains. The variation
in size is not exactly comparable to the structural
variation in the exine, but her results, those of
Praglowski and Erdtman (1969), and ours, indicate
that one plant, or even one inflorescence, is capable
of producing variable or dimorphic grains.

To summarize the results of pollen analysis in
this family — the striking difference in the two types
designated as “Armeria” and “Plumbago,” the dis-
tinction of the Plumbago type from all other pollen
examined to date in these studies, the existence of
the two Armeria forms within a single inflorescence
raise more questions than they answer. The exist-
ence of heterostyly and dimorphic pollen grains in
the Plumbaginaceae, however, in no way detracts
from the conclusion of this study: None of the
pollen examined in the Plumbaginaceae is simi-
lar or related to the common type found in the
Centrospermae.

POLYGONACEAE
Figures 82-173

This is a much larger family than either the
Plumbaginaceae or Primulaceae. The possibility of
a relationship with the Centrospermae is more
widely held and thus was examined in greater
depth. The 85 species (Table 4) representing 36
genera were examined by light microscopy and
SEM; 32 of these species were sectioned and ex-
amined by TEM.

The Polygonaceae are one of the most palyno-
logically diverse families in the dicotyledons, and

this extensive variation may have great taxonomic
potential at all levels, particularly that of generic
definition.

The aperture structure is more complex and vari-
able than in the Primulaceae and much more so
than in the Centrospermae or Plumbaginaceae.
While the most common type in the Polygonaceae
is 3- (rarely 4—) colporate, the endoaperture, readily
delimited in light microscopy, is variable and in-
cludes some zonorate types. Also found in the
family are taxa with pantoporate (Figures 113, 114,
134) and pantocolpate (Figures 112, 118, 120, 133)
apertures. Some collections of 3-colporate taxa had
occasional grains that were syncolporate.

The surface of the ektexine varies widely and in-
cludes tectate and nontectate (reticulate) forms. The
tectate forms can be punctate (Figures 82-85);
punctate-striate (Figures 94-99); prominently spinu-
lose (Figures 133, 134); or perforate and small-
spinulose (Figures 136, 138); and there are some
unique types that are difficult to describe accurately
in text (Figures 98, 106, 108, 110). The nontectate
forms are finely to coarsely reticulate (Figures 112-
123). In some taxa the gi'ains had one type of sur-
face outlining the colpi, and another on the poles
and mesocolpial ridges (Figures 130, 140).

The most common surface pattern is punctate.
The size and distribution alone of these punctae
vary widely, but an additional characteristic is pres-
ent in many taxa — the punctae are connected by
grooves or striae which are also variable in depth,
width, and placement (Figures 89, 91, 93, 95, 97, 99).
In Ruprechtia laxifiora (Figure 106), the tectum be-
tween the striae is variously upraised, producing an
uneven surface. In Ruprechtia ramifiora (Figures
108, 109), the intergroove tectum is more promi-
nently upraised. In Atraphaxis buxifolia (Figures
98, 99), the connecting striae are larger, tend to be
parallel, and result in, at least superficially, a striate-
like surface pattern. These distinctions are not
as conspicuous in TEM (Atraphaxis, Eigure 143;
Ruprechtia laxifiora. Figure 171; R. pallida, Figure
172), but this, of course, is frequently the case when
cross-sections are compared with surfaces having
irregular variations.

Within the broad category of punctate ektexines,
the following six subtypes with some representative
species can be recognized:

1. Ektexine finely punctate: Chorizanthe breiueri.

NUMBER 37

9

Eriogonum correlUi, and Nemacaulis denudata
(Figure 82).

2. Ektexine with clustered punctae: Centrostegia
thurberi, Chorizanthe fimbriata (Figure 84), C. pani-
culata (Figure 85), Eriogonum parishii, and E. race-
mosum.

3. Ektexine punctate, the punctae connected by
small striae: Eriogonum inflatum, E. ynarijoliurn
(Figures 90, 91), E. thomasii, Mucronea californica
(Figures 88, 89), Oxytheca trilobata (Figure 83).

4. Ektexine more prominently punctate-striate:
Antigonurn guatemalense, Calligonum comosurn
(Figures 94, 95), Gymnopodium antigonoides (Fig-
ures 92, 93), Harfordia macroptera (Figures 96, 97),
Muehlenbeckia chilensis (Figures 102, 103), M. poly-
botryar, Neomillspaughia paniculata, and Triplaris
americana (Figure 86).

5. Ektexine with prominent sunken punctae:
Fagopyrum esculentum (Eigure 87), Muehlenbeckia
tamnifolia (Eigures 104, 105).

6. Ektexine microreticulate: Oxygonum drege-
anum, O. zeyheri, and Podopterus mexicanus (Fig.
ure 132).

The subtypes listed above should be treated with
reserve since the distinction between consecutive
ones is not very great, particularly between subtypes
3 and 4. Also, variation within a sample made
classification of some taxa arbitrary or impossible.

Observations of exine wall with TEM indicate
that the six subtypes established by SEM have con-
siderable structural diversity as well. Sixteen of
these taxa were examined in sections: Chorizanthe
breweri (Eigure 145) and Nemacaulis denudata (Fig-
ure 157) of subtype 1, Chorizanthe pajiiculata (Fig-
ure 146) and Eriogonum parishii (Figure 149) of
subtype 2, Eriogonum marifolium (Figure 148) and
Oxytheca trilobata (Figure 158) of subtype 3,
Antigonum guatemalense (Figure 142), Calligonum
comosurn (Figure 144), Harfordia macroptera (Fig-
ure 152) and Muehlenbeckia polybotryar (Figure
156) of subtype 4, and Muehlenbeckia tamnifolia
(Figure 154) and Fagopyrum esculentum (Figure
150) of subtype 5 are all essentially similar in pos-
sessing a moderate to thick tectum, long, narrow
columellae {Calligonum comosurn, Figure 144, ex-
cepted), a very reduced (absent?) foot-layer, and a
distinguishable endexine. The TEM’s of some of
the above taxa revealed an unusual characteristic
that may be of phylogenetic significance: Chori-

zanthe spp. (Eigures 145, 146), Eriogonum parishii
(Figure 149), Nemacaulis (Figure 157), and Oxytheca
trilobata (Figure 158) all had a tectum in which the
inner face was conspicuously granular.

Two of these 16 taxa examined by TEM, Mueh-
lenbeckia chilensis (Figure 153) and Podopterus
mexicanus (Figure 160), are distinguished from the
preceeding ones by a prominently developed foot
layer and an endexine well developed only in the
region of the aperture. Gymnopodium (Figure 151)
and Triplaris (Figure 173), with a thin foot-layer
and endexine, appear intermediate between the
above gioups.

The second most common surface pattern is a
reticulum formed by larger and fewer columellae,
arranged in a network and distally fused, thus pro-
ducing incomplete tectum. Differences in the size
and placement of the primary columellae and in
the surface of the lumina produce wide varieties
and some recognizable pollen types. Grains with
this ektexine were 3-colpate (colporate?) (Figures
116, 122), pantocolpate (Figures 112, 118, 120), or
pantoporate (Figures 113, 114). In Polygonum capi-
tatinn (Figures 122, 123) and P. dielsii (Figures 116,
117) the columellae are massive, and a high, con-
spicuous reticulum is the result. The lumina are
larger in P. dielsii, but in both species they are
almost free of any columella material. In Poly-
gonum acuminatum (Figure 113) and P. orientale
(Figures 114, 115), both pantoporate, the muri alter-
nate with large lumina filled with smaller, free
columellae. In three pantocolpate taxa, Persicaria
coccinea (Figure 112), Polygonum amphibium (Fig-
ures 120, 121), and P. virginianum (Figures 118,
119), the columellae which form the reticulum are
smaller in length and diameter, closely placed, and
a finer reticulum results. In all three the lumina
are filled with small, free columellae.

Thin-sections of Polygonum orientale (Figure
159), P. dielsii (Figure 166), P. amphibiinn (Figures
162, 163), and P. virginianutn (Figure 169) empha-
size the SEM observations.

The formation of the reticulum in the above
Polygonaceae is unusual: the ridges or muri are
formed by distal fusion along the midline of two
rows of columellae, which are opposite or sometimes
alternate, producing a “staggered” effect liest illus-
trated in a “planar” view (Eigures 115, 117).

Four species of Polygonum, P. affine (Figure 124),

10

SMITHSONIAN CONTRIBUTIONS TO BOTANY

P. bistorta (Figures 128, 129), P. vaccinifolium (Fig-
ures 126, 127) and P. vivipariim (Figure 125), which
belong, among others, to the Bistorta complex
(Hedberg, 1947), and an Australian species of
Muehlenbeckia, M. cimninghamii (Figures 100,
101), have an ektexine pattern that could be de-
scribed as sparsely spinulose and sparsely punctate.
Grains of all five taxa were 3-colporate, the endo-
aperture well defined. In thin-section, Muehlen-
beckia cunninghamii, (Figure 155) is very similar to
the other species of Muehlenbeckia examined by
TEM (Figures 153, 154, 156).

It is difficult to reconcile the TEM of P. bistorta
(Figure 164) with the sparsely spinulose and sparsely
punctate ektexine depicted in SEM (Figures 128,
129). The lack of a continuous tectum may be due
in part to an oblique section and perhaps to the
densely packed columellae. The SEM’s of P. vac-
cinifolium (Figures 126, 127), with the larger per-
forations in the tectum, are more compatible with
the cross-section illustrated in TEM (Figure 168).

Koenigia islandica. Polygonum cyanandrium
(Figures 134, 135), and P. forrestii (Figure 133) had
grains that were either pantoporate or pantocolpate
with a prominently spinulose ektexine. Comparison
of these giains (Figures 133-135) with the common
type in the Centrospermae (Figures 1-12) indicates
that the above species could not be related paly-
nologically to that order. Polygonum forrestii (Fig-
ure 167) was the most distinctive Polygonum of
those examined by TEM. The exine consists of a
thin endexine and an ektexine containing abun-
dant holes or internal foramina. The surface of the
tectum was composed of very large, solid spines
with numerous, small projections in between.

Hedberg (1947), in a classic study of pollen
morphology in the genus Polygonum , sensu lato,
recognized ten pollen types for which he gave
descriptions and listed species. He segregated Poly-
gonum convolvulus (Figures 140, 141), P. cristatum,
and P. dumetorum as slightly aberrant forms of his
Tiniaria type. These three species have almost
identical pollen giains: 3-colporate with a zonorate
endoaperture, the ektexine echinate near the colpi
and psilate at the poles and the mesocolpial re-
gions. The above characteristics make the gi'ains
distinct, not only from the more typical Tiniaria
pollen, but from the remaining species examined in
the present study. This particular morphology is

paraded by that of Polygonella fimbriata (Figure
130) and closely related species (Horton, 1963.
181-183): both groups have grains with similar
apertures and a dimorphic ektexine in which the
two surface patterns have the same distribution.
Examination of P. convolvulus in TEM (Figure
165) revealed a remarkably uniform tectum
(throughout the psilate area of the ektexine) and
foot layer, underlain by a prominent endexine.
This rare combination of well-developed foot layer
and endexine was also found in Polygonella (Fig-
ure 161), which reinforces the parallel exomor-
phologies. Polygonum cilinode, P. cuspidatum, and
P. multiflorum, which Hedberg lists as typical
Tiniaria, have 3-colporate grains and a uniform
ektexine, punctate-striate or prominently punctate.

Four species of Rumex were examined: R.
aquaticus, R. acetosa (Figures 138, 139), R. aceto-
sella, and R. scutatus, all of which had a perforate
and very finely and evenly spinulose ektexine (Fig-
ure 139). Two species of Emex had an ektexine
(Figure 136) similar to that of Rumex, but with a
distinctive aperture structure. The ectoaperture is
very reduced in length, and its polar margins
almost coincide with those of the endoaperture.
The close similarity of the ektexine in Emex and
Rumex and its restricted occurrence reinforce Ham-
mer’s (1893) consecutive placement of the two
genera. Comparison of the ektexine of Rumex
(Figure 139) with the common Centrospermae type
(Figures 7-12) indicates that there is not a close
relationship between the two groups. The section
of Rumex acetosa (Figure 170) reveals a thin exine
in which the columellae and especially the foot-
layer-endexine unit are greatly reduced.

The ektexine of seven species in the large New
World genus Coccoloba illustrates a continuous
variation. All taxa are 3-colporate, and Coccoloba
cordata (Figures 110, 111) has an ektexine best de-
scribed as columnar-pyramidal on a base of ran-
domly oriented small rods; C. barbeyana has an
ektexine similar to Ruprechtia rarniflora (Figures
108, 109); C. diversifolia is similar to Calligonum
comosum (Figures 94, 95); Coccoloba venosa and
C. obovata are pi'ominently punctate; and C. pari-
mensis has a microreticulate ektexine similar to
Podopterus mexicanus (Figure 132) and Polygonella
polygama (Figure 131), but with slightly smaller
lumina.

NUMBER 37

11

The internal morphology of Coccoloba cordata
pollen (Figure 147) also appears unique for the
family. The ektexine surface consists of prominent
irregular protuberances supported by a moderately
thick tectum underlain by notably reduced colu-
mellae, greatly thickened foot-layer, and an ex-
tremely thin endexine, if any.

The genus Lastarriaea has been regarded as con-
sisting of two endemic species, L. chilensis Remy in
Chile, and L. coriacea (Goodman) Hoover in Baja
and Southern California. The California taxon has
been treated as L. chilensis ssp. californica Gross,
or as a species of Chorizanthe, C. coriacea Good-
man, who had to select another specific epithet
since “californica” was already applied. The pollen
of Lastarriaea chilensis (Figure 137), L. coriacea,
and Chorizanthe species (Figures 84, 85) are all very
similar, 3-colporate with a punctate ektexine. This
is the most common morphology in the family,
however, and the similarity does not necessarily
indicate that Lastarriaea is congeneric with Chori-
zanthe or is closely related. For the same reason,
the close similarity of the pollen in the two species
of Lastarriaea does not, a priori, indicate a single
species. Careful observation of the floral morphol-
ogy when sampling for pollen, however, revealed
that the flowers of the plants from Chile and from
California are nearly identical in structure. Good-
man (1934), in a revision of the North American
species of Chorizanthe that included the California
taxon, described the genus as having nine stamens,
rarely six or three. The California and Chilean
species of Lastarriaea have only three stamens.

Within limits of the taxa investigated (Table 4),
a number of pollen types were found in only one
species, and such cases are discussed separately here.

Polygonella fimbriata (Figure 130). Grains 3-
colporate, the endoaperture zonorate; the tectum
variable; finely punctate in the region of the col-
pus, conspicuously reticulate at the mesocolpial
ridges and more so at the poles. The TEM also
indicates that P. fimbriata is distinctive for the
family. This conclusion is based on the presence of
a well-developed foot-layer and a thickened en-
dexine (Figure 161), two characteristices that were
rarely combined. See also the earlier discussion of
Polygonum convolvulus and allied species.

Polygonella polygama (Figure 131). Grains 3-
colporate, the endoaperture zonorate, the nexine

thickened at the margins; the ektexine ± micro-
reticulate.

Comparison of the major subdivisions of the
Polygonaceae (Dammer, 1893; Roberty and Vau-
tier, 1964; Reveal and Howell, 1976) with pollen
morphology reveals little correlation. This lack of
correlation is due primarily to two complementary
phenomena: the wide distribution of the punctate-
striate pollen type, which cuts across subfamily and
tribal lines, and the wide array of pollen types
within one genus. Polygonum. Thus far, the pollen
morphology would support Reveal’s concept (Reveal
and Howell, 1976) of the subfamily Eriogonoideae.
Twelve of the 13 genera that he assigned to this
subfamily, have been examined by light microscopy
and/or SEM, and all are 3-colporate with either
punctate or punctate-striate ektexines, the most
common pollen type in the Polygonaceae. However,
an unusual characteristic illustrated by TEM, a
granular inner surface of the tectum, is known thus
far to be restricted to the Eriogonoideae; Chori-
zanthe (Figures 145, 146); Eriogonum (Figures 148,
149); Nemacaulis (Figure 157); Oxytheca (Figure
158). Another genus in this subfamily, Harfordia
(Figure 152), is slightly gi'anular. Examination of
additional genera by TEM might well reinforce the
value of this characteristic and also the validity of
the subfamily.

Primulaceae

Figures 174-200

Twenty-nine species (Table 5) representing 22
genera were examined by light microscopy and
SEM, and 9 of these were sectioned and examined
by TEM.

The structure of the aperture in the Primulaceae
is more complex than in the Plumbaginaceae and
in the Centrospermae. If the number is three or
four, the apertures are generally compound, and
the endoaperture is easily seen in light microscopy.
A number of taxa with 3-colporate grains had a
bridge over the endoaperture formed by the exten-
sion of the two lateral margins of the colpus (Eig-
ures 175, 180-184). One species, Lysimachiopsis
hillebrandia, had 4-colporate grains in which each
side of an endoaperture frequently terminated in a
V-shaped process. Sometimes the V from one endo-

12

SMITHSONIAN CONTRIBUTIONS TO BOTANY

aperture and that of the adjacent one formed the
outline of a diamond. Grains with 5-8 apertures,
e.g., some Primula species, are generally zonocol-
pate (Figure 186). In some taxa, e.g., Cortusa
matthioli L. (Figure 188), the colpi fuse at the poles
to form a triangular apocolpial field.

Different surface patterns were due largely to
variation in the perforation of the tectum. In some
grains the surface was finely punctate (Lysimachia
hybrida, Figure 175; Douglasia montana, Figure
182), in others prominently punctate {Coris mon-
speliensis, Figure 174; Cyclamen neopolitanum.
Figure 184), irregularly perforate (Naurnbergia
thrysiflora, Figure 176), or microreticulate {Hot-
tonia palustris, Figure 185; Primula veris, Figures
186, 187; Omphalogramma vinciflorum, Figure
191). Still others {Lysimachia terrestris, Figure 177;
Stimpsonia ch am ae dry aides, Figures 178, 179) might
be described as finely reticulate. Additionally, in
some grains the tectal perforations were most pro-
nounced in the mesocolpial regions {Coris, Figure
174; Naurnbergia, Figure 176; Lysimachia terrestris.
Figure 177; Anagallis, Figure 183).

Transmission electron microscopy indicates a
variability of exine structure within the family.
Most taxa examined have a well-developed foot-
layer and tectum, e.g., Anagallis (Figure 192),
Glaux (Figure 194), Nainnbergia (Figure 196),
Omphalogramma (Figure 197), ± short columellae,
e.g., Glaux (Figure 194), Naurnbergia (Figure 196),
and an endexine that is massive in the region of the
colpus and thinner but still prominent in the meso-
colpial regions, e.g., Anagallis (Figure 192), Glaux
(Figure 194), Namnbergia (Figure 196), Omphalo-
gramma (Figure 197), Stimpsonia (Figure 199). In
Lysimachia hybrida (Figure 195) the irregular tec-
tum, incomplete columellae, and possibly the exten-
sive endexine, are due to oblique section. Two
species of Prnnula, P. officinalis (Figure 198) and
P. veris (Figure 200), are exceptional, the foot-layer
seemingly absent.

Heterostyly and dimorphic pollen grains are
known to occur in the Primulaceae. Punt, et el.
(1974) acknowledge this condition in some species
of Primula and in Hottonia palustris, as well as
dimorphic pollen grains in Trientalis europaea,
due apparently to polyploidy. In Glaux maritima
the collection of Redfield s.n. from Maine was dis-
tinct enough from that illustrated by Punt, et al.

(1974, pi. 14: figs. 1, 2) that the identification of
Redfield s.n. was rechecked and verified, but still a
second sample, Haakana s.n. from Finland, was
examined, and these grains were more similar to
those illustrated by Punt, et al. (1974). The differ-
ences in Glaux pollen may be due to heterostyly,
not reported in the literature to our knowledge,
or the pollen grains of this species may be simply
rather variable. There are also some differences
between our illustration of Hottonia palustris (Fig-
ure 185) and that of Punt, et al. (1974, pi. 4: figs.
11, 12; pi. 5: figs. 1-6), but the distinctly prolate
shape shown in Figure 185 may be an artifact of
preparation, i.e., collapse. We acknowledge the
existence of heterostyly in the Primulaceae and
dimorphic pollen (as in the Plumbaginaceae) but
did not want the present study to be diluted by a
detailed examination of this phenomenon. Whether
it is the long- or short-styled form that is illustrated
for Hottonia, Primxda, and other Primulaceae, both
authors feel strongly that the alternate form will
not be shown to have the ektexine characteristic of
so many of the centrospermous taxa (Figures 1-12).
For detailed descriptions and measurements, the
reader is advised to consult the article by Punt,
et al. (1974).

Discussion and Conclusions

Of the three families examined in this study, the
Polygonaceae is the most frequently considered as
being related to the Centrospermae. Of the families
that comprise this order, the Caryophyllaceae ap-
pears to be the most likely family from which the
Polygonaceae could be derived or related to, since
both have the anthocyanin pigments. The data
from plastid structure, however, does not support
a relationship between the Polygonaceae and Caryo-
phyllaceae; the Caryophyllaceae not only have
the protein or P-type plastid characteristic of the
Centrospermae, but the plastids have a central crys-
talloid with a distinctive polygonal shape Behnke,
1976:42); the Polygonaceae have the more common
starch or S-type plastid, but admittedly this char-
acterization is based on a small number of species.

The data presented in this palynological study
indicate that the Polygonaceae and Caryophyllaceae
are not related. Although the Polygonaceae is a
large and palynologically diverse family, examina-

NUMBER 37

13

tion of 85 species representing 36 genera (Figures
82-173), revealed no pollen types similar to those
in the Caryophyllaceae. (For additional information
on Caryophyllaceae, see Nowicke, 1975; Skvarla and
Nowicke, 1976). The pollen data do not support
a relationship between the subtribe Illecebrinae
(Caryophyllaceae) or the larger Illecebraceae and
any of the Polygonaceae examined in this study.
The grains of Illecebnan verticillaturn are almost
identical to those of Herniaria glabra (Figure 15):
in both species the pollen has a distinctive tetra-
hedal shape with a large aperture on each of the
three (four?) faces. Cardionema ramosissima (Figure
16) has pantoporate grains, roughly cube-shaped
and with a spinulose and tubuliferous/punctate
ektexine. The unusual grains of Illecebnim verii-
cillatum and Herniaria glabra have not been found
in any other taxa examined to date, including
those remaining in the order Centrospermae, as well
as the Plumbaginaceae, Polygonaceae, and Primu-
laceae. These results would support Hutchinson’s
(1959) inclusion of Herniaria in the Illecebraceae,
but not necessarily the family status since the re-
maining species examined, including another species
of Herniaria, H. hirsuta, all have a common panto-
porate grain.

An argument could be raised that the pollen of
the Bistorta complex in Polygonum (Figures 124-
129) and that oi Muehlenbeckia cunninghamii (Fig-
ures 100, 101) provide some evidence for a rela-
tionship to the Centrospermae. However, we think
it much more likely that the occurrence of this
ektexine surface in a limited number of species is
a reflection of the enormous palynological diversity
in the Polygonaceae: ektexines that are ± psilate,
punctate, punctate-striate, conspicuously spinulose,
perforate with small spines, microreticulate, finely
reticulate to coarsely reticulate; apertures that are
porate, pantocolpate or colporate with variable
endoapertures; and grains that are probably unique
(limited to the Polygonaceae), such as those of Poly-
gonella fimbriata (Figure 130) and related species.
Polygonum convolvulus (Figures 140, 141) and re-
lated species, or Coccoloba cordata (Figures 110,
111).

It is also clear that a diversity of structural forms
exist in the Polygonaceae. These observations par-
allel the sculpturing pleomorphism illustrated by
SEM. The TEM reveals wide variation in the de-

velopment of two layers — the endexine and foot-
layer. The endexine can be absent (apparently) as
in Podopterus (Figure 160), thin but recognizable
as in Chorizanthe paniculata (Figure 146), well de-
veloped as in Atraphaxis (Figure 143) and Nerna-
caulis (Figure 157), and greatly enhanced as in
Eriogoniwi spp. (Figures 148, 149). The foot-layer
has a similar range of variation: from apparently
absent in Chorizanthe breweri (Figure 145) and
Nemacaulis (Figure 157) to prominently thickened
in Coccoloba cordata (Figure 147) and Podopterus
(Figure 160), with many intermediate stages repre-
sented in other taxa.

Another unusual characteristic is the granular
inner surface of the tectum found in a number of
closely related genera: Chorizanthe spp. (Figures
145, 146), Eriogonum (Figure 149), Nemacaulis
(Figure 157), and Oxytheca (Figure 158).

While the tectum tends to be thin in the above
taxa, in others, Fagopyrum (Figure 150) and Tri-
plaris (Figure 173), it is much thicker.

The structure of the exine in the Bistorta com-
plex of Polygonuju (Figures 164, 168), densely
packed columellae, is unique in all the taxa exam-
ined with TEM in this study or previously (Skvarla
and Nowicke, 1976) and thus diminishes the pos-
sibility tliat the Centrospermae and the Poly-
gonaceae could be linked by this group. In fact, it
seems quite notable that in spite of the diverse
exine structures in this family, there is very little
similarity to the Centrospermae.

Tlie spinulose and punctate ektexine surface
found in the Polygonaceae or in the Centrospermae,
for that matter, is simple enough to have arisen
independently in any of these taxa. However, the
significance attached to this ektexine in the Cen-
trospermae is that in every betalain family, as well
as in tlie Caryophyllaceae and Molluginaceae, the
overwhelming majority of the species examined had
pollen grains with a spinulose and tubuliferous/
punctate ektexine, which, moreover, were some-
times identical between families. Insofar as pollen
structure is concerned, a more defensible relation-
ship could be proposed between the Polygonaceae
and any dicot family with a preponderance of
punctate ektexines. Pollen grains with a promi-
nently reticulate ektexine are widely (and ran-
domly?) distributed among dictoyledon families,
but this may be due to parallel evolution and does

14

SMITHSONIAN CONTRIBUTIONS TO BOTANY

not necessarily mean that these families are related
to each other or to the Polygonaceae.

This study reinforces current opinion (Cronquist,
1968; Philipson, 1975:74; Takhtajan, 1969:220)
that the Primnlaceae are not related to nor derived
from the Centrospermae. According to Croncjuist
(1968:177, 223) the only special feature in common
between the Caryophyllales (Centrospermae) and
the Primulales is the free-central placenta tion. The
results of pollen analysis indicate that the ancestors
and/or relatives of the Primnlaceae are not in the
centrospermous families. Although the pollen of the
Primnlaceae exhibits some variation (Figures 174-
200), none of the taxa examined had grains with a
counterpart in the Centrospermae.

Unlike the Primnlaceae, the Plumbaginaceae are
still regarded as being related to the Centrospermae
(Takhtajan, 1969:215), although Cronquist (1968:
185) refers to the family as “somewhat more iso-
lated but may also be derived from the Caryo-
phyllales” and Philipson (1975:74) considers the
Plumbaginaceae-Centrospermae relationship as more
debatable (than that of the Polygonaceae-Centro-
spermae). The pollen of the Plumbaginaceae
(Figures 29-81) consists of the two major types,
designated previously as Armeria and Plumbago,
each of which can be dimorphic in heterostyled
species. This results in four distinctive types, all
3- (or 4-) colpate (Ceratostigma xvilbnottianum and
Limoniastrum monopetalum notwithstanding):
finely and coarsely reticulate Armeria forms (Figures
41-58), and pointed-verrucae and rounded-verrucae

Plumbago forms (Figures 29-40). Admittedly the
two forms of the Plumbago type are based on re-
sults from only two species, but the Plumbago
ektexine type has not been found in any other
taxon in this study or in earlier related ones
(Nowicke, 1975; Skvarla and Nowicke, 1976). The
coarsely reticulate form of the Armeria type has a
superficial resemblance to some reticulate grains
in the Polygonaceae, but the formation of the muri
in the Polygonaceae is distinctive, alternate dupli-
baculate (Figures 112-117, 122, 123), versus sim-
plibaculate in the Armeria forms (Figures 41-58).
In reticulate grains found in the Centrospermae
and in the Polygonaceae (Figures 113-116), the
lumina usually have at least some free columellae
(or bacida), whereas in both Armeria forms (Figures
42, 44, 46, 48, 51, 55, 56), the lumina are consistently
free of any columellar material (the data of Praglow-
ski and Erdtman, 1969, notwithstanding).

In the Plumbaginaceae the pollen data, plastid
structure, and pigmentation (unless the tie is to
the Caryophyllaceae and/or Molluginaceae) suggest
that this family is not related to the Centrospermae
and that its phylogenetic position needs to be
reevaluated.

Like the Plumbaginaceae, the Primnlaceae has the
anthocyanin pigments, and has the starch type
plastid structure (Behnke, pers. comm.). However,
most authors (Cronquist, 1968; Philipson, 1975;
Takhtajan, 1969) do not regard the Primnlaceae as
related to the Centrospermae and the pollen data
presented here reinforces that view.

Literature Cited

Baker, H. G.

1948. Dimorphism and Monomorphism in the Plumba-
ginaceae, I; A Survey of the Family. Annals of
Botany, 12:207-219.

1953. Dimorphism and Monomorphism in the Plumba-
ginaceae, II: Pollen and Stigmata in the Genus
Limonium. Annals of Botany, 17:433-445.

1966. The Evolution, Functioning and Breakdown of
Heteromorphic Incompatability Systems, I: The
Plumbaginacea. Evolution, 20:349-368.

Behnke, H. D.

1971. Sieve-tube Plastids of Magnoliidae and Ranun-
culidae in Relation to Systematics. Taxon, 20:723-
730.

1975. Hectorella caespitosa: Ultrastructural Evidence
against Its Inclusion into Caryophyllaceae. Plant
Systematics and Evolution, 124:31-34.

1976. Delimitation and Classification of the Order Caryo-
phyllales (Centrospermae) According to Ultrastruc-
tural Data from Sieve-Element Plastids: A Survey
Based on 146 Species. Plant Systematics atid Evolu-
tion, 126:31-54.

Behnke, H. D., and B. L. Turner

1971. On Specific Sieve-tube Plastids in Caryophyllales:
Further Investigations with Special Reference to the
Bataceae. Taxon, 20:731-737.

Cronquist, A.

1968. The Evolution and Classification of Flowering
Plants. X 396 pages. Boston: Houghton Mifflin Co.

Dammer, U.

1893. Polygonaceae. Pages 1-36 of volume la in Engler
and Prantl, Die Naturlichen Pflanze^ifamilien III.
Leipzig: Englemann.

Dulberger, R.

1975. Intermorph Structural Differences between Stig-
matic Papillae and Pollen Grains in Relationship
to Incompatibility in Plumbaginaceae. Proceedings
of the Royal Society (London), series B, 188:257-274.

Eckhardt, T.

1964. Engler’s Syllabus der Pflanzenfamilien. 12th edition,
volume 2. Berlin: Gebruder Borntraeger. [Centro-
spermae, pages 79-101].

Erdtman, G.

1966. Pollen Morphology and Plant Taxonomy: Angio-
sperms. xii q- 553 pages. Stockholm: Almcjuist &
Wiksell.

1970. Dber Pollendimorphie in Plumbaginaceae-Plumba-
gineae. Svensk Botanisk Tidskrift, 64:184-188.

Erdtman, G., anci A. Dunbar

1966. Notes on Electron Micrographs Illustrating the

Pollen Morphology in Armeria maritima and
Armeria sibirica. Grana Palynologica, 6:338-354.

Goldblatt, P., J. W. Nowicke, T. J. Mabry, and H. D. Behnke

1976. Gyrostemonaceae: Status and Affinity. Botanica
Notiser, 129:201-206.

Goodman, G.

1934. A Revision of the North American Species of the
Genus Chorizanthe. Annals Missouri Botanical Gar-
den, 21:1-102.

Hedberg, O.

1947. Pollen Morphology in the Genus Polygonum L. s.
lat. and Its Taxonomical Significance. Svensk
Botanisk Tidskrift, 40:371-404.

Horton, J. H.

1963. A Taxonomic Revision of Polygonella (Polygona-
ceae). Brittonia, 15:177-203.

Hutchinson, J.

1959. Dicotyledons. Volume 1 of Families of Floivering
Plants, xi -t- 510 pages. London: Oxford University
Press.

Kohler, E.

1976. Pollen Dimorphism and Heterostyly in the Genus
Waltheria (Sterculiaceae). Pages 147-161 in 1. K.
Ferguson and J. Muller, editors. The Evolutionary
Significance of the Exine. London: Academic Press.

Mabry, T. J., A. Taylor, and B. L. Turner

1963. The Betacyanins and Their Distribution. Phyto-
chemistry, 2:61-64.

Nowicke, J. W.

1975. Pollen Morphology in the Order Centrospermae.
Grana. 15:51-77.

Pax, F., and K. Hoffman

1934. Caryophyllaceae. Pages 275-364 of volume 16c in
Engler and Prantl, Die Naturlichen Pflanzen-
familien II. Leipzig: W. Engelmann.

Philipp, M.

1974. Morphological and Genetical Studies in the Armeria
maritima .Aggregate. Botanisk Tidsskrift, 69:40-51.

Philipson, W. R.

1975. Evolutionary Lines within the Dicotyledons. New
Zealand fournal of Botany, 13:73-91.

Philipson, W. R., and J. P. Skipworth

1961. Hectorellaceae; A New Family of Dicotyledons.
T ransactions of the Royal Society of New Zealand,
1:31.

Praglowski, J., and G. Erdtman

1969. On the Morphology of the Pollen Grains in Armeria
sibirica in Specimens from between Longitude 30° W
and 60° E. Grana Palynologica, 9:72-91.

Punt, W., J. De Leeuw van Weenen, and W. A. P. van Oostrum

1974. Primulaceae. In Janssen, Punt, and Reitsma, editors.

15

16

SMITHSONIAN CONTRIBUTIONS TO BOTANY

The Northwest European Pollen Flora, 3. Review
of Paleobotany and Palynology, 17(3/4):31-70.

Reveal, J. L., and J. T. Howell

1976. Dedeckera (Polygonaceae), a New Genus from Cali-
fornia. Brittonia, 28:245-251.

Roberty, G., and S. Vautier

1964. Les Genres de Polygonacees. Boissiera, 10:7-128.

Roland, F.

1968. fitude de L’ultrastucure des Aperures, II: Pollens a
Sillons. Pollen et Spores, 10:479-519.

Skvarla, J. J.

1973. Pollen. Pages 456-459 fn P. Gray, editor. Encyclo-
pedia of Microscopy and Microtechnique. New
York: Van Nostrand Reinhold.

Skvarla, J. J., and D. A. Larson

1965. An Electron Microscopic Study of Pollen Morphol-
ogy in the Compositae with Special Reference to
the Ambrosiinae. Grana Palynologica, 6:210-269.

Skvarla, J. J., and J. W. Nowicke

1976. The Structure of the Exine in the Order Centro-
spermae. Plant Systematics and Evolution, 126:55-78.

Takhtajan, A.

1969. Flowering Plants, Origin and Dispersal, x -f 310
pages. Washington, D. C.: Smithsonian Institution
Press.

Thorne, R. F.

1968. Synopsis of Putatively Phylogenetic Classification of
the Flowering Plants. Aliso, 6:57-66.

Table 1. — Results of pollen analysis of taxa examined for ektexine characteristics

Spinulose -t-
Tubuliferous/

Taxa

Species

Genera

Punctate

Other

Plurabaginaceae

20

9

0

21

Polygonaceae

85

36

4?

81

Primulaceae

Betalain families

29

22

0

29

plus Caryophyllaceae
and Molluginaceae

21?

184

33

Table 2. — Centrospermae specimens examined, arranged alphabetically by family (pollen types
I, II, and III are 3-colpate, pantojxirate, and pantocolpate respectively, all with a spinulose and
tubuliferous or punctate ektexine; Spec. = specialized, any type not included in I-III)

Taxa

Collector

Location

Pollen

description

Figure

numbers

AIZOCEAE

Geoaarpon minimum Mackenzie

Palmer 5517

Missouri

II

9

Mesembryanthemum variabile Hawortn....

Walther s.n.

California

I

1, 8, 19

Tetragonia arbusoula Fenzl

Bayliss 2171

S. Africa

I

AMARANTHACEAE

Aahyranthes aspera L

Philipson 10510

New Zealand

II

Chamissoa altissima (Jacquin) H.B.K

Curtiss 269

Cuba

II

Dicraurus altemifolius (Watson) Uline & Bray.

Carter et al. 2061

Mexico

II

Psilotriohum amplum Suessenguth

Burger 3248

Ethiopia

II

13, 20

Ptilotus aorymbosum Gaudichaud-Beaupre. .......

Pritzel 58

Australia

II

P. obovatum Gaudichaud-Beaupre

Lazarides 4333

Australia

II

NUMBER 37

17

Table 2 — Continued

Taxa Collector

BASELLACEAE

Anredera saandens Moquin. Nelson 1666

Location Pollen Figure

description numbers

Mexico

II

3, 21

CACTACEAE

Borsiaaatus tenuiserpens (Rauh & Backeberg)
Kimnach

Hamatoaaatus setispinus Britton & Rose

Opuntia lindheimeri Engelmann. . .

Hutchison & Wright
4448

Pratt 9483
Griffiths 9031

Peru

Texas

Texas

I 11

III 12

Spec. 17, 22

CARYOPHYLLACEAE

Aohyronyohia aooperi A. Gray

Cardionema ramosissima A. Nelson & Macbride,..

Gymnoaarpos frutiaosum Per soon

demiaria glabra L

H. hirsuta L

Illeaebrum vertiaellatum L

Polliohia aampestris Alton.

Silene armeria L

Siphonyahia ameriaana (Nuttall) Torrey &

A. Gray.

Tunica striata (Bunge) Fischer & Meyer

CHENOPODIACEAE

Chenopodium ambrosioides L.

Parker et al. 7768

Arizona

II

Fosberg 22540

Ecuador

II

16

Chevallier 28

Algeria

II

4, 23

Staszkiewicz in
Plantae Poloniae,

Poland

527

Spec .

15, 24

Mandaville 139

Saudi Arabia

II

MO 176804

France

Spec.

Wood 591

S. Africa

II

Klein 2621

Brazil

II

Thorne 15254

Florida

II

14

Goloskokov s.n.

USSR

II

Guinea 112

Spain

II

5, 25

HECTORELLACEAE

Lyallia kjcrguelensis Hooker

Transit of Venus Expd. New Zealand II

MOLLUGINACEAE

Limeum viscosum Fenzl

Mollugo oppositifolius L

NYCTAGINACEAE

Abronia angustifolia Greene.

Bougainvillea peruviana H.B.K.

Mirabilis alipes Watson. ......................

Selinooarpus diffusus A. Gray

Seydel

751

SW Africa

I

Gandhi

2092

India

I

Peebles & Harrison

Arizona

Spec

3949

Hutchison 1415

Peru

Spec

Atwood 4550

Utah

II

Woo ton s.n.

New Mexico

II

2, 7, 26

18, 27

PHYTOLACCACEAE

Agdestis alematidea Mocino & Sesse. .
Miarotea muypurensis (H.B.K.) G. Don
Stegnosperma aubense A. Richardson..

Pringle 3276
Bang 1589
Goldsmith 99

Mexico I

Bolivia II

Mexico I

PORTULACACEAE

Lenzia ahamaepithys Philippi

Lewisia Columbiana (Howell) Robinson. .

Montia meridensis Friedrich.

Naioarene parvifolia (Moquin) Rydberg.

Johnston 6091
Jones 214
Hammen 1194
Applegate 2556

Argentina I

Idaho I

Colombia Spec.

Oregon m

6, 10, 28

18

SMITHSONIAN CONTRIBUTIONS TO BOTANY

Table 3. — Plumbaginaceae specimens examined (all 3-colpate except as noted; A = Armeria
type, AC = coarsely reticulate Armeria form, AF = finely reticulate Armeria form, P = Plum-
bago type, PP = pointed verrucae Plumbago form, PR = rounded verrucae Plumbago form)

Species

Collector

Location

Pollen

description

Figure

numbers

Acantholimon glumaoeum

Boissier

Unknown collector

Cultivated Hort.

AC

Brussels

A. Olivieri Jaubert & Spach. .

Koelz 15794

Iran

AC

65,66

A. phrygium Boissier

Bornmuller 5511

Turkey

AF

Armeria maritima (Miller)

Willdenow

Mathiesen & Pedersen s.n.

Denmark

AC

45, 46, 67

Svenson s.n.

' Scandinavia'

AF

43, 44, 68

•

Thedenius s.n.

Sweden

AC

A. maritima var. sibirioa

(Turczaninow) Lawrence

Bartlett 147

Greenland

A

Calder 2014

Canada

A

Cody 1794

Greenland

A

Ekblaw 610

Greenland

A

Lindager s.n.

Greenland

A

Stein 175

Greenland

A

A. saabra Pallas

H. Smith s.n.

Sweden

AC

Ceratostigma griffithvi

C. B. Clarke

Rock 6577

China

PP

31, 32, 70

Rock 11578

China

PR

29, 30, 69

C. minus Stapf

Schneider 2584

China

P

C. willmottianum Stapf

Kingdon-Ward 12530 MO

India

PP

61, 62

Wilson 1373

China

ppl

59, 60, 71

Dyerophytum afriaanum

(Lamarck) 0. Kuntze

Goldblatt 2250 MO

S. Africa

PR

Werdermann & Oberdieck 594

S. Africa

P

63, 64, 72

Seyel 4025 MO

Namibia

P

Goniolimon aollinum

(Grisebach) Boissier

Rechinger f. 10375

Greece

AC

53, 54, 73

Limoniastrum guyonianum

Durieu de Maisonneuve

Chevallier 82

Algeria

AF

74

L. monopetalvjn Boissier......

Puech s.n.

France

Ac2

Limoniwn viaiosoi Pan.

Vicioso s.n.

Spain

AC

41, 42, 75

L. vulgare Miller

Holm-Nielsen s.n. MO

Denmark

AF

52

Hubbard 13206 MO

Britain

AC

Moss MO

Britain

AC

Lawalree 3820 MO

Belgium

AC

51

Plumbago avcriaulata

Lamarck

Bayliss 1187 GH

S. Africa

PR

39, 40

Bayliss 3106 MO

S. Africa

P

P. europaea L

Raine s.n. GH

'Almunarro'

PP

37, 38

Sporiks s.n. GH

Persia

PR

35, 36

P. rosea L

Urban 5490 GH

Puerto Rico

PP

33, 34

P. saandens L

Woytkowski 5592

Peru

P

76

Statiae sinuata L. .........

Maire & Wilczek 1048

Morocco

AF

57, 58, 77, 78

Vaccari 495

Greece

AC

55, 56, 79, 80

S. tenella Turczaninow

Chaney 517

Mongolia

AC

47, 48, 81

Chaney 517a

Mongolia

AF

49, 50

^ Pantocolpate

2

4-5 zonocolpate

NUMBER 37

19

Table 4. — Polygonaceae specimens examined (3C = 3-colporate, PC = pantocolpate,

PP = pantoporate)

Species

Collector

Location

Pollen

description

Figure

numbers

Afrobmnniahia erecta

Hutchison & Dalziel

Antigonon guatemalense

Baldwin 11489

Liberia

3C, punctate

142

Meisner

Atraphaxis buxi folia

Laughlin 2635

Mexico

3C5 punctate-striate

Jaubert & Spach

Woronow s.n.

USSR

3C, striate-punctate

98, 99, 143

Brunniahia airvhosa Banks . .
Calligonwn aomosim

L' Heritier

Centrostegia thurberi

Biltmore Herb. 1678 B

Alabama

3C, punctate-striate

Mandaville 69

Saudi Arabia

3C, punctate-striate

94, 95, 144

A. Gray

Chorizanthe bvewevi S.

Hoover 4373

California

3C, clustered punctate

Watson

Hoover 11393

California

3C, finely punctate

145

C. fimbi-iata Nut tall

Wiggins 9908

Mexico

3C, clustered punctate

84

C. membranaoea Bentham. ....

Bruce 155

California

3C, clustered punctate

C. paniaulata Bentham

Bro. Claude Joseph 2997

Chile

3C, punctate

85, 146

Chorizanthe sp

Simon 289

Chile

3C, punctate

Coaaoloba bavbeyana Lindau.

Schunke 76

Peru

3C, deeply punctate-
striate

C. oordata Chamisso

Hatschbach et al. 13352

Brazil

3C, columnar-pyramidal

110, 111, 147

C. diversifolia Jacquin....

Bro. Clemente s.n.

Cuba

3C, punctate-striate

C. obovata H.B.K

Duke 4988

Panama

3C, prominently punctate

C. parimensis Bentham

Ducke 1289

Brazil

3C, microreticulate

C. swavtzii Meisner.

Webster et al. 10296

Bahamas

3C5 punctate-striate

C. venosa L

Dedeakera eurekensis

Harris & Britton 10756

Jamaica

3C, punctate

Reveal & Howell.

Reveal et al. 3909

California

3C, punctate

Emex australis Steinheil...

Scheepers 928 MO

S. Africa

3C, perforate and
spinulose

Seydel 2975

SW Africa

3C, perforate and
spinulose

136

E. spinosa (L.) Campdera. . .

Koelz 14304

Iran

3C(?), perforate and
spinulose

Eriogonum oorrellii Reveal.

E. inflation Torrey &

Reveal & Davidse 883

Texas

3C, finely punctate

Fremont

E. mari folium Torrey &

Gentry 14437

Mexico

3C, clustered punctate

A. Gray

Holmgren & Reveal 2706

Oregon

3C, finely punctate-
striate

90, 91, 148

E. parishii S. Watson.

Wiggins & Demaree 4913

Mexico

3C, clustered punctate

149

E. raaemosum Nut tall. ......

Eggleston 5847

Colorado

3C, clustered punctate

E. thomasii Torrey

Fagopynm esaulentum

Jones s.n.

California

3C, punctate-striate

Moench

Gilmania luteolum

Braun s.n.

Ohio

3C, deeply punctate

87, 150

(Coville) Coville. .......

Gymnopodium antigonoides

Gilman 1520

California

3C, punctate

(Robinson?) Blake

Gaumer & Sons 23207

Mexico

3C, punctate-striate

Harfordia maaroptera

Goldman 743

Mexico

3C, punctate-striate

92, 93, 151

(Bentham) Greene & Parry.
Hollisteria lanata S.

Wiggins 7601

Mexico

3C, punctate-striate

96, 97, 152

Watson

Hoover 3496

California

3C, punctate

Koenigia islandioa L. .....

Koelz 6424

India

PP, spinulose

Lastarriaea ahilensis Remy.

Worth & Morrison 16270 UC

Chile

3C, punctate

L. aoriaaea (Goodman)

Worth & Morrison 16338 UC

Chile

3C, punctate

137

Hoover

Muoronea oalifomioa

Leiberg 3260

Howell 32083

California

California

3C, punctate

3C, punctate

Bentham

Muehlenbeakia ahilensis

Grinnell s.n.

California

3C, finely punctate-
striate

88, 89

Meisner.

E. & M. Holway 199

Chile

3C, punctate-striate

102, 103, 153

U. aunninghamii Mueller. . . .

Aston 984

Constable 18440

Lazarides 5745

Australia

Australia

Australia

3C, sparsely spinulose
and punctate

3C, sparsely spinulose
and punctate

3C, sparsely spinulose
and punctate

100, 101, 155

20

SMITHSONIAN CONTRIBUTIONS TO BOTANY

Table 4 — Continued

Species

Collector

Location

Pollen

description

Figure

numbers

M. polybotryar Meisner.....

Pritzel 399

Australia

3C, punctate-striate

156

M. tamni/blia (HBK) Meisner .
Neomillspaughia paniculata

Balls 5804

Ecuador

SC, punctate-striate

104, 105, 154

(Donnell-Smith) Blake....
Nemaaaulis denudata

Edwards P 675

Honduras

3C, punctate-striate

Nuttall. .................

Oxygonum atpipliaifolium

Cleveland s.n.

California

3C, finely punctate

82, 157

Martelli.

Piemeisel & Kephart 56

Kenya

3C, punctate and
microreticulate

0. dregecmm Meisner .......

Bayliss 4515

S. Africa

3C, microreticulate

0. seyhepi Sender. .........

Galpin 13075

S. Africa

SC, microreticulate

Oxypia digynia (L.) Hill...

Oxy theca dendroidea

Neilson 902

Canada

3 colpate-very thin
walled

Nuttall. .................

Eastwood & Howell 7353

Wagenknecht 18118

Nevada

Chile

3C, finely punctate-
striate

3C, finely punctate-
striate

0. luteola Parry. ..........

Pringle s.n.

Walther 14379

California

California

3C, finely punctate-
striate

3C, finely punctate-
striate

0. parishii Parry. .........

S. & W. Parish 993

California

SC, punctate

0. trilobata A. Gray.......

Abrams 3807

California

3C, finely punctate-
striate

83, 158

Persiaaria aooainea Greene.

Eastwood 1427

California

PC, finely reticulate

112

Podoptepus mexicanus Kunth.
Polygonella fimbrdata

Langlassi 922

Mexico

3C, microreticulate

132, 160

(Elliot) Horton. .........

P. polygama (Ventenat)

Ward & Smith 2236

Florida

3C, finely punctate and
reticulate

130, 161

Engelmann & A. Gray. .....

Bright 6309

S. Carolina

3C, microreticulate

131

Polygonwn acuminatum Kunth.

Irwin & Souza 9565

Brazil

PP, reticulate

113

P. affine D. Don. ..........

Staunton et al. 1328

Nepal

3C, finely punctate and
spinulose

124

P. amphibium L. ...........

Braun 29

Indiana

PC, finely reticulate

120, 121, 162, 163

P. bistopta L. ............

Cantlon & Gillis 57-439

Alaska

3C, finely punctate and
spinulose

P. capitatum D. Don........

Cantlon & Malcolm 580085

Calder & Billard 3833

Alaska

Canada

3C, finely punctate and
spinulose

3C, finely punctate and
spinulose

128, 129, 164

Tsiang 4503

China

3C, reticulate

122, 123

C'il'inode Michaux

Hardin 722

N. Carolina

3C, finely punctate-
striate

P. convolvulus L. .........

P. cristatum Engelmann

Horr & McGregor E495

Kansas

3C, psllate and spinulose

140, 141, 165

& Gray. ..................

P. auspidatum Siebold &

Graham 25

Texas

3C, psllate and spinulose

Zuccarini. ...............

Makino 13368

Japan

3C, deeply punctate

P. cyanandpium Diels. ......

Rock 5935

China

PP, spinulose

134, 135

P. dielsii Leville (?).....

Henry 9379

China

3C, coarsely reticulate

116, 117, 166

P. dumetonan L. ...........

Makino 13491

Japan

3C, psilate and spinulose

P. foppestii Diels.........

Rock 9697

China

PC, spinulose

133, 167

P. multiflopum Thunberg....

Makino 15020

Japan

3C, deeply punctate

P. opientale L. ...........

Johnson s.n.

New York

PP, reticulate

114, 115, 159

P. vaaainifolium Wallich. . .

Koelz 976

Ind ia

3C, punctate

126, 127, 168

P. virginianum L. .........

Standley 8342

Missouri

PC, finely reticulate

118, 119, 169

P. vivipavum L. ...........

Ptepopypum auchepi

Aim Son*

Sweden

3-4C, very finely
spinulose

125

Jaubert & Spach. .........

Ptepostegia dpymopioides

Rechinger 27660

Pakistan

3C, punctate-striate

Fischer & Meyer. .........

Gillespie 5495

Arizona

3C, punctate-striate

Rheum delavayi Franchet ....

Rock 16256

China

3C, punctate

Rumex aaetosa L. ..........

Greuter & Hainard 567

Switzerland

3(-4)C, perforate and
spinulose

138, 139, 170

R. aoetosella L. ..........

Rouleau 67

Canada

3(-4)C, perforate and
spinulose

NUMBER 37

21

Table 4~Continued

Species

Collector

Location

Pollen

description

Figure

numbers

R. aquations L

Heikkinen s.n.

Finland

3(-4)C, perforate and
spinulose

R. arispus L

Standley 18083

Montana

3C, perforate and
spinulose

R. sautatus L

Rupreohtia laxifLova

Pfister 1158/2

France

3C, perforate and
spinulose

Meisner

Hatschbach 13161

Brazil

3C, deeply punctate-
striate

106, 171

R. pallida Standley

R. rami flora (Jacquin)

Newman 17

Mexico

3C, perforate-rugulose

107, 172

Meyer.

Syrmeria paniaulata

Haught 3914

Colombia

3C, deeply punctate-
striate

108, 109

Bentham.

Ducke 650 MO

Brazil

3C, i psilate

Triplaris amerioana L

Plttier 12204

Venezuela

3C, punctate-
striate

86, 173

22

SMITHSONIAN CONTRIBUTIONS TO BOTANY

Table 5. — Primulaceae specimens examined (3C = 3-colporate, 4C = 4-colporate)

Species

Collector

Location

Pollen

description

Figure

numbers

Anagallis linifolia L

Reverchon 388

Spain

3C, prominently punctate

183, 192

Androsaae septentrionalis L.

Tarleton 8

Alaska

3C, punctate

Apoahoris pentapetala Duby..
Ardisandra wittsteinii R.

Zimmermann 257

China

3C, microreticulate

Wagner

Centunaulus pentandrus R.

Greenway & Kanuri 15044

MO

Kenya

3C, punctate

Brown

Standley 12941

Florida

3C, microreticulate

Coris monspeliensis L. .....

Tidestrom 13554

France

3C, sculptured 4 psilate

174, 193

Cortusa matthioli L

Cyalamen neapolitanwn

Koelz 4994

India

3-syncolporate , granular

188

Tenore.

Stefani s.n.

Corsica

3C, punctate

184

Dionysia mira Wendelbo

Dodeaatheon alpinvm

Mandaville 3600

Oman

6-8 zonocolpate, deeply
punctate

(A. Gray) Greene

Maguire & Holmgren 26449

Oregon

3C, punctate-granular

180, 181

Douglasia montana A. Gray...

Thompson 13766

Idaho

3C, punctate

182

Glaux maritima L

Haakana s.n.

Finland

3C, punctate-striate

Redfield s.n.

Maine

3C, irregularly punctate-
striate

190, 194

Hottonia palustris L

Cernohorsky 4 Kraj ina

286

Czechoslovakia

3C, microreticulate

185

Lysimaohia ailiata L

Cronquist 4 Jones 5956

Idaho

3C, psilate-f inely
punctate

L. hybrida Michaux.

L. terrestris (L.) Britton,

Leonard 4 Killip 871

Maryland

3C, finely punctate

175, 195

Stern & Poggenberg

Ly simaohiopsis hillebrandii

Mac Kenzie 2720

New Jersey

3C, microreticulate

177

Heller

Naumbergia thyrsiflora

Degener 21278 MO

Hawaii

4C, punctate-psilate

(L.) Reichenbach

OmphaZo gramma vinaae flora

Andersen s.n.

Norway

3C, irregularly
microreticulate

176, 196

Franchet

Rock 3573

China

3C, microreticulate

191, 197

Primula farinosa L.

St. John 90675

Quebec

4-syncolporate ,
microreticulate

P. inaisa Franchet

Rock 17474

China

3C, microreticulate

P. offioinalis (L.) Jacquin.

Ludera s.n.

Poland

6-8 zonocolpate,
microreticulate

198

P. veris L

Charpin et al. s.n.

France

6-10 zonocolpate,
microreticulate

186, 187

Skvortsoo s.n.

USSR

6-8 zonocolpate,
microreticulate

200

P. vulgaris Hudson. .........

Pfister s.n.

Switzerland

6-8 zonocolpate,

microreticulate (many
abnormal grains)

Samolus ebracteatus Kunth. . .

Francis 8a

Florida

3C, punctate

Soldanella alpina L

Treffer, 1362 in Flora
Exsiccata Austro-
Hungarica

Austria

3-colpate (syncolpate? ) ,
scabrate-spinulose

189

S. austriaaa Vierhapper

Stimpsonia ahamaedryoides C.

Keller s.n.

Austria

3-colpate (syncolpate?),
scabrate-spinulose

Wright. ...................

Trientalis ameriaana

Ying Hu 10001

Hong Kong

3C, finely reticulate

178, 179, 199

(Persoon) Pursh

Palmer 1370

Canada

3C, punctate

NUMBER 37

23

Figures 1-6. — Centrospermae pollen, SEM: 1, Aizoaceae, Mesembryanthemum variabile Haworth,
equatorial view, x 4100; 2, Molluginaceae, Lirneum viscosum Fenzl, equatorial view, X 3900; 3,
Basellaceae, Anredera scandens Moquin, X 3550; 4, Caryophyllaceae, Gymnocarpos fruticosum
Persoon, X 3350; 5, Chenopocliaceae, Che7Topodium a7nbrosioides L., X 3600; 6, Portulacaceae,
Naiocrene paTrvifolia (Moquin) Rydberg, x 1900. (Photo reduced to 81 percent.)

24

SMITHSONIAN CONTRIBUTIONS TO BOTANY

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NUMBER 37

25

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26

SMITHSONIAN CONTRIBUTIONS TO BOTANY

NUMBER 37

27

28

NUMBER 37

29

30

SMITHSONIAN CONTRIBUTIONS TO BOTANY

Figures 29-34. — Plumbaginaceae pollen, SEM. Ceratostigma griffithii C. B. Clarke: 29, equatorial
view, X 1250; 30, ektexine surface, X 7500 (29, 30, Rock 11578, rounded verrucae Plumbago
form); 31, equatorial view, X 1450; 32, ektexine surface, X 7500 (31, 32, Rock 6577, pointed
verrucae Plumbago form). Plumbago rosea L.: 33, equatorial view, X 1500; 34, ektexine surface,
X 7500 (33, 34, pointed verrucae Plumbago form). Photo reduced to 74 percent.)

NUMBER 37

31

Figures 35-40. — Plumbaginaccae pollen, SEM. Plumbago eiiropaea L.: 35, polar view, X 1700; 36,
ektexine surface, X 7500 (35, 36, Sporiks sm. GH, rounded verrucae Plumbago form); 37, polar
view, X 1700; 38, ektexine surface, X 7500 (37, 38, Raine s.n. GH, pointed verrucae Plumbago
form). Plumbago auriculata I.amarck: 39, polar view, x 1250; 40, ekte.xine surface, X 7500 (39,
40, Bayliss 1187 GH, rounded verrucae Plumbago form). (Photo reduced to 76i^ percent.)

32

SMITHSONIAN CONTRIBUTIONS TO BOTANY

Figures 41-46. — Plumbaginaceae pollen, SEM. Limonium viciosoi Pan: 41, equatorial view, X 1250;
42, ektexine surface, X 3000 (coarsely reticulate Armeria form). Armeria maritima (Miller)
Willdenow: 43, equatorial view, X 1150; 44, ektexine surface, X 5000 (43, 44, Svenson s.n., finely
reticulate Armeria form); 45, polar view, X 1250; 46, ektexine surface, X 3000 (45, 46, Mathiesen
& Pedersen s.n., coarsely reticulate Armeria form). (Photo reduced to 77 percent.)

NUMBER 37

33

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34

SMITHSONIAN CONTRIBUTIONS TO BOTANY

Figures 53-58. — Plumbaginaceae pollen, SEM. GonioUmon collinum (Griscbach) Boissier; 53,
polar view, X 1600; 54, ektexine surface, X 5000 (coarsely reticulate Armeria form). Statice
sinuata L.: 55, polar view, X 1500; 56, ektexine surface, X 4000 (55, 56, Vaccari 495, coarsely
reticulate Armeria form); 57, polar view, X 1550; 58, ektexine surface, X 7500 (57, 58,
Maire it Wilczek 1048, finely reticulate Armeria form). (Photo reduced to 76 percent.)

NUMBER 37

35

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36

SMITHSONIAN CONTRIBUTIONS TO BOTANY

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NUMBER 37

37

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SMITHSONIAN CONTRIBUTIONS TO BOTANY

Figures 69-74. — Plumbaginaceae pollen, TEM. Ceratostigma griffithii: 69, the section is some-

NUMBER 87

39

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SMITHSONIAN CONTRIBUTIONS TO BOTANY

NUMBER 37

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SMITHSONIAN CONTRIBUTIONS TO BOTANY

Figures 82-87. — Polygonaceae pollen, SEM; 82, Nemacaulis denudata Nuttall, mesocolpial view,
X 3100; 83, Oxytheca trilobata A. Gray, mesocolpial view, X 1850; 84, Chorizanthe fimbriata
Nuttall, mesocolpial view, X 1950; 85, C. paniculata Bentham, equatorial view, X 2750; 86,
Triplaris americana L., equatorial view, X 2650; 87, Fagopyrum esculentum Moench, equatorial
view, X 1600 (Photo reduced to 77i/2 percent.)

NUMBER 37

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SMITHSONIAN CONTRIBUTIONS TO BOTANY

Figures 94-99. — Polygonaceae pollen, SEM: 94, Calligonum comosum L’Heritier, equatorial view,
X 2500; 95, ektexine surface, X 7500; 96, Harfordia macroptera (Bentham) Greene & Parry,
equatorial view, X 3050; 97, ektexine surface, X 7500; 98, Atraphaxis buxifolia Jaubert &
Spach, equatorial view, X 2800; 99, ektexine surface, X 7500. (Photo reduced to 84 percent.)

NUMBER 37

45

Figures 100-105. — Polygonaceae pollen, SEM: 100, Muehlenbeckia cuniiinghamii Mueller, equa-
torial view, X 2730 (Aston 984)-, 101, ektexine surface, X 7500 (Aston 984); 102, M. chilensis
Meisner, equatorial view, x 3570; 103, ektexine surface, X 7500; 104, M. tamnifolia (H.B.K.)
Meisner, equatorial view, X 3600; 105, ektexine surface, X 7500. (Photo reduced to 83i^ percent.)

46

SMITHSONIAN CONTRIBUTIONS TO BOTANY

Figures 106-11 1. — Polygonaceae pollen, SEM: 106, Ruprechtia laxiflora Meisner, equatorial view,
X 3400; 107, R. pallida Standley, equatorial view, X 2500; 108, R. ramiflora (Jacquin) Meyer,
equatorial view, X 2700; 109, ektexine surface, X 5000; 110, Coccoloba cordata Chamisso, equa-
torial view, X 2520; 111, ektexine surface, X 7500. (Photo reduced to 79 percent.)

NUMBER 37

47

Figures 112-117. — Polygonaceae pollen, SEM: 112, Persicaria coccinea Greene, X 1575; 113,
Polygonurn acuminatum Kunth, x 1575; 114, P. orientate L., X 1600; 115, ektexine surface,
X 7500; 116, P. dielsii Leville(?), oblique view, colpus horizontal in lower half, X 2200; 117,
ektexine surface, colpus in center, X 5000. (Photo reduced to 76 percent.)

48

SMITHSONIAN CONTRIBUTIONS TO BOTANY

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Figures 118-123. — Polygonaceae pollen, SEM: 118, Polygonum virginianum L., x 2000; 119,
ektexine surface, X 5000; 120, P. amphibium L., X 1400; 121, ektexine surface, colpus lower
right, X 5000; 122, P- capitatum D. Don, polar view, X 2500; 123, ektexine surface, X 7500.
(Photo reduced to 77i/2 percent.)

NUMBER 37

49

Figures 124-129. — Polygonaceae pollen, SEM. Polygonum affine D. Don: 124, equatorial view,
X 2200. P. viviparum L.: 125, slightly oblique equatorial view, X 1700. P. vaccmifolium Wallich:
126, equatorial view, X 1900; 127, ektexine surface X 5000. P. bistorta L.: 128, mesocolpial view,
X 2300; 129, ektexine surface, x 5000 (128, 129, Caution ir Malcolm 580085). (Photo reduced
to 83 percent.)

50

SMITHSONIAN CONTRIBUTIONS TO BOTANY

Figures 130-135. — Polygonaceae pollen, SEM: 130, Polygonella fimbriata (Elliot) Horton, meso-
colpial view, X 3400; 131, P. polygama (Ventenat) Engelmann & A. Gray, equatorial view,
X 5750; 132, Podopterus mexicanus Kunth, equatorial view, X 2970; 133, Polygonum forrestii
Diels, X 2900; 134, P. cyanandrium Diels, X 3600; 135, ektexine surface, X 8500. (Photo re-
duced to 74 percent.)

NUMBER 37

51

Figures 136-141. — Polygonaceae pollen, SEM: 136, Emex australis Steinheil, equatorial view,
X 2990 (Seydel 2975); 137, Lastarriaea chilensis Remy, mesocolpial view, X 3400 {Worth ir Morrison
16338 UC); 138, Rumex acetosa L., equatorial view, X 4400; 139, ektexine surface, X 10,000; 140,
Polygonum convolvulus L., mesocolpial view, X 3050; 141, ektexine surface, X 7500. (Photo
reduced to 78i/^ percent.)

52

SMITHSONIAN CONTRIBUTIONS TO BOTANY

Figures 142-149. — Polygonaceae pollen, TEM: 142, Antigonon guatemalense Meisner, the
endexine is narrow and plate-like; 143, Atraphaxis buxifolia, section somewhat oblique but

NUMBER 37

53

54

SMITHSONIAN CONTRIBUTIONS TO BOTANY

Figures 150-156.— Polygonaceae pollen, TEM: 150, Fagopyrum esculentum, the section is

NUMBER 37

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Figures 157-165. — Polygonaceae pollen, TEM: 157, NemacauUs denudata, the section illustrates

NUMBER 37

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SMITHSONIAN CONTRIBUTIONS TO BOTANY

Figures 166-173 — Polygonaceae pollen, TEM: 166, Polygonum dielsii, section ± oblique, solid

NUMBER 87

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Figures 174-179. — Primulaceae pollen, SEM: 174, Coris monspeliensis L., equatorial view,
X 2500; 175, Lysimachia hybrida Michaux, equatorial view, X 5000; 176, Naumbergia thyrsifiora
(L.) Reichenbach, mesocolpial view, X 4620; 177, Lysimachia terrestris (L.) Britton, Stern, &
Poggenberg, mesocolpial view, X 4400; 178, Stimpsonia chamaedryoides C. Wright, equatorial
view, X 3200; 179, ektexine surface, x 7500. (Photo reduced to 72 percent.)

NUMBER 37

61

Figures 180-185. — Primulaceae pollen, SEM: 180, Dodecatheon alpinum (A. Gray) Greene, equa-
torial view, X 4000; 181, ektexine surface, X 7500; 182, Douglasia montana A. Gray, meso-
colpial view, X 4250; 183, Anagallis linifolia L., equatorial view, X 3900; 184, Cyclamen
neapolitanum Tenore, mesocolpial view, X 6500; 185, Hottonia palustris L., equatorial view,
X 5250. (Photo reduced to 78 percent.)

62

SMITHSONIAN CONTRIBUTIONS TO BOTANY

Figures 186-191. — Primulaccae pollen, SEM: 186, Primula veris L., oblique view, X 3200; 187,
ektexine surface, X 7500 (186, 187, Charpin et al. s.n.); 188, Cortusa matthioli L., polar view
with triangular apocolpial field, x 5000; 189, Soldanella alpina L., equatorial view, X 6060; 190,
Glaux maritima L., equatorial view, X 4300 (Redfield s.n.); 191, Omphalogramma vincaeflora
Franchet, equatorial view, X 3550. (Photo redut ed to 76 percent.)

NUMBER 37

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