SMITHSONIAN CONTRIBUTIONS TO BOTANY 0 NUMBER 37 

Pollen Morphology and the 

Relationship of the 

Plumbaginaceae, Polygonaceae, 

and Prirnulaceae to the 

Order Centrospermae 

Joan W. Nowicke 

and John 3. Skvnrln 

SMITHSONIAN INSTITUTION PRESS 

City of M7ashington 

1977 
ABSTRACT 

Nowicke, Joan W., and John J. Sklarla. Pollen Morphology and the Relationship of the Plumbaginaceae, Polygonaceae, and Primulaceae to the Order Centrospermae. Siiiz1hsoniun Contl-zbzitions to L’olany, number 37, 64 pages, 200 figures, 

3 tables, 197’i.-Three families, Plumbaginaceae, Poljgonaceae, and Pi imulaceae, 

are consiclei ed to be related to or dei i\ ed from the 01 der Centrospei mae by 1 arious authors. These three families hai e anthocyanin pigments in contrast to the 

betalains found in all but tu o families in the Centrospermae. In addition, all 

three are known to hale staich-tjpe sieie-tube plastids in contrast to the piotein 

type found in all centiospermous families. Examination of the pollen of 134 

species by SEN, TEN, and light micioscopy reiealed g.reat dii ersity, especially in 

the Polygonaceae, but not the spinulose and tubuliferous/punctate ektexine, 

which characterizes the i ast majoritj of the centrospermous taxa. Recent ei idence 

argues against a close relationship of the Plumbaginaceae, Pol) gonaceae, and 

Primulaceae with the Centrospei mae, and the absence of any pollen tjpes common to the three families further suggests that they are not closely related to each 

other. 

OFFICIAL PVBLICATIOS DATE is handstamped in a limited number of initial copies and is recorded 

in the Institution’s annual report, Smithsoninn Year. SERIFS COVER DESIGN: Leaf clearing from the 

katsura tree Cercidiphylltcm japoniczcm Siebold and Zuccarini. 

Library of Congress Cataloging in Publication Data 

Sorvicke, Joan LV. 

Pollen morphology and the relationship of the Plumbaginaceae, Polygonaceae, and Primulaceae 

(Smithsonian contributions to botany ; no. 37) to the order Centrosperrnae. 

Bibliography: p. 

1. Plumbaginaceae. 2. Pol>-gonaceae. 3. Primulaceae. 4. Centrospermae. 5. Palynotaxonomy. 

6. Pollen morpholorry. I. Sktarla, Tohn T., ioint author. 11. Title. 111. Series: Sinithsonian 

Institution. SkithsoGian contributioGs to 6ota”uy ; no. 37. 

QKl.S2747 no. 37 [QK495.A12] 581’.08s [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 

... 

111 

Pollen Morphology and the 

Relationship of the 

Plumbaginaceae, Polygonaceae, 

and Primulaceae to the 

Order Centrospermae 

Joan W. Nowicke 

and John 3. Skvarla 

Introduction 

The Centrospermae (Caryophyllales) represents 

one of the most controversial orders in the angiosperms. 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 characteristics, has unique N-containing pigments, the betalains, and a distinctive structure in the sieve-tube 

plastids. Both Cronquist (1968) and Takhtajan 

(1969) unite the betalain families, Aizoaceae, Amaranthaceae, Basellaceae, Cactaceae, Chenopodiaceae, 

Didiereaceae, Nyctaginaceae, Phytolaccaceae, and 

Portulacaceae, as well as two anthocyanin families, 

Caryophyllaceae and Molluginaceae, in the order 

Caryophyllales ( k Centrospermae). Both authors 

include Halophytum ameghinoi Spegazzini (a betalain 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 

Gyrostemonaceae as part of the Phytolaccaceae, and 

Takhtajan separates it as a distinct family. Takhtajan recognizes the Tetragoniaceae as a separate 

family while Cronquist includes it in the Aizoaceae. 

Dysphania is treated as a member of the Chenopodiaceae by both authors; and finally Takhtajan also 

places the Hectorellaceae and the Bataceae in this 

order. 

Thorne (1968) in a synopsis of angiosperm classification has placed the betalain families in a suborder, Chenopodiinae, one of two constituting the 

order Chenopodiales. He recognized the Gyrostemonaceae 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, Caryophyllinae, consists of two families, the Caryophyllaceae and the Polygonaceae. The next order, Batidales, consists of only the Batidaceae, but Thorne 

apparently regards this family as somewhat related 

tithe Chenopodiales since both orders are united 

Joan W. Nowicke, Department of Botany, National Museum under a superorder, Chenopodiiflorae. 

of Natural History, Smithsonian Institution, Washington, 

D. C. 20560. John J. Skvarla, Department of Botany and Microbiology, University of Oklahoma, Norman, Oklahoma 73019. 

Mabry, and Turner and Behnke 

and Turner (1971) have treated the betalain fami1 
2 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

lies as a distinct group, separate from, yet closely 

allied to, the two anthocyanin families, Caryophyllaceae and Molluginaceae. 

Evidence from ultrastructural research on sievetube 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 Caryophyllaceae and the Molluginaceae. In an investigation 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 designated as the P-type. The betalain families and the 

Caryophyllaceae and the Molluginaceae all had the 

P-type plastid in which proteinaceous filaments 

formed a peripheral ring usually around a crystalloid. IVithin the above-mentioned families, Behnke 

(1976) was able to distinguish three groups based on 

the crystalloid: globular, the most common; polygonal, found in the Caryophyllaceae and in two 

other genera, Limeum (Molluginaceae) and Stegnosperma (Phytolaccaceae); and crystalloid-free, having only the ring, found in two families, the 

Amaranthaceae and the Chenopodiaceae. 

Examination of the pollen of 177 species (Nowicke, 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 pattern in every betalain family and in the Caryophyllaceae 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, Gyrostemonaceae, 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 structure indicates that these three families are not 

closely related to the Centrospermae. The relationship 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 subject of a multidisciplinary study (Goldblatt, et al., 

1976) in which all lines of evidence, including cytological, 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 Centrospermae, i.e., to include or exclude the Caryophyllaceae and Molluginaceae, may never be resolved to everyone’s satisfaction. Of more interest 

in view of the accumulating data on the Centrospermae (sensu lato) is the question of the derivation of the families Plumbaginaceae, Polygonaceae, 

and Primulaceae from this order. 

Takhtajan (1969) considers the Polygonales, containing only the Polygonaceae, as near the Caryophyllales, especially the Portulacaceae and Basellaceae, and probably derived from the same stock as 

the Caryophyllales (Centrospermae). His views on 

the Plumbaginaceae are similar: nearest to Portulacaceae 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. 

A4ccording 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 compound, 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 Caryophyllales by similarities in the pollen and by a more 

or less transitional group of genera which are variously referred to the Caryophyllaceae or treated as 

a separate family, Illecebraceae. The Plumbagina- 
NUMBER 31 3 

ceae are somewhat more isolated but may also be 

derived from the Caryophyllaceae.” 

In one respect the system of Thorne (1968) parallels those of Cronquist (1968) and Takhtajan (1969) 

in that all three authors consider the Polygonaceae 

as related to the Caryophyllaceae. 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 evolutionary 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 Polygonaceae as at least peripheral to the group. The 

position of the Plumbaginaceae remains more debatable” (Philipson, 1975:74). 

Hutchinson (1959: 117) regarded the Caryophyllales (Elatinaceae, Molluginaceae, Caryophyllaceae, 

Ficoidaceae, and Portulacaceae) as “a prolific herbaceous group which has given rise to apetalous orders 

such as Polygonales, Chenopodiales, and perigynous 

petaliferous families as Lythrales, besides sympetalous 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 Hem iayia, 

Paronychia, and Scleranthus, as well as rllecebrzirn, 

among the genera listed at the end of the family 

description. He united the Primulaceae and Plumbaginaceae as the order Primulales, closely related 

to the Caryophyllaceae and perhaps the Saxifragaceae. The Primulaceae, Polygonaceae, and Plumbaginaceae contain the anthocyanin pigments, and the 

last two, at least, are known to have the starch-type 

of plastid (Behnke and Turner, 1971) and apparently 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 samples were removed from herbarium material and 

acetolyzed according to procedure outlined in Erdtman (1966). Samples for the SEM were vacuumcoated 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 0.125y0 OsO, in 0.1 M sodium cacodylate buffer for two hours prior to agar 

incorporation. Thin-sections were made with diamond knives, collected on uncoated grids and 

stained with uranyl acetate and then lead citrate. 

Electron microscope observations were made with 

a Philips model-200 transmission electron microscope. Slides of all samples are deposited at the 

Palynological Laboratory, Department of Botany, 

Smithsonian Institution. 

This study is a comparison of the pollen morphology of three families, Plumbaginaceae, Polygonaceae, 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 annotations. The identifications would have to be mis- 
4 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

taken at the level of family to be significantly misleading 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, Polygonaceae, and Primulaceae revealed diverse forms within 

each family, especially the Polygonaceae, but no distinctive pollen type(s) was common to all three 

families. For this reason each family will be discussed separately following a review of the Centrospermae. 

CENTROSPERMAE 

FIGURES 1-28 

The common pollen types in the Centrospermae 

are illustrated in Figures 1-6: 3-colpate, pantoporate, 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 wriable class of apertures is that of pantoporate. Anredera scandens 

(Figure 3) is usually 6-porate, with one pore on each 

plane of the roughly cube-shaped grain. Gymnocarpos frziticosum pollen (Figure 4), with large, sunken 

pores in a geometric pattern, contrasts sharply 

with that of Chenopodizrm ambrosioides (Figure 5), 

which has numerous small pores. 

Figures 7 through 12 illustrate at high magnification (X 7500) the ektexine surface, spinulose, and 

tubuliferous/punctate, which characterizes the T ast 

majority (85%) of the centrospermous taxa. Hamatocactus septispinus pollen (Figure 12) has the 

largest perforations among the species examined 

by SEM. 

Figures 13 through 18 illustrate specialized or unusual forms, but at least one species, Cal-dioizerna 

ramosissima (Figure 16), has the typical ektexine. 

Two other taxa, Psilotrichtim 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 ~ersion of it-the spines and perforations are much 

reduced. 

The internal structure of the exine in various 

centrospermous pollen types is illustrated in Figures 19-28. Characteristically in the centrospermous 

taxa, the ektexine is well developed while the endexine, with few exceptions (e.g., Mesembryanthemum 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 Opuntia (Figure 22), Limeuni (Figure 26), and Gymnocarpos (Figure 23). Representative pollen from nine families of the Centrospermae 

were examined by TEM. Comparison of these data 

with those of earlier TEM work in the Centrospermae (Skvarla and Nowicke, 1976) allows us to 

expand our knowledge of this order, and a brief 

summary for each family is presented below. 

AIZoAcEAE.-Mesem bryanthemum variabile pollen (Figure 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 narrow but consistent endexine. Mesembrganthemum, 

however, is a large and unsatisfactory genus and 

structural yariation is not unexpected. The exine 

stratification of M. varia bile bears a striking similarity to that of Boerhavia erecta of the Nyctaginaceae (Sklai-la and Nowicke, 1976, fig. 30). 

been shown with SEhI and TEM to be highly pleomorphic (Nowicke, 1975; Skvarla and Nowicke, 

1976). The exine structure of Psilotrichurn amplum 

(Figure 20) extends this pleomorphism by indicating a complete absence of endexine. 

BASELLACEAE.-some pollen in this family is 

unique in having a cuboidal shape (Nowicke, 1975; 

Skvarla and Nowicke, 1976). The structure of 

Anl-edera scandens pollen (Figure 21), is quite different from that of Basella alba (Skvarla and 

Kowicke, 1976, fig. 19) but similar to that of 

Bozissingniiltia gracilis (Roland, 1968, pl. 8: fig. 4), 

a more typical member of the family. 

CACTACEAE.-Tlle structure of Opzintia Zindheimeri pollen (Figure 22) is consistent with other examples of the Cactaceae. Comparison of this micrograph with one of Allztaz~clia (Skvarla and Nowicke, 

1976, fig. 28) again underscores the close relationship of the Cactaceae and Didieriaceae. 

AMARANTHACEAE.-The pollen of this family has 
NUMBER 37 5 

CARYOPHYLLACEAE.-The two taxa examined, 

Gymnocarpos fruticosum (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. 

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 

Limeum viscoszim (Figure 26) appears similar to the 

pollen of Cometes surattensis of the Caryophyllaceae (Skvarla and Nowicke, 1976, fig. 41), as well 

as to Mesembryanthemum crystallinurn of the 

Aizoaceae (Skvarla and Nowicke, 1976, fig. 13). 

These relationships are of interest in the context 

of the introductory remarks concerning the Molluginaceae. 

NYCTAGINACEAE.-~ bronia angzistifolia (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 morphology. 

Naiocrene pal-vifolia (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. Geocarpon m inimzim 

(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, 196l), or 

included in the Caryophyllaceae (Eckhardt, 1964). 

Material of Hectorella was not available, and a 

pollen sample of Lyallia kerguelensis 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 appears characteristic of the Centrospermae, but inconclusive with regard to status or family affinity 

CHENOPGDIACEAE.-The exine structure Of ChePORTL'LACACEAE.-The exine morphology Of 

since these types are of wide occurrence in the 

order. The evidence from plastid 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 plastid 

but lacks the polygonal central crystalloid that characterizes the Caryophyllaceae (Behnke, 1975). 

The pollen types in Figures 15 and 16 are representative of a small group of genera that have 

been treated as a subtribe, the Jllecebrinae, in the 

Caryophyllaceae (Pax and Hoffman, 1934) or given 

family status, the Illecebraceae (Hutchinson, 1959). 

Pax and Hoffman regarded the subtribe as consisting 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, Paronychiinae, with the Illecebrinae as the second. Cronquist (1968:185) regards these genera as a transitional group between the Caryophyllaceae and the 

Polygonaceae. Figure 15 is a single grain of Herniaria glabra and is extremely similar to those of 

Illecebrum verticillatum in scanning electron microscopy 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). 

PLUMB ACIN ACEAE 

FIGURES 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 willmottianzim (Figure 59) the grains were pantocolpate. The ektexine 

was either reticulate or verrucose, two sharply distinct 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 

2940, 59-64) has a well-defined foot layer, highly 

irregular columellae and a continuous tectum. Extending 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 TEN. The Armeria type (T’g ‘1 ures 

41-58) has a foot layer supporting straight, regular 

columellae, which are fused distally into an incomplete tectum of the reticulate configuration. 

Heterostyly, frequently associated with dimorphic 

stigmas and dimorphic pollen grains, is a welldocumented phenomenon in the Plumbaginaceae 

(Baker, 1948, 1953, 1966; Dulberger, 1975; Philipp, 

1974). Although the collections at the US. h’ational 

Herbarium are too limited and valuable to permit 

a study of heterostyly per se, and this is not our intention, 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 longstyled 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 Plumbago type pollen can be illustrated by two collections of Ceratostigma grifithii from China (Figures 

29-32) and two collections of Plum bag0 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). TZ’hether 

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 SEAM have been 

designated as having pointed verrucae or rounded 

verrucae in Table 3. Collections having the pointed 

verrucae were found in Plum bag0 rosea (Figures 33, 

34) and in Ceratostigma willmottianum (Figures 

59-62). A collection identified as Plumbago auriculata (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 maritima (Figures 43-46), Limonium 

uiilgare (Figures 51, 52), and Statice sinziata (Figures 

55-58) and one collection of Statice tenella (Figures 

47-50) with several unattached branches, presumably from different plants. In each of the above 

species there is a prominently (or coarsely) reticulate form (Figures 45-48, 51, 55, 56) and a finely 

reticulate form (Figures 43, 44, 49, 50, 52, 57, 58). 

ilihile each form exhibits subtle interspecific differences, i.e., the coarsely reticulate grain of Armeria 

maritima 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 

SEN are Goniolinum collinum (Figures 53, 54) and 

Limonium viciosoi (Figures 41, 42). In both species 

the collection sampled had the coarsely reticulate 

form. 

TEA1 observations corroborate the dimorphism 

shown by SEM. The verrucose type is represented 

by Ceratostigma grifithii (Figure 69): the fine 

bristle-like columellae heads correspond to the 

rounded verrucae illustrated in Figure 30 by SEM; 

in 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 Armeiia maritima 

(Figures 67, 68) and Statice sinziata (Figures 77-80), 

representative of the reticulate morphology, TEN 

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 structure between our TEMs (Figures 67, 68) and those 

of Erdtman and Dunbar (1966, figs. 1, 2) are obvious. 

In all samples examined the endexine pollen wall 

unit that characteristically occurs beneath the foot 

layer in the majority of angiosperm poIlen 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 transmission 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 willmottianurn, a collection from 

India had 3-colpate grains (Figures 61, 62), and a 

collection from China had pantocolpate grains (Figures 59, 60). The pantocolpate grain has larger clusters 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 Ceratostigma grifithii and Plumbago europaea, but dimorphic pollen grains based on aperture structure 

are not unknown (Kohler, 1976). Additional samples 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 Plurnbaginaceae 

with dimorphic pollen. We are aware that a detailed 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 anthers 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 vuZgare, examination of four collections (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. I), 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 (Figures 33, 34). It seems unlikely that both major 

pollen types, which are morphologically and structurally very different, would be found in one species, 

but our results do not exclude this possibility. 

Another problematical taxon is Armeria maritima 

var. sibiricu (Turczaninow) Lawrence. This particular 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 incIude an “A” and “B” that correspond 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 inflorescence on each of six collections (Table 3). The 

SEhIs, taken at a range of low magnifications, illustrate mixtures of the finely and coarsely reticulate 

Armeria forms. Unfortunately this variety was examined 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 distinction between the two forms is not as striking 

as in the other dimorphic Armeria taxa. Philipp 

(1974, fig. 1) illustrates grains from dimorphic 
a SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Danish Armeria maritima (1974, fig. IA,B) which 

agree very well with our Figures 4346, but more 

importantly the two SEWS from monomorphic 

Greenland plants (1974, fig. Ic,D) also show a possible dilution of the distinction: the mesh of what 

could be the finely reticulate form (fig. lc) is larger, 

and the mesh of what could be the coarsely reticulate form (fig. 1~) 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. maritima 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 24y0 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 (1 969), 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 distinction 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 existence 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 similar or related to the common type found in the 

Centrospermae. 

POLYCONACEAE 

FICURE~ 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 

SERI; 32 of these species were sectioned and examined by TEM. 

The Polygonaceae are one of the most palynologically 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 variable than in the Primulaceae and much more so 

than in the Centrospermae or Plumbaginaceae. 

IVhile the most common type in the Polygonaceae 

is 3- (rarely 4-) colporate, the endoaperture, readily 

delimited in light microscopy, is variable and includes 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 includes tectate and nontectate (reticulate) forms. The 

tectate forms can be punctate (Figures 82-85); 

punctate-striate (Figures 94-99): prominently spinulose (Figures 133, 134); or perforate and smallspinulose (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 grains had one type of surface 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 present 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 laxiflora (Figure 106), the tectum between the striae is variously upraised, producing an 

uneven surface. In Ruprechtia ramiflora (Figures 

108, log), the intergroove tectum is more prominently upraised. In Atraphaxis buxifolia (Figures 

98, 99), the connecting striae are larger, tend to be 

parallel, and result in, at least superficially, a striatelike surface pattern. These distinctions are not 

as conspicuous in TERI (Atraphaxis, Figure 143; 

Ruprechtia laxiflora, 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 breweri, 
NUMBER 31 9 

Eriogonum correllii, and Nemacaulis denudata 

(Figure 82). 

2. Ektexine with clustered punctae: Centrostegia 

thurberi, Chorizanthe fimbriata (Figure 84), C. paniculata (Figure 85), Eriogonum parishii, and E. racemosum. 

3. Ektexine punctate, the punctae connected by 

small striae: Eriogonum inflatum, E. marifolium 

(Figures 90, 91), E. thomasii, Mucronea californica 

(Figures 88, 89), Oxytheca trilobata (Figure 83). 

4. Ektexine more prominently punctate-striate: 

Antigonum guatemalense, Calligonum comosum 

(Figures 94, 95), Gymnopodium antigonoides (Figures 92, 93), Harfordia macroptera (Figures 96, 97), 

Muehlenbeckia chilensis (Figures 102, 103), M. polybotryar, Neom illspaugh ia panicu Eata, and Trip laris 

americana (Figure 86). 

5. Ektexine with prominent sunken punctae: 

Fagopyrum esculentum (Figure 87), Muehlen beckia 

tamnifolia (Figures 104, 105). 

6. Ektexine microreticulate: Oxygonum dregeanum, 0. 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 considerable structural diversity as well. Sixteen of 

these taxa were examined in sections: Chorizanthe 

breweri (Figure 145) and Nemacaulis denudata (Figure 157) of subtype 1, Chorizanthe paniculata (Figure 146) and Eriogonum parishii (Figure 149) of 

subtype 2, Eriogonum marifolium (Figure 148) and 

Oxytheca trilobata (Figure 158) of subtype 3, 

A n tigo n u m guatema lense (Figure 142), Ca 11 igon um 

comosum (Figure 144), Harfordia macroptera (Figure 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 possessing a moderate to thick tectum, long, narrow 

columellae (Calligonum comosum, Figure 144, excepted), 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: Chorizanthe spp. (Figures 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, MuehEenbeckia 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 groups. 

The second most common surface pattern is a 

reticulum formed by larger and fewer columellae, 

arranged in a network and distally fused, thus producing 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 Polygonurn capitatum (Figures 122, 123) and P. dielsii (Figures 116, 

117) the columellae are massive, and a high, conspicuous reticulum is the result. The lumina are 

larger in P. dielsii, but in both species they are 

almost free of any columella material. In Polygonum acuminatum (Figure 113) and P. orientale 

(Figures 114, 115), both pantoporate, the muri alternate with large lumina filled with smaller, free 

columellae. In three pantocolpate taxa, Persicaria 

coccinea (Figure 112), Polygonurn amphibium (Figures 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. amphibium (Figures 

162, 163), and P. virginianum (Figure 169) emphasize 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 best illustrated in a “planar” view (Figures 115, 117). 

Four species of Polygonum, P. afine (Figure 124), 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

P. bistorta (Figures 128, 129), P. vaccinifolium (Figures 126, 127) and P. uiviparum (Figure 125), which 

belong, among others, to the Bistorta complex 

(Hedberg, 1947), and an Australian species of 

Muehlenbeckia, M. cunninghamii (Figures 100, 

101), have an ektexine pattern that could be described as sparsely spinulose and sparsely punctate. 

Grains of all five taxa were 3-colporate, the endoaperture well defined. In thin-section, Mziehlenbeckia cunninghumii, (Figure 155) is very similar to 

the other species of Muehlenbeckia examined by 

TEN (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. vaccinifolium (Figures 126, 127), with the larger perforations in the tectum, are more compatible with 

the cross-section illustrated in TEM (Figure 168). 

Koenigia islandica, Polygonum cyanandyium 

(Figures 134, 135), and P. forrestii (Figure 133) had 

grains that were either pantoporate or pantocolpate 

with a prominently spinulose ektexine. Comparison 

of these grains (Figures 133-135) with the common 

type in the Centrospermae (Figures 1-12) indicates 

that the above species could not be related palynologically to that order. Polygonzim f orrestii (Figure 167) was the most distinctive Polygonum of 

those examined by TEM. The exine consists of a 

thin endexine and an ektexine containing abundant 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 Polygonzim convolvultis (Figures 140, 141), P. cristatunz, 

and P. dumetorum as slightly aberrant forms of his 

Tiniaria type. These three species have almost 

identical pollen grains: 3-colporate with a zonorate 

endoaperture, the ektexine echinate near the colpi 

and psilate at the poles and the mesocolpial regions. The above characteristics make the grains 

distinct, not only from the more typical Tiniaria 

pollen, but from the remaining species examined in 

the present study. This particular morphology is 

paralled 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 (Figure 161), which reinforces the parallel exomorphologies. Polygonzim cilinode, P. czispidatum, and 

P. mzilfiporiim, 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. 

aqziaticzis, R. acetosa (Figures 138, 139), R. acetosella, and R. sczitafus, all of which had a perforate 

and very finely and evenly spinulose ektexine (Figure 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 

Rzimex and its restricted occurrence reinforce Dammer’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 Riimex acetosn (Figure 170) reveals a thin exine 

in which the columellae and especially the footlayer-endexine unit are greatly reduced. 

The ektexine of seven species in the large New 

TVorld genus Coccoloba illustrates a continuous 

variation. ,411 taxa are 3-colporate, and Coccoloba 

cordafn (Figures 110, 111) has an ektexine best described as columnar-pyramidal on a base of randomly oriented small rods; C. bm-beyuna has an 

ektexine similar to Riiprechtia ramifEora (Figures 

108, 109); C. diven~ifolin is similar to Calligonzim 

comosiim (Figures 94, 95); Coccoloba uenosa and 

C. obovata are prominently punctate; and C. @rimensis has a microreticulate ektexine similar to 

Podopfertis mexicanzis (Figure 132) and Polygonella 

polygama (Figure 131), but with slightly smaller 

lumina. 
NUMBER 97 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 moderateIy 

thick tectum underlain by notably reduced columellae, greatly thickened foot-layer, and an extremely thin endexine, if any. 

The genus Lastarriaea has been regarded as consisting 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 Goodman, 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 Chorizanthe 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 morphology when sampling for pollen, however, revealed 

that the flowers of the plants from Chile and from 

California are nearly identical in structure. Goodman (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 colpus, 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 endexine (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 -+ microreticulate. 

Comparison of the major subdivisions of the 

Polygonaceae (Dammer, 1893; Roberty and Vautier, 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 punctatestriate 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: Chorizanthe (Figures 145, 146); Eriogonum (Figures 148, 

149); Nemacaulis (Figure 157); Oxytheca (Figure 

158). Another genus in this subfamily, Harfordia 

(Figure 152), is slightly granular. 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 

SEhI, 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 extension of the two lateral margins of the colpus (Figures 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 zonocolpate (Figure 186). In some taxa, eg., 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 monspeliensis, Figure 174; Cyclamen neopolitanum, 

Figure 181), irregularly perforate (Naum bergia 

thrysifio~~, Figure 176), or microreticulate (Hottonia palustris, Figure 185; Primula veris, Figures 

186, 187; Omphalogramma vinciflorum, Figure 

191). Still others (Lysimachia terrestris, Figure 177; 

Stimpsonia chamaedryoides, Figures 178, 179) might 

be described as finely reticulate. Additionally, in 

some grains the tectal perforations were most pronounced in the mesocolpial regions (Coris, Figure 

174; Nazimbergia, 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 footlayer and tectum, e.g., Anagallis (Figure 192), 

Glaztx (Figure 194), Naiimbergia (Figure 196), 

Onzphalog?-amma (Figure 197), k short columellae, 

e.g., Glntix (Figure 194), Naumbergia (Figure 196), 

and an endexine that is massive in the region of the 

colpus and thinner but still prominent in the mesocolpial regions, e.g., Anagallis (Figure 192), Glaux 

(Figure 194), Nazimbergia (Figure 196), Onzphalogramma (Figure 197), Stimpsonia (Figure 199). In 

Lysimachia hy brida (Figure 195) the irregular tectuin, incomplete columellae, and possibly the extensive endexine, are due to oblique section. Two 

species of Primzila, P. officinalir (Figure 198) and 

P. ueris (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 Primiila and in Hottonia palust?-is, as well as 

dimorpliic pollen grains in Trientalis europaea, 

due apparently to polyploidy. In Glaux maritima 

the collection of Redfield s.n. from Maine was distinct enough from that illustrated by Punt, et al. 

(1974, pl. 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 differences 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 (Figure 185) and that of Punt, et al. (1974, pl. 4: figs. 

11, 12; pl. 5: figs. 1-6), but the distinctly prolate 

shape shown in Figure 185 may be an artifact of 

preparation, i.e., collapse. JVe 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. JVhether 

it is the long- or short-styled form that is illustrated 

for Hottonia, Primula, 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 

Pollgonaceae is the most frequently considered as 

being related to the Centrospermae. Of the families 

that comprise this order, the Caryophyllaceae appears to be the most likely family from which the 

Po1)gonaceae could be derived or related to, since 

both have the anthocjanin pigments. The data 

from plastic1 structure, however, does not support 

a relationship between the Polygonaceae and Caryophjllaceae; the Caryoph)llaceae not only have 

the protein or P-type plastid characteristic of the 

Centrospermae, but the plastids have a central crystalloid with a distinctive polygonal shape Behnke, 

1976:42); the Polygonaceae have the more common 

starch or S-t) pe plastid, but admittedlj this characteriration is based on a small number of species. 

The data presented in this palynological study 

indicate that the Polygonaceae and Car)ophyllaceae 

are not related. Although the Polygonaceae is a 

large and paljnologically diverse family, examina- 
NUMBER 37 18 

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 Illecebrum verticillatum are almost 

identical to those of Herniaria glabra (Figure 15): 

in both species the pollen has a distinctive tetrahedal 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 tubuIiferous/punctate 

ektexine. The unusual grains of Illecebrum verticillatum 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 Primulaceae. These results would support Hutchinson’s 

(1959) inclusion of Herniaria in the Illecebraceae, 

but not necessarily the family status since the remaining species examined, including another species 

of Herniaria, H. hirsuta, all have a common pantoporate grain. 

An argument could be raised that the pollen of 

the Bistorta complex in Polygonum (Figures 124- 

129) and that of Muehlenbeckia cunninghamii (Figures 100, 101) provide some evidence for a relationship 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 3- 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 Polygonella fimbriata (Figure 130) and related species, 

Polygonum convolvulus (Figures 140, 141) and related species, or Coccoloba cordata (Figures 110, 

It is also clear that a diversity of structural forms 

exist in the Polygonaceae. These observations parallel the sculpturing pleomorphism illustrated by 

SEM. The TEM reveals wide variation in the de11 1). 

velopment of two layers-the endexine and footlayer. The endexine can be absent (apparently) as 

in Podopterus (Figure 160), thin but recognizable 

as in Chorizanthe paniculata (Figure 146), well developed as in Atraphaxis (Figure 143) and Nemacaulis (Figure 157), and greatly enhanced as in 

Eriogonum 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 represented 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), Eriogonzirn (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 Triplaris (Figure 173), it is much thicker. 

The structure of the exine in the Bistorta complex of Polygonurn (Figures 164, 168), densely 

packed columellae, is unique in all the taxa examined with TEM in this study or previously (Skvarla 

and Nowicke, 1976) and thus diminishes the possibility that the Centrospermae and the Polygonaceae 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. 

The 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 Centrospermae is that in every betalain family, as well 

as in the Caryophyllaceae and Molluginaceae, the 

overwhelming majority of the species examined had 

pollen grains with a spinulose and tubuliferous,’ 

punctate ektexine, which, moreover, were sometimes identical between families. Insofar as pollen 

structure is concerned, a more defensible relationship could be proposed between the Polygonaceae 

and any dicot family with a preponderance of 

punctate ektexines. Pollen grains with a prominently reticulate ektexine are widely (and randomly?) 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 Primulaceae are not related to nor derived 

from the Centrospermae. According to Cronquist 

(1968:177, 223) the only special feature in common 

between the Caryophyllales (Centrospermae) and 

the Primulales is the free-central placentation. The 

results of pollen analysis indicate that the ancestors 

and/or relatives of the Primulaceae are not in the 

centrospermous families. Although the pollen of the 

Primulaceae exhibits some variation (Figures 174- 

200), none of the taxa examined had grains with a 

counterpart in the Centrospermae. 

Unlike the Primulaceae, the Plumbaginaceae are 

still regarded as being related to the Centrospermae 

(Takhtajan, 1969:2 15), although Cronquist (1968: 

185) refers to the family as “somewhat more isolated but: may also be derived from the Caryophyllales” and Philipson (1975:74) considers the 

Plumbaginaceae-Centrospermae relationship as more 

debatable (than that of the Polygonaceae-Centrospermae). 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 (Ceratostignza willmottianum and 

Limoniastrum monopetalunz 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 results 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 Pol>gonaceae, but the formation of the muri 

in the Pollgonaceae is distinctive, alternate duplibaculate (Figures 112-1 17, 122, 123), versus simplibaculate in the Armeria forms (Figures 41-58). 

In reticulate grains found in the Centrospermae 

and in the Polygonaceae (Figures 113-1 IS), the 

lumina usually have at least some free columellae 

(or bacula), 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 Praglowski and Erdtman, 1969, notwithstanding). 

In the Plumbaginaceae the pollen data, plastid 

structure, and pigmentation (unless the tie is to 

the Caryophyllaceae and ’or llolluginaceae) suggest 

that this family is not related to the Centrospermae 

and that its phylogenetic position needs to be 

reevaluated. 

Like the Plumbaginaceae, the Primulaceae has the 

anthoqanin pigments, and has the starch type 

plastid structure (Behnke, pers. comm.). However, 

most authors (Cronquist, 1968; Philipson, 1975; 

Takhtajan, 1969) do not regard the Primulaceae as 

related to the Centrospermae and the pollen data 

presented here reinforces that view. 
Baker, H. G. 

Literature Cited 

Pollen Morphology in Armeria maritima and 

1948. 

1953. 

1966. 

Behnke, 

1971. 

1975. 

1976. 

Behnke, 

1971. 

Dimorphism and Monomorphism in the Plumbaginaceae, I: A Survey of the Family. Annals of 

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Dimorphism and Monomorphism in the Plumbaginaceae, 11: Pollen and Stigmata in the Genus 

Limonium. Annals of Botany, 17:433-445. 

The Evolution, Functioning and Breakdown of 

Heteromorphic Incompatability Systems, I: The 

Plumbaginacea. Evolution, 20:349-368. 

H. D. 

Sieve-tube Plastids of Magnoliidae and Ranunculidae in Relation to Systematics. Taxon, 20:723- 

730. 

Hectorella caespitosa: Ultrastructural Evidence 

against Its Inclusion into Caryophyllaceae. Plant 

Systematics and Evolution, 124:31-34. 

Delimitation and Classification of the Order Caryophyllales (Centrospermae) According to Ultrastructural Data from Sieve-Element Plastids: A Survey 

Based on 146 Species. Plant Systematics and Evolution, 126:31-54. 

H. D., and B. L. Turner 

On Specific Sieve-tube Plastids in Caryophyllales: 

Further Investigations with Special Reference to the 

Bataceae. Taxon, 20:731-737. 

Cronquist, A. 

1968. The Evolution 

Plants. x + 396 

and Classification of Floulering 

pages. Boston: Houghton Mifflin Co. 

1893. Polygonaceae. Pages 1-36 of volume la in Engler 

and Prantl, Die Naturlichen Pflanzenfamilien Ill. 

Leipzig: Englemann. 

Dulberger, R. 

Dammer, U. 

1975. Intermorph Structural Differences between Stigmatic Papillae and Pollen Grains in Relationship 

to Incompatibility in Plumbaginaceae. Proceedings 

of the Royal Society (London), series B, 188:257-274. 

Engler's Syllabus der Pflanzenfamilien. 12th edition, 

volume 2. Berlin: Gebruder Borntraeger. [Centrospermae, pages 79-101 J. 

Eckhardt, T. 

1964. 

Erdtman, G. 

1966. Pollen Morphology and Plant Taxonomy: Angiosperms. xii + 553 pages. Stockholm: Almquist & 

Wiksell. 

1970. Uber Pollendimorphie in Plumbaginaceae-Plumbagineae. Svensk Botanisk Tidskrift, 64: 184-188. 

1966. Notes on Electron Micrographs Illustrating the 

Erdtman, G., and A. Dunbar 

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. 

A Revision of the North American Species of the 

Genus Chorizanthe. Annals Missouri Botanical Garden, 21:l-102. 

Pollen Morphology in the Genus Polygonurn L. S. 

lat. and Its Taxonomical Significance. Svensk 

Bo tan isk Tidskrift , 40: 3 7 1-404. 

Goodman, G. 

1934. 

Hedberg, 0. 

1947. 

Horton, J. H. 

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

Hutchinson, J. 

1959. Dicotyledons. Volume 1 of Families of Flowering 

Plants. xi + 510 pages. London: Oxford University 

Press. 

Pollen Dimorphism and Heterostyly in the Genus 

Waltheria (Sterculiaceae). Pages 147-161 in I. K. 

Ferguson and J. Muller, editors, The Evolutionary 

Significance of the Erine. London: Academic Press. 

1963. The Betacyanins and Their Distribution. PhytoKohler, E. 

1976. 

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

c h em is try, 2: 6 1-64. 

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 Pflanzenfamilien 11. Leipzig: W. Engelmann. 

hforphological and Genetical Studies in the Armeria 

maritima Aggregate. Botanisk Tidsskrift, 69:40-51. 

Philipp, 31. 

1974. 

Philipson, W. R. 

1975. Evolutionary Lines within the Dicotyledons. New 

Zealand Journal of Botany, 13:73-91. 

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

1961. Hectorellaceae: A New Family of Dicotyledons. 

Transactions of the Royal Society of New Zealand, 

1:31. 

On the Morphology of the Pollen Grains in Armeria 

sibirica in Specimens from between Longitude 30° W 

and 60' E. Gram Palynologica, 9:72-91. 

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

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

Praglowski, J., and G. Erdtman 

1969. 

1974. 

15 
16 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

The Northwest European Pollen Flora, 3. Review 

of Paleobotany and Palynologp, 17(3/4):31-70. 

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

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

Etude de L'ultrastucure des Aperures, 11: Pollens a 

Sillons. Pollen et Spores, 103479-519. 

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

1976. 

Roberty, G., and S. Vautier 

1964. 

Roland, F. 

1968. 

Skvarla, J. J. 

1973. Pollen. Pages 456459 in P. Gray, editor, Encyclopedia of Microscopy and Microtechnique. New 

York: Van h'ostrand Reinhold. 

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

1969. An Electron Microscopic Study of Pollen hlorphology in the Cornpositae with Special Reference to 

the Ambrosiinae. Grana Palynologica, 6:210-269. 

Skvarla, J. J., and J. XV. Nowicke 

1976. The Structure of the Exine in the Order Centrospermae. Plant Systematics and Evolution, 126355-78. 

1969. Flowering Plants, Origin and Dispersal. x + 310 

pages. Washington, D. C.: Srnithsonian Institution 

Press. 

Synopsis of Putatively Phylogenetic Classification of 

the Flowering Plants. Aliso, 6:57-66. 

Takhtajan, A. 

Thorne, R. F. 

1968. 

TABLE 1 .-Results of pollen analysis of taxa examined for ektexine characteristics 

Taxa 

Plumbaginaceae ........... Polygonaceae ............. Frimlaceae .............. Betalain fanilies 

plus C aryop hy llac eae 

and Molluginaceae ...... 

Spinolose + 

Tubulif erous/ 

Species Genera Punc tat e Otner 

20 9 0 

85 36 4? 

29 22 0 

217 184 

21 

El 

29 

33 

TABLE 2.-Centrosperrnae specimens examined, arranged alphabetically by family (pollen types 

I, 11, and I11 are 3-colpate, pantoporate, and pantocolpate respectively, all with a spinulose and 

tubuliferous or punctate ektexine; Spec. = specialized, any type not included in 1-111) 

Collector Location Pollen Figure 

description numbers Taxa 

~ ~~ 

AIZOCEAE I 

I 

Geocarpon minim Mackenzie ................... Mesembryanthemm variabile Haworth ............ Tetragonia arbuscuZa Fenzl .................... 

AMARANTHACEAE 

Achgranthes aspera L. ........................ Chamissoa altissima (Jacquin) H.B.K. ......... Dicraurus aZternifoZius (Watson) Uline & Bray. 

PsiZotrichm ampZm Suessenguth ............... PtiZotus corymbosm Gaudichaud-Beaupre.. ...... P. obouatm Gaudichaud-Beaupre ................ 

Palmer 5517 

Wal ther s . n. 

Bayliss 2171 

Missouri I1 9 

California I 

S. Africa I 

1, 8, 19 

Philipson 10510 New Zealand I1 

Curtiss 269 Cuba I1 

Carter et al. 2061 Mexico I1 

Burger 3248 Ethiopia I1 

Pritzel 58 Australia I1 

Lazarides 4333 Australia I1 

13, 20 
NUMBER 37 17 

TABLE 2-Continued 

Taxa 

BASELLACEAE 

Anredera scandens Moquin ...................... 

CACTACEAE 

Borzicactus tenuiserpem (Rauh & Backeberg) 

Kimnach..................................... 

Hamatocactus setispinus Britton & Rose........ 

Gpuntia lindheheri Engelmann ................. 

CARYOPHYLLACEAE 

Achyronychia cooperi A. Gray .................. Cardionema ramosissha A. Nelson L Macbride,.. 

Gymocarpos fruticosm Persoon................ 

Hemiaria glabra L. .......................... 

H. hirsuta L. ................................ Illecebm verticellatm L. . . .. .. . .. .. .. . . . .. Pollichia campestris Aiton.................... 

SiZene meria L. ............................ Siphonychia americana (Nuttall) Torrey L 

A. Gray ..................................... Tunica stricta (Bunge) Fischer L Meyer... ..... 

CHENOPODIACEAE 

Chenopodim ambrosioides L. .............. 

HECTORELLACEAE 

LyalZia kergueZensis Hooker............. ... 

MOLLUGINACEAE 

Limem viscosm Fenzl..... .................... Mollugo oppositifolius L. .................... 

NYCTAGINACEAE 

Abronia angustifolia Greene................... 

Bougainvillea peruviana H.B.K. ............... MirabiZis alipes Watson....................... 

Selinocarpus diffusus A. Gray ................. 

PHYTOLACCACEAE 

Agdestis clematidea Mocino & Sesse.... ........ Microtea mypurensis (H.B.K.) G. Don .......... Stegnosperma cubense A. Richardson ............ 

PORTULACACEAE 

Lenzia chumaepithys Philippi .................. Lewisia colmbiaw (Howell) Robinson. ......... Montia meridensis Friedrich.... ............... Naiocrene parvifollia (Moquin) Rydberg ......... 

Collector Location Pollen Figure 

description numbers 

Nelson 1666 Mexico 

Hutchison L Wright Peru 

Pratt 9483 Texas 

Griffiths 9031 Texas 

4448 

Parker et al. 7768 Arizona 

Fosberg 22540 Ecuador 

Chevall ier 28 Algeria 

Staszkiewicz in Poland 

Mandaville 139 Saudi Arabia 

MO 176804 France 

Wood 591 S. Africa 

Klein 2621 Brazil 

Thorne 15254 Florida 

Goloskokov s.n. USSR 

Plantae Poloniae, 527 

Guinea 112 Spain 

Transit of Venus Expd. New Zealand 

Seydel 751 

Gandhi 2092 

I1 3, 21 

1 

111 

Spec. 

I1 

I1 

I1 

Spec. 

I1 

Spec. 

I1 

I1 

I1 

I1 

I1 

I1 

SW Africa I 

India I 

Peebles L Harrison Arizona 

Hutchison 1415 Peru 

Atwood 4550 Utah 

Wooton s.n. New Mexico 

3949 

Pringle 3276 Mexico 

Bang 1589 Bolivia 

Goldsmith 99 Mexico 

Johnston 6091 Argentina 

Jones 214 Idaho 

Hammen 1194 Colombia 

Applegate 2556 Oregon 

Spec. 

Spec. 

I1 

I1 

I 

I1 

1 

I 

I 

Spec. 

I11 

11 

12 

17, 22 

16 

4, 23 

15, 24 

14 

5, 25 

2, 7, 26 

18, 27 

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 = Plumbag0 type, PP = pointed verrucae Plumbago form, PR = rounded verrucae Plumbago form) 

Species 

Acantho limon glurnacem 

Boissier ................... 

A. olivieri Jaubert & Spach.. 

A. phrygiwn Boissier ......... Armeria maritima (Miller) 

Willdenow .................. 

A. maritima var. sibirica 

(Turczaninow) Lawrence.. ... 

A. scabra Pallas..... ........ Ceratostigma griffithti 

C. B. Clarke ............... 

C. minus Stapf ............... C. willmottianurn Stapf ....... 

Dyerophytm africanm 

(Lamarck) 0. Kuntze ........ 

Goniolimon collinm 

Limoniastrwn guyonianm 

(Grisebach) Boissier ....... 

Durieu de Maisonneuve...... 

Limoniwn viciosoi Pau ........ L. monopetalurn Boissier.. .... 

L. vulgare Miller ........... 

Plumbago auriculata 

Lamarck ................... 

?. europaea L. ............. 

P. rosea L. ................ ?. scandens L. ............. Statice sinuata L. ......... 

S. tenella Turczaninow...... 

Collector Location Pollen Figure 

description numbers 

Unknown collector 

Koelz 15794 

Bo rnmu Ile r 551 1 

Mathiesen & Pedersen s.n. 

Svenson s.n. 

Thedenius s.n. 

Bartlett 147 

Calder 2014 

Cody 1794 

Ekblaw 610 

Lindager s.n. 

Stein 175 

H. Smith s.n. 

Rock 6577 

Rock 11578 

Schneider 2584 

Kingdon-Ward 12530 MO 

Wilson 1373 

Goldblatt 2250 MO 

Werdermann & Oberdieck 594 

Seyel 4025 MO 

Rechinger f. 10375 

Chevallier 82 

Puech s.n. 

Vicioso s.n. 

Holm-Nielsen s.n. MO 

Hubbard 13206 MO 

Moss MO 

Lawalree 3820 MO 

Bayliss 1187 GH 

Bayliss 3106 MO 

Raine s.n. GH 

Sporiks s.n. GH 

Urban 5490 GH 

Woytkowski 5592 

Maire & Wilczek 1048 

Vaccari 495 

Chaney 517 

Chaney 517a 

Cultivated Hort. 

Iran 

Turkey 

Denmark 

'Scandinavia' 

Sweden 

Greenland 

Canada 

Greenland 

Greenland 

Greenland 

Greenland 

Sweden 

China 

China 

China 

India 

China 

S. Africa 

S. Africa 

Namibia 

Greece 

Algeria 

France 

Spain 

Denmark 

Britain 

Britain 

Belgium 

S. Africa 

S. Africa 

'Almunarro' 

Persia 

Puerto Rico 

Peru 

Morocco 

Greece 

Mongolia 

Mongolia 

Brussels 

AC 

AC 

AF 

AC 

AF 

AC 

A 

A 

A 

A 

A 

A 

AC 

PP 

PR 

P 

PP 

PPl 

PR 

P 

P 

AC 

AF 

AC2 

AC 

AF 

AC 

AC 

AC 

PR 

P 

PP 

PR 

PP 

P 

AF 

AC 

AC 

AF 

65,66 

45, 46, 67 

43, 44, 68 

31, 32, 70 

29, 30, 69 

61, 62 

59, 60, 71 

63, 64, 72 

53, 54, 73 

74 

41, 42, 75 

52 

51 

39, 40 

37, 38 

35, 36 

33, 34 

76 

57, 58, 77, 78 

55, 56, 79, 80 

47, 48, 81 

49, 50 

Pan tocolpa te 

4-5 zonocolpate 
NUMBER 37 19 

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

PP = pantoporate) 

Species 

Afrobmnnichia erecta 

Antigonon guatemalense 

AtraphLwris buxifolia 

Jaubert & Spach .......... Brtolnichia cirrhosa Banks.. 

Calligonum cornosum L'Heritier ............... Centrostegia thurberi 

A. Gray .................. Chorizanthe breweri S. 

Watson ................... 

C. rnembranacea Bentham..... 

Chorizanthe sp. ........... Coccoloba barbeyana Lindau. 

C. cordata Chamisso........ 

C. obovata H.B.K. ......... C. parimensis Bentham...... 

C. swartzii Meisner........ 

C. venosa L. .............. Dedeckera eurekensis 

Reveal & Howell .......... hex austraZis Steinheil... 

Hutchison & Dalziel...... 

Meisner .................. 

C. fimb24ata Nuttall.. ..... 

C. paniculata Bentham ...... 

C. diversifolia Jacquin.. .. 

E. spinosa (L.) Campdera.. . 

Eriogowm come 1 lii Reveal. 

E. inflaturn Torrey & 

Fremont .................. E. marifoliurn Torrey & 

A. Gray .................. 

E. parishii S. Watson...... E. racernosum Nuttall ....... 

Fagopyrwn esculentum 

Gilrnania luteolum 

Gymnopodium antigonoides 

E. thomasii Torrey........ . 

Moench ................... 

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

(Rob ins on? ) Blake. ....... 

Harfordia macroptera 

(Bentham) Greene & Parry. 

Hollisteria lanata S. Watson ................... Koenigia islandica L. ..... Lastarriaen chilensis Remy. 

L. coriacea (Goodman) 

Hoover ................... 

kcrowa californica 

Bentham... ............... 

MuehZenbeckia chitensis 

Meisner .................. M. cunninghamii Mueller.... 

Collector Location Pollen 

description 

Figure 

numbers 

Baldwin 11489 

Laughlin 2635 

Woronow s. n. 

Biltmore Herb. 1678 B 

Mandaville 69 

Hoover 4373 

Hoover 11393 

Wiggins 9908 

Bruce 155 

Bro. Claude Joseph 2997 

Simon 289 

Schunke 76 

Hatschbach et al. 13352 Bro. Clemente s.n. 

Duke 4988 

Ducke 1289 

Webster et al. 10296 

Harris & Britton 10756 

Reveal et al. 3909 

Scheepers 928 MO 

Seydel 2975 

Koelz 14304 

Reveal & Davidse 883 

Gentry 14437 

Holmgren & Reveal 2706 

Wiggins & Demaree 4913 

Eggleston 5847 

Jones s.n. 

Braun s.n. 

Gilman 1520 

Gaumer & Sons 23207 

Goldman 743 

Wiggins 7601 

Hoover 3496 

Koelz 6424 

Worth & Morrison 16270 UC 

Worth & Morrison 16338 UC 

Leiberg 3260 

Howell 32083 

Grinnell s.n. 

E. & M. Holway 199 

Aston 904 

Constable 18440 

Lazarides 5745 

Liberia 

Mexico 

USSR 

Alabama 

Saudi Arabia 

California 

California 

Mexico 

California 

Chile 

Chile 

Peru 

Brazil 

Cuba 

Panama 

Brazil 

Bahamas 

Jamaica 

California 

S. Africa 

SW Africa 

Iran 

Texas 

Mexico 

Oregon 

Mexico 

Colorado 

California 

Ohio 

California 

Mexico 

Mexico 

Mexico 

California 

India 

Chile 

Chile 

California 

California 

California 

Chile 

Australia 

Australia 

Australia 

3C, punctate 

3C, punctate-striate 

3C, striate-punctate 

3C, punctate-striate 

3C, punctate-striate 

3C, clustered punctate 

3C, finely punctate 

3C, clustered punctate 

3C, clustered punctate 

3C, punctate 

3C, punctate 

3C, deeply punctatestriate 

3C, columnar-pyramidal 

3C, punctate-striate 

3C, prominently punctate 

3C, microreticulate 

3C, punctate-striate 

3C, punctate 

3C, punctate 

3C, perforate and 

spinulose 

3C, perforate and 

spinulose 

3C(?), perforate and 

spinulose 

3C, finely punctate 

3C, clustered punctate 

3C, finely punctate3C, clustered punctate 

3C, clustered punctate 

3C, punctate-striate 

3C, deeply punctate 

3C, punctate 

3C, punctate-striate 

3C, punctate-striate 

3C, punctate-striate 

3C, punctate 

PP, spinulose 

3C, punctate 

3C, punctate 

3C, punctate 

3C, punctate 

3C, finely punctatestriate 

striate 

3C, punctate-striate 

3C, sparsely spinulose 

3C, sparsely spinulose 

3C, sparsely spinulose 

and punctate 

and punctate 

and punctate 

142 

98, 99, 143 

94, 95, 144 

145 

84 

85, 146 

110, 111, 147 

136 

90, 91, 148 

149 

87, 150 

92, 93, 151 

96, 97, 152 

137 

88, 89 

102, 103, 153 

100, 101, 155 
20 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Species 

.M. polgbotryar Meisner.. ... M. tmifolia (HBK) Meisner. 

Neomi 1 lspaughia paniculata 

Nemacaulis denudata 

Oqgonun atriplicifolium 

(Donnell-Smith) Blake. ... 

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

Martelli ................. 

3. dregeanm Meisner....... 

3. zeyheri Sonder .......... Oxyyria digynia (L.) Hill.. . 

Oxytheca dendroidea 

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

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

0. parishii Parry.. ........ 0. trilobata A. Gray ....... 

Persicaria coccinea Greene. 

Podopterus mexicanus Kunth. 

Polygonel la fimbriata 

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

P. polygma (Ventenat) 

Engelmann L A. Gray...... 

Polygonun acwninatwn Kunth. 

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

P. amphibiiu? L. ........... P. bistorta L. ............ 

P. capitatum D. Don ........ P. ailinode Michaux ........ 

P. convolvulus L. ......... P. cristatm Engelmann 

L Gray ................... P. cuspidatun Siebold L 

Zuccarini ................ P. cyanandrim Diels ....... P. dielsii Leville (?). .... P. dumetonvn L. ........... P. forrestii Diels ......... P. rnuZtiflorwn Thunberg.. , . P. orientale L. ........... P. vaccinifolium Wallich.. . P. virginianun L. ......... P. viviparm L. ........... 

Pteropym aucheri 

Jaubert 6 Spach. ......... Pterostegia drymariaides 

Fischer & Meyer .......... A?hem delavayi Franchet.... 

Fwnex acetosa L. .......... 

R. acetosella L. .......... 

TABLE +Continued 

Collector 

Pritzel 399 

Balls 5804 

Edwards P 675 

Cleveland s.n. 

Piemeisel & Kephart 56 

Bayliss 4515 

Galpin 13075 

Neilson 902 

Eastwood L Howell 7353 

Wagenknecht 18118 

Pringle s.n. 

Walther 14379 

S. L W. Parish 993 

Abrams 3807 

Eastwood 1427 

Langlassi 922 

Ward L Smith 2236 

Bright 6309 

Irwin & Souza 9565 

Staunton et al. 1328 

Braun 29 

Cantlon L Gillis 57-439 

Cantlon & Malcolm 580085 

Calder & Billard 3833 

Tsiang 4503 

Hardin 722 

Horr L McGregor E495 

Graham 25 

Makino 13368 

Rock 5935 

Henry 9379 

Makino 13491 

Rock 9697 

Makino 15020 

Johnson s.n. 

Koelz 976 

Standley 8342 

Alm s.n. 

Rechinger 27660 

Gillespie 5495 

Rock 16256 

Greuter L Hainard 567 

Rouleau 67 

Location Pollen 

description 

Figure 

numbers 

Au s t r a1 ia 

Ecuador 

Honduras 

California 

Kenya 

S. Africa 

S. Africa 

Canada 

Nevada 

Chile 

California 

California 

California 

California 

California 

Mexico 

Florida 

S. Carolina 

Brazil 

Nepal 

Indiana 

Alaska 

Alaska 

Canada 

China 

N. Carolina 

Kansas 

Texas 

Japan 

China 

China 

Japan 

China 

Japan 

New York 

India 

Missouri 

Sweden 

Pakistan 

Arizona 

China 

Switzerland 

Canada 

3C, punctate-striate 

3C, punctate-striate 

3C, punctate-striate 

3C, finely punctate 

3C, punctate and 

microreticulate 

3C, microreticulate 

3C, microreticulate 

3 colpate-very thin 

walled 

3C, finely punctatestriate 

3C, finely punctatestriate 

3C, finely punctatestriate 

3C, finely punctatestriate 

3C, punctate 

3C, finely punctatestriate 

PC, finely reticulate 

3C, microreticulate 

3C, finely punctate and 

reticulate 

3C, microreticulate 

PP, reticulate 

3C, finely punctate and 

spinulose 

PC, finely reticulate 

3C, finely punctate and 

spinulose 

3C, finely punctate and 

spinulose 

3C, finely punctate and 

spinulose 

3C, reticulate 

3C, finely punctate3C, psilate and spinulose 

3'2, psilate and spinulose 

3C, deeply punctate 

PP, spinulose 

3C, coarsely reticulate 

3C, psilate and spinulose 

PC, spinulose 

3C, deeply punctate 

PP, reticulate 

3C, punctate 

PC, finely reticulate 

3-4C, very finely 

striate 

sp inulo se 

3C, punctate-striate 

3C, punctate-striate 

312, punctate 

3(-4)C, perforate and 

spinulose 

3(-4)C, perforate and 

sp inu lo se 

156 

104, 105, 154 

82, 157 

83, 158 

112 

132, 160 

130, 161 

131 

113 

124 

120, 121, 162, 163 

128, 129, 164 

122, 123 

140, 141, 165 

134, 135 

116, 117, 166 

133, 167 

114, 115, 159 

126, 127, 168 

118, 119, 169 

125 

138, 139, 170 
NUMBER 37 

TABLE +Continued 

21 

Species 

R. aquaticus L. ........... 

R. crispus L. ............. 

R. scutatus L. ............ 

Ruprechtia lmiflora 

Meisner .................. 

R. pallida Standley ........ R. ramiflora (Jacquin) 

Meyer .................... 

Symeria paniculata 

Bentham .................. Triplaris americana L. .... 

Collector Location Pollen Figure 

description numbers 

Heikkinen s.n. Finland 3(-4)C, perforate and 

Standley 18083 Montana 3C, perforate and 

Pfister 1158/2 France 3C, perforate and 

spinulose 

s p i n u l o se 

spinulose 

Hatschbach 13161 Brazil 

Neman 17 Mexico 

Haught 3914 

Ducke 650 MO 

Pittier 12204 

3C, deeply punctate- 106, 171 

3C, perforate-rugulose 107, 172 

striate 

Colombia 3C, deeply punctate- 108, 109 

striate 

Brazil 3c, 2 psilate 

Venezuela 3C, punctatestriate 

86, 173 
22 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

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

Species 

Anagallis iinifolia L. ..... Androsace septentrionalis L. 

Apochoris pentapetala Duby.. 

Ardisandra wittsteinii R. 

Wagner .................... 

Centunciilus pentandrus R. 

Brown ..................... Coris monspeliensis L. ..... Cortusa natthioli L. ....... Cyclamen neapolitanm 

Tenore .................... 3ionysia nira Wendelbo ...... 

Dodecatheon alpinm 

(A. Gray) Greene... ....... Douglasia montuna A. Gray.. . Glaw maritima L. .......... 

Hottonia palustris L. ...... 

Lysimachia ciliata L. ...... 

L. hybrida Michaux .......... L. terrestris (L.) Britton, 

Stern & Poggenberg ........ Lysimachiopsis hillebrandii 

Heller .................... .Vambergia thyrsiflora 

(L.) Reichenbach .......... 

Omph~logr~~n~a vincaeflora 

Franchet.... .............. Prirnula farinosa L. ........ 

P. incisa Franchet. ......... P. officinalis (L.) Jacquin. 

?. veris L. ................ 

P. vulgaris Hudson....... 

Samolus ebracteatus Kunth... 

SoZdaneZZa alpina L. ....... 

S. austriaca Vierhapper ..... 

Stimpsonia chmaedryoides C. 

Wright .................... Trientalis americana 

(Persoon) Pursh ........... 

Collector Location 

Reverchon 388 

Tarleton 8 

Zimmermann 257 

Greenway & Kanuri 15044 

MO 

Standley 12941 

Tidestrom 13554 

Koelz 4994 

Stefani s.n. 

Mandaville 3600 

Maguire & Holmgren 26449 

Thompson 13766 

Haakana s.n. 

Redfield s.n. 

Cernohorsky & Kraj ina 

Cronquist & Jones 5956 

Leonard & Killip 871 

Mac Kenzie 2720 

Degener 21278 NO 

Andersen s.n. 

286 

Rock 3573 

St. John 90675 

Rock 17474 

Ludera s.n. 

Charpin et al. s.n. 

Skvortsoo s.n. 

Pfister s.n. 

Francis 8a 

Treffer, 1362 in Flora 

Exsiccata AustroHungar ica 

Keller s.n. 

Ying Hu 10001 

Palmer 1370 

Spain 

Alaska 

China 

Kenya 

Florida 

France 

India 

Corsica 

Oman 

Oregon 

Idaho 

Finland 

Maine 

Czech0 Slovakia 

Idaho 

Maryland 

Yew Jersey 

Hawaii 

Norway 

China 

Quebec 

China 

Poland 

France 

USSR 

Switzerland 

Florida 

Austria 

Austria 

Hang Kong 

Canada 

Pollen 

description 

Figure 

numbers 

3C, prominently punctate 

3C, punctate 

3C, microreticulate 

3C, punctate 

183, 192 

3C, microreticulate 

3C, sculptured & psilate 

3-syncolporate, granular 

3C, punctate 

6-8 zonocolpate, deeply 

punctate 

3C, punctate-granular 

3C, punctate 

3C, punctate-striate 

3C, irregularly punctate3C, microreticulate 

3C, psilate-finely 

3C, finely punctate 

3C, microreticulate 

4C, punctate-psilate 

3C, irregularly 

striate 

punctate 

microreticulate 

3C, microreticulate 

4-syncolporate, 

microreticulate 

3C, microreticulate 

6-8 zonocolpate, 

microreticulate 

6-10 zonocolpate, 

microreticulate 

6-8 zonocolpate, 

microreticulate 

6-8 zonocolpate, 

microreticulate (many 

abnormal grains) 

3C, punctate 

3-colpate (syncolpate?), 

scabrate-spinulose 

3-colpate (syncolpate?), 

scabrate-spinulose 

312, finely reticulate 

3C, punctate 

174, 193 

188 

184 

180, 181 

182 

190, 194 

185 

175, 195 

177 

176, 196 

191, 197 

198 

186, 187 

200 

189 

178, 179, 199 
FIGURES 1-6.4entrospermae pollen, SEM: 1, Aizoaceae, Mesembryanthemum variabile Haworth, 

equatorial view, x 4100; 2, Molluginaceac, Limeum viscosum Fenzl, equatorial view, x 3900 3, 

Basellaceae, Anredera scandens Moquin, x 3550; 4, Caryophyllaceae, Gymnocarpos fruticosum 

Pcrsoon, X 3350; 5, Chenopodiaceae, Chenopodium ambrosioides L., x 3600; 6, Portulacaceae, 

Naiocrene pawifoh (Moquin) Rydberg, x 1900. (Photo reduced to 81v2 percent.) 
FIGURES 7-12.-centrospermae pollen, SEM: 7, Molluginaceae, Limeum uiscosum, ektexine surface, X 7500; 8, Aizoceae, Mesembryanthenium variabile, ektexine surface, x 7500; 9, Aizoaceae, 

Geocarpon minimum Mackenzie, ektexine surface, x 7500; 10, I’ortulacaceae, Naionene paruifolia, X 7500; 11, Carlaceae, Borricuctur tem~iserpens (Rauh & Backeberg) Kimnach, ektexine 

surface, X 7500; 12, Cactaceae, Ilarnatocuclu5 seti\pinus Britton & Rose, ektexine surface, x 7500. 

(Photo reduced to 7GS percent.) 

n 

0 

z 

4 

W 

c: 

0 

z 

z 

2 

m 

4 

0 
FICURFS 13-18.-Centrospermae pollen, SEM: 13, Amaranthaceae, Psilotrichurn arnpZum Sussenguth. X 4000; 14, Caryophyllaceae. Siphonychia arnericana (Nuttall) Torrey & A. Gray, x 4600; 

15, Caryophyllaceae, Herniaria @bra L., x 6000; 16, Caryophyllaceae, Cardionema ramosissirna 

A. Nelson & Macbride, x 5500; 17, Cactaceae, Opuntia Zindheimeri Engelmann, x 840; 18, 

Nyctaginaceae, Abronia angustifolia Greene, equatorial view, x 2870. (Photo reduced to 81 

percent .) 

FICURE~ 19-23.4entrospermae pollen, TEM: 19, Aizocaceae, Mesembryanthemum variabile, 

the foot layer is extremely thickened and is underlain by a thin but consistent endexine; the 

break in the endexine at extreme right may be a ruptured area; 20, Amaranthaceae, Psilotrichurn amplum, an endexine is entirely absent; the thin-layered aperture contains a large pointed 

fragment of ektexine; 21, Basellaceae, Anredera scandens, a wedge of endexine is prominent 

beneath the colpus, but in the mesocolpial regions it is less distinct and has an electron density 

similar to that of the foot layer; however, the endexine is fibrillar and the inset (arrows) shows 

the fibrils to at least superficially resemble unit membranes; 22, Cactaceae, Opuntia lindheimeri, 

section is through one of the large apertures (Figure 17) with numerous flecks of ektexine; 23, 

Caryophyllaceae, Gyrnnocarpos fruticosurn, it is difficult to distinguish a foot layer or endexine; 

the inset suggests that there are two layers with an irregular separation. (Scales equal 1 em.) 

u W 

FIGURES 24-28.4entrospermae pollen, TEM: 24, Caryophyllaceae, Herniuriu glubra, an irregular 

but notablc endexine is characteristic of this taxon; the absence of spines or pcrforations in a 

section is consistent with the surface depicted by SEM (Figure 15); 25, Chenopodiaceae, 

Chenopodium nmbrorioides, the equatorial section and that contained in the inset (polar section) 

indicate a thin foot-layer, a highly irregular and granular endexine, and “internal foramina-like” 

openings in the thickened tectum; 26, Molluginaceae, Lirneuni uiscosum, micrograph includes 

sections through mesocolpial (upper) and apcrtnral (lower) regions; aperture has numerous 

flecks of ektexine; 27, Nyctaginaceae, Abronia ungustifoliu, the irr~gular structures are due to 

an oblique section through a coarsely reticulate ektcxine; an endexine is not evident in this 

taxon; 28, Portulacaceae, Naiocrene parvifolia, the endexine is evident in the apcrturc region 

while in the mesocolpial regions it is either reduced or absent. (Scales equal 1 am.) 
FIGURFS 2%34.--Plumbaginaceae pollen, SEM. Cerutostigma gri@thii C. B. Clarke: 29, equatorial 

view, X 1250; 30, ektexinc surface, x 7500 (29, 30, Rock 11578, rounded verrucac Plumbago 

Iorm); 31, equatorial view, x 1450; 32, ektexine surface, x 7500 (31, 32, Rock 6577, pointed 

verrucac Plumbago form). Plumbago rosen L.: 33, equatorial view, x 1500; 34, ektexine surface. 

X 7500 (33, 34, pointed verrucae Plumbago form). Photo reduced to 74 percent.) 

4 

0 
FIGURFS 35-40.-Plumbaginaccae pollen, SEM. PZurnbago europaea L.: 35, polar view, X 1700; 36, 

ektexirie surface, x 7500 (35, 36, Sporiks s.n. GH, rounded verrucac Plurnbago form); 37, polar 

view, X 1700; 38, ektexine surface, x 7500 (37, 38, Rnine 5.n. GH, pointed verrucac Plumbago 

form). Plumbago auriculala Lamarck: 39, polar view, x 1250; 40, ektexine surface, x 7500 (39, 

40, Bayliss 1187 GH, rounded verrucae Plumbago form). (Photo reduced to 7694 percent.) 
FlGURFS 414G.-Plnmhaginaceae pollen, SEM. Liinoniunl uiciosoi Pau: 41, equatoiial view, X 1250; 

42, ektexine surface, x 3000 {coarsely reticulate Armeria form). Arrneria rnaritirna {Miller) 

Willdenow: 43, equatorial view, x 1150; 44, ektexine surface, x 5000 (43, 44, Suenson s.n., finely 

reticulate Armeria form); 45, polar view, x 1250; 46, cktexine surface, x 3000 (45, 46, Mathiesen 

Q Pedersen s.n., coarsely reticulate Armeria form). (Photo reduced to 77 percent.) 

4 

0 
FIGURE 47-52.-Plumbaginaceae pollen, SEM. Statice tenella Turczaninow: 47, polar view, X 1500; 

48, ektexine surface, x 5000 (47, 48, Chaney 517, coarsely reticulate Armeria form); 49, polar 

view, x 1500; 50, ektexine surface, x 7500 (49, 50, Chaney 517a, finely reticulate Armeria form). 

Lirnonium vulgare Miller: 51, ektexine surface, x 5000 (Lawalree 3820 MO, coarsely reticulate 

Armeria form); 52, ektexine surface, x 7500 (Holm-Nielsen s.n. MO, finely reticulate Armeria 

form). (Photo reduced to 79 percent.) 
FIGURES 53-58.-Plumbaginaceae pollen, SEM. Goniolitnon collinuin (Grisebach) Boissier: 53, 

polar view, X 1600; 54, ektexine surface, x 5000 (coarsely reticulate Armeria form). Statice 

sinuatu L.: 55, polar view, x 1500; 56, ektexine surface, x 4000 (55, 5G, Vuccari 495, coarsely 

reticulate Armeria form); 57, polar view, x 1550; 58, ektexine surface, x 7500 (57, 58, 

Maire 6. WilczeR 1048, finely reticulate Armeria form). (Photo reduced to 76 percent.) 

n 

0 

z 

4 

0 
2 

c! 

W U 

FIGURES 5!?-64.-Plumbaginaceae pollen, SEM. Ceratostigma willmottianurn Stapf 59, X 1150; 

60, ektexine surface, x 5000 (59, 60, Wilson 1373, pantocolpate Plumbago form); 61, polar view, 

X 1300; 62, ektexine surface, x 5000 (61, 62, Kingdon-Ward I2530 MO, 3-colpate Plumbago 

€ortn). Dyerofihyturn africanurn (Lamarck) 0, Kuntze: 63, polar view, x 2000; 64, ektexine surface, x 5000 (Werderrnann 6 Oberdieck 594, pointed? verrucae Plumbago form). (Photo reduced 

to 79% percent.) 

FICURE~ 65-68.-Plumbaginaceae pollen, TEM. Acantholimon olivieri Jaubert & Spach: 65, 

oblique section in aperture region; note thin but distinct endexine and random internal 

foramina (small holes) in the ektexine; 66, oblique section emphasizing thin endexine (arrow) 

and numerous internal foramina (coarsely reticulate Armeria form). Arrneria maritirna: 67, a 

line network of sporopollinen surrounds the muri and the inset, a tangential section, clarified 

this interpretation and correlates with images obtained from SEM (Figures 44 and 46); arrows 

point to internal foramina partially filled with electron dense material (Mathiesen dr Pedersen 

s.n., coarsely reticulate Armeria form); 68, in this section of an aperture the endexine is evident 

although highly fragmented; note that the outer surface of the muri appears somewhat ridged 

due to an off-center cut through the smaller spines (Svenson s.n., finely reticulate Armeria 

form). (Scales equal 1 am.) 

FIGURES 6%74.-Plumbaginaceae pollen, TEM. Cerutostigrnu grifithii: 69, the section is somewhat oblique and therefore the two levels of columellae appear incomplete; note that the 

terminal portions of the incompletely depicted outer collumellae consist of fine bristle-like 

structures; the endexine is absent in the mesocolpial areas (Rock 11578, rounded verrucae 

Plumbago form); 70, morphology similar to Figure 69 except that the terminal portion of outer 

columellae appears more massive (Rock 6577, pointed verrucae Plumbago form). C. wihottianurn: 71, note thin, plate-like endexine (arrow) in aperture region (Wilson 1373, pantocolpate 

Plumbago form). Dyerophyturn ofricanum: 72, radial section illustrates characteristics of inner 

columellae, smaller in diameter but almost equal in length to the outer columellae (Werdermann & Oberdierk 591, pointecl? verriicae Plumbago form). Goniolirnon collinurn: 73, in contrast 

to Figures 69-72, all one form or the other of the Plumhago type, this section illustrates the 

single layer of columellae characteristic of the Armeria type; the foot layer has perforations 

but at the present time they are not considered to be internal foramina; arrow points to 

endexine in the region of the aperture; the granular material surrounding the outer part of the 

exine appears to be acetolysis resistant and is thought to be a component of the anther rather 

than the pollen wall (coarsely reticulate Armeria form). Lirnoniactrurn guyonionurn Durieu de 

Maisonueuve: 74, arrows point to endexirie in regions of apertures (finely reticulate Armeria 

form). (Scales equal 1 @m.) 

FIGUWS 75-81.-Plumbaginaceae pollen, TEM. Limonium viciosoi: 75, arrows indicate internal 

foramina in ektexine; the lower part of the foot layer is somewhat lamellar and may be a thin 

endexine (coarsely reticulate Armeria form). Plumbago scandens L.: 76, oblique section (rounded? 

verrucae Plumbago form). Statice sinuata: 77, very oblique section; 78, section -I- tangential 

(77, 78, Maire d. Wilcxek 1048, finely reticulate Armeria form); 79, section +- tangential; 80, note 

fragmented but visible endexine (79, 80, Varcari 495, coarsely rcticu!ate Armeria form). Statice 

tenella: 81, oblique section, fragmented area on right indicates aperture region (Chaney 517, 

coarsely reticulate Armeria form). (Scales equal 1 Gm.) 

6 
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 Moeuch, equatorial 

view, X 1600 (Photo reduced to 77v2 percent.) 
d 

E 

P 

W u 

F~CURES 88-93.-Polygonaceae pollen, SEM: 88, Mucronea californica Bentham, equatorial view, 

X 2525; 89, ektexine surface, x 7500; 90, Eriogonum marifolium Torrey k A. Gray, equatorial 

view, X 3250; 91, ektexine surface, x 7500; 92, Gymnopodiunz antigonoides (Robinson?) Blake, 

equatorial view, x 2425 (Goldman 743); 93, ektexine surface, x 7500 (Goldman 743). (Photo 

reduced to 76 percent.) 
FIGURES 9499.-Polygonaceae pollen, SEM: 94, Calligonum comosum L’Heritier, equatorial view, 

X 2500; 95, ektexine surface, X 7500; 96, Hurfordia mucropleru (Bentham) Greenc & Parry, 

equatorial view, x 3050; 97, cktexine surface, x 7500; 98, Atraphaxis buxifolia Jaubert & 

Spach, equatorial view, x 2800; 99, ektcxine surface, x 7500. (Photo reduced to 84 percent.) 

4 

0 
FICURFS 100-105.-Polygonaceae pollen, SEM: 100, Muehlenbeckiu cunningkamii Mueller, equatorial 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 83% percent.) 
FIGURES lOC%111.-Polygonaceae pollen, SEM: 106, Ruprechtia laxiflora Meisner, equatorial view, 

X 3400; 107, R. pullida Standlcy, equatorial view, x 2500; 108, R. romiflora Uacquin) Meyer, 

equatorial view, X 2700; 109, ektexine surface, x 5000; 110, Coccoloba cordata Chamisso, equatorial view, X 2520; 111, ektexine surface, x 7500. (Photo reduced to 79 percent,) 

+l 

0 
FIGURES 112-117.-Polygonaceae pollen, SEM: 112, Persicaria coccinea Greene, X 1575; 113, 

Polygonurn acuminaturn Kunth, x 1575; 114, P. orientale 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.) 
FIGURES 118-123.-Polygonaceae pollen, SEM: 118, Polygonurn virginianurn I>., x 2000; 119, 

ektexine surface, x 5000; 120, P. arnphibiurn L., X 1400; 121, ektexiue surface, colpus lower 

right, X 9000; 122, P. capilutum D. Don, polar view, x 2500; 125, ektcxiue surface, x 7500. 

(Pholu reduced to 77v2 pcrceiit.) 

4 

0 
FIGURES 124-129.-Polygonaceae pollen, SEM. Polygonurn a@ne D. Don: 124, equatorial view, 

X 2200. p. viviparurn L.: 125, slightly oblique equatorial view, x 1700. P. vaccinifoliurn 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, Cantlon 6. Malcolm 580085). (Photo reduced 

to 83 percent.) 
FIGURES 130-135.-Polygonaceae pollen, SEM: 130, Polygonella fimbriata (Elliot) Horton, mesocolpial view, X 3400; 131, P. polyguma (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 Dick, x 3600; 135, ektexine surface, x 8500. (Photo reduced to 74 percent.) 

+I 

0 
FIGURES 136--14l.-Polygonaceae pollen, SEM: 136, Emex australis Steinheil, equatorial view, 

X 2990 (SeydeZ2975); 137, Lastarriaea chilensis Remy, mesocolpial view, x 3400 (Worth Q Morrison 

16338 UC); 138, Rumex acetosa L., equatorial view, x 4400; 139, ektexine surface, x 10,000; 140, 

Polygonurn conuol~lus L., mesocolpial view, x 3050; 141, ektexine surface, x 7500. (Photo 

reduced to 78y2 percent.) 

FIGWRFS 142-149.-Polygonaceae pollen, TEM: 142, Antigonon guaternalense Meisner, the 

endexine is narrow and plate-like; 143, Atraphaxis buxifolia, section somewhat oblique but 

endexine is thick and partially lamellar; 144, Calligonurn cornosurn, the tectum and columellae 

are well developed but the foot layer is greatly reduced; 145, Choriranthe breweri S. Watson, 

the tectum is thin and the lower part highly globular; the foot layer is rednced (or absent?) and 

the endcxine narrow but uniform; 146, C. paniculata, this section is very similar to that of 

Figure 145, but there is a very thin foot layer and endexine; 147, Coccoloba cordata, section 

somewhat oblique; foot layer is extremely thick while the columellae and tectum are of smaller 

dimensions: extending from the tectum are large, irre\gularly shaped components; a vestigial 

endexine is suggested but additional work is necessary for confirmation: 148, Eriogonurn rnarifoliurn, foot layer is very reduced as in Figures 144 and 145, but the endexine appears massive 

due largely to section through aperture: 149, E. parishii, S. Watson, very similar to Figure 145. 

(Scales equal 1 pm.) 

FIGURES 150-156.-Polygonaceae pollen, TEM: 150, Fagopyrurn esculenturn, the section is 

similar to Antigonon guatemalense (Figure 142) in the thickened tectum with funnel-like 

depressions (Figure 87), reduced foot layer, and thin but recognizable endexine; 151, Gymnopodium nntigonoides, the endexine is not clearly evident in this somewhat oblique view, but, 

the inset of a section including the colpus reveals uniform endexine (Goldman 743); 152, 

Harfordia macroptera, section through the colpus with prominent endexine; the lower surface 

of the tectum is irregular (Figures 145, 146, 149, 157, 158); 153, Muehlenbeckia chilensis, 

except for the aperture regions the endexine is highly reduced; 154, M. tarnnifolia, the endexine 

appears to be more uniform than in Figure 153; 155, M. cunninghamii, the tectum is thick 

with few perforations and the highly lamellar endcxine and foot layer are approximately equal 

in thickness (Aston 984); 156, M. polybotryar Meisner, section somewhat oblique; the foot layer 

reduced and endexine prominent. (Scales equal 1 am.) 

FIGURES 157-165.-Polygonaceae pollen, TEM: 157, Nemacaulis denudata, the section illustrates 

several characteristics of the punctate type, i.e., tectum with granular lower snrface, very 

reduced foot layer, and an endexine which at least in the region of the colpus is very prominent; 

158, Oxytheca trilobata, same as 157, the double arrows indicate the extremely thin foot layer, 

single arrows indicate acctolysis resistant material, which can be readily mistaken for exine 

layering; 159, Polygonurn orientale, long columellae fused at the apex to form the muri alternate 

with smaller free columellac in the lumina (Figures 114, 115); a thin endexine can be recognized 

in the aperture region to the left; 160, Podopterus mexicanus, an endexine cannot he recognized 

but the lower part of the foot layer appcars to consist of a thin, less electron-dense zone; 161, 

Polygonella firnhriata, the section reveals a well-developed foot layer and thickened endexine, 

an unusual combination in the family; 162, Polygonurn arnphibium, section at right angle 

through muri; 163, P. arnphibium, section parallel through murus; 164, P. bistorta, section, 

which includes entire pollen grain, is oblique; the densely packed columellae are rare and 

limited to Bistorta complex (Cantlon Q Malcolm 580085); 165, P. conuolvulus, section through 

polar region where tectum is psilate; this taxon and Polygonella fimbriata (Figure 161) are the 

only two in which both the foot layer and endexine are well developed; this supports their 

parallel exomorphology observed in SEM (Figures 130, 140, 141) and discussed in text. (Scales 

equal 1 am.) 

FIGURES 166-173-Polygonaceae pollen, TEM: 166, Polygonum dielsii, section 5 oblique, solid 

tectum indicates section is through muri; 167, P. jorrestii, the most distinctive member of the 

genus in TEILI; the endexine is thin, the ektexine contains abundant holes or internal foramina, 

and the tectum is composed of very large, solid spines on an irregular or granular base; 168, 

P. vaccinijolium, very similar to Figure 164. densely packed columellae and also a member of 

the Bistorta complex; 169, P. virginianum, the discontinuous tectum reflects reticulate sculpture; 

170, Rumex acetosa, the foot layer is either greatly reduced or absent; endexine thickness and 

rupture indicate colpus; 171, Ruprechtia laxiflora, foot layer is consistent and well developed; 

arrows point to what appears to be a lamellar endexine in colpial region; 172, R. pdida, 

similar to Figure 171 but endexine somewhat better developed; the accumulation of apparent 

foot layer material in colpus region is unusual; 173, Triplaris americana, similar to Figure 151, 

thickened tectum, thin foot layer and endexine. (Scales equal 1 @m.) 
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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 thyrsiflora 

(I..) Reichenbach, mesocolpial view, x 4620; 177, I.ysimachia terrcstris (L.) Britton, Stern, & 

Poggenberg, mcsocolpial view, x 4400; 178, Stimpsonia chamaedryoides C. Wright, equatorial 

view, X 3200; 179, ektexine surface, x 7500. (Photo reduced to 72 percent.) 

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FIGURES 180-185.-Primulaceae pollen, SEM: 180, Dodecatheon alpinum (A. Gray) Greene, equatorial view, x 4000; 181, ektexine surface, x 7500; 182, Douglasia montana A. Gray, mesocolpial view, x 4250; 183, AnagaZZis Zinifolia L., equatorial view, x 3900; 1% Cyclamen 

neapolitanum Tenore, mesocolpial view, x 6500; 185, Hottonia palwtris L., equatorial view, 

x 5250. (Photo reduced to 78 percent.) 
FIGURES 18&191.-Primulaceae pollen, SEM: 186, Prirnula veris L., oblique view, x 3200; 187, 

ektexinc 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, Omphalograrnma vincaeliora 

Frauchet, equatorial vicw, x 3550. (Photo redut crl to 76 perccrit.) 

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FIGURES 192-195.-Primulaceae pollen, TEM: 192, Anagallis linifolia, a uniform endexine is notable as is the prominent foot layer; 193, Cork 

rnonsfieliensis, the section includes the entire grain and reflects the variation of the sculpture as seen in SEM (Figure 174); the endexine is greatly 

thickened in the region of the apertures; 194, GZaux rnaritirna, the foot 

layer and endexine are both prominent and approximately equal in thickness (Redfield s.n.); 195, Lysimachia hybrida, the oblique section enhances 

the thickness of the foot layer and endexine. (Scales equal 1 Gm.) 
FIGURES 196-200.-Prirnulaceae pollen, TEM: 196, Nuumbergiu thrysifloru, 

thickened tectum is supported by reduced columellae on a well-defined 

foot layer and prominent endexine; 197, Omphalogrammu uincuelloru, similar to Figure 196 except for more massive columellae; 198. Primula 

oficinulis (L.) Jacquin, in contrast to all other taxa of Primulaceae examined in TEM, those in the genus Prirnulu are without a foot layer; 

however in light microscopy and SEM, the samples appeared at least partially sterile; 199, Stimpsoniu chamaedryoides, oblique section of entire 

grain illustratirig consistcnt foot layer, and an endexine that is massive in 

aperture regions and thin but recognizable in the mesocolpial areas; 200, 

Primulu ueris L., see Figure legend 198 (Skuortsoo s.n.). (Scales equal 1 Gm.) 

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