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

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Pollen Morphology and the Relationship of the Plumbaginaceae, Polygonaceae, and Prirnulaceae to the Order Centrospermae 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 Relation- ship of the Plumbaginaceae, Polygonaceae, and Primulaceae to the Order Centro- spermae. 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 01der Centrospei mae by 1 ari- ous 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 iast 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 com- mon to the three families further suggests that they are not closely related to each other. OFFICIALPVBLICATIOS DATE is handstamped in a limited number of initial copies and is recorded in the Institution’s annual report, Smithsoninn Year. SERIFSCOVER 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 to the order Centrosperrnae. (Smithsonian contributions to botany ; no. 37) 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 Gyrostemonaceae as part of the Phytolaccaceae, and Takhtajan separates it as a distinct family. Takh- The Centrospermae (Caryophyllales) represents tajan recognizes the Tetragoniaceae as a separate one of the most controversial orders in the angio- family while Cronquist includes it in the Aizoaceae. sperms. This group of at least 10 families, which in Dysphania is treated as a member of the Chenopo- the past has been cited as one of the few natural diaceae by both authors; and finally Takhtajan also orders based mostly on morphological characteris- places the Hectorellaceae and the Bataceae in this tics, has unique N-containing pigments, the beta- order. lains, and a distinctive structure in the sieve-tube Thorne (1968) in a synopsis of angiosperm classi- plastids. Both Cronquist (1968) and Takhtajan fication has placed the betalain families in a sub- (1969) unite the betalain families, Aizoaceae, Ama- order, Chenopodiinae, one of two constituting the ranthaceae, Basellaceae, Cactaceae, Chenopodiaceae, order Chenopodiales. He recognized the Gyrostemo- Didiereaceae, Nyctaginaceae, Phytolaccaceae, and naceae and the Halophytaceae as distinct families Portulacaceae, as well as two anthocyanin families, Caryophyllaceae and Molluginaceae, in the order but treated the Molluginaceae as a subfamily in the Aizoaceae, the Achatocarpaceae as a subfamily in Caryophyllales ( k Centrospermae). Both authors include Halophytum ameghinoi Spegazzini (a beta- the Phytolaccaceae, and Dysphania as a member of the Chenopodiaceae. The other suborder, Caryo- lain taxon) and the Gyrostemonaceae in this order: phyllinae, consists of two families, the Caryophylla- Cronquist (1968) treats Halophytum as a monotypic ceae and the Polygonaceae. The next order, Bati- genus in the Chenopodiaceae while Takhtajan (1969) gives it family status; Cronquist regards the dales, 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 Micro- Mabry, and Turner and Behnke biology, University of Oklahoma, Norman, Oklahoma 73019. and Turner (1971) have treated the betalain fami- 1 2 SMITHSONIAN CONTRIBUTIONS TO BOTANY lies as a distinct group, separate from, yet closely from palynology, pigmentation, and plastid struc- allied to, the two anthocyanin families, Caryo- ture indicates that these three families are not phyllaceae and Molluginaceae. closely related to the Centrospermae. The relation- Evidence from ultrastructural research on sieve- ship of the Achatocarpaceae, a family unknown tube plastids (Behnke and Turner, 1971; Behnke, with regard to pigmentation, to the Centrospermae 1976) and pollen morphology (Nowicke, 1975; is more obscure-the pollen morphology does not, Skvarla and Nowicke, 1976) has reinforced the close in Nowicke’s opinion (1975) support a close tie, but tie between the betalain families and the Caryo- on the other hand the two genera that comprise the phyllaceae and the Molluginaceae. In an investiga- family, Achatocarpus and Phaulothamnus, do have tion of these plastids (colorless leucoplasts found in the P-type of plastid (Behnke, 1976). the sieve-tubes) of the above-mentioned families The Gyrostemonaceae have in fact been the sub- (Behnke and Turner, 1971) and the Magnoliidae ject of a multidisciplinary study (Goldblatt, et al., and Ranunculidae (Behnke, 1971), two main types 1976) in which all lines of evidence, including cyto- were found: plastids accumulating only starch and logical, argue strongly against any relationship of designated as the S-type; and plastids accumulating this family to the Centrospermae. at least some protein, variously deposited, and desig- Although there is almost universal agreement on nated as the P-type. The betalain families and the treating the betalain families as a closely related Caryophyllaceae and the Molluginaceae all had the group, the precise definition of the order Centro- P-type plastid in which proteinaceous filaments spermae, i.e., to include or exclude the Caryo- formed a peripheral ring usually around a crystal- phyllaceae and Molluginaceae, may never be re- loid. IVithin the above-mentioned families, Behnke solved to everyone’s satisfaction. Of more interest (1976) was able to distinguish three groups based on in view of the accumulating data on the Centro- the crystalloid: globular, the most common; polyg- spermae (sensu lato) is the question of the deriva- onal, found in the Caryophyllaceae and in two tion of the families Plumbaginaceae, Polygonaceae, other genera, Limeum (Molluginaceae) and Stegno- and Primulaceae from this order. sperma (Phytolaccaceae); and crystalloid-free, hav- ing only the ring, found in two families, the Takhtajan (1969) considers the Polygonales, con- Amaranthaceae and the Chenopodiaceae. taining only the Polygonaceae, as near the Caryo- Examination of the pollen of 177 species (No- phyllales, especially the Portulacaceae and Basella- wicke, 1975) by light microscopy and SEM revealed ceae, and probably derived from the same stock as that 85% had a similar type of ektexine pattern or the Caryophyllales (Centrospermae). His views on sculpture, which was described as spinulose and the the Plumbaginaceae are similar: nearest to Portula- tectum perforate, the openings described as tubules caceae and Basellaceae and probably derived from or punctae. This was the predominant ektexine pat- the same stock as Caryophyllales. He does admit tern in every betalain family and in the Caryo- that the pollen morphology of the Plumbaginaceae phyllaceae and the Molluginaceae; in some families, is different-apparently from that of Portulacaceae i.e., the Phytolaccaceae and the Molluginaceae, this and Basellaceae. pattern was the only one found. Examination of A4ccording to Cronquist (1968: 185), “both the selected taxa by TEM (Skvarla and Nowicke, 1976) Polygonaceae and Plumbaginaceae are pretty clearly revealed that the predominant pollen-type in the related to the Caryophyllales. Both have a single, betalain families and in the Caryophyllaceae has a basal, bitegmic, crassinucellate ovule in a com- similar, sometimes
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