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Comparative Morpholog of the Lens on Legume

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Comparative Morpholog of the Lens on Legume firy!{^M/ , States rtment of Agriculture Comparative Morpholog Agricultural Research of the Lens on Legume Service Technical (Fabaoeae) Seeds, with Bulletin Number 1791 Emphasis on Species in Subfamilies Caesalpinioideae and l\/limosoideae in cooperation with Iowa State University Abstract Lersten, Neis R., Charles R. Gunn, and Curt L. The lenses of the Caesalpinioideae showed greater Brubaker. 1992. Comparative Morphology of the variability in surface appearance and in Lens on Legume (Fabaceae) Seeds, with Emphasis anatomical features than those of the on Species in Subfamilies Caesalpinioideae and Mimosoideae. Both subfamilies contain some Mimosoideae. U.S. Department of Agriculture species with pop-off lenses—that is, lenses whose Technical Bulletin No. 1791. 44 pp. epidermis (plus subepidermal tissue in one species) forms a loose blister that is easily dislodged upon The three subfamilies (Caesalpinioideae, disturbance. The caesalpinioid genus Bauhinia Mimosoideae, and Faboideae) of the Fabaceae showed a lenslike structure, which we termed a differ in certain morphological and anatomical pseudolens. features of their seeds. One of these features is alleged to be the presence of a lens on most The caesalpinioid and mimosoid lenses show more faboid seeds and the absence of a lens on variation in gross morphology than the faboid caesalpinioid and mimosoid seeds. In this report, lenses but, internally, are much simpler in terms however, the lens, a biconvex bulge along the of cell types and tissue layering. A consistent sagittal midline, is shown to also be a widespread feature of obvious lenses is the upward curving of but not ubiquitous feature of seeds of the the hilar vascular bundle to occupy a superficial Caesalpinioideae and Mimosoideae. to subepidermal position beneath the lens, beyond which the bundle typically dips downward again. A comprehensive review of the occurrence of the The tracheid bar (found in faboid seedcoats) does lens on seeds in all three subfamilies is presented. not occur in any of the caesalpinioid or mimosoid This review is followed by original scanning species that were studied. electron microscope observations of the surface and interior of seedcoats from 13 species of 13 KEYWORDS: anatomy, Faboideae, hilum, genera representing all 5 tribes of the Leguminosae, micropyle, Papilionoideae, pop-off Caesalpinioideae and from 16 species of 14 genera lens, pseudolens, raphe, seedcoat, strophiole, testa. representing all 5 tribes of the Mimosoideae. United States Department of Comparative Morphology Agriculture Agricultural of the Lens on Legume Research Service (Fabaceae) Seeds, with Technical Bulletin Number 1791 Emphasis on Species in Subfamilies Caesalpinioideae and Mimosoideae by Neis R. Lersten, Charles R. Gunn, and Curt L Brubaker Acknowledgments We acknowledge the support of curators and Botany and Mycology Laboratory, Beltsville, MD, directors of numerous herbaria who loaned us assisted in the preparation of the typescript. critically needed material. The typescript was Karen Parker, botanical illustrator, Bowie, MD, reviewed by Duane Isely, Botany Department, prepared the plates for publication and the cover Iowa State University, Ames, lA, and by Joseph illustration. Microscopy and ancillary procedures H. Kirkbride, Jr., and John H. Wiersema, both of were done in the Bessey Microscopy Facility, the Systematic Botany and Mycology Laboratory, Department of Botany, Iowa State University. Beltsville, MD. Carole A. Ritchie, Botanist, Laurel, MD, and Judith R. Purman, Systematic Contents Procedures 2 Caesalpinioideae Kunth 13 Lens in the Fabaceae Lindley 3 Amherstieae Bentham 14 Function of the lens 6 Caesalpinieae Bentham 15 Faboideae Kunth 7 Cassieae Bronn 15 Abreae (Wight & Arnott) Hutchinson 7 Cercideae Bronn 16 Aeschynomeneae (Bentham) Hutchinson 8 Detarieae A.P. de CandoUe 16 Cicereae Alefeld 8 Mimosoideae Kunth 17 Crotalarieae (Bentham) Hutchinson 8 Acacieae Bentham 19 Galegeae (Bronn) Torrey & Gray 8 Ingeae Bentham 19 Genisteae (Adanson) Bentham 9 Mimoseae Bentham 20 Indigofereae (Bentham) Rydberg 9 Mimozygantheae Burkart 21 Phaseoleae A.P. de CandoUe 10 Parkieae (Wight & Arnott) Bentham 21 Psoraleeae (Bentham) Rydberg 11 Literature cited 22 Robinieae (Bentham) Hutchinson 12 Scientific name index 25 Sophoreae Sprengel 12 Thermopsideae Yakovlev 12 Trifolieae (Bronn) Bentham 12 Vicieae (Adanson) A.P. de CandoUe 13 Trade names are used in this publication solely for the purpose of providing specific information. Mention of a trade name does not constitute a guarantee or warranty of the product by the U.S. Department of Agriculture or an endorsement by the Department over other products not mentioned. Copies of this publication may be purchased from the National Technical Information Service, 5285 Port Royal Road, Springfield, VA 22161. ARS has no additional copies for free distribution. January 1992 11 Comparative Morphology of the Lens on Legume (Fabaceae) Seeds, with Emphasis on Species in Subfamilies Caesalpinioideae and Mimosoideae Neis R. Lersten, Charles R. Gunn, and Curt L. Brubaker The Fabaceae Lindley (Leguminosae A.L. de (Suborder III). Taubert (1894) retained this Jussieu of authors, including Isely and Polhill tripartite division, treating the suborders as 1980) is the largest flowering plant family after subfamilies and moving the Swartzieae A.P. de the Asteraceae Dumortier and Orchidaceae A.L. Candolle (containing taxa with a curved de Jussieu. However, only the Poaceae Barnhart embryonic axis) from the Faboideae to the rivals the Fabaceae in agricultural importance. Caesalpinioideae Kunth. Corner (1976) recognized The past, present, and future value of the four subfamilies—the fourth being the Fabaceae (the legume family) has been Swartzioideae Kunth. The Swartzieae was returned documented by Skerman (1977), the National to the Faboideae by Cowan (1981); later, however. Academy of Sciences (1979), Summerfield and Cowan et al. (1989) treated the Swartzieae in the Bunting (1980), Duke (1981), and Isely (1982). Caesalpinioideae. Cowan et al. noted that "the Swartzieae is treated here where it has De CandoUe (1825), who prepared the earliest traditionally been placed, even though it is comprehensive taxonomic treatment of the probably more accurately assigned to the Fabaceae, divided the family into two suborders. Papilionoideae [=Faboideae], and in fact is so His simplistic one-character division separated the aligned in...[(Cowan 1981)]." Although El-Gazzar Curvembriae A.P. de Candolle (which has a and El-Fiki (1977) concluded that the bipartite curved embryonic axis) from the Rectembriae division of de Candolle is superior to the A.P. de Candolle (which has a straight embryonic tripartite division, the latter division, axis). The basis of the separation, namely the reaffirmed by Polhill and Raven (1981), has curvature of the embryonic axis, foreshadowed the remained stable. Some authors (e.g., Hutchinson idea that seed characters might be important in 1964, Cronquist 1981, and Cowan et al.) legume systematics. Although the curvature of the promoted the Fabaceae (Leguminosae) to an order, embryonic axis is no longer regarded as the best Fabales Bromhead, with three families: Mimosaceae character for a primary division of the family, it R. Brown, Caesalpiniaceae R. Brown, and Fabaceae has a role in protecting the radicle and may be (Papilionaceae Giseke). one of a combination of seed characters (especially hilar characters) that may be used to There are two generalizations concerning the separate the Faboideae Kunth from the other seedcoat (or testa) of the subfamilies of the subfamilies. Burkart (1952) used such a Fabaceae. The first is that the tracheid bar is a combination in the subfamily portion of his character of only the Faboideae (Lersten 1982). legume seed key. The second generalization is that the lens—a seedcoat structure of varied size and shape found Bentham (1865) used floral and vegetative along the hilum/micropyle midline and typically characters instead of seed characters (shown in separated from the micropyle by the hilum—is fig. 1) to create a third suborder by dividing the found only in faboid seeds and not in Rectembriae (which was based on a seed caesalpinioid and mimosoid seeds. Although both character) into the Caesalpinieae Bentham generalizations are based on scanty knowledge, (Suborder II) and the Mimoseae Bentham our new data support the first generalization and disprove the second. Neis R. Lersten is a professor of botany, Iowa State University, Ames, lA 50011; Charles R. Gunn is a research botanist. Systematic Botany & Mycology Laboratory, Agricultural Research Service, U.S. Department of Agriculture, Beltsville, MD 20705; Curt L. Brubaker is a graduate student, Iowa State University, Ames, lA. Procedures The purposes of this bulletin are to 1) survey the Seeds used in this study are deposited in the U.S. pertinent literature about the lens on seeds National Seed Herbarium, Beltsville, MD. The representing the three legume subfamilies, and 2) scientific names and their authors used in this present original scanning electron microscope bulletin were checked using the following (SEM) observations of the lenses from seeds of a publications in this order: Wiersema et al. (1990), small but taxonomically representative number of Polhill and Raven (1981), Gunn (1983), Cronquist species of Caesalpinioideae and Mimosoideae (1981), Mabberley (1988), and Kew Index (1895- Kunth. This survey of the lens is the most 1988). ambitious attempted for these subfamilies.
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