Testa Characters in Tribe Vicieae, with Notes About
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&:.i ~F 8 ~ w ~1112.5 1.0 W ~ IIIII~ 1.(' ~ - wW 2.2 Ii& I&.l Iii Iii 122~ ~W ~ W === Ilol Ilol ::: ~ : ~ .0 .. k ... k 1.1 ....k 1.1 ...... --- -- I I 111111.25 111111.4 111111.6 111111.25 111111.4 111111.6 MICROCOPY RESOLUTION TEST CHART MICROCOPY RESOLUTION TEST CHART NATIONAL BUREAU Of ST~NDARDS-1963·A NATIONAL BUREAU or STANDARDS·1963·A United States Department of Agriculture Testa Characters in Agricultural Research Tribe Vicieae, With Service Technical Notes About Tribes Bulletin Number 1667 Abreae, Cicereae, and Trifolieae (Fabaceae) by Nels R. Lersten and Charles R. Gunn AcitnowlE>dgments We express our appreciation to the Plant Introduction U.S. National Arboretum, Washington, D.C.; and for the staff of the U.S. Department of Agriculture, curators of technical assistance of Lynda E. Chandler, Department herbaria, and Steven Broich, Department of Botany, of Botany, University of Maryland, College Park, and Oregon State University, Corvallis, for supplying some Seanna R. Rugenstein and Larry D. Hufford, Depart of the seed S\amples. We are grateful for the technical ment of Botany, Iowa State University, Ames. advice of Frances Kupicha, Botany School, Cambridge, Microscopy and ancillary procedures were done in the England; Frank Bisby, Biology Department, Tile Univer Bessey Microscopy Facility, Department of Botany, sity, Southampton, England; Duane Isely, Department Iowa State University. This research was partly sup of Botany, lov-~a State University, Ames; J. Stuart ported by National Science Foundation Grant 420-21-74 Lassetter, Department of Biological Sciences, Eastern to N. R. Lersten. Kentucky University, Richmond; Frederick G. Meyer, Iii Contents Page Materials J.nd methcds ...•...••...•..•....•..•••.• _..................... 2 Page Tribes linked with Vicieae·.............................................. 10 Vicieae seed morphology··•• ...••._•..••....•. _....................... 2 Vicieae testa anatomy•••..•...••••••..••.•••._........................... 4 Abreae Wight & Arnott .•...••...•.....•........•......••...•...•.. 10 .. Literature survey ...••..••...•.....•.....•. _........................... 4 Cicereae Alefeld·····.................................................. 11 Trifo/ieae Bronn •...•.•.••.•.......•..••...•.......•...•...•..•....•..• 11 Observations •.••.••.•.•...•....•..••..•.•......••...•...••......•.....•. 5 Literature cited··............................................................... 13 Vicieae hilar anatomy-•..•..•••..••..•••..•...._......................... 6 Appendix······..................................................................... 16 Vicieae lens anatomy ..........•... _...................................... 7 Genera of Vicieae-··......................................................... 7 Historical references······......................................... 16 Lathyrus Linroaeus .••.•..•••.•....••....•...••.•••..••..••......••... 7 Taxonomic list of Vicieae taxa in table 1········..··•• 16 Lens Miller .••...••...•.....•....•.......•...••..••....••..........•....•.. 8 Table 1.-Summary of testa characters of Pisum Linnaeus······.................................................. 8 Vicieae species in SEM survey .••.....••...•...18 Vavilovia A. Fedorov ................................................. 9 Table 2.-Summary of testa characters of Vicia Li n na eu s.....••...•.......••...•......••..••...•........•........• 9 Trifolieae species in SEM survey ..........••.21 \ Issued September 1982 Iv Testa Characters In Tribe Vlcleae, With Notes About Tribes Abreae, Clcereae, .. and Trlfolleae (Fabac8ae) by Nels R. Larsten and Charles R. Gurm1 The Vicieae (Adanson) de Candolle is 1 of 41 tribes in pressed, without externally discernible radicular lobe, the legume family (Fabaceae or Leguminosae) and 1 of with elongate to circular hilum, with smooth to varl· 31 tribes in the subfamily Faboideae (Papllionoideae of ously rough·textured testa at x 10 or above x 30 Polhill, 1981).2 This tribe is homogeneous for most taxo· paplilose, and with extremely reduced hilar rim. Endo· nomically important characters. It consists of 5 genera sperm absent or nearly so. Seedlings hypogeous, with with the following number of species: Lathyrus Lin· radicle and hypocotyl triarch (rarely tetrarch). First naeus ± 140, Lens Miller 5, Pisum Linnaeus 2, Vavilovia cataphyll borne on side of plumule away from cotyle· A. Fedorov 1, and Vicia Linnaeus ± 150. Two other dons. x = 7 (6, 5), polyploidy rare. Canavanine some· genera placed in the Vicieaa by many taxonomists in times present in Vicia, pro parte. Type genus Faba the past and now in monotypic tribes are Abrus Adan· Miller (a synonym of Vicia Linnaeus). son in the Abreae W;ght & Arnott (Gunn, 1969; Hutchin· son, 1964; Polhill, 1981) and Cicer Linnaeus in the Seed characters seldom have been used to delimit the Cicereae Alefeld (Clarke and Kupicha, 1976; Kupicha, Vicieae and to assess its relationships to other tribes. 1977 and 1981). Except to identify weedy or crop speCies, especially in the genus Vicia, only the recent publications of Some Vicia species (vetches) are economically impor· Kupicha (1977), Lersten (1979), and Gunn (1981) have tant crops used for forage, hay, green manure, seeds, used seed characters to support taxonomic and phylo· and soil improvement. In addition, Vicia faba Linnaeus genetic hypotheses. Kupicha suggested that certain (faba·bean), Pisum sativum Linnaeus (pea), Lens seed features are important and used them both in culinaris Medikus (lentil), and some Lathyrus species delimiting the Vicieae and as a partial basis for trans (wild·peas) are grown throughout the temperate zone. ferring Cicer to the monotypic tribe Cicereae. Lersten Some Lathyrus are used as ornamentals. A few suggested that additional seed characters, especially Lathyrus and Vicia are considered weeds. In Morocco anatomical characters of the testa, may have tax and Poland, Vicia species are statutory noxious weeds. onomic and phylogenetic value. In this bulletin a com· prehensive treatment is given of morphological and Taxa In Vicieae share the following characters (after anatomical seed characters in the Vicieae and three Kuplcha, 1977): Perennial and annual herbs with erect tribes that have been linked with it. or more usually climbing or sprawling habit. Leaves epulvinate, exstipellate, alternate, distichous, pari pin· nate with rachis ending in tendril or mucro or rarely imparipinnate; leaflets entire (rarely dentate), many· paired to unijugate; rarely (in Lathyrus) leaves phyllodic or reduced to tendril and two stipules. Stipules semi· sagittate or hastate or variously divided. Inflorescence usually a secund axillary raceme or sometimes one· flowered, rarely a panicle. Bracteoles rarely present. Staminal tube diadelphous, but vexillary stamen with flattened filament lightly adhering to its neighbors (in all species); mouth of tube truncate or oblique. Anthers introrse, versatile, of equal size, rarely (Pisum and Vavilovia) alternate filaments dilated at apex. Style borne at right angles to ovary, usually dorsally or later ally compressed, usually pubescent (distribution of pubescence various), sometimes spathulate andlor con· torted. Stigma terminal, rarely double (Lathyrus). Legume more or less linear, laterally compressed, (one) two· to many-seeded, usually dehiscent, occasionally winged, sometimes with 'wooly' or rarely membranous partitions between seeds. Seeds spherical to com· 'Professor, Department of Botany, Iowa State University, Ames 50011, and botanist, Plant Exploration and Taxonomy Laboratory, Beltsville Agricultural Research Center, Beltsville, Md. 20705, respectively. 2The year In italic after the author's name refers to Literature Cited, p. 13. 1 Materials and Methods Vicieae Seed Morphology Seed samples used in this study are deposited in the Like seeds of most other legumes, seeds of the Vicieae U.S. National Seed Herbarium. Most of them also are consist of a testa, hilum, lens, micropyle, and an em vouchered by herbarium specimens, and these data are bryo composed of two cotyledons and an axis with a available from the second author. radicle, hypocotyl, mesocotyl, and epicotyl (figs. 1 and 2). Seeds of some Lathyrus and Vicia species have an External and internal topographical data were collected indurate funiculus covering the hilum. in a study of mature seeds from about 70 species of Lathyrus, 3 of the 5 species of Lens, both species of External and internal morphological characters of Pisum, the single species of Vavilovia, and over 100 seeds in this tribe have beE\n used to identify species, species of Vicia. Four species of Abrus (Abreae), 7 of but they have seldom been used in taxonomic consider Cicer (Cicereae), and over 100 species from all 7 genera ations. In the genus Vicia, for example, only Gunn in the tribe Trifolieae Bronn were studied. (1970) has used external topographic features to iden tify 100 of the 150 species in the world and in a sepa Anatomical studies were based on mature seeds repre rate publication all 36 native and naturalized species of senting 88 species from all 5 genera of Vicieae, 4 North America (Gunn, 1971). Other seed morphology species of Abrus Adanson (Abreae), 7 species of Cicer studies of Vicia species have been restricted to a few Linnaeus (Cicereae), and 27 species from all 7 genera crop or weedy species (Brouwer and Stahlin, 1955; of Trifolieae. Four seeds per species usually were used. Capit.aine, 1912; Dudik and Kondratyuk, 1970; Iannelli, Two seeds were left intact for surface