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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 _...... 7 Genera of Vicieae-··...... 7 Historical references······...... 16 Lathyrus Linroaeus .••.•..•••.•....••....•...••.•••..••..••......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 examination; the 1964; Justice, 1952; Korsmo, 1935; Kostrakiewicz, 1951; other two were cut through the hilum with a razor Lhotska and Chrtkova, 1978; Musil, 1963; Pierpoint, blade, one in transection and the other in longitudinal 1949; Quartley, 1968a and 1968b; Red'kina and section. Seed specimens were dipped in 100 percent Khoroshailov, 1974; Swederski, 1924; Tupikova, 1926; ethanol, mounted on a brass disk with silver paste, Utkin, 1965; Vodak, <1956; Wittmack, 1922; and Zertova, coated with carbon then gold or gold-palladium in a 1962). Similar studies of Lathyrus, Lens, Pisum, and vacuum evaporator or a sputtering device, and ex­ Vavilovia seeds have not been undertaken, except in amined using a JEOL JSM-353 scanning electron micro­ those seed identification manuals that may include scope (SEM). Photographs were taken using Polaroid­ crop or weed species. type 665 film. Seeds of the Vicieae have the following morphological characters, whi~h are illustrated in figures 1-35 and 61-87.

Primary seed outlines are circular, lenticular, oblong, oval, pyramidal, and square and either terete or com­ pressed in transection. Occasionally seeds may be compressed-spherical or oblong, with one or both ends truncated or angular. These modified shapes are usu­ ally caused by pressure from adjacent seeds in the leg­ y ume (pod). Seeds usually mature without pressure, because some ovules remain unfertilized or abort. When all, or nearly all, the ovules mature, seeds may be compressed from one or both sides. Lathyrus angulatus Linnaeus, L. niger (Linnaeus) Bernhardi (fig. 4), and Vicia lathyroides Linnaeus seeds are excep­ tions because their cuboidal shape is the result of pressure from adjacent seeds as well as the legume wall.

The surface of the testa is usually smooth. Vicia species that occasionally have the optical illusion of a velvetlike testa surface include V. benghalensis Linnaeus, V. lutea Linnap.us, V. sativa Linnaeus, and 3Mention of a trademark or proprietary product does not con­ V. villosa Roth. The testa of Pisum sativum may be stitute a guarantee or warranty of the product by the U.S. wrinkled. The testa of some Lathyrus and rarely Vicia Department of Agriculture and does not Imply its approval to species may be ornamented (figs. 2-29): Aculeate in the exclusion of other products that may also be suitable. V. lathyroides, rugose in L. cirrhosus Seringe, ruminate

2 with large interstices in L. pusillus Elliott, ruminate The lens is all reinforcements of normal tissue of the with small interstices in L. latifolius Linnaeus, testa, situated between the hilum and chalaza on the tuberculate in L. hirsutus Linnaeus, and tuberculate­ trajectory of the principal ovule vascular bundle. Exter­ ridged (some tubercula coalesced) in L. annuus nally it is usually a lens-shaped, raised structure near Linnaeus. On some of the ruminate testae with small the hilum (fig. 70). The term 'chalaza' is often incor­ interstices, the ornamentation may be more a factor of rectly used for this lens area. According to Pitot (1936), testa color (a mottled appearance) than elevation of the the chalaza may be close to the lens but is never found testa surface. under it. He and other authors defined the chalaza as a "point at which the vascular bundles reach the Seed and hilum colors are whitish, straw, yellowish, nucellus." orange red, reddish, ocher, green, greenish, brown, chocolate, gray, purplish, blackish, and combinations The lens is located near the narrow end of the hilum, of these colors. The seeds may be monochrome or the end opposite the micropyle, except when the lens marked. Mar~;ed seeds (fig. 2) may have mottles (larger is on the opposite side of the seed, e.g., Vicia hybrida, blotches or irregular shapes), points (point-shaped V. lutea, and V. pannonica Crantz. The positions of the dots), and streaks (lines of color). lens relative to the hilum are as follows: Lens on the opposite side of the seed from the center of the hilum, Seeds are classified by length (parallel to hilum) as center of lens removed (more than 0.5 mm) from the small (less than 3 mm), medium (3 mm to less than hilum, center of lens near the hilum (less than 0.5 mm) 6 mm), or large (6 mm or more). but the discolored area not touching the hilum, and lens confluent with the hilum (discolored area touching The hilum is usually the color of the testa, but funicular the hilum). remnants occasionally persist, making the hilum ap­ pear silvery, viz, Lathyrus nissolia Linnaeus (fig. 3) and Within the testa are two well-developed equal cotyle­ Vicia hybrida Linnaeus. The funiculus (or most of it) dons and a curved embryonic axis. The absence of a rarely persists as it does on V. benghalensis and clearly visible endosperm in mature seeds is one of the V. hirsuta (Linnaeus) S. F. Gray, or as the rim-aril of characters that unites the taxa. V. grandiflora Scopoli (fig. 7). The hilum of most of the Faboidea8 has a fine central groove. This is easily seen in Vicieae seeds, except where the funiculus persists. Either the lips of the groove may be the same color as the hilum or the whitened lips may be rather prominent.

A few species have a definite rim around the hilum, viz, Lathyrus pusillus, L. sulphureus Brewer ex Gray, and L. tingitanus Linnaeus. Others have a halo, a discolored area surrounding the hilum, that is not built up like a rim, viz, Vicia articulata Hornemann and V. villosa subsp. varia (Host) Corbiere.

The five hilum outlines are circumlinear, linear, oblong, wedge-shaped, and oval (elliptical) (figs. 3-8). A circum­ linear hilum is more than 10 times longer than wide, occupies 50 percent or more of the seed circumfer­ ence, and is always linear. A linear hilum is 5 to 7 (rarely 9 to 10) times as long as wide, occupies less than 50 percent of the seed circumference, and has parallel margins. An oblong hilum is less than five times but always more than twice as long as wide, has slightly curved margins, and occupies less than 40 per­ cent of the seed circumference. A wedge-shaped hilum has margins converging toward the lens. In all other respects it is similar to the oblong hilum. An oval hilum is never longer than twice its width, is rounded in out­ line, and occupies less than 20 percent of the seed circumference~

3 Vicleae Testa Anatomy

Literature Survey is an incomplete loop found in the raphe." Kuhn (1927) Pammel (1899) summarized 19th century work on testa reexamined C. arietinum and V. faba and agreed with anatomy of legumes, including Vicieae. He and certain Le Monnier's observations. earlier workers were aware that epidermal (malpighian) cells in some species had a conical distal tip and that Pitot (1936) included several Vicieae and described ~~e surface was papillose as a result. Pammel noted it them as having "... two lateral recurrent bunales for 10 species of Vicieae; he found it elsewhere only in more or less developed, occasionally consisting only of Medleago Linnaeus and Melilotus Adanson but not in phloem...." Trifolium Linnaeus of Trifolieae. Chalon (1875) included 32 species of Vicieae in his survey. He remarked that Kupicha (1977) followed the course of vasculature in "in most of the vetches, the external extremity of the seeds of about 30 Vicieae species and found all to [malpighian cells], instead of being flat, is strongly con­ have a to••• simple, unbranched vascular bundle form­ vex, which gives the surface a finely mammillate ap­ ing a loop round the periphery of the seed." She did not pearance." Chowdhury and Buth (1970) reported 'den­ list Vieia faba as a species examined but said that she tate' cuticle in two Vicieae species, and Patel (1976) did not agree with Kuhn (1927) and therefore also included similar observations for a few edible species disagreed with Le Monnier (1872). of Vicieae and Cieer arietinum Linnaeus. A quite different observation was made by Hardham Malpighian cells in Vicieae seeds were reported by (1976) in Pisum sativum. He reported that, in addition to Pammel to range from 40 to 200 J!m in length among 14 the single unbranched bundle that extends about half­ species measured, with no distinct tendencies in way around the seed, two bundles conSisting only of Lathyrus, Lens, Pisum, or Vieia. Various structural de­ phloem branch off just above where the seed is attach­ 4 tails (as light line and pore canals) have been de­ ed to the funiculus and extend on either side of the scribed in these cells; they are not taxonomically sig­ testa toward the radicle of the embryo. Hardham's nificant at the tribal level. observations and the disagreement of Kupicha (1977) with Kuhn (1927) and Le Monnier (1872) raise questions The sclereid layer consists of osteosclereids ranging about the uniformity of the testa vascular pattern in from 10 to 60 J!m in height (fig. 2). Pammel reported that Vicieae. Harz (1885) said this layer varied from 58 to 140 J!m among varieties of Vieia faba. Rowland (1977) verified Since Pammel (1899), some reports have included infor­ this. mation on the testa of Vicieae, mostly of Pisum and Vieia faba. Alexandrov and Alexandrova (1935) de­ Beneath the sclereid layer, several layers of cells are scribed the testa of several species and varieties of compressed and more or less completely crushed at Pisum. All were similar in anatomy. Reeve (1946) stud­ maturity. Pammel and earlier authors called these col­ ied developing and mature testa in P. sativum. He lectively the 'nutrient layer,' thinking that it contributed pOinted out that the tracheid bar, via transitional cells, something to the developing embryo. This remnant connects with the xylem of the ovule bundle. He also layer of the inner integument and nucellus is similar to showed that there is no strophiolate (= lens) groove that in other tribes (fig. 2). and that in this area the malpighian cells are merely greatly elongated. Le Monnier (1872) included many legumes in his survey. Concerning Vicieae he commented: "One can see [in Spumy, in several reports (see Append!>c), explored the Cieer arietinum] a simple bundle which, departing from chemical composition of malpighian cells of Pisum the hilum, follows the contours of the seed; arriving at sativum and concluded that each cell has a "starlike" a point almost directly opposite the hilum, it bifurcates suberized cap at its external tip. He attached a movie to give rise to branches which curve onto each face of camera to a microscope to demonstrate this while the seed. In [Vieia faba] there is an analogous disposi­ watching the cellulosic part of the cell dissolve tion, except that the bifurcations are very short; but in (Spumy, 1963). A similar conclusion was reached by the majority of Vicia, these lateral branches have disap­ Stein von Kamienski-Jancke (1957), who studied peared completely. Finally, in Pisum and Lathyrus there P. sativum and Vieia faba among others.

4The use of the term 'light line' in a discussion of malpighian The first published SEM photograph of Vicieae testa cells In the testa of legume seeds should not be confused surface at high magnification was in an article by with the pleurugram, or so-called 'light line,' which is a sur­ Marbach and Mayer (1974), who showed the character­ face feature of some seeds in the Mimosoideae and istic crimped papillose surface in Pisum sativum Caesalpinioideae. subsp. elatius (M. Bieberstein) Ascherson & Graebner

4 seeds but did not mention it in the text. McEwen et al. The malpighian cells in most species extend above the (1974) showed surface and internal features of Vicia surface and give the testa a papillose appearance at faba seeds. The surface was not papillose in their high magnifications (figs. 9, 10, 30, 64, 80). Variations on SEM, but they did not indicate the location of their this conical theme also occur (figs. 11-15), with a few observations. One micrograph included cells of the species exhibiting faint papillae (figs. 16, 17). Mounds tracheid bar. It was sufficiently magnified to show pits occur in some species of most genera (figs. 18-22, 23, as small, bordered, rather uniform in size and arrange­ 24, 26) as a result of local elongation of a group of ment, and nonvestured. malpighian cells (figs. 21, 26, 32). Isolated ridges (fig. 35) or reticulately continuous ridges (figs. 34, 35) Kupicha (1977) included seed characters in delimiting rarely occur, also as a result of malpighian cell exten­ the Yicieae, emphasizing seed and hilum shape and the sion (fig. 29), never by additional cell layers of the epi­ unbranched ovule bundle in the mature seed. She men­ dermis. A taxonomic list of Vicieae taxa studied by tioned that ~he seed surface may be "smooth or SEM and topographic features summarized in table 1 variously rough-textured." are in the Appendix.

Saint-Martin (1978) included 12 species of Lathyrus and There is some variation in the relative proportions of Vicia and 23 species of Trifolieae in a SEM survey of malpighian and sclereid layers. Figures 55 and 69 illus­ seeds from 4 faboid tribes. She listed most species of trate the most common condition. More rarely the Vicieae and Trifolieae under the heading of "rounded malpighian layer is extremely thick and the sclereid protuberances," but her SEM of Lathyrus sylvestris layer thin (fig. 56) or vice versa (fig. 57). Linnaeus shows much more conspicuous papillae than the SEM of Medicago turbinata (Linnaeus) Allioni. In The sclereid layer is usually most prominent near the Saint-Martin's survey, seeds of Vicieae and Trifolieae hilum, becoming inconspicuous, or even seemingly ,. are distinctive but similar to each other in topograph­ crushed, progressively toward the midseed area. The ical features. representative condition is shown in figures 54 and 66. This aspect of the testa was observed in 65 species, of Trivedi et al. (1978) examined six species of Vicieae which 61 showed a reduced sclereid layer away from and also Cicer arietinum and Abrus precatorius the hilum, including all species of Lathyrus, Pisum, and Linnaeus, using a SEM. We agree with their results, Lens. Two species of Vicia maintained a prominent except they show a low, irregularly ridged, and undu­ sclereid layer all over the seed (fig. 57), and another late pattern in Pisum sativum that is unlike the sharply pair of Vicia species had a small or nonexistent papillose pattern we have seen and that has been sclereid layer all around the seed. The decrease in shown by others using a SEM. sclereid height away from the hilum is also the com­ mon condition in other tribes. It probably occurs Lersten (1979) pointed out that, in seeds of Viceae, the because of the pressure of embryo expansion during reduction of the hilar rim and the. papillose seed sur· seed development. face when added to the usually elongate hilum provide y a trio of characters that strengthen the definition of the tribe presented by Kupicha (1977).

In summary, earlier anatomical studies have provided information that to varying degrees has helped to de· velop a set of characteristics for the seeds of Vicieae. Some of these may be unique to this tribe. Nineteenth century work of lesser importance is included in the Appendix.

Observations Except in the hilum, the testa of a mature seed con­ sists of the single layer of slender, elongate malpighian cells, a subtending layer of columnar sclereids (vari­ ously termed macrosclereids, asterosclereids, hour­ glass cells) with prominent intercellular spaces, and below this a poorly defined zone of partially or com­ pletely crushed cells, the remnant layer, which may consist of remnants of endosperm (fig. 2).

5 Vicieae Hilar Anatomy

A definite counter palisade is present in most Vicieae Because pits in the tracheid bar of other tribes are usu­ seeds (figs. 37, 38, 41, 6f'\, but in a few it is obscure ally warty or vestured (Lersten, 1982), these pits were (figs. 36, 40). A tracheid bar is always present, extend­ investigated in 76 Vicieae species. In 35 species (46 ing the length of the hilum just beneath the hilar percent), all pits seen were nonvestured (figs. 48-50, 77, groove. This bar is always embedded in a mass of 86). Included here were both Pisum species and Lens rather spongy parenchyma, which forms a thicker layer culinaris. In 31 species (41 percent), the pits were non­ here than elsewhere in the seed (figs. 36-41, 66, 72, 85). vestured to warty, mostly slightly so (fig. 68). Warts are simple, usually small, wall protuberances (figs. 51, 52). In transection the tracheid bar is usually elliptical, A third group of 10 Vicia and Lathyrus species (13 per­ which Is variously narrow or broad (figs. 36-40, 66, 76, cent) had a range of nonvestured to warty to simply .. 85); more rarely it appears circular (fig. 41). No trends vestured, or all pits were warty to vestured. The ves­ were evident, although Vicia and Pisum are apparently tures were never elaborate and were more knobbed more uniform than Lathyrus. One might assume that than branched (fig. 53). True vestured pits, therefore, the transectional area of the tracheid bar would be pro­ such as those reported in the secondary xylem of portional to the size of the seed, but there was no several dicot families (Miller, 1977; van Vliet, 1978; strong correlation between them. The tracheid bar Baas and Zweypfenning, 1979) are essentially absent in abuts on the hilar groove, and if the groove is open, as Vicieae. A uniformly regular pit of somewhat distinctive ... it usually is in mature dry seeds, the uppermost size and shape, usually nonvestured or with a few tracheids are exposed. small warts on the pit borders, is the rule in the Vicieae. Individual tracheids in the bar are always much larger than tracheary elements in true vascular bundles. This size difference is easily seen in figures 43 and 45. The tracheids resemble more the 'tracheoid idioblasts' that have been described in vegetative organs, mostly leaves, of several families (Lersten and Bender, 1975; Rao and Das, 1976). The enlarged tracheid bars of Lathyrus cfymenum Linnaeus (fig. 42) and L. fatitofius (fig. 67) are representative; they show some detail of cell size, shape, and orientation and illustrate the numerous pits.

In a longitudinal section the tracheid bar always ex­ tends from the micropylar edge (fig. 44) at one end of the hilum to the ovule vascular bundle at the other end, with which it seems to merge (figs. 43, 45). The tracheid bar thus appears to be a curious 'idioblastic' extension of the ovule bundle. Figures 45-47 illustrate some ex­ y amples of tracheids; those in figure 47 have a consider­ ably thicker wall than the others. The tracheid wall is always of the pitted type, never helical, reticulate, or scalariform. This is unlike tracheary elements in the ovule bundle (fig. 45).

Pits in the tracheid bar of Vicieae offer features of in­ terest when compared to those of other faboid tribes. The pits are surprisingly uniform in size and shape (figs. 48-50, 68, 77, 86); larger, more Irregularly shaped pits are rare.

The primary wall or pit membrane is absent at maturity from virtually all pits in all Vicieae species. Only rarely is the primary wall seen to be even partially intact (fig, 48, upper right). This is also common in other tribes, but an intact primary wall is often seen in tracheary elements of the ovule vascular bundle (Lersten, 1982).

6 Vicieae Lens Anatomy Genera of Vicieae

In the Vicieae, the lens (boss, chalaza, or strophiole of Lathyrus Linnaeus various authors) is confluent to separate from the non­ The ± 140 species in this genus are indigenous in micropylar end of the hilum. The lens appears in sur­ Europe, Asia, North America, and Mexico, extending face view as a roughly circular swelling (figs. 1, 70). The into temperate South America and tropical East Africa, ovule vascular bundle occurs directly beneath it. with most species in the Mediterranean and Irano­ Anatomically the lens is not distinctive in Vicieae. The Turanian regions (Kupicha, 1981). Lathyrus has not malpighian cells are unusually elongate, but otherwise been monographed in this century. Bassler (1966) the testa appears as elsewhere, with no evidence of a reorganized the subgenus Orobus (Linnaeus) Bassler, disrupted cuticle. Mattirolo and Buscalioni (1892) but the subgeneric hierarchy elsewhere in Lathyrus is showed this in Vicia faba as did Kondo (1913) in patchy and doubtful (Ball, 1968; Davis, 1970), especially Lathyrus sativus Linnaeus and Pitot (1936) and Reeve New World taxa (Hitchcock, 1952). The Lathyrus section (1946) in Pisum sativum. Zimmermann (1936) reported of the taxonomic list of Vicieae (Appendix) variously that this was the general condition in Vicieae, based on follows Bassler (1966), Simola (1968), and Ascherson examination of Lathyrus japonicus Willdenow subsp. and Graebner (1909). Unlike Vicia there are no general maritimus (Linnaeus) P. W. Ball, Lens culinaris, P. treatments of Lathyrus seed morphology or anatomy. sativum, and two species of Vicia. Matlirolo and Seeds of 65 species from throughout the range of the Buscalioni concluded that "In Vicia faba, in Pisum, in genus were studied for morphological features, and 44 Gicer, and in Ervum Linnaeus [Lens pro parte] ..., the species were studied for anatomical features of the [lens] is formed exclusively by the malpighian cells testa. which elongate exaggeratedly •..." Seed shapes reflect the range of shapes for the tribe, Lens anatomy was seE:.,n in seven Vicia and Lathyrus with a larger number of species with truncate or species; figures 58-60 show representative examples. A angular sides or compressed in Lathyrus (fig. 61) than disadvantage of using SEM and simple razor-sectioned in Vicia. seeds is that one does not obtain a series of sections, but instead is dependent mostly on the luck of the blade. Because of this one cannot follow, for example, The surface of the testa is smooth and unwrinkled, but the path of the ovule vascular bundle in the testa. In at x 10 may be aculeate (L. nissolia), rugose addition, details of individual cells exposed by section­ (L. cirrhosus), ruminate with large (L. pusillus) or small ing are usually ubscure, so that one cannot clearly interstices (L. latifolius), tuberculate (L. hirsutus), and record wall thickness or the occurrence of the light line tuberculate-ridged (L. annuus). Testa of some species, in the malpighian layer. Such information must be i.e., L. latifolius, L. sylvestris, and L. tuberosus obtained by other methods. Linnaeus may vary from ruminate (fig. 62) to smooth.

Testa colors primarily range through the browns to blaCkish, with a few sper;ies exhibiting ocher, but no orange red, reddish, or green occurs. Less than 20 per­ cent of the studied seeds are mottled, pointed, or streaked. More than three-quarters of the studied seeds are from 3 mm to less than 6 mm in length, with the remainder equally divided between small and large.

The hilum is conspicuous, usually beca.use of the white to whitened lips of the hilar groove and often the color of the testa. Hilar shapes are as recorded for the tribe, except circumlinear, which is not present. The oval out­ line (figs. 61, 63) is more commonly found in Lathyrus than in Vicia.

Some species of Lathyrus, such as L. pusillus, L. sulphureus, and L. tingitanus, have a Ililar rim; this is absent in seeds of the other genera. Occasionally the hilum is rather encompassed by a discolored area, a halo, which is usually darker than the rest of the testa (L. blepharicarpus Boissler).

7 Because the testa is usually brown or a derivative hue, Seed shape is strictly lenticular. The surface of the the black lens is generally conspicuous and often is a testa is smooth at x 10. Testa colors range from mound of tissue near the hilum, sf.:lldom confluent with monochrome to mottled, streaked, or pointed browns to the hilum. blackish. Seed sizes vary from small to large. The linear hilum (fig. 70) is the color of the testa and con­ The internal structures reflect the structures for the spicuous because of the whitened lips of the hilar tribe. groove. There is no hilar rim. A halo is present in L. ervoides. A linear micropyle adnate to the hilum was The testa of Lathyrus mostly conforms to the general observed in all examined species (fig. 71). The black or features of the tribe, but more variation was noted than reddish-brown lens is a mound of tissue to the right of in other genera. More than one-half (26/44) of the spe­ the hilum in figure 70 and is conspicuous on seeds cies had standard papillae in the hilar region (figs. 9, with either brown or black testa. The internal structures 10, 64), and 10 of them showed attenuation and fading of Lens seeds reflect the structures for the tribe out of the papillae at various distances from the hilum. (fig. 72). Four species had low papillae near the hilum (figs. 14, 17). The testa of Lens resembles that of most Vicieae, dif­ fering only in that the papillae are shorter than in most Mounds or ridges were fairly common in Lathyrus, other taxa (figs. 15, 73). occurring along with the papillae (figs. 19, 23, 24, 26, 29). Twelve species (about 30 percent of the sample) had such ornamentation, including all 8 representatives Pisum Linnaeus of the section Lathyrus (figs. 62, 65). Two species had Like Lens, the generic concept of Pisum has remained pits (fig. 25) along with papillae, but no mounds or stable, but the number of species varies from author to ridges (fig. 30); this pattern was not found elsewhere in author. Kupicha (1981) recognized two species: P. the tribe. fulvum Sibthorp & Smith and P. sativum, both in­ digenous in the Mediterranean region. The hiklm in Lathyrus shows anatomical features as described for the tribe. The tracheid bar in transec­ Seed shapes range from spherical to subspherical to tional view (figs. 42, 66) seemed to show a greater angular or almost cuboidal. The surface of the testa range in size and shape than in other genera, but this varies from smooth to wrinkled (unique to Pisum) at x is merely a qualitative judgment. 10 (fig. 2). Testa colors reflect those for the tribe. Most testae are monochrome; some are dichrome because of When pits in the tracheid bar were viewed at high mag­ mottles or points. Seed sizes range from medium to nification (40 species in the sample), 21/40 (52 percent) large. The oval hilum is usually conspicuous (fig. 74), were smooth and nonvestured (fig. 67). In 15/40 species because it may be either darker than the light-colored (38 percent), pits were smooth or sligh!Jy warty (fig. 68), testa or silvery because of a funicular residue, an and in 4/40 species (10 percent), pits ranged from epihilum. If the hilum is the color of the testa and the smooth to warty (fig. 52) and vestured. The only trend epihilum is absent, then the hilum may be inconspic­ noted was that North American species tended to have uous. The lips of the hilar groove may be the color of warts or vestured pits. the hilum to yellC'wish. There is neither a hilar rim nor a halo. As in other Vicieae genera, the elongate In the 34 species in which the testa was observed in micropyle is ad nate to the hilum (fig. 74). The lens, a transectional view, the sclereid layer was always con­ mound of tissue near the hilum, is the color of the spicuous near the hilum (fig. 66), becoming reduced in testa or nearly so and thus inconspicuous. height (fig. 69) or crushed completely toward midseed.

Lens Miller The papillose testa of Pisum conforms anatomically to The generic concept of Lens has remained stable, but that of the tribe in general. The papillae on all samples the number of species varies from author to author. had a characteristic crimped appearance (figs. 12, 13, Kupicha (1981) recognized five species: L. culinaris, 18, 75); (Lersten, 1979; Marbach and Mayer, 1974). Ir­ L. montbretii (Fischer and Meyer) Davis & PI itmann, L. regular mounds also were seen on P. sativum subsp. nigrlcans (M. Bieberstein) Godron, L. orientalis elatius (fig. 18), similar to those on seeds of some Vicia (Boissier) Handel-Mazzetti, all indigenous in the Medi­ and Lathyrus species. The internal structures of Pisum terranean region, and L. ervoides (Brignoli) Grande, seeds also reflect the structures for the tribe (figs. 37, believed to be indigenous in Uganda, Ethiopia, and 44, 76). Tracheid bar pits are smooth and regularly Zaire. distributed (fig. 77).

8 Vavilovia A. Fedorov Seed and hilum are tribal colors. Seeds may be Vavilovia is a segregate genus of Pisum and contains monochrome or mottled, pointed, or streaked. More one Near East species, V. formosa (Steven) A. Fedorov. than three-quarters of the studied seeds range from 3 mm to less than 6 mm in length, with the remainder Seed shape is oblong. The surface of the testa is equally divided between small and large seeds. smooth at x 10 and monochrome or mottled brown. Seed size is medium. The short, wedge-shaped hilum The hilum, often the color of the testa, is conspicuous, (fig. 78) is the color of the testa but made conspicuous usually because of the white to whitened lips of the by the whitish lips of the hilar groove. There is a partial hilar groove. Hilar shapes are as recorded for the tribe, hilar rim, most prominent on each side of the micropyle with oval and circumlinear occurring less frequently (figs. 78, 79). A halo is absent. The elongate micropyle, than other shapes (figs. 2, 7, 81). Vieia koeieana, the although ad nate with the hilum, exhibits a slight bifur­ only species in section Anatropostylia, possesses cation not seen in other Vicieae seeds (fig. 79). The unusual characters as do the species in the seven lens, a mound of tissue near the hilum, is the color of other monotypic sections (see taxonomic list in Appen­ the testa and thus inconspicuous. The testa surface is dix). Even seeds of V. koeieana are unusual, reinforcing typically papillose (figs. 78-80). The internal structures the section recognition-the circular hilum and of Vavilovia seeds reflect the structures for the tribe. funicular remnant (figs. 33, 34). The other seven species have typical seed characters.

Vicia Linnaeus The hilar rim is absent on Vieia seeds. Occasionally the The ± 150 species in this genus are indigenous in hilum is encompassed by a discolored area, a halo, Europe, Asia, North America, Mexico, and Central which is usually darker than the rest of the testa as in America, extending to temperate South America, V. articulata. Hawaii, tropica.1 East Africa, and the Canary Islands. Most species occur in the Mediterranean and Irano­ As is characteristic for the tribe, the micropyle is linear Turanian regions. Kupicha (1976) reviewed the and extends shortly into the hilum (fig. 82). parameters of Vieia and created a worldwide in­ frageneric structure for the species. Gunn (1969) Because the testa is usually brown or brownish, the discussed the 25 genera that have been segregated black lens is generally conspicuous and often is a from Vieia. All now are synonyms of Vieia. The mound of tissue near the hilum, seldom confluent with segregate genus Anatropostylia (PI itmann) Kupicha was the hilum or on the opposite side of the seed from the created and accepted by Kupicha (1973, 1976) and hilum (V. hybrida, V. lutea, and V. pannoniea). reduced to a section of Vieia by Kupicha (1981). The internal structures reflect these structures for the There has been no recent study of Old World Vieia tribe. species, except as part of the recent floras of Europe and Turkey (Ball, 1968; Davis, 1970). All Vieia species in With few exceptions (e.g., fig. 11), the testa has the North America and Hawaii have been studied since standard papillae, which occur over the entire surface the 1950's (Gunn, 1968, 1971; Hermann, 1960; Lassetter (fig. 83). Vieia faba has reduced papillae (fig. 16), and 1972, 1975; and Lassetter and Gunn, 1979). The species six species have the papillae restricted to the area of South America have not been studied. Unlike around the hilum. Only two species Ilave mounds or Lathyrus, the morphology of Vieia seeds has been short ridges accompanying the papillae (figs. 33-35, studied both in North America (including Mexico) and 84). worldwide (Gunn, 1969, 1971). There have been no • similar anatomical studies. Seeds of 100 species from The tracheid bar in the hilum is broadly elliptical in thlOughout the range and from all 23 sections of the transectional view in most species (figs. 41, 85), but it genus were studied for morphological features, and 40 approaches the circular in V. sylvatiea Linnaeus, V. species from throughout. the range and from 15 sec­ faba, and V. bithyniea Linnaeus, and it is more narrowly tions were studied for anatomical features of the testa. elliptical in such species as V. semiglabra Boissier and V. sativa. The tracheid bar is noticeably smaller in • Seed shapes reflect the range of shapes for the tribe . transectional area in such species as V. unijuga A. The surface of the testa is smooth, not wrinkled. Brown, V. eassubiea Linnaeus, and V. nigrieans Hooker However, at x 10 it rarely may have a velvetlike testa & Arnott subsp. gigantea (Hooker) Lassetter & Gunn. surface (V. benghalensis, V. lutea, V. sativa, and V. vil/osa) or rarely be pusticulate (V. lathyroides and V. The pits in the tracheid bar are mostly nonvestured (fig. koeieana Rechinger f.). 86) or with slight warts (fig. 51) to more obviously warty.

9 Tribes Linked With Vicieae

Only six species have simply vestured pits of the type Abreae Wight & Arnott shown in figure 53. It is of interest that four of these This monotypic tribe containing the genus Abrus with - occur either in the first section (V. sylvatica, V. unijuga) up to 13 species was included in Vicieae by several or in the second section (V. cassubica, V. semiglabra), 19th and 20th century authors. Recent investigators, in­ the two sections considered least advanced by Kupicha cluding Pol hill (1981), have found nluch evidence for (1976) on the basis of other characters (see taxonomic removing it, and Kupicha (1977) has discussed and list in Appendix). Vicia faba is another species with tabulated differences between Abrus and. Vicieae. vestured pits. Because seed characters were not tabulated by Kupicha, we examined Abrus species seeds. In the nonhilar part of the testa, 22 of 26 species showed a reduced size, sometimes obliteration, of the The seed outlines vary from oval or oblong to ellipsoid­ sclereid layer from near the hilum to midseed (fig. 87). globose and compressed (Abrus aureus Viguier) or terete (A. precatorius) in transection. Verdcourt (1970) Of the two large genera of Vicieae, Vicia appears to be used seed transection shape (terete versus com­ the most uniform in testa characters, with little varia­ pressed) as a major key character for the East African tion from the standard. taxa of Abrus. Compressed seeds are either monochrome or mottled light to dark brown (brown Kupicha (1977) mentioned that the ovule vascular bun­ black) with a rim- andlor tongue-aril around a sunken dle in Vicia continues past the chalaza and remains un­ elliptical hilum, which is not covered by a funicular branched. The techniques used in our study did not residue. Terete seeds are dichrome black around the allow us to follow the course of the vascular bundle in hilum area and scarlet or monochrome black, brown, the testa. tan, cream, or white without a dry aril around the sunken hilum, which is covered by a white funicular residue. The testa is usually glossy and smooth to minutely shagreen. Seeds range from 4 to 7 mm long and 2 to 5 mm wide. Like the Vicieae, the radicle lobe is not externally visible. Internally there is a thin layer of endosperm ad nate to the testa.

Figures 88-93 illustrate the testa features of Abrus and show little similarity with Vicieae. The hilum is longer than broad (fig. 88), but the funicular remnant is rare in Vicieae and the conspicuous hilar rim (figs. 88, 89) does not occur. The micropyle in Abrus is deltoid and rather removed from the hilar edge (fig. 89). In Vicieae the micropyle is linear and adnate to the hilar edge.

The seed surface shows no hint of papillae (fig. 90), and at higher magnification one sees only small cuticular ., wrinkles and excrescences (fig. 91), which were never noted in Vicieae. Trivedi et al. (1978) noted and il­ lustrated by SEM the same differences between seeds of Abreae and Vicieae, but they retained Abrus in Vicieae. +. The hilum in transectional view is seen to be sunken (fig. 92); in the Vicieae the hilum is fiush with the sur­ face or raised, except slightly sunken in Vicia vil/osa subsp. varia. Other features of the hilum transection are similar to those in Vicieae, but they are also com­ mon to virtually all faboid seeds.

Tracheid bar pits are unlike those of Vicieae because of their reduced border, irregular shape, and the almost universal retention of the pit membrane (fig. 93). The pits are smooth to slightly warty.

10 We were unable to see the lens area properly, but the The tracheid bar, as represented by Gicer arietinum, ex­ excellent paper by Bariola (1912), whose observations tends the length of the hilum (fig. 98) and merges with and illustrations agree with ours in other respects, the ovule vascular bundle (fig. 99). The tracheid bar ap­ .. shows lens anatomy that is different from that of pears similar to those of Vicieae in having rather Vicieae. He shows no noticeable elongation of the regular, conspicuously bordered pits. The pits are most­ malpighian layer in the lens area, but subtending this ly nonvestured (fig. 100), with at most only a few slight layer are three layers of shorter, more slender warts (fig. 101). The hilum in transection has the usual palisadelike cells, 50 to 60 cells wide, with the lower appearance (fig. 97) and resembles the drawing of the layer narrower and centered just above the ovule bun­ hilar transection by Kondo (1913). Kondo shows the dle. A conc;oicuous fissure extends through all four cell hilar parenchyma as small and closely packed; in figure layers. There is no sclereid layer subtending the epider­ 97 the parenchyma also appears less spongy than that mis in the lens. Pitot (1936) also illustrated a transec­ of observed Vicieae seeds (figs. 36-42). tion of the lens of Abrus precatorius. His figure agrees in general with that of Bariola but was drawn in less Kondo (1913) illustrated the lens of Gicer arietinum in detail. sectional view. The malpighian cells are exaggeratedly elongate, and a vascular bundle occurs directly After studying the microscopic features of a large sam­ beneath. In contrast to Vicieae, however, he showed ple of Vicieae seeds, it is patently clear to us that that the sclereid layer is interrupted by parenchyma Abrus is distinct and thus supports the correct view tissue, which occurs between the vascular bundle and that this genus belongs in a separate tribe, the Abreae. the epidermis. Pitot (1936) showed a similar illustration, but with less detail. Behl and Tiagi (1977) studied fruit and seed development in Gicer. They mentioned that Cicereae Alefeld the mature seed has ridges, beak, and sunken hilum This monotypic tribe containing the genus Gicer was not found in Pisum or Lathyrus. reestablished (\S a tribe by Kupicha (1977, 1981). Prior to the article by Glarke and Kupicha (1976), most Trifolieae Bronn authors regarded Gicer as a member of the tribe This tribe contains 7 genera: Factorovskya Eig (1 sp.), Vicieae. Medicago Linnaeus (± 50 spp.), Melilotus Adanson (± 20 spp.), Ononis Linnaeus (70-80 spp.), Parochetus Gicer seeds, as described by van der Maesen (1972), are Buchanan-Hamilton ex D. Don (1 sp.), Trifolium Lin­ "bilobular to subglobular, conspicuously beaked, hilum naeus (± 300 spp.), and Trigonel/a Linnaeus (± 70 spp.). rather deep [shorter than most Vicieae], [lens] present; surface smooth, wrinkled, tuberculate or with smooth Seed shapes in the tribe vary. Medicago and some or echinate spinelets; in various shades of brown, grey, Trigonel/a species vary as to subreniform shape, black, white, yellowiSh-orange or green." He included because the radicle lobe is about one-half the line drawings of seeds of 26 of the 40 species in the cotyledon lobe length and not parallel to the length of genus. this lobe. Angularity and twisting are caused by pressure from closely adjacent seeds in the multi­ Our examination of seeds of seven Gicer species and seeded legumes as well as from the shapes of the the seed drawings and descriptions in van der legumes, which may be coiled, arched, or curved. Seed Maesen's monograph revealed nonvicioid features, shapes of the other genera as well as some Trigonel/a especially the circular to elliptical sunken hilum sur­ species are regular because of lack of pressure from rounded by a thick hilar rim (figs. 94, 97). The testa sur­ the legume wall or adjacent seeds, and they are face consists of large irregular plates often con­ mitiform or submit:;orm. The radicle lobe is parallel and "'.. spicuously elevated to conical protuberances and about as long as or longer than the cotyledonary lobe. separated by narrow channels (fig. 95). At higher magnification (fig. 96), there is no evidence of the The surface of the testa is either smooth or infre­ typical papillose pattern of Vicieae. Trivedi at al. (1978) quently tuberculate except in Ononis and Trigonel/a, noted and Illustrated by SEM the same differences be­ where it is mostly tuberculate. A newly named species, tween seeds of Gicereae and Vicieae but retained Gicer Trifolium chilaloense TrlUlin, has a finely warty testa as In Vicieae. Seeds of the other six species revealed shown in a SEM photograph (Thulin, 1976). bizarre testa topography when compared with seeds of Vicieae or other tribes. The unique seeds of Gicer are Testa color patterns range from monochrome to rarely described in detail in Lersten and Gunn (1981). The mottled, and colors are various shades of ocher to micropyle is deltoid and ad nate to the hilum, not the brown for genera other than Trifolium. Seeds of this linear shape of Vicieae. genus range in color through most shades and hues.

11 Seeds as classified by length (parallel to hilum) are shorter than the typical Vicieae papillae. Figure 103 usually small (less than 3 mm in length, rarely to 5 mm shows the typical hilum and surrounding area in T. long). arabica Delile, and figure 104 illustrates mounds and the adjacent papillose surface in more detail. .. The hilum and lens are the color of the testa and the hilum is punctiform; thus both are inconspicuous. Both In Meli/otus the testa surface patterns are further are situated between the cotyledon and radicle lobes. removed from tho typical Vicieae pattern. Papillae are The hilar groove is present but not conspicuous. There low or obscure and rather irregular (fig. 106). Mounds is neither a persistent funiculus nor a funicular were noted in three species (table 2). remnant. No mounds were seen in the five Medicago species ex­ Seeds of the Medicago species often have a partial rim amined, and the testa surface was mostly of low (fig. around the hilum, which is lacking in other genera. 107) or obscure (figs. 108, 109) papillae. Mounds were also lacking in Trifolium, and either the testa had Within thp. testa are two well-developed equal obscure papillae or no pattern could be discerned (fig. cotyledons and a curved embryonic axis. Factorovskya 110). aschersoniana (Urban) Eig is unusual in this tribe because the radicle "is thicker, wider and longer than Two little-known genera of Trifolieae are Factorovskya the cotyledons, so that the outstanding and visible part and Parochetus. Factorovskya aschersoniana in gross of the seed, which in other Trifolieae corresponds to appearance resembles other Tr!folieae (fig. 111), with a the cotyledons, represents ... the radicle and the lower circular hilum, hilar rim, and deltoid micropyle (fig. 112). and thinner part represents the cotyledons" (E!g, 1927). The testa surface exhibits low, rounded papillae similar The endosperm is easily discerned. to those seen in Trigol/e/la (fig. 113). Seeds of Parochetus communis Buchanan-Hamilton ex D. Don Testa characters that appear to celimit the Vicieae can also show typical Trifolieae features of seed shape and be seen to merge in various ways with the Trifolieae. hilum (figs. 114, 115). The testa surface, however, was Ononis and Trigone/la are most similar to Vicieae and almost completely smooth and featureless (figs. 115, Trifolium and Parochetus the least. 116), similar to that of Medicago and Trifolium seeds.

Seeds of 27 species of 7 genera of Trifolieae were ex· In Trifolieae the hilum and testa as seen in transec­ amined, using the same methods as for Vicieae seeds. tional view showed nothing different from these Table 2 summarizes our observations, and figures characters in Vicieae. The pits of the tracheid bar were 102-110 and 111-116 illustrate representative features. mostly warty, with usually some nonvestured pits and, in seven species, some mildly vestured pits. Figure 105 Concerning the hilum, one species (Ononis pubescens of Meli/utus messanensis (Linnaeus) Allioni shows the Linnaeus) had a slightly oval hilum, a greatly reduced maximum vesturing seen, which is about equal to that hilar rim, and a micropyle reminiscent of those seen in noted in Vicieae (fig. 53). Overall, pits in Trifolieae were Vicieae (fig. 102). All other Trifolieae species had a cir· somewhat more ornamented than in Vicieae except cular hilum, conspicuous hilar rim (figs. 103, 108), and a Trigone/la. broad deltoid micropyle (fig, 108), features not seen in Vicieae.

The six Ononis species examined had conspicuous papillae identical to the standard papillae of Vicieae seeds (fig. 9). LaSota et al. (1979) have also illustrated .­ this in O. mitissima Linnaeus and O. pubescens. In ad­ dition, three species had mounds similar to those of Vicieae (figs. 22, 24). They have been illustrated by LaSota et al. using SEM and earlier by Watson (1948) using light microscopy. Rowson (1952) showed ',hat these "epidermal protuberances" in O. arvensis Un­ naeus resulted from locally elongated malpighian cells, as in Vicieae.

All four Trigone/la species had papillae, and three also had mounds (table 2). The papillae were broader and

12 Literature Cited

ALEXANDROV, W. G., and ALEXANDROVA, O. G. 1935. ANATOMY OF FLOWER, FRUIT, AND SEED OF PEAS 1970. A KEY AND DIAGRAMS FOR THE SEEDS OF ONE HUN· (GENUS PISUM TOURN.). Bul. Appl. Bot., Genet., DRED SPECIES OF VICIA (LEGUMINOSAE). Internatl. and Plant Breeding, Leningrad, Ser. 3, 9: 7-151. Seed Testing Assoc. Proc. 35: 773-790. ASCHERSON, P., and GRAEBNeR, P. 1909. SYNOPSIS DER MITTELEUROPAISCHEN FLORA. V.6 1971. SEEDS OF NATIVE AND NATURALIZED VETCHES OF (2),1,093 pp. W. Engelmann, Leipzig. NORTH AMERICA. U.S. Dapt. Agr. Agr. Handb. 392, BAAS, P., and ZWEYPFENNING, R. C. V. J. 42 pp. 1979. WOOD ANATOMY OF THE LYTHRACEAE. Acta Bot. Neerland. 28: 117-155. 1981. SEEDS OF LEGUMINOSAE. In Polhill, R. M., and BALL, P. W. Raven, P. H., eds., Advances in Legume 1968. VICIA L. In Tutin, T. G., et aI., Flora Europaea, v. Systematics, pt. 2, pp. 913-925. Internatl. 2, pp. 129-136. Cambridg'3 Univ. Press, Cam· Legume Conf., Kew, Proc. 1978, v. 2. Min. Agr., bridge, England. Fisheries and Food, Richmond, England. BARIOLA, R. HARDHAM, A. R. 1912. SULL 'ANATOMIA DEL SEME DELL' ABRUS 1976. STRUCTURAL ASPECTS OF THE PATHWAYS OF PRECATORIUS L. (JEQUIRITY) E DEI SEMI USATI PER NUTRIENT FLOW TO THE DEVELOPING EMBRYO AND SOF!STICARLO. Atti 1st. Bot. Univ. Pavia, Ser. II, COTYLEDONS OF PISUM SATIVUM L. Austral. Jour. 16:1-16. Bot. 24: 711-722. BASSLER, M. HARZ, C. D. 1966. DIE STELLUNG DES SUBGENUS OROBUS (L.) BAKER IN 1885. LANDWIRTHSCHAFTLICHE SAMENKUNDE. V. 2, DER GATTUNG LATHYRUS L. UNO SEINE SYSTEM· Vicieae, pp. 639-688. Paul Parey Verlag, Berlin. ATISCHE GLIEDERUNG. Feddes Repert. 72: 69-97. HERMANN, F. J. BEHL, H. M., and TIAGI, B. 1960. VETCHES OF THE UNITED STATES-NATIVE, 1977. DEVELOPMENTAL ANATOMY OF SEED AND FRUIT IN NATURALIZED, AND CULTIVATED. U.S. Dept. Agr. CICER ARIETINUM LINN. Indian Bot. Soc. Jour. 56: Agr. Handb. 168, 84 pp. 313-321. HITCHCOCK, C. L. BROUWER, W., and STAHLlN, A. 1952. A REVISION OF THE NORTH AMERICAN SPECIES OF 1955. HANDBUCH DER SAMENKUNDE. 650 pp. DLG· LATHYRUS. Wash. Univ. Pubs. BioI. 15: 1-104. Verlags·GmbH., Frankfurt. HUTCHINSON, J. CAPITAINE, L. 1964. THE GENERA OF FLOWERING PLANTS. 516 1912. LES GRAINES DE LEGUMINEUSES. 455 pp. Emile pp. Oxford Univ. Press, Oxford, England. Larose, Paris. IANNELLI, P. CHALON, J. 1964. VARIETY TESTING OF VETCHES. Internatl. Seed 1875. LA GRAINE DES LEGUMINEUSES. Soc. Sci., Arts, et Testing Assoc. Proc. 29 (4): 887-907. Let. Hainaut, v. 3, No. 10, pp. 55-115. JUSTICE, O. L. CHOWDHURY, K. A., and BUTH, G. M. 1952. TESTING AGRICULTURAL AND VEGETABLE 1970. SEED COAT STRUCTURE AND ANATOMY OF INDIAN SEEDS. U.S. Dept. Agr. Agr. Handb. 30, 440 pp. PULSES. Unn. Soc. London, Jour., Bot. 63 (sup. KONDO, M. 1): 169-179. 1913. DER ANATOMISCHE BAU EINIGER AUSLANDISCHER CLARKE, G. C. S., and KUPICHA, F. K. HULSENFRUCHTE, DIE JETZT VIEL IN DEN HANDEL 1976. THE RELATIONSHIPS OF CICER L. (LEGUMINOSAE): KOMMEN. Ztschr. f. Untersuch. der Nahr. u. THE EVIDENCE FROM POLLEN MORPHOLOGY. Linn. Genussmtl. 25: 1-56. Soc. London, Jour., Bot. 72: 35-44. KORSMO, E. DAVIS, P. H. 1935. WEED SEEDS. 175 pp. Gyldendal Norsk Forlag, 1970. FLORA OF TURKEY. V. 3, 628 pp. Edinburgh Oslo. Univ. Press, Edinburgh. KOSTRAKIEWICZ, K. DUDIK, N. M., and KONDRATYUK, E. N. 1951. STUDIA SYSTEMATYCZNE NAD POLSKIMI GATUljKAMI 1970. FRUIT AND SEED ATLAS OF LE:GUMINOSAE OF RODZAJU VICIA L. Polska Akad. Umiejetnosci, • NATURAL FLORA IN THE UII.. rllNIAN SSR. 215 pp. Physiographica Polon. Doc. 27: 1-71. Naukola Dumka, Kiev, Russia. KUHN, G. EIG, A. 1927. BElT RAGE ZUR KENNTNIS DER INTRASEMINALEN 1927. A SECOND CONTRIBUTION TO THE KNOWLEDGE OF LEITBUNDEL BEl DEN ANGIOSPERM EN. Bot. Jahrb. THE FLORA OF PALESTINE. Inst. Agr. and Nat. 61: 325-379. His!., Agr. Expt. Sta. Bul. 6, 88 pp., 6 pis, KUPICHA, F. K. GUNN, C. R. 1973. STUDIES IN THE VICIEAE I.: THE NEW GENUS 1968. THE VICIA AMERICANA COMPLEX (LEGUMINOSAE). ANATROPOSTYLIA. Roy. Bot. Gard., Edinb. Notes Iowa State Jour. Sci. 42 (3): 171-214. 32: 247-250.

1969. GENERA, TYPES AND LECTOTYPES IN THE TRIBE 1976. THE INFRAGENERIC STRUCTURE OF VICIA. Roy. VICIEAE. Taxon 18: 72S~733. Bot. Gard.., Edinb. Notes 34: 287-326.

13 KUPICHA, F. K. MUSIL, A. F. 1977. THE DELIMITATION OF THE TRIBE VICIEAE 1963. IDENTIFICATION OF CROP AND WEED SEEDS. U.S. (LEGUMINOSAE) AND THE RELATIONSHIPS OF CICER Dept. Agr. Agr. Handb. 219, 171 pp. L. Linn. Soc. London, Jour., Bot. 74: 131-162. PAM MEL, L. H. 1899. ANATOMICAL CHARACTERS OF THE SEEDS OF 1981. TRIBE VICIEAE and TRIBE CICEREAE. In Polhill, R., LEGUMINOSAE, CHIEFLY GENERA OF GRAY'S and Raven, P., eds., Advances in Legume MANUAL. Acad. Sci. St. Louis, Trans. 9: 91-263, Systematics, pt. 1, pp. 377-382. Internatl. 7 tables, 35 pis. Legume Conf., Kew, Proc. 1978, v. 2. Min. Agr., PATEL, J. D. Fisheries and Food, Richmond, England. 1976. COMPARATIVE SEED COAT ANATOMY OF SOME IN­ LASOTA, L. R., LINK, C. B., and GUNN, C. R. DIAN EDIBLE PULSES. Phyton (Austria) 17: 1979. SEM IN THE STUDY OF COMPARATIVE TESTA MOR­ 287-299. PHOLOGY. SEM 1979, III: 231-236. PIERPOINT, M. LASSETTER, J. S. 1949. CERTAIN VETCHES AND SIMILAR PLANTS. In Assoc. 1972. A BIOSYSTEMATIC STUDY OF THE VICIA LUDOVICIANA Off. Seed AnalystlO, Contribution to the Hand­ COMPLEX (LEGUMINOSAE). 73-9458, Univ. Micro­ book on Seed Testing, 7 pp. Queen's Printer, films, Ann Arbor, Mich. Ottawa. PITOT, A. 1975. TAXONOMIC STATUS OF VICIA HASSEl 1936. ISOLEMENT ET CHUTE DE LA GRAINE A MATURITE (LEGUMINOSAE). Madroiio 23: 73-78_ CHEZ LES LEGUMINEUSES. 233 pp. Impr. de la __ and GUNN, C. R. Charite, Montpellier. 1979 VICIA MENZIESII SPRENGEL (FABACEAE) RED IS­ POLHILL, R. M .• [1980J. COVERED: ITS TAXONOMIC RELATIONSHIPS. Pacific 1981. PAPILIONOIDEAE and ABREAE. In Polhill, R., and S"Ci. 33: 85-101. Raven, P., eds., Advances in Legume System­ LE MONNIER, G. atics, pt. 1, pp. 191-208, 243-244. Internatl. 1872. RECHERCHES SUR LA NERVATION DE LA GRAINE. Legume ConL, Kew, Proc. 1978, v. 2. Min. Agr., Ann. des Sci. Nat., Bot., Ser. V, 16: 233-305. Fisheries and Food, Richmond, England. L:;:gSTEN, N_ R. QUARTLEY, C. E. 1979. A DISTINCTIVE SEED COAT PATTERN IN THE VICIEAE 1968a. IDENTIFICATION OF VICIA SPP. Internatl. Seed (PAPILIONOIDEAE; LEGUMINOSAE). Iowa Acad. Sci. Testing Assoc. Proc. 33 (2): 329-330. Proc. 86: 102-104. 1968b. KEY TO 7 SPECIES OF VICIA and LATHYRUS. Inter­ 1982. TRACHEID BAR AND VESTURED PITS IN LEGUME natl. Seed Testing Assoc. Proc. 33 (2): 331-334. SEEDS (LEGUMINOSAE: PAPILIONOIDEAE). Amer. RAO, T. A., and DAS, S. Jour. Bot. 69: 98-107. 1976. ON IDIOBLASTS IN A FEW TAXA OF PENAEACEAE. __ and BENDER, C. G_ Cur. Sci. 45: 750-752. 1975. TRACHEOID IDIOBLASTS IN CHENOPODIACEAE: A RED'KINA, Z. V., and KHOROSHAILOV, N. G. • REVIEW AND NEW OBSERVATIONS ON SALICORNIA 1974. A GUIDE FOR THE IDENTIFICATION OF SEEDS OF THE VIRGINICA. Iowa Acad. Sci. Proc. 82: 158-162. USSR WEEDY VETCH SPECIES. Appl. Bot., Genet., __ and GUNN, C. R. and Plant Breeding Trans. 51: 3-46. 1981. SEED MORPHOLOGY AND TESTA TOPOGRAPHY IN REEVE, R. M. CICER (FABACEAE, FABOIDEAE). System. Bot. 6: 1946. ONTOGENY OF THE SCLEREIDS IN THE INTEGUMENT 223-2.30. OF PISUM SATIVUM L. Amer. Jour. Bot. 33: LHOTSKA, M., and CHRTKOVA, A. 806-816. '. 1978. KARPOLOGIE A DIASPOROLOGIE: CESKOSLOVEN­ ROWLAND, G. G. SKYCH ZASTUPCU CELEDI FABACEAE. 296 pp. 1977. SEED COAT THICKNESS AND SEED CRUDE FIBRE IN Academia, Prague. FABA BEANS (VICIA FABA). Canad. Jour. Plant Sci. McEWEN, T. J., DRONZEK, B. L., and BUSHUK, W. 57: 951-953. 1974. A SCANNING ELECTRON MICROSCOPE STUDY OF ROWSON, J. M. FABABEAN SEED. Cereal Chem. 51: 750-757. 1952. THE HYPODERMAL ZONE IN THE TESTA OF CERTAIN MARBACH, I., and MAYER, A. M. LEGUMINOUS SEEDS. Roy. Micros. Soc. Trans. 72: 1974. PERMEABILITY OF SEED COATS TO WATER AS 46-55. RELATED TO DRYING CONDITIONS AND METABOLISM SAINT-MARTIN, M. OF PHENOLICS. Plant Physiol. 54: 817-820. 1978. OBSERVATIONS SEMINOLOGIOUES AU MICROSCOPE MATTIROLO, 0., and BUSCALIONI, L. ELECTRONIOUE A BALAYAGE DE DIVERSES ESPECES 1892. RICERCHE ANATOMO-FISIOLOGICHE SUI TEGUMENTI DE PAPILIONACEES. [ParisJ Acad. des Sci. Compt. SEMINALI DELLE PAPILIONACEE. R. Accad. delle Rend., Ser. D, 287: 927-930. ScI. Torino II Mem. 42: 223-318, 359-445. SIMOLA, L. K. MILLER, R. B. 1968. COMPARATIVE STUDIES ON NUMBER OF LEAFLETS, 1977. VESTURED PITS IN BORAGINACEAE. IAWA Bui. VENATION, AND EPIDERMAL STRUCTURE IN THE 1977: 43-48. GENUS LATHYRUS. Canad. Jour. Bot. 46: 71-84.

14 SPURNY, M. ZIMMERMANN, K. 1963. CELL WALL STRUCTURE OF EPIDERMAL CEI.LS OF 1936. ZUR PHYSIOLOGISCHEN ANATOMIE DER LEGUMINo­ THE PEA SEED COAT (PISUM SATIVUM L.) STUDIED BY SENTESTA. Landw. Verso Sta. 127: 1-56. MICROCINEMATOGRAPHY. Mikroskople 18: 272-279. STEIN VON KAMIENSKI·JANCKE, I. 1957. UNTERSUCHUNGEN UBER BAU, CHEMISMUS UNO SUB­ MIKROSKOPISCHE STRUKTUR DER PALISADENSCHICHT VON LEGUMINOSENSAMENSCHALEN. Mikroskopie 12: 357-392. SWEDERSKI, W. 1924. SUR LA DETERMINATION Di:S GRAINES DES ESPECES DE VICIA L. D'APRES LA GI~ANDEUR DE L'HILE. Rocz. Nauk Rolnicz. i Lesnych 11 (1): 44-51. THULIN, M. 1976. TWO NEW SPECIES OF TRIFOLIUM FROM ETHIOPIA. 'r Bot. Notiser 129: 167-171. TRIVEDI, B. S., BAGCHI, G. D., and BAJPAI, U. 1978. SPORODERM PATTERN IN SOME TAXA OF VICIEAE (PAPILIONA TAE-LEGUMINOSAE). Phytomorphology 28: 405-410. TUPIKOVA, A. 1926. BOTANICO-AGflONOMICAL INVESTIGATIONS OF AN­ NUAL VETCHES. Bul. Appl. Bot., Genet., and Plant Breeding 16 (1): 151-246. UTKIN, W. W. 1965. SEEDS OF WILD VETCHES FROM THE CRIMEA. Nauch. Dok. Vysshei Shkoly BioI. Nauk. 2: 101-110. VAN DER MAESEN, L. J. G. 1972. CICER L., A MONOGRAPH OF THE GENUS, WITH SPECIAL REFERENCE TO THE CHICKPEA (CICER ARIETINUM L.), ITS ECOLOGY AND CULTIVATION. 342 pp. H. Veenman & Zonen, Wageningen. VAN VLIET, G. J. C. M. 1978. VESTURED PITS OF COMBRETACEAE and ALLIED FAMILIES. Acta Bot. Neerland. 27: 273-285. VERDCOURT, B. 1980. STUDIES IN THE LEGUMINOSAE-PAPILIONOIDEAE FOR THE FLORA OF TROPICAL EAST AFRICA: II. Kew Bul. 24: 235-253. VODAK, A. 1956. SEMENA NEBO PLODY NASICH KULTURNICH ROSTLIN A NEJCASTEJSICH PLEVELU. 241 pp. Ceskoslov. Akad. Zem~del., Prague. WATSON, D. P. 1948. STRUCTURE OF THE TESTA and ITS RELATION TO • GERMINATION IN THE PAPILIONACEAE TRIBES TRIFOLIEAE and LOTEAE. Ann. Bot. 12: 385-409. WITTMACK, L. 1922. LANDWIRTSCHAFTLICHE SAMENKUNDE. 58'l pp. Paul Parey, Berlin. ZERTOVA, A. 1962. EIN SCHUSSEL ZUR BESTIMMUNG DER TSCHECHOSLowAKISCHEN ARTEN DER GAT'TUNG VICIA L. NACH DEN MORPHOLOGISCHEN MERKMALEN DEA SAMEN. Acta Hort. Bot. Pragensls 1962; 113-118,

15 Appendix

Historical References Pammel, L. H. 1899. Anatomical characters of the In addition to the authors cited in the text, other seeds of Leguminosae, chiefly genera of Gray's workers have reported on other aspects of Vicieae manual. Acad. Sci. St. Louis, Trans. 9: 91-263, 7 testa anatomy. Nineteenth century work is cited in tables, 35 pis. Pammel and in Mattirolo and Buscalionl, but neither of Pringsheim, N. 1848. De forma et incremento these reports is easily obtained, and some of their stratorum crassiorum in plantarum cellula, obser­ references are cited incompletely or incorrectly. vationes quedam novae. Linnea 21: 145-180. (Not Coupled with the preceding Literature Cited, these an· seen. Pammel says he described testa of notated citations form a comprehensive (although prob­ unspecified number of Vicieae.) ably not complete) listing of work on the anatomy of Reeve, R. M. 1946. Structural composition of the Vicieae testa. sclereids in the integument of Pisum sativum L. Beck, G. 1878. Vergleichende Anatomie der Samen Amer. J. Bot. 33: 191-204. von Vicia und Ervum. Sitzber. K. Akad. der Wiss. Sempolowski, A. 1874. Ueber den Bau der Schale land­ Wi en 78: 93-94. (Does not indicate number of wirtschaftlich wichtiger Samen .. Landw. Jahrb. 3: species studied. Found a light line in both genera, 823-866. (Includes Pisum and Vicia.) notably a double one in Vicia bivonea Rafinesque.) Spurny, M. 1954a. Cell wall structure of the epidermal Dahmen, M. 1892. Anatomisch-Physiologische Unter­ cells of the pea seed coat (Pisum sativum L.). suchung uber den Funiculus der Samen. Jahrb. f. Preslia 26: 79-88. Wiss. Bot. 23: 441-478. (Pisum, Vicia included. Spurny, M. 1954b. The "light line" of the epidermal Calcium oxalate crystals may occur in funiculus.) cells of the pea seed coat. (Pisum sativum L.) Godfrin, J. 1880. Etude histologique sur les teguments Preslia 26: 139-142. seminaux des angiospermes. Soc. des Sci. de Spurny, M. 1954c. The permeability of testa and the in­ Nancy Bul. 5: 109-219. (Includes Lathyrus (Orobus) equality in the swelling velocity of the pea seeds vernus (Linnaeus) Bernhardi and Vicia faba.) (Pisum sativum L.). Preslia 26: 239-262. Holtert, J. 1890. Die Nahrschichte der Samenschalen. Spurny, M. 1964. Changes in the permeability of the Flora 73: 279-313. (Brief description of Pisum seed coat in connection with the development of sativum, Vicia faba, Lathyrus pratensis.) suberin adcrustations of the macrosclereids from Junowicz, R. 1878. Die Lichtlinie in den Prismenzellen the seed coat of the pea (Pisum sativum L.). Flora der Samenschalen. Sitzber. K. Akad. der Wiss. 154: 547-567. Wien, Abt. 1a, 76: 335-352. 2 pis. (Includes Streicher, O. 1902. Beitrage zur vergleichenden Lathyrus verna: Malpighian cells-shows light Anatomie der Vicieen. Beih. Bot. Centbl. 12: line.) 483-598. (Not illustrated. Mentions species of Lindemuth, K. 1924. Beitrag zur 8iologie von Vicia hir­ Gicer, Vicia, Lens, Lathyrus, Pisum. Only 2 pages suta Koch und ihre Bedeutung als land­ on seeds, mostly referring to the work of others.) wirtschaftliches Unkraut. Bot. Arch. von Mez 7: Yocum, L. E. 1922. Some phases o~ structure and 195-251. (Brief section on testa.) development of garden pea and white sweet clover Macchiati, L. 1891. Nota preventiva sulla morfologia seeds as related to hardness. Elisha Mitchell Sci. ed anatomia del ~eme della Vicia narbonensis. Soc. Jour. 38: 76-83. .. Nuovo. Gior. Bot. Ital. 23: 150-157. (Testa said to be essentially like that described for other .. legumes. No illustrations.) Taxonomic List oU Vicieae Taxa in Table 1 Mattirolo, O. 1885. La linea lucida nelle cellule malpighiane degli integumenti seminali. Roy. Vicia Linnaeus Acad. Sci. Torino Mem. II, 37: 219-246. (Reports a Subg. Vicilla (Schur) Rouy light line from species of Pisum and Vicia, in­ Sect. Vicilla (Schur) Ascherson & Graebner cluding Ervum.) amoena Seringe Mattirolo, 0., and Buscalioni, L. 1889. Sulla strutiura amurensis Oettel degli spazii intercellular! nei tegumenti seminali dumetorum Linnaeus delle Papilionaceae. Malpighia 3: 143-159. (In­ pal/ida Turczaninow cludes Pisum, Vicia, Lathyrus.) sy/vatica Linnaeus Mattirolo, 0., and Busf'alloni, L. 1892. Ricerche unijuga A. Brown anatomo-fisiologiche sui tegumenti seminali delle Sect. Gassubicae Radzhi Papilionacee. Roy. Acad. Sci. Torino Mem. 11,42: cassubica Linnaeus 223-318, 359-445, (Review of all previous work. In­ nigricans Hooker & Arnott cludes original work on a few cultivated members subsp. gigantea (Hooker) Lassetter & Gunn of Vicleae.) semig/abra Bolssier

16 Sect. Cracca Oumort odoratus Linnaeus benghalensis Linnaeus sylvestris Linnaeus caroliniana Walter tingitanus Linnaeus cracca Linnaeus tuberosus Linnaeus disperma de Candolle Sect. Cicercula Medikus f/oridana S. Watson annuus Linnaeus hirsuta (Linnaeus) S. F. Gray cicera Linnaeus leavenworthii Torrey & Gray gorgoni Parlatore ludoviciana Nuttall hirsutus Linnaeus meyeri Boissier numidicus Battandier villosa Roth sativus Linnaeus subsp. varia (Host) Corbiere Sect. Clymenum Adanson Sect. Variegatae Radzhi articulatus Linnaeus megalotropis Ledebour clymenum Linnaeus Sect. Volutae Kupicha ochrus (Linnaeus) de Cando lie biennis Linnaeus Sect. Aphaca Adanson Sect. Ervum (Llnna9us) Taubert aphaca Linnaeus tetrasperma Moench Sect. Nissolia Adanson Sect. Ervilia (Link) W. Koch nissolia Linnaeus ervilia Willdenow Sect. Orobastrum Taubert Sect. Anatropostylia Plitmann cassius Boissier koeieana. Rechinger f. inconspicuus Linnaeus Sect. Australes Kupicha japonicus Willdenow graminea Smith subsp. maritimus (Linnaeus) P. W. Ball Sect. Mediocinctae Kupicha pubescens Hooker & Arnott leucophaea Greene Subg. Orobus (Linnaeus) Baker Subg. Vicia Sect. Orobus Sect. Atossa (Alefeld) Ascherson & Graebner Ser.Orobi sepium Linnaeus aureus (Steven) Brandza Sect. Vi cia gmelinii (Fischer) Fritsch grandiflora Scopoli Ser. Verni Fritsch lathyroides Linnaeus venetus (Miller) Wohlfahrt sativa Linnaeus vernus (Linnaeus) Bernhardi subsp. amphicarpa (Oorthes) Ser. Nigri Fritsch Ascherson & Graebner niger (Linnaeus) Bernhardi saliva Linnaeus Ser. Pisiformes B~ssler subsp. sativa pisiformis Linnaeus Sect. Faba (Miller) Ledebour Sect. Platystylis (Sweet) B~ssler bithynica Linnaeus boissieri Sirjaev faba Linnaeus digitatus (M. Bieberstein) Fiori Sect. Hypechusa (Albfeld) Ascherson & Graebner Sect. Pratensis B~ssler galatea Boissier pratensis Linnaeus lutea Linnaeus Sect. Orobon Tamamschjan melanops Sibthorp & Smith roseus Steven • pannonica Crantz Sect. "North America" Sect. Peregrinae Kupicha delnorticus C. L. Hitchcock michauxii Sprengel holochlorus (Piper) C. L. Hitchcock Sect. unknown jepsoni Greene costata Ledebour nevadensis S. Watson Lathyrus parvifolius S. Watson Subg. Lathyrus polyphyl/us Nuttall ex Torrey & Gray Sect. Lathyrus pusillus Elliott clrrhosus Seringe vestitus Nuttall ex Torrey & Gray grandiflorus Sibthorp & Smith subsp. puberulus (White ex Greene) heterophyllus Linnaeus C. L. Hitchcock latifolius Linnaeus Sect. "South America"

17 ...

magel/anicus Lamarck Ascherson & Graebner Sect. unknown sativum Linnaeus drummondii Hortus subsp. sativi.lm leptophyllus M. Bieberstein Lens Miller Pisum Linnaeus culinaris Medikus fulvum Sibthorp & Smith Vavilovia A. Fedorov sativum Linnaeus formosa (Steven) A. Fedorov subsp. elatius (M. Bieberstein)

Table 1.-Summary of testa characters of Vicieae species in SEM surveyl

Surface Tracheid Taxa Near hilum Midseed bar pits Comment

Lathyrus annuus ------Areolate mounds ------Areolate mounds ------Nonvestured------Unusual surface pattern. aphaca ------PapiIlose ------Papillose------do------Lens of elongate malpighian cells. articulatus ------do------do------Nonvestured to slightly warty. aureus ------Papillose (irregular) ------do------Not observed ------­ boissieri ------Papillose------Fades near hilum ------Nonvestured ------­ cassius------Mounds, ridges ------Mounds, ridges ------do------Papillae mOdified. ~ cicera ------Papi lIose ------Papi lIose (fai nt Iy) ------do------­ cirrhosus ------Papillose, mounds ------Papillose, mounds ------do------Papillae sharply pointed. '( clymenum ------Papillose------Papillose------Nonvestured to warty ----­ delnorticus------do------do------Nonvestured to rather warty. digitatus------do------Nonpapillose ------Warty to slightly vestured. drummondii ------do------Papillose------Nonvestured to slightly vestured. gmelinii ------Papillose (irregular) ------Fades near hilum ------Nonvestured------Lens of elongate malpighian cells. gorgon i ------Pap i lIose, pi tted ------Papi lIose, pi tted ------do------­ grandiflorus ------Papillose, mounds ------Papillose, mounds ------do------­ heterophyllus ------do------do------do------­ hlrsutus ------Papillose------PapilIose (faintly) ------Not observed ------­ holoch lorus ------do------Papi Ilose ------Nonvestured to rather warty. inconspicuus------do------do------Nonvestured to warty -----Papillae of unusual appearance. japonicus ------do------Papillose (faintly) ------Not observed ------­ iepsoni ------do------PapilIose ------Nonvestured to rather warty. latifolius------Papillose, faint mounds ---Papillose, faint mounds ---Nonvestured------­ lep toph yllu s------PapiII ose ------PapiIlose------N onves tu red to slightly vestured. magellanicus-----Papillose (faintly) ------Fades close to hilum ------Nonvestured------­ nevadensls------PapilIose ------Papi 1I0se ------Nonvestured to rather warty. niger ------do------NonpapiIlose ------Nonvestured------Lens of elongate malpighian cells. nlssolla------Mounds, pits, papillose Mounds, pits------Warty------Unusual surface pattern. In narrow zone near hilum. See footnote at end of table.

18 Table 1.-Summary of testa characters of Vicieae species in SEM survey'-Continued

Surface Tracheid Taxa Near hilum Midseed bar pits Comment

numidfcus ------Papillose, pits------Papillose, pits------Nonvestured ------Papillae sharp pointed. ochrus ------Papi Ilose ------Pap ill ose -----·-----·------do------­ odoratus------Papillose, mounds ------Papillose, mounds ------do------Malpighian cells large, thin walled; lens of elongate mal pig hi an cells. parvifolius ------PapilIose ------Papillose------Nonvestured to slightly vestured. pisfformis------Papillose (low, crimped) ---Fades close to hilum ------Nonvestured to warty -----Papillae resemble those of Pisum. polyphyllus ------Pap ill ose ------PapiIlose ------do------­ pratensis ------do------Nonpapillose ------Nonvestured ------­ pubescens ------do------Papillose (faintly) ------Norwestured to warty ----­ pusillus------Mounds, ridges ------Papillose, many ridged ------do------Unusual surface pattern. roseus------Papi Ilose ------Non papi Ilose ------Nonvestu red ------­ sativus ------do------do------Nonvestured to warty -----Lens of elongate malpighian cells. sylvestris ------Papillose, mounds ------Papillose, mounds ------Nonvestured------­ tingitanus------Papillose, faint mounds ---Papillose------do------Lens of elongate tuberosus------Papillose, mounds ------Papillose, mounds ------Nonvestured to slight malpighian cells. mounds. venetus------PapiIlose ------PapilIose ------Not observed ------­ vernus------do------do------Nonvestured to warty ----­ vestitus------do------do------Nonvestured ------Lens culinaris ------Papillose (low) ------Papillose (low) ------do------­ Pisum f u Ivu m ------Pap ill ose ------Pap i Ilose ------do------­ sativum subsp. elatius --Papillose, mounds ------Papillose, mounds ------do------­ sativum subsp. sa tivum - Papi Ilose ------PapiIlose ------do------­ Vavilovia formosa ------do------do------Not observed ------­ V{cia amoena ------do------Papillose (faintly) ------do------­ am urensis ------do------·------do------Nonves tu red ------­ benghalensis ------d0 ------do------N ot observed ------­ biennis ------do------Papi Ilose ------N onves tu red ------­ bithynica ------do------do------Nonvestured to slightly warty. caroliniana------c.'o------do------Nonvestured to warty ----­ cassublca ------do------do------Nonvestured to slightly vestured. costata ------do------do------Nonvestured to slightly warty. cracca------do------do------Nonvestured to warty ----­ dlsperma ------do------do------Nonvestured to slightly warty. dumetorum ------do------do------Not observed ------­ ervllia ------do------do------Nonvestured to slightly warty. faba ------Papillose (falntly)------Nonpapillose ------Nonvestured to slightly vestured. See footnote at end of table.

19 Table 1.-Summary of testa characters of Vicieae species in SEM survey1-Continued -.------Surface Tracheid Taxa Near hilum Midseed bar pits Comment

floridana------do------do------Nonvestured------­ galatea------do------do------do------­ graminea ------do------do------Nonvestured to slightly warty. granditlora ------do------do------Not observed ------Rim-aril associated with hilum. hlrsuta ------do------do------do------­ koeieana ------Papillose, mounds ------Papillose, mounds ------Nonvestured------Circular hilum, heavy hilar rim. lathyroides------do------do------Not observed ------­ leaven worthii-----Papillose ------PapilIose------Nonvestured to sl ightly warty. leueophaea ------do------do------do------­ ludovieiana ------do------do------do------­ lu tea ------do------do------Nonvestured ------­ megalotropis ------do------do------do------­ melanops ------do------do------Nonvestured to slightly warty. meyeri------do------do------do------­ miehauxii------do------do------Nonvestured------­ nigr{eans subsp.gigantea------do------do------do------­ pallida ------do------do------Nonvestured to slightly warty. pannoniea ------do------do------Not observed ------­ sativa subsp. amphiearpa ------do------do------Nonvestured to slightly vestured. sativa subsp. sativa ------do------do------Nonvestured to slightly warty. semiglabra ------do------Nonpapillose ------Nonvestured to vestured--­ seplum ------do------Papillose------Not observed ------­ sylva t lea ------do------d0 ------Nonves tu red to vestu red --­ letrasperma ------do------do------Nonvestured to slightly warty. unljuga ------do------Papiliose (faintly) ------do------­ vII losa ------do------PapiIlose ------Nonvestu red ------­ subsp. varia ------do------do------·------do------­ subsp. villosa------do------do------do------­

'For taxonomy of Vlcleae taxa, see p. 16.

20 Table 2.-Summary of testa characters of Trifo/ieae species in SEM survey

Surface Tracheid Taxa Near hilum Midseed bar pits

Faclorovskya aschersoniana ------Papi1I0se (low) ------Papillose (Iow)------Not observed. Medicago cancel/ala ------do ------do ------Do. mesopolamica------Papillose (obscurely) ------Nonpapillose------Nonvestured to warty. pironae ------Papillose (Iow)------Papillose (Iow)------Warty to slightly vestured. plalycarpa ------do ------do ------Nonvestured to slightly warty. sa /iva ------Papillose ------NonpapiIlose------Nonvestured to warty. Melilotus alba ------Papillose (irregular) ------Papillose (irregular) ------Nonvestured to slightly warty. hirsuta ------Papillose (Iow)------·------Nonpapillose------Nonvestured to slightly vestured. italica ------Papillose (obscurely), mounds--- Nonpapillose, mounds------Do. messanensis ------Papillose (low), mounds ------Papillose (low), mounds------Nonvestured to vestured. neapolitana ------do ------do ------Do. Ononis adenolricha ------Papillose ------Papillose ------Not observed. bit/ora ------Papillose, mounds ------Papillose, mounds ------Nonvestured to slightly vestured. hirsula ------Papillose ------··------Nonpapillose------Nonvestured. mitissima ------Papillose, mounds ------Papillose, mounds ------Nonvestured to slightly vestured. natrix ------do ------do ------Do. pubescens ------Papillose ------Papillose ------Nonvestured to slightly warty. " Parochelus communis ------Nonpapillose------Nonpapillose------Not observed. Trifolium a/exl1ndrinum------Papillose (obscurely) ------do ------Nonvestured to slightly warty. ochroleucon------Nonpapillose------do ------Do. pannonicum------do ------do ------Not observed. repens------Papillose (obscurely) ------do ------Nonvestured to slightly warty. semipilosum ------Nonpapillose------do ------Do. Trigonel/a arabica ------Papillose, mounds ------Papillose, mounds ------Not observed. caerulea ------do ------do ------Nonvestured. foenum-graecum ------do ------do ------Do. pubescens ------Papillose ------Papillose ------Nonvestured to slightly warty.

21 MORPHOLOGY EXTERNAL INTERNAL

HYPOCOTYL MICROPYLE TESTA RADICLE HILUM HILAR GROOVE EPICOTYL MESOCOTYL LIPS HI LAR GROOVE HALO RIM COTYLEDON LENS ..

HILUM LENS

Figure 1.-External and internal morpho­ logical features of seeds in the Vicieae. External views are of different Vicia spp.

22 ORNAMENTATIONS HILA

OBLONG LINEAR ACULEATE RUGOSE

CIRCUMLINEAR TUBERCULATE RIDGED & TUBERCULATE

RUMINATE RUMINATE OVAL WEDGE (LARGE (SMALL INTERSTICES) INTERSTICES) SURFACES

TISSUES

SMOOTH WRINKLED

MARKS MALPIGHIAN LAYER ~ ~ I!.!JJ '[j _SCLEREID STREAKED __-- LAYER REMNANT ----~=: :..:.;.. ,'. ,- :.":.....:::.~. L. AYER S -~ ...... _., •.I'"! .: ----=::::::::::­=-- .~ ~.. ~:.:. MOTTLED POINTED

Figure 2.-Testa and hilar characters of the Vlcleae.

23 I 6

".:it.

FIgures 3-8. -Lathyrus and Vicia seeds selected to Illustrate hilar varia lion (x 20). 3. L nissa/fa Unnaeus: 4, L. niger (Un­ ,. naeus) Bernhardl; 5, L hirsutus Unnaeus; 6. L Japan/cus Wilidenow; 7, V grandiflara Scopoli with nm·aril; 8, L. sy/vestns Un­ naeus

24 Figures 9-1S.-Testa surface patterns in Vicleae ( x 1.000). 9, La thyrus odora tus Llnnaeus, standard papillose pattern; 10, L. numtdlcus Battandler. large conical papillae: 11. Vlcia mlchauxil Sprengel, angular papillae with dimpled top; 12, PlsLlm satlvum Llnnaeus. and 13, P. fulvum Slbthorp & Smith, both with crimped papillae characteristic of this genus; 14. Lathyrus mconspicuus L,nnaeus, low angulM paptllae;IS. L"!ns cu/mans Medikus, low. Irregular. domelike papillae.

25 Figures 16-22.-Testa surface pattern in typical papillose cells between (x 400). L. Vicleae. 16. Vlcia faba Linnaeus. low, odoratus Linnaeus: 19, Low, separate rounded papillae (x 440); 17, Lathyrus mounds (x 60); 20, mounds with papillose bOlssien Slrjaev, inconspicuous papillae cells between (x 200). V, lathyroides lin· restricted to a few eells adjacent to hilum naeus: 21, One mound with interior expos­ (x 1,000): 18, Pisum sativum Linnaeus ed to show elongate malpighian cells (x sub::;p. elatius (M,Bieberstein) Ascherson & 720); 22. surface with mounds and typical Graebner, numerous small mounds with papillose cells between (x 200),

26 Figures 23-29. ~-Mounds and rtdges on (x 1,000); 26, transection through two Lathyrus seeds L mssolta linnaeus. 23, mounds shOWing areas of mc,'pighian cell Part I)f hilum and adjacent testa with small elongation (x 400), L. pus;lIus Eltiott; 27, but abruptly rising mQunds (x 54); 24, Testa surface covered with mostly inter­ enlarqed area of testa showing several connected ridges (x 30); 28. enlarged view mounds and one of the many pits scat with papillose cells (arr .....N) in more com­ tered over 'lead (x 200) 25, enlarged View monly veined Interridge area (x 200): 29. of PIt. whlr.h conSIsts of progressively 10ngisectIOn through one ridge to show shorter malplghlan cells forming locally un~ elongate malpighlan layer with widely Gharar.tBrlstlr. nonpapiltose surface pattern separated sclereids Just below.

27 \

\ t ~

FH1'j!f~" 10 1°) P:t'), mounff,; dntl mlqes f)!1 elongate malplghlan cells t x 180) V l dlhyru" ,11. l Vu;,a ';Hf~(h 10, L numldlCus koeleana ReChtnger f 33. Testa surface Bd!L]ndij.~r ,,,,,td <.!lda!.!' wtlh numerous {x 30).34, enlarqerJ view of Circular hilum ~ jt~ hid HI) rnfJUfld!:) 1)( ndqp<) I x 240) L Govered wIth funicular remnant (X 100).35. donUUS l,nru"q', 11 Hilum dnd ddJd';ent erJqe !)f hilum with adjacent testa 5howtng l p ',ld c .r.JI.,kH"'J d'(junrj (',Jch 1Ti11t)rvJ i :< 40" tYPIcal Pi)plli()<;e surface between mClUncJs "1,( t";JrF)f~(,b,.n (j! r}np rnoqn{j ,,,hch'Vtnq and ',hOft nrlqWi (X 300)

28 Flqures 36-42 ~- Representative hllar transec­ mgncans HOoker & Arnott subsp, gigantea tlons In VICleae, 36. Lens culmans (Hooker) Lassetter & Gunn, flat hilum with Medlkus. extreme elevation with narrowly small tracheid bar (arrow) (x 36); 41, V. elliptical tracheld bar (X 100); 37, Pisum faba Linnaeus, flat hilum with circular [ulvum Slbthorp & Smith. raised hilum tracheld ba r (x 40); 42, La thyrus clymenum ( x 60). 38. VICW cassublca lInnaeus, Llnnaeus, closeup of tracheid bar with sllqhtly ralserJ hIlum (X 120); 39, P sallvum tractlelds short, broad. and uniformly pittee l 'lnaeU5, almost level hilum WIth broadly (X 150), ell.,.\lcal lrachmd bar (x 30); 40, V

29 -

FlijlJres 43-47- Tracheld bars In longitudinal VI,!W tn VI';lf!de 43. Vlcla pall/da Turc­ l"lrlHlO'N ,mJdJqed VI!)W ul hllar edge show mq "vule va<;cular bundle (arrow) In con· telet wIth trdGhf~lcJ bar (x 200). 44. Pls(lm sd/tVum LmnafWS. trm:helrJ bar. extenrJlnq full lenqlh Ilf hilum up It) rnl!:mpyle (arrowl 'x flO!. 4'1. Ld!hyrus qr,md/flor(ls Slblhorp ~. Snllih ';l",HlUP of ovulE! bundle (arrow) In ·.'Jntact With tfactwl{J bar (X 300).46. V VIII()~i1 Af)lh. f!nldrqecJ vlnw .,f Ir.\chmrl b,u ",tll.wtrH) plttlnq pclttnrn (x 4001 47, V IU(jov/cIJn.j Nuttall. tr(1(;heHJ bdr r;AII" with lJl1\1'ltJdlly ttlJt;k wall I x 6001

30 /'V'~ t ---,...~ Jfil"·...~ 48 49

Figures 48-53.-Enlarged views of tracheid nonvestured, regularly distributed; 50, P. bar pits in Vicieae (x 1,000). 48, Pisum sativum Linnaeus, extremely slitlike, non­ (u/vum Slbthorp & Smith, regularly bordered pit occasionally seen on some distributed, conspicuously bordered, tracheids; 51, Vicla pal/ida Turczaninow, nonvestured pits with primary wall rem­ slightly warty pits; 52, L. inconspicuus Lin­ nants on two extreme upper right pits; 49, naeus, warty pits; 53, V. sylvatlca Linnaeus, Lathyrus bo/ssierl Sirjaev, another view of modestly vestured pits representing the the common pit pattern, uniform, most ex.treme condition in Vicleae.

31 Figures 54-60.-Nonhilar testa of Vlcieae. from hilum, an extreme example of thick 54, Lathyrus gmelinii (Fischer) Fritsch, malpighian layer (x 300); 57, V. faba Lin­ overview showing sclereid layer con­ naeus, transection about midseed with thin spicuous near hilum (at top) and becoming malpighian layer and conspicuous elongate progressively shorter toward midseed, the sclereid cells (x 180); 58, V. ludoviciana usual pattern (x 26); 55, Vicia biennis Lin­ Nuttall, transection through lens (x 100); naeus, typical testa transection at 0.5- 1 59, V. sativa Linnaeus, transection through mm distance from hilum (x 200); 56, V. lens (x 150); 60, V. villosa Roth, transection cracca Linnaeus, transection about 1 mm through lens (x 48).

32 ..

• Figures 61-66.-Summary of testa features and papillose surface (x 66); 64, L. hir­ of La thyrus. 61, L. cicera Linnaeus, with sutus Linnaeus, at edge of hilum (across angular, somewhat compressed seed and top) (x 480); 65, L. c;rrhosus, enlarged oval hilum (X 20); 62, L c;rrhosus Seringe, hilum area (across bottom) showing with mounds, ridges, and oblong hilum papillose pattern with mounds (x 100); 66, (X 20); 63, L c;cera, hilum area showing L. tingitanus Linnaeus, hilum transection typical linear micropyle, lack of hllar rim, (X 40).

33 ~~.­ .. '. ~~....- ..--­- ~ ...'.­ ~ ...... ~,

"

Figures 67-73. -Summary of testa features Greene) C. L. Hitchcock, testa transection of Lathyrus spp. and Lens Gulwaris, 67. away from hilum (x 300), Lens GuJinaris Lathyrus lallrO/IUS linnaeus. traeheld bar Medlkus: 70, Hilar view with lens to far right transection showing small regularly distrib­ (X 30); 71, enlarged view of micropylar uted Pits (x 600): 68, L. polyphyllus Nuttall area of hilum showing typical micropyle ex Torrey & Gray. with smooth and warty and papillose surface (x 240); 72. hilum Pits (x 4.000): 69, L. veslltus Nuttall ex Tor­ transection with no hilar rim (x 100): 73, rey & Gray subsp, puberulus (White ex view of irregular papillae of testa (X 1,000).

34 Figures 74-BO.-Summary of testa features transection (x 30); 77, P. fulvum Sibthorp of Pisum spp. and Vavilovia formosa. P. & Smith, typical Vicieae tracheid bar pits sa/ivum Unnaeus subsp. sativum: 74, (x 4,000). V. formosa (Steven) A. Fedorov: Hilum with typical linear micropyle and no 7B, Hilum with slight hilar rim near hilar rim (x 26); 75, enlarged view of micropyle (x 72); 79, enlargement of 7B to characteristic crimped papillae (x 3,000). show hilar rim and unusual bifurcated 76, P. satlvum Unnaeus subsp. elatius (M. micropyle near top (x 300); BO, testa sur­ Biebersteln) Ascherson & Graebner, hilum face showing typical papillae (x 1,000).

35 Flqures 8187 -Summary of Vlcla testa (x 1,000); 84. V. lalhyroldes Ltnnaeus. fea tures 81 V cassub/ca Llnnae:us. seed papillose testa with small mounds (x 200): wdh elonqate micropyle and no hilar (1m 85, V d/sperma de Candolle, hilum transec­ I x 78) 82. V ludov/c/ana Nuttall, tYPical tion (x 60): 86, V. blennts Ltnnaeus. View of linear micrOpyle and papillose surface typical nonvestured tracheid bar pits (x l x 400). 83, V mgflcans Hooker & Arnott 4,000); 87. V. sylva/lca Ltnnaeus. transec­ subsp glganlea (Hooker) Lassetter & tIOn of testa away from hilum (x 400) GU!)/). enlarqed VieW 01 papllll)se surface

36 88

";" ,. ""f~;j:t ~.:r ~'.

Figures 88-93,-Testa of Abrus precatorius (x 1,000); 91, enlarged part with numerous Linnaeus. 88, Sunken hilum with promi­ cuticular excrescences and fine cuticular nent hilar rim and attached rim-arll (x 30); reticulum in between (x 3,000); 92, sunken 89, enlarged micropylar end 01 hilum with hilum in transection (x 40); 93, pits in conspicuous hilar rim and V-shaped tracheid bar showing irregular shape, micropyle separated from hilum at far right poorly bordered, mostly with primary wall (x 100); 90, testa surface at edge of hilum intact, and with slight warts (x 4,000).

37 ~,,,:..:.ti.;f" Ji

';,\:'~.~~';t.(. ...-_~"~' ' •.w;(~,,~

" ~101 '.'.'.~.'1<.., .~. Figures 94-101.-Testa of Cicer arietinum linnaeus. 94. Hilum with heavy hilar rim and micropyle at far right appearing to be within hilum (x 54); 95, testa surface at midseed (x 100): 96, closeup of testa near hilum (x 1.000); 97. Iransectlon of sunken hilum (x 48); 98. longisection of hilum with tracheld bar (right arrow) extending length to prominent micropyle (Ief! arrow) (x 40); 99. enlarged view of tracheid bar merger with ovular vascular bundle a t left (x 260); 100. traeheid bar Pits lacking warts (x 4.000); 101, slightly warty pits, some wilhremnants of primary wall (x 4,000).

38 •

Figures 102-110.-Testa of Trifolieae. 102, Linnaeus subsp. caeru/a (Lessing ex Ononis pubescens Linnaeus, reduced hilar Ledebour) Schmalhaussen, low papillae rim and linear micropyle in contact with near hilum (x 1,000); 108, Medicago ova! hilum (>< 120). Trigone/la arabica pironae Visiani, hilum with conspicuous Delile: 103, Circular, sunken hilum and micropyle (above) and testa with low papillose testa with mounds (x 200); 104, papillae (x 150); 109, Medicago sativa Lin· enlargement of part (x 1,000). 105, naeus subsp. sativa, obscure papillose pat· Meli/otus messanensis (Linnaeus) Allioni, tern near hilum (x 1,000); 110, Trifolium vestured pits from tracheid bar (x 4,000); ochroleucr:Jn Hudson, no pattern on testa 106, Melilotus alba Medikus, seed testa near hilum (x 1,000). near hilum (x 1,000); 107, Medicago sativa

39 r

Figures 111-116.-Testa of Trifolieae. Factor­ ovskya aschersoniana (Urban) Eig: 111, Prom­ inent lens area (right arrow) and dam­ aged area (left arrow) (x 32); 112, hilum, hilar rim, deltoid micropyle at right, and papillose surface (x 200); 113, testa sur­ face enlarged to show low papillae (x 1,000). Parochetus communis £3uchanan­ Hamilton ex D. Don: 114, Lens (arrow) (x 40); 115, hilum, hilar rim, and deltoid micropyle (x 160); 116, testa at hilar edge (top) showing almost smooth surface (x 1,000).

40

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