Genus Vicia with Notes About Tribe Vicieae (Fabaceae) in Mexico and Central America

Home , Vicia, Vicia americana, Vicia andicola, Vicia hassei, Vicia hirsuta, Vicia nigricans

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GENUS VICIA WITH NOTES ABOUT TRIBE VICIEAE (FABACEAE) IN MEXICO AND CENTRAL AMERICA

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/^^ UNITED STATES TECHNICAL PREPARED BY ((iàM DEPARTMENT OF BULLETIN SCIENCE AND ^SF AGRICULTURE NUMBER 1601 EDUCATION ADMINISTRATION ABSTRACT Gunn, Charles E. 1979. Genus Vicia with notes about tribe Vicieae (Fabaceae) in Mexico and Central America. U.S. Department of Agriculture, Technical Bulletin No. 1601,41 pp. Ten native and naturalized species and two native subspecies of Vicia in Mexico and Central America are keyed, described, discussed, and illustrated. Two new subspecies are named. All genera in the tribe Vicieae are keyed and discussed. KEYWOKDS : Lathyrus^ Lens^ Pisvm, Vicia, Vicieae, Mexico, Central Amer- ica, Guatemala, Fabaceae, Leguminosae. GENUS VICIA WITH NOTES ABOUT TRIBE VICIEAE (FABACEAE) IN MEXICO AND CENTRAL AMERICA

by CHARLES R. GUNN

/^^ UNITED STATES TECHNICAL PREPARED BY ààM DEPARTMENT OF BULLETIN SCIENCE AND ^F ACRirULTURE NUMBER 1601 EDUCATION '^'-"^^^ ADMINISTRATION

For sale by the Superintendent oí Documents, U.S. Ooverament Printing OfBce Wasliington, D.C. 20402 Stock Number 001-000-04034-1 ACKNOWLEDGMENTS I appreciate the courtesy of the curators in extending the duration of the herbarium loans so that this bulletin could be completed. The discussion of Vicia setifolia was enhanced by the loan of a flower from the type and the notes of Alicia Lourteig, Museum National d'Histoire Naturelle Laboratoire de Phanerogamic, Paris, France. I am grateful for the technical advice of Frances Kupicha, Botany Department, British Museum (Natural History), London, England; Stuart Lassetter, Biology Department, Eastern Kentucky Univer- sity, Richmond ; and John Wurdack, Botany Department, Smithsonian Insti- tution, Washington, D.C. The illustrations of Vicia species were prepared by Regina O. Hughes, Botany Department, Smithsonian Institution.

CONTENTS Page Tribe Vicieae 1 Key to Vicieae genera ^ Lathyrus Linnaeus ^ Lens P. Miller ^ Pisum Linnaeus ^ Vicia Linnaeus ^ Key to Vicia species ^ Descriptions of Vicia species 6 1. Vida villosa Roth (subgenus Vicilla, section Cracca) 6 2. Vicia hirsuta (Linnaeus) S. F. Gray (subgenus Vicilla^ section Cracca) ^ 3. Vicia hmnilis Humboldt, Bonpland, Kunth (subgenus Vicilla^ section Cracca) 1^ 4. Vicia hassei S. Watson (subgenus Vicilla^ section Cracca) 12 5. Vicia Ivdovidana Nuttall subspecies ludoviciana (subgenus Vicilla^ section Cracca) 14 6. Vicia pulchella Humboldt, Bonpland, Kunth subspecies pulchella (subgenus Vicilla^ section Cracca) 1Î 7. Vicia jmlchella Humboldt, Bonpland, Kunth subspecies mexicana (Hemsley) C. R. Gunn (subgenus Vimlla^ section Cracca) 1^ 8. Vicia americana Muhlenberg ex Willdenow subspecies ameri- camu ( subgenus Vicilla^ section A mericanae ) 21 9. Vicia americana Muhlenberg ex Willdenow subspecies mexicana C. R. Gunn (subgenus VicUIa^ section Americanae) 25 10. Vima leucophaea Greene (subgenus ViciXla^ section Medio- cinctae) 26 11. Vicia sativa Linnaeus (subgenus Vicia^ section Vicia) 28 12. Vicia jaba Linnaeus (subgenus Vicia^ section F aha) 32 Excluded species ^^ Literature cited ^^ Issued December 1979 GENUS VICIA WITH NOTES ABOUT TRIBE VICIEAE (FABACEAE) IN MEXICO AND CENTRAL AMERICA By Charles R. Guirn ' Vicia species (vetches) are economically im- CandoUe. This bulletin on Mexican and Central pedant temperate zone crops used for forage hay, American taxa in the genus Vicia expands and green manure, seeds, and soil improvement. These . species and such essential human seed-food crops Partially updates Agriculture Handbook 168, as Lens cidinajis Medikus and Pisvm sativwm which treats the vetches of the United States Linnaeus are in the tribe Vicieae (Adanson) de (Hermann, 1960) .=

TRIBE VICIEAE The relationship of the genus Vida to the three lary filament free to rather connate with others other genera (Z«í%riís,Ze«^, and Pwîîm)Mn the into an adaxially split sheath, filament apices tribe Vicieae has been discussed (Hutchinson, linear, anthers uniform and versatile, style hairy 196^; Gunn, 1969; Gunn and Kluve, 1976; to glabrous, ovules 2 or more, legumes 2-valved, Kupicha, 1^5; Radzhi, 1971). Oicer has been re- seeds often spherical to subspherical to lenticular moved from the Vicieae and placed in its own tribe with funiculus expanded above hilum, germina- Cicereae (Kupicha, 1977). Hutchinson also has tion usually hypogeal, most seedlings triarch, first adequately described all five genera. leaves cataphylls (trifid scales), n = 7, 6, 5. Type Viciea« is a natural tribe. Its taxa usually ex- genus Vicia. hibit these characters: Low or clhnbing (not The generic key is designed to quickly inform twining) herbs, stipules usually well developed, the user as to whether an unknown is a Vicia leaves often paripinnate (rarely imparipinnate or species or a member of another genus of the tribe. reduced to phyllodes in Lathyrm species) with A key to the Vicia species is included with the rachis terminating in a tendril or bristle, stipels genus discussion. The genera are presented alpha- absent, ñowers raxîemose or solitary in leaf axils, betically, and 10 Vicia species .are given according corolla papilionaceous, standard usually glabrous to the phylogeny of Kupicha {1976). or rarely pubescent (F. aruitolica TxxmW, V. Measurements with one number without a range hyirida Linnaeus, V. pannonica Crantz), vexil- should be considered as averages.

KEY TO VICIEAE GENERA 1. Abaslal surface of style pubescent or hairs encircling style or style glabrous (fig. 1) y.^^ 1. Adaxial surface of style pubescent. 2. Stipules as large as or larger than leaflets ; style grooved (easily seen at base of style) p^^^ 2. Stipules much smaller than leaflets ; style not grooved. 3. Flowers less than 10 mm long ; androecial sheath apex oblique Lens 3. Flowers more than 10 mm long (for North American species) ; androecial sheath apex truncate (for Mexican species) Lathyrus

* Botanist, Plant Taxonomy Laboratory, Beltsville Agri- ture Cited, p 35 cultural Research Center,.Beltsville. Md. 20705. = Kupicha (pers. commun.) recognizes a fourth genus, me year in italic after authors' names refers to Litera- Vavllovia A. Fedorov. 1 TECHNICAL BULLETIN 1601, U.S. DEPT. OF AGRICULTURE

LATHYRUS LINNAEUS

Lathyrus Linnaeus, Sp. PL : 729. 1753 et Gen. PL ed. 5 : tingitanu^ Linnaeus (including L. mexicanus 326. 1754. LECTOTYPE: L. sylvestris Linnaeus Schlechtendal) (Mexico). (vide Britton et Brown, 111. FI. N. U.S. and Can. ed. 2, Two species, L, longipes and Z. parvifoUus^ are 2: 412. 1913). About 150 species. endemic only to Mexico. Their status is open to question because they are morphologically similar The most recent North American Lathyrus to each other and to L. ari^onicus Britton, Z. monograph has Mexican distribution maps for six graminifolius^ and L. pauciflorm Femald. The native species (Hitchcock, 1952), Two cultivated type of Z. longipes is not significantly different and now naturalized species, named by Linnaeus, from many collections of Z. parvifoUus^ according also have been collected in Mexico. The following to Hitchcock. However, other collections are sufficiently different to merit specific recognition. species occur in Mexico and all except ,L, odoratus Lathyrus longipes and Z. parvifoUus may be sep- are illustrated in Hitchcock : L. eucosmus Butters arated from Z. paucifiorus by their range and such & St. John (Sonora), i. graminifolius (S. Wat- minor characters as leaflet and flower length, num- son) White (Sonora and Chihuahua), i. Ueti- ber of flowers per raceme, and length of calyx fiorus Greene subsp. alefeUii (White) Jepson teeth. (northern Baja California), L, longipes White Lathyrus is not native to Central America, and (Chihuahua and Nuevo León), Z. odoratus no native Mexican species have been collected Linnaeus (Michoacán, Puebla, and Chiapas), ,L. south of the northern Oaxaca border. I have seen parvifoUus S. Watson (from Sinaloa west to no herbarium sheets of naturalized or introduced Tamaulipas and south to Oaxaca), L. splendens Lathyrus species from Central America. There are Kellogg (northern Baja California), and L. endemic Lathyrus species in South America.

LENS P. MILLER

tens P. Miller, Gard. Diet. Abridg. ed. 4. 1754, nom, cons. America, this species is cultivated in both regions. [non Lens Linnaeus, Herb. Amboin. 18. 1754 et in Westphal (197i, pp, 109-llJf, fig. 10) presents a Amoen. Acad. 4:128 et 134. 1759, nom. invalid., at- lucid discussion and an excellent illustration of tributed in Kew Index to Stickman]. LECTO- Z. culinaris. Its flowers are self-pollinating and the TYPE : L. culinaris Medikus, Tories. Churpf. Phys. Ges. 2:361. 1787 (Ervum lens Linnaeus, Sp. PL : plants produce many seeds. Like other self-pol- 738. 1753). Five species. linating seed crops, this species has many races based on seed, pod, and plant characters as well Although I have not seen an herbarium sheet of as geography. For example, Barulina {1930) rec- L, culinaris (lentil) from either Mexico or Central ognized 58 varieties, now regarded as races.

PISUM LINNAEUS

Pisum Linnaeus, Sp. PI.: 727. 1753 et Gen. PI. ed. 5: naeus (chickpea), which is now in the monogeneric 324. 1754. LECTOTYPE: P. sativum Linnaeus tribe Cicereae, and Lens culinaris, it is a self- {vide Alefeld in Bonplandia 9: 126. 1861). Five pollinating seed crop. Its seed characters have been species. used to erect lower taxa. Cultivar names are more widely used in this species than in the other two Pisum sativum (pea) is cultivated in Mexico species. Excellent illustrations of P. sativum are and Central America. Like Oieer arietinum Lin- in Westphal {197i, figs, 20-22), GENUS VICIA WITH NOTES ABOUT TRIBE VICIEAE

VICIA LINNAEUS Vicia Linnaeus, Sp. PL : 734. 1753 et Gen. PI. ed. 5: 327. boldt, Bonpland, Kunth of South America appear 1754. LECTOTYPE: V, sativa Linnaeus (vide Brit- to be closely related. ton and Brown, IlL Fl. N. U.S. and Can. ed. 2, 2: 408. 1913). About 150 species. A recently developed combination of tested morphological characters is used to establish Vicia species are primarily distributed in the species parameters. These characters include stylar hair patterns, external seed topography, north temperate zone and are also found in Hawaii, Canary Islands, and central Africa (map flower topography and placement on and number per raceme, presence of nectary pits on stipules, 1). The only large concentration in the south tem- presence of fibrous cross ribs (between seeds) on perate zone is in South America. The intrusion inner valve surfaces of legumes, and life forms. I into tropical areas is primarily based on the amelioration of temperature by elevation. Moun- used these characters in a general survey of U.S. Vicia species, and I was able to identify native and tain chains are an important factor in the distribu- naturalized North American Vicia to species by tion of these species in Mexico, Guatemala, north- their seeds alone (Gunn, 1968^ 1971). Some of ern South America, Hawaii, southern China, these characters were used by Lassetter {1972) in northern India, and central Africa. In these areas, a revision of the southwestern U.S. Vicia species, native Vicia species are found at higher elevations, the F. exigua-leavenworthii-ludoviciana complex. 2,100-3,050 meters, than in temperate zones. The only published treatment of U.S. Vicia species In North America there are 16 endemic and 17 (Hermann, 1960) was completed before these data naturalized species of Vicia, Six of the sixteen were coordinated. species are also native to Mexico, and the range of Stylar hair patterns of more than 100 species of 2 of these 6, F. humilis Humboldt, Bonpland, Vicia studied thus far may be placed in one of five Kunth and F. pulchella Humboldt, Bonpland, illustrated categories (fig. 1) : Style glabrous to Kunth subsp. pulchella^ extends into adjacent glabrate ; style with apical hairs restricted to abax- Guatemala. Vicia hirsuta (Linnaeus) S. F. Gray ial surface, or adaxial surface bearing a few scat- and F. sativa Linnaeus have become naturalized tered small hairs ; style with encircling apical hairs in Mexico, whereas F. sativa and F. villosa Roth that are brushlike on the abaxial surface; style have become naturalized in Guatemala. Although with encircling apical hairs that are uniform in F. faha Linnaeus may not be naturalized in Mexico length and density on both surfaces; and style or Guatemala, it has been included because of the with encircling medial hairs that are uniform in number of collections from field plantings in length and density on both surfaces. A sixth cate- southern Mexico and Guatemala and the wide- gory with hairs restricted to the adaxial surface spread use of its seeds as human food. (not illustrated) is found in F. koeieana Rechin- This is the first systematic treatment of Vicia ger f., a Near East species, and some specimens of species in Mexico and Central America and is a F. ervilia (Linnaeus) Willdenow (Gunn and precursor to a study of these species in North and Kluve, 1976', Kupicha, 1973,1976), South America. Central America, with its lower Five hilum shapes, based on a study of more than elevations, generally does not provide a favorable 100 Vicia species, are circumlinear, linear, oblong, habitat for native or naturalized Vicia species. wedge, and oval (fig. 1). A circumlinear hilum is Therefore, any gene flow between North and more than 10 times longer than wide, has parallel South America has been broken by the lower eleva- margins, and occupies about 75 percent of the seed tions in Central America south of Guatemala. One circumference. A linear hilum is 5 to 7 (rarely 9 to species complex is native to both North and South 10) times longer than wide, has parallel margins, America, F. nigricans Hooker & Amott subsp. and oceupies less than 50 percent of the seed cir- nigricans and F. nigricans Hooker & Amott sub- cumference. An oblong hilum is usually less than sp. gigantea (Hooker) Lassetter & Gunn. The five times but usually greater than twice as long vicarial species F. americana Muhlenberg ex Will- as wide, has slightly curved margins, and occupies denow of North America and F. andicola Hum- less than 50 percent of the seed circumference. A TECHNICAL BULLETIN 1601, U.S. DEPT. OF AGRICULTURE

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wedge hilum has margins that converge to the (MICH) Herbarium of the University of lens ; in all other respects it is similar to the oblong Michigan, Ann Arbor hilum. An oval hilum has a length that is never (MO) Herbarium of Missouri Botanical more than twice its width, is rounded in outline, Garden, St. Louis and occupies less than 20 percent of the seed cir- (NA) Herbarium of U.S. National Ar- cumference. boretum, Washington, D.C. The other morphological characters used in tax- (ND) The J. A. Nieuwland Herbarium of onomic determinations are depicted in figures University of Notre Dame, Notre 2-14. Dame, Indiana The following abbreviations from Holmgren (NDG) The E. L. Greene Herbarium of and Keuken {197^) are used in this bulletin : University of Notre Dame, Notre Dame, Indiana (ARIZ) The University of Arizona Her-

barium, Tucson (NY) Herbarium of the New York Botan- ical Garden, Bronx ( B ) Botanischer Garten und Botanisches (OKL) Bebb Herbarium of University of Museum Berlin-Dahlem, Berlin, Oklahomia, Norman Federal Republic of Germany (P) Museum National d'Histoire Natu- (BM) British Museum (Natural History), relle, Paris London (PH) Herbarium of Academy of Natural (CAS) California Academy of Sciences Sciences, Philadelphia Herbarium, San Francisco (POM) Herbarium of Pomona College, (ENCB) Escuela Nacional de Ciencias Bio- Claremont, California lógicas Herbarium, Mexico (RM) Rocky Mountain Herbarium of (F) John G. Searle Herbarium of Field University of Wyoming, Laramie Museum of Natural History, Chi- (RSA) Rancho Santa Ana Botanic Garden cago Herbarium, Claremont, Califor- (GH) Gray Herbarium of Harvard Uni- nia versity, Cambridge, Mass. (SD) San Diego Museum of Natural His- (ISC) Iowa State University Herbarium, tory Herbarium, San Diego, Cal- Ames ifornia (JEPS) Jepson Herbarium and Library (SMU) Herbarium of Southern Methodist of University of California, University, Dallas, Texas Berkeley (TEX) University of Texas Herbarium, (K) The Herbarium and Library of Austin Royal Botanic Gardens, Kew, (UC) Herbarium of the University of Richmond, England California, Berkeley (US) U.S. National Herbarium of Smith- (LINN) The Linnean Society of London, sonian Institution, Washington, London D.C. (MA) Instituto "Antonio José Cavanilles," ( VT) Pringle Herbarium of University of Madrid Vermont, Burlington (MEXU) Universidad Nacional Autónoma de (WTU) Herbarium of University of Wash- Mexico, Mexico ington, Seattle TECHNICAL BULLETIN 1601, U.S. DEPT. OF AGRICULTURE KEY TO VICIA SPECIES 1. stipules bearing nectary patches. 2. Tendrils absent (leaves ending in cusps) ; plants erect and blackening on drying 12. V. faha 2. Tendrils present ; plants not erect and remaining green on drying 11. V. sativa 1. Stipules without nectary patches. 3. Corolla 5 mm or less; style nearly glabrous; hilum concealed by permanent funiculus : 2. V, hirsuta 3. Corolla more than 6 mm long; style pubescent; hilum visible. 4. Stylar hairs encircling and of same length. 5. Racemes 1-2 flowered ; stylar hairs medial 10. F. leucophaea 5. Racemes usually 3 or more flowered ; stylar hairs apical. 6. Annual; flowers usually 1-5 (1-17 in subsp. ludoviciana phase laxiflora) ; calyx teeth unequal 5. V. ludoviciana 6. Perennial ; flowers 8-26 ; calyx teeth subequal. 7. Corolla up to 10 mm long 6. V. pulchella subsp. pulchella 7. Corolla 12-18 mm long 7. V. pulchella subsp. mexicana 4. Stylar hairs encircling or on one surface but always forming an abaxial brush. 8. Standard less than 10 mm long. 9. Style without hairs on adaxial surface ; calyx teeth equal ; hilum occupying less than 20 percent of seed circumference ; restricted to Baja Cali- fornia 4. y. hassei 9. Style without hairs on adaxial surface ; salyx teeth equal ; hilum occupying about 30 percent of seed circumference; no farther north and west than Sinaloa 3. V. humilis 8. Standard more than 11 mm long. 10. Calyx gibbous at base; racemes densely 10-14 flowered 1. V. villosa 10. Calyx not gibbous at base; racemes loosely (2)-3-10 flowered. 11. Calyx teeth unequal, upper 2 much shorter than lower 3 8. V. americana subsp. americana 11. Clayx teeth equal or nearly so 9. V. americana subsp. mexicana

DESCRIPTIONS OF VICIA SPECIES

1. Vicia villosa Roth (subgenus Vicilla^ section Cracca) Hairy Vetch

(Fig. 2, Map 2)

Vicia villosa Roth, Tent. Fl. Germ. 2:182. 1795. Cracca V. hoissieri Heldreich & Sartorelli in Boissier, Diagn. Ser. villosa (Roth) Godron & Grenier, Fl. Fr. 1:470. II 2 : 40. 1856. 1848. Ervum villosum (Roth) Trautvetter, Acta V. reuteriana Boissier & Buhse, Nouv. Mem. Soc. Nat. Horti Petrop. 3:47. 1875. TYPE: UNKNOWN. Mose. 12 : 73. 1860. V. pseudocracca Bertoloni, Rar. Ital. PI. Dec. 3: 58. 1805. V. plumosa Martin-Donos, Fl. Tarn. 179. 1862. V. polyphylla Waldstein & Kitaibel, PI. Rar. Hung. 3: 282. V. pseudovillosa Schur, Enum. PI. Transs. 166. 1866. 1812. V. vulcanica Huet ex Nyman, Consp. 207. 1878. V. consentina Sprengel, Pugill. 2: 74. 1815. V. glahrescens Heimerl, Oest. Verb. Zool. Bot. Gesell. V. littoralis Salzmann, Flora 4:110. 1821. Wien. 31:173. 1881. F. microphylla Urville, Mem. Soc. Linn. Par. 1: 343. 1822. V. elegantissima Shuttleworth ex Rouy, Rev. Soc. Nat. V. Uvonae Seringe in de CandoUe, Prod. 2:357. 1825. Ser. Ill 1: 229. 1883. V. amUgua Gussone, Fl. Sic. Prod. 2: 435. 1828. V. glahrescens A. Kerner, Sched. Fl. Austro-Hung. 4:1. V. elegans Gussone, Fl. Sic. Prod. 2:438. 1828. 1886. V. dasycarpa Tenore, Abruzz. 81. 1829. Cracca dasy- V. godroni Rouy in Rouy & Foucaud, Fl. Fr. 5: 237. 1899. ca/rpa (Tenore) Alefeld, Bonplandia 9:121. 1861. V. eriocarpa Halacsy, Consp. Fl. Graec. 1: 489. 1901. V. varia Host, Fl. Aust. 2: 332. 1831. Cracca varia (Host) Grenier & Godron, Fl. Fr. 1:469. 1848. Annual or biennial, rarely perennial. Her- V. littoralis Tenore, Fl. Nap. 4:103. 1838. V. salaminia Heldreich & Sartorelli in Boissier, Diagn. baceous vine, up to 1.3 m long, drying green, Ser. II 2: 39. 1856. villose to glabrous. Trichomes white. !Leaves pin- GENUS VICIA WITH NOTES ABOUT TRIBE VICIEAE

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FIGURE 2.—Yida villosa drawn from greenhouse-grown plants : A, Habit (X 0.5) ; B, leaflets ( X 2.5) ; C, flower ( X 2.5) : D, androecial sheath and style (X 6) ; E, stylar hair pattern (X 11) ; i^, mature legumes (x 1) ; Ö, seeds ( X 3).

299-212 O - 80 - 2 8 TECHNICAL BULLETIN 1601, U.S. DEPT. OF AGRICULTURE nately compound, terminating in a ramified pre- Guatemalan specimens examined.—CHIMAL- hensile tendril. Leaflets 8-20, entire, thin, oblong- TENANGO: Koadside thicket, elev. 1,800 m ovate to linear-lanceolate or linear, 8-35 by 1-10 (F) : Tecpan, Johnston UO (F, US) ; ^ near mm, apex acute to obtuse and mucronulate, base Chimaltenango, elev. 1,800 m, Standley 80^81 (F). abruptly tapered. Stipules inconspicuous, egland- GUATEMALA: Aguilar 2U (F). QUEZAL- ular, usually entire. Racemes elongate, with (3)- TENANGO: Scrub forest of Baccharia and 5-40 closely spaced flowers, generally longer than Quercus^ elev. 3,000 m, Williams^ Molina Ä., & subtending leaf. Calyx one-half standard length, Williams 26631 (US). teeth unequal with 2 upper shortest poorly de- Vicia villosa is a polymorphic species that is veloped and curved and 2 lateral and especially usually cross-pollinated. Some self-pollination basal straight and long attenuate, gibbous at base, may occur, but seed set is greatly increased when usually partially bluish purple toward apex. bees are available to pollinate flowers. The species Corolla varying shades of bluish purple to pink or varies in degree of stem and legume pubescence, rarely white, with standard 12-20 mm long. An- width of leaflets, number of flowers per raceme, droecial sheath oblique at apex. Stigma globular. length and color of corolla, size of calyx teeth, and Style apex encircled by hairs with abaxial surface legumes. These and other characters have been bearing elongated hairs forming brush. Ovary used as bases for describing many species and glabrous to puberulent. Legume 20-40 mm long, lower taxa. The list of synonyms is not complete dark to light straw color, glabrous, reticulate and and includes only species epithets. punctate, oblong, flattened, obliquely short acumi- Although the subspecific taxonomy may be satis- nate at both ends (occasionally almost rounded), factorily established for Europe (Ball, 1968)^ it on 2-3 mm stipe, bearing 4-5 ovules, inner valve perhaps does not reflect all elements found in the surface smooth, twisting tightly to loosely during New World, especially in the United States. The dehiscence. Seed 3.4-5 mm in diameter, spherical description here is for V, villosa in a broad sense. to nearly sublenticular, smooth, dull dark reddish The six Guatemalan specimens seen during this brown to greenish brown densely mottled and study represent V. villosa subsp. villosa^ because pointed with blackish brown. Hilum color of seed- their stems are villous. This subspecies is often coat to darker, oblong, 3 times longer than wide, parallel to length of seed, occupying 14 percent of found in the cooler parts of the species range. seed circumference, hilar lips color of hilum. Lens Other subspecies that are glabrous or appressed- darker than seedcoat, with raised center 1-1.3 mm pubescent may be expected in warmer areas as in- from hilum. troduced crops or naturalized plants. Distribution.—Vicia villosa is a widely planted Vicia villosa is an important green manure crop fodder and green manure plant that has become that apparently has not been widely planted in naturalized in the united States. No herbarium Mexico or Central America. One reason may be the specimens were seen from Mexico, though it vetch bruchid {Bruchus hrachialis Fahraeus). Its has been collected in Guatemala where it is larvae hollow out the seeds and thus decrease their naturalized. viability and value.

2. Vicia hirsuta (Linnaeus) S. F. Gray (subgenus Vicilla^ section Cracca) Tiny Vetch

(Fig. 3, Map 2) Vicia hirsuta (Linnaeus) S. F. Gray, Nat. Arr. Brit. PI. 2 : Endiusa hirsuta (Linnaeus) Alefeld, Oest. Bot. Zeit. 9: 614. 1821. 360. 1859. Cracca hirsuta (Linnaeus) Genn., Nuo. Ervum hirsutum Linnaeus, Sp. PI. 138. 1753. Gior. Bot. Ital. 2 : 137.1870. TYPE : EUROPE (HOLO- V. mitchellii Rafinesque, Prec. Decouv. 37. 1814. V. TYPE : LINN, International Documentation Centre michellii Desvaux, Jour. Bot. 4: 269. 1814, sphalm. microfiche 907.5). Ervilia vulgaris Godron, Fl. Lorr. 1: 175. 1845. Cracca minor Grenier & Godron, Fl. Fr. 1: 473. 1848. Ervilia hirsuta (Linnaeus) Opiz, Sez. Rost. Kvst. * Collector's name and collection number in italic ; Ceske 41. 1852. herbarium symbol in parentheses (see p. 5). GENUS VICIA WITH NOTES ABOUT TRIBE VICIEAE 9

FIGURE 3.—Vicia hirsuta drawn from greenhouse-grown plants : A, Habit (x 0.5) ; B, stipule (X 4) ; C, leaflets (X 2) D, flowers {X 2) ; E, androecial sheath and style ( X 4) ; F, stylar hair pattern (X S) ; G, mature legumes (X 2) IT, seeds (X 5). 10 TECHNICAL BULLETIN 1601, U.S. DEPT. OF AGRICULTURE Vicioides hirsuta Moench, Meth. 137. 1794. monochrome). Hilum concealed by shiny choco- Vicia parviflora Lapeyrouse, Hist. Abr. Pyren. 418. 1813. late-colored f uniculus attached at one end of hilum Ervum sardoum Moris ex Sprengel, Syst. Veget. 4: 346. (if exposed), hilum darker than seedcoat, linear to 1827. slightly wedge-shaped, 6-9 times longer than wide, Lathy rus angustifolius Sessé, Mociño, Castillo, & Mal- parallel to length of seed, occupying 14-21 percent donado, based on their collection 3640 (name not intended to be published here). (Negative 46447, F) of seed circumference, with lips of hilar groove V. linearis Sessé, Mocifio, Castillo, & Maldonado, based on ocher. Lens darker than seedcoat, with raised their collection 3641 (name not intended to be pub- center 0.6-1 mm from hilum. lished here). (Negative 46448, F) Distribution.—Vicia hirsuta is a native of Eu- rope. It has now been naturalized in North Amer- Annual.—Herbaceous vine, up to 1 m long, dry- ica along roadsides and waste places. Plants in ing green, glabrate or sparingly pubescent. flower or fruit are inconspicuous and often over- Trichomes white. Leaves pinnately compound, looked. terminating in a simple or ramified prehensile Mexican specimens examined.—STATE UN- tendril. Leaflets 4-16, entire, thin, linear, narrowly KNOWN: ^^Lathyras angmtifoliics^^^ Sesse^ Ho- oblong or narrowly elliptical, 5-20 by 1-4 mm, cino^ Castillo^, & Maldonado SdJfi (F) ; Vicia lin- apex usually truncate or emarginate and mucron- earis^ Sessé^ Mociño^ Castillo^ & Maldonado 3ßJf.l ulate, base tapering and rounded. Stipules incon- (fragment F). MWKICO: Pringle (F) ; Temas- spicuous, eglandular, sharply serrate to toothed. caltepec, elev. 2,700 m, Hinton 4895 (ARIZ, US). Racemes elongate, with 2-6-(8) closely spaced Vicia hirsuta has two characters that separate it flowers, as long as to shorter than subtending leaf. from most other Vicia species, a glabrate stylar Calyx about one-half standard length, teeth sub- apex and a persistent funiculus. For these and equal straight attenuate, not gibbous at base, green other reasons, this species has been placed in six throughout. Corolla whitish blue with standard genera, and it is the lectotype of two segregate 2-5 mm long. Androecial sheath not oblique at genera, Endiusa Alefeld and Ervilia Opiz. Al- apex. Stigma globular. Style apex bearing only a though it is generally regarded as a weed or ig- few scattered hairs on abaxial side. Ovary pubes- nored in the New World, it is cultivated in the cent. Legume 6-11 by 2.5-4 mm, brown to blackish, Near East. Like other self-pollinating Vicia pubescent, obscurely reticulate, oblong, flattened, species, it is polymorphic, exhibiting pronounced obliquely short, acuminate to nearly rounded at variation in stature, density and number of leaves both ends, without a stipe, bearing (l)-2-(3) and leaflets, leaflet size, legume pubescence, and ovules, with inner valve surface smooth, twisting seed size. Plitmann {1968) described a form, prob- loosely to tightly during dehiscence. Seed 1.5-2.8 ably cultivated, with stems about 13 dm long and mm in diameter, subspherical to spherical to sub- leaflets 10-20 by 3-4 mm. Although it is not known lenticular, smooth, glossy greenish straw to red- whether a lack of stylar hairs enhances self-polli- dish straw", lightly to densely mottled with dark nation or not, apparently hairs are not needed for brown (some seeds so densely mottled as to appear this species because selfing approaches 100 percent.

3. Vicia hutnilis Humboldt, Bonpland, Kunth (subgenus Vicilla^ section Cracca) Low Vetch

(Fig. 4, Map 2)

Vicia humilis Humboldt, Bonpland, Kunth, Nov. Gen. et nately compound, terminating in a simple pre- Sp. PI. 6: 498. 1824. TYPE : MEXICO : Humboldt hensile tendril. Leaflets 2-6, entire, thin, linear, á Bonpland, Moran, elev. 2,^00 m (HOLOTYPE: P). 10-40 by 0.9-3 mm, apex tapered and mucronulate, Lathyrus parvißorus Sessé, Mociño, Castillo, & Maldonado, based on their collection 1903 (name not intended Lo base tapered. Stipules inconspicuous, eglandular, be published here). (Negative 46449, F) entire. Racemes subsessile, with 1 or rarely 2 closely spaced flowers, shorter than subtending Perennial.—Herbaceous vine, up to 1 m long, leaf. Calyx about one-half standard length, teeth dry green, glabrate. Trichomes white. Leaves pin- essentially equal straight attenuate, not gibbous at GENUS VICIA WITH NOTES ABOUT TRIBE VICIEAE 11

FIGURE 4.—Vicia humilis drawn from greenhouse-grown plants: A, Habit (X 0.5) ; B, leaflets with stipules and flower (X 2) ; (7, flower (X 4) ; Z), androecial sheath and style ( X 6) ; JE?, stylar hair pattern (X 14) ; F, mature legumes (X 1) ;G^, seeds (X 4). 12 TECHNICAL BULLETIN 1601, U.S. DEPT. OF AGRICULTURE base, green throughout. Corolla varying shades of US). MEXICO: Temascaltepec, Hinton 8897 bluish purple, with standard 5-10 mm long. An- ( F, MO, N Y, US ). MICHO AC AN : Elev. 2,500 droecial sheath oblique at apex. Stigma conical. m, Moore^ Hernandez X., Porras H, 5586 Style apex with abaxial surface bearing a brush (MEXU). NUEVO LEON: Sierra Madre and adaxial surface nearly glabrous. Ovary gla- Oriental, elev. 2,286-2,438 m, Mueller and Mueller brous to pubescent. Legume 30-40 by 4-7 mm, Uß (F, MEXU, MICH, TEX). OAXACA: straw color, glabrous to pubescent, strongly reticu- Teposcolula, Seler 1555 (US). PUEBLA : Boca late, linear-falcate, terete to partially flattened, del Monte, elev. 2,300 m, Arsène 7092 (US). oblique and acute at both ends, on 1 mm stipe, SAN LUIS POTOSÍ: Alvarez, Palmer lU (F, bearing 7-8 ovules, with inner valve surface NA, US). SINALOA : Penseos, elev. 1,675-1,980 smooth, twisting tightly during dehiscence. Seed m. Gentry 7215 (MICH). TAMAULIPAS: 2-2.3 mm in diameter, spherical to subspherical, Elev. 1,500 m, Runyon 1049 (US). VERA- smooth, blackish gray to greenish ocher, densely CRUZ : Boca del Monte, Nelson 2U (US). mottled with black so as to nearly mask base color. Guatemalan specimens examined.—HUEHUE- Hilum color of seedcoat or silvery (due to funic- TENANGO : Aguaoatan, elev. 1,950 m, Standley ular remnants), oblong, 4 times longer than wide, 81298 (F). QUEZALTENANGO : Cerro la Ped- parallel to length of seed, occupying 30 percent of rera, elev. 2,400 m, Standley 65539, 65561 (F). seed circumference, with lips of hilar groove color TOTONICAPAN: Chui-quisis, elev. 2,500-2,800 of hilum. Lens darker than seedcoat, with raised m, Standley 8U01 (F). center 0.5-0.7 mm from hilum. Vicia humilis is one of the most common and Distribution.—Vida humilis is found from widespread native Mexican vetches, and is one of Sinaloa and Coahuila south to northern Guate- two vetches endemic to Central America. The other mala, though apparently absent from the main is F. pulchella subsp. pulchella. Although the spe- axis of the Sierra Madre Occidental Mountains. It cies was originally described as possessing solitary is usually found above 2,000 meters in wet mead- flowers, a few widely scattered plants with two ows, pinewoods, and wooded ravines. flowers per raceme have been found. These plants Selected Mexican specimens examined.— occur throughout the range of the species, includ- STATE UNKNOWN: Lathyrus parvi-ßonis, ing northern Guatemala. These twinned-flowered Sessé^ Mociño^ Castillo^ cß Maldonado 1903 and i. plants do not exhibit other characters that could angustifolim, 1905 (F). CHIAPAS: Tenejapa, be used as a basis for separating one- and two- elev. 2,310 m, Breedlove 7013 (US). COA- flowered plants. Mature legumes may be glabrous HUILA: Sierra de Parras, Purpm 4580 (F, or pubescent ; there is no correlation between this UC, US). DISTRITO FEDERAL: Exclava, character and distribution or other studied Pnngle 7121 (MEXU, VT). HIDALGO : Sierra morphological characters. Although it is wide- de Pachunca, elev. 2,740 m, Pringle 781S (MEXU, spread in Mexico, little is known about this species. US, VT). JALISCO: Sierra del Tigre, elev. Greenhouse tests have shown that it is self- 2,100 m, McVmigh 13012 (MEXU, MICH, TEX, pollinating.

4. Vicia hassei S. Watson (subgenus Vicilla, section Cracca) Slender Vetch (Fig. 5, Map 3)

Vicia hassei S. Watson, Proc. Amer. Acad. Arts and Sei. Annual.—Herbaceous vine, approaching 1 m 25:129. 1890. long, drying green, pubescent. Trichomes white. V. exigua var. hassei (S. Watson) Jepson, Fl. W. Mid. Leaves pinnately compound, terminating in a Calif. 296. 1901. TYPE : UNITED STATES : Hasse, 1888, hiUs about Los Angeles (LECTOTYPE : GH; simple or ramified prehensile tendril. Ijeaflets SYNTYPE: PH ; PARATYPE S : Anderson, Santa 4-12, entire, thin, linear-lanceolate to elliptic, 8-^0 Cruz, GH, and Bigelow, Benicia, GH, NY). F. exigua by 1.5-9 mm, apex usually truncate to retuse or var. californica Torrey, Pacif. RR. Rep. 4: 76. 1856. emarginate, base rounded. Stipules inconspicuous, TYPE : UNITED STATES : Bigelow, Benicia (HOLO- TYPE : GH ; ISOTYPE : NY). eglandular, usually sharply serrate. Kacemes elon- GENUS VICIA WITH NOTES ABOUT TRIBE VICIEAE 13

FIGURE 5.—Vicia hassei drawn from herbarium specimens: A, Habit (X 0.75) ; B, leaflets (X 2) ; (7, flower (x 3) ; D, androecial sheath and style (X 4) ; i?, sty lar hair pattern (X 12) ; F, mature legumes ( X 1) ; G, seeds ( X 6). 14 TECHNICAL BULLETIN 1601, U.S. DEPT. OF AGRICULTURE gate, with 1-2 closely spaced flowers, shorter than islands to Guadalupe Island. Plants occur near subtending leaf. Calyx less than one-half standard sea level in a variety of habitats, infrequently length, teeth essentially equal straight short- along roadsides. attenuate, not gibbous at base, green throughout. Mexican specimens examined.—BAJA CALI- Corolla white to faint bluish or lavender, with FORNIA : Coronado Islands, elev. 150 m, Cowles standard 6.5-8.5 mm long. Androecial sheath 7 (POM) ; Ejido, elev. 300 m, Raven^ Lewis^ & oblique at apex. Stigma globular. Style apex en- Thompson 122Í8 (UC) ; Ensenada, Norland circled by hairs with abaxial surface bearing elon- (SD) ; South Todos Santos Island, Mœ^an 2799 gated hairs forming brush. Ovary pubescent. (UC) ; Valle Denos Creek, Bra.ndegee (UC) ; Val- Legume 22-38 by 5-7 mm, straw color, pubescent, lecito, Orcutt (MO). GUADALUPE ISLAND: strongly reticulate, oblong, partially flattened, Elev. 200 m, Moran 6643 (SD); Palmer 8It7 oblique and curved upward at apex, tapering to (MEXU, US) ; Wiggins & Ernst 200 (SD, UC). base, on 2-3 mm long stipe, bearing 4-10 ovules, Vicia hassei^ often regarded as a variety of V, with inner valve surface smooth, twisting during exigua^ has been a poorly understood mono- dehiscence. Seed 2.5-3 mm in diameter, sub- morphic species that is restricted in Mexico to the spherical, smooth, dull medium to dark brown or Pacific coast of northern Baja California and ad- gray brown, sparingly to densely mottled with jacent islands. This species shares many characters darker brown or uniformly gray brown. Hilum and its range with the much wider ranging poly- color of seedcoat or lighter, slightly wedge-shaped, morphic species complex V, ludoviciana^ which 6 times longer than wide, parallel to length of now includes V. exigua. Although the circum- seed, occupying 14—17 percent of seed circumfer- scriptions of V, hassei and V. ludovieiana have ence, with lips of hilar groove lighter than hilum. Lens inconspicuous, blackish, with raised center been controversial, Lassetter {1975) presents evi- about 0.5 mm from hilum. dence herein accepted that clearly separates them. Distribution.—Vicia hassei is essentially a Vicia hassei has an apical abaxial stylar brush and coastal species, which may be found from the San pubescent ovaries and legumes. Vicia liodoviciana Francisco Bay area (an isolated collection has been has the apex of its style encircled by hairs of es- made in southeastern Oregon) south to northern sentially equal length and glabrous ovaries and Baja California and westward through the coastal legumes.

5. Vicia ludovieiana Nuttall subspecies ludovieiana (subgenus Vieilla, section Craeea) Deerpea Vetch (Fig. 6, Map 3)

Vicia ludovieiana NuttaU subsp. ludovieiana in Torrey & Ervum tridentatum Alefeld, Oest. Bot. Zeit. 9: 363. 1859, Gray, Fl. X. Amer. 1: 271. 1838. Craeea ludovieiana Craeea tridentata Alefeld, Bonplandia 9: 119. 1861 (NuttaU) Alefeld, Bonplandia 9: 119. 1861. V. Non Vieia tridentata Schweinitz et V. trífida Dietrich ludovieiana var. typiea Shinners, Field and Lab. 16 : Ervum pilosum Engelmann ex Alefeld, Oest. Bot. Zeit. 9 23. 1948. TYPE : UNITED STATES : Leavenworth, 363. 1859. near Nagodoches . . . and Natchitoches (HOLO- V. thurheri S. Watson, Proc. Amer. Acad. Arts and Sei. 25 TYPE: NY); Tainturier, Louisiana (PARATYPE: 129. 1890. TYPE: UNITED STATES: Thurler PH). 299, Dona Ana (LECTOTYPE: GH ; ISOTYPES V. exigua Nuttall in Torrey & Gray, Fl. N. Amer. 1: 272. GH, NY; PARATYPES: Thurher 150, GH ; Wright 1838. Craeea exigua (Nuttall) Alefeld, Bonplandia 1350, GH, US ; Parry 33, ISC, NY). 9: 119. 1861. TYPE : UNITED STATES : Nuttall, V. produeta Rydberg, Bull. Torr. Bot. Club 28: 500. 1901. Columbia Plains (HOLOTYPE: BM, photo; ISO- TYPE : UNITED STATES : Rydberg d Vreeland 6006, TYPE: PH). Huérfano Co., Colo. (LECTOTYPE: NY; ISOTYPE: V. earoliniana var. texana Torrey & Gray, Fl. N. Amer. 1 : 271. 1838. V. texana (Torrey & Gray) Small, Fl. NY). Se. U.S. 656. 1903. V. ludovieiana var. texafia V. ludovieiana var. laxiflora Shinners, Field and Lab. 16 : (Torrey & Gray) Shinners, Field and Lato. 16: 23. 22. 1948. TYPE: UNITED STATES: Whitehouse 1948. TYPE: UNITED «STATES: Leavenivorth, 15275, Wise Co., Tex. (HOLOTYPE: SMU ; ISO- Texas (LECTOTYPE: NY). TYPES: MO, NY). GENUS VICIA WITH NOTES ABOUT TRIBE VICTEAE 15

lyt^^ Q. y\AA^JxjLo-

FiGURE 6.—Vicia ludoviciana drawn from herbarium specimens: A, Habit (X 0.5) ; B, leaflets (X 5) ; C, inflorescence and subtending leaf ( X 1) ; D, flower {X 5) ; E, androecial sheath and style (X 9) ; F, stylar hair pattern ( x 27) ; G, mature legumes (X 1) ; H, seeds ( X 7).

9-212 O - 80 - 3 16 TECHNICAL BULLETIN 1601, U.S. DEPT. OF AGRICULTURE V. Icarcnicorthil var. occidcntalis Shinners, Field and Shreve 7139 (MICH, NA, ITC). CHIHUAHUA : Lab. 16: 22. 1948. TYPE: UNITED STATES: Martizen & Fechu 160 (TEX). COAHUILA: Mulle i\ Chisos Mountains, Boot Spring (HOLO- Muzpuiz, Marsh 2137 (F, OKL, TEX). NUEVO TYPE: SMT ; ISOTYPES: MO, NY). LEON: Monterrey, Dodge (US). Winter annual.—Herbaceous vine, up to 2 m Vicia ludoviciana subsp. hidoviciana is part of long, drying green, glabrous to puberulent. Tri- what has been a poorly understood alliance of chories white. Leaves pinnately compound, termi- southwestern ITnited States and northern Mexico nating in a ramified prehensile tendril. Leaflets Viciu species that included F. exigua^ F. hassei, F. 5-17, entire, thin, linear-oblong or elliptic to oval, leavenwovtKii^ and F. ludoviciana (Shinners, 5-18 by 1.2-11 mm, apex acute to truncate or 1948; Turner, 1959; Hermann, 1960). Lassetter emarginate and sometimes mucronulate, base {1972) recorded the characters that have intrinsic tapered. Stipules inconspicuous, eglandular, en- taxonomic significance and concluded that two tire. Racemes with 1-19 closely spaced flowei-s, varieties and seven geographical races (phases) shorter than to exceeding subtending leaves. Calyx were recognizable and definable in the F. ludo- less then one-half standard length, teeth subequal viciana complex. Although a variety of nomencla- with 2 upi^er averaging 0.2 mm shorter than lower ture has been formalized in accordance with the 3 and all lance subulate, not gibbous at base, "International Code of Botanical Nomenclature," green. Corolla deep blue to bluish purple or whit- the phase nomenclature for geographical races is ish with standard (4.2)-5-6.5-(8.2) mm long. An- informal and is used for guidance. Vicia hassei^ droecial sheath oblique at apex. Stigma globular. now reestablished as a species, is an element of the Style apex encixxled by hairs of approximately Baja California flora, and the other three taxa equal length. Ovary glabrous. Legume 16-30 by are now part of the polymorphic species complex, •4-6 mm, straw color to brown, glabrous, reticulate, F. ludoviciana (Lassetter, 1975), rhombic-oblong, partially flattened to nearly The two subspecies of F. ludoviciana are ludo- terete, obliquely acuminate at both ends, on 1 mm viciana and leavenworthii (Torrey & Gray) Las- stipe, bearing 4-7 ovules, with inner valve surface setter and Gunn stat, nov.^ based on F. ludoviciana smooth, twisting tightly during dehiscence. Seed Torrey and Gray {1838, p, 271). Subspecies leav- 2 mm in diameter, spherical to subspherical, enworthii has two phases, leavenworthii and smooth, light greenish ocher to reddish brown, louisiana. Although neither phase has been col- mottled and pointed with dark brown. Hilum lected in ^léxico, phase leavenworthii likely may color of seedcoat, oblong, 5 times longer than wide, be in Mexico along the Rio Grande, based on its parallel to length of seed, occupying about 30 per- U.S. distribution. cent of seed circumference, with lips of hilar Subspecies ludoviciana has five phases, with groove usually pale straw color. Lens brown to three in Mexico. Vicia ludoviciana var. ludo-viciana blackish, with raised center about 0.5 mm from phase exigua is restricted to Baja California and hilum. adjacent islands. This taxon is sympatric with F. Distribution.—Vicia ludoviciana subsp. Ivdo- hassei. Phase laxiflora is found in northern Coa- viciana is w^idely scattered in northeiTi Arkansas, huila and Nuevo Leon. Phase producta is known becoming more frequent in extreme southern Mis- from one specimen collected in Chihuahua. Phase sissippi and adjacent Alabama, in southern Loui- texana likely may be found in northern Mexico siana, in eastern Texas, in northern Nuevo León to along the Rio Grande, based on its U.S. distribu- Chihuahua, and in Baja California and adjacent tion. Phase ludoviciana is not expected to be in islands. Mexico, based on its U.S. distribution. Selected Älexican specimens examined.— Table 1 summarizes some of the characters that STATE UNKNOWN: Woolson, May 1851, may be used to identify the Mexican members of (MICH). BAJA CALIFORNIA: San Quentin this species complex. Lassetter {1972) presents ad- Bay, Palnwr 60S (F, MEXU, MICH, US) ; ditional charactei-s and a discussion. GENUS VICIA WITH NOTES ABOUT TRIBE VICIEAE 17

TABLE 1.—Characters used to identify members 0/Vicia ludoviciana complex ^

Occurrence and characters

V. ludoviciana subspecies T-H 10 0 CO 1 1 1 rH

0) 1 e cC 1 Î internode elongation peduncle and present during initial flowering 3, 1 terminal and pair below each from 0 0 separate point Immature fruits Flowering before Flowers usually E Flowers 1-3,

leavenworthii phase leavenworthii. No X X X X X ludoviciana phase exigua Yes____ X X X ludoviciana phase laxiflora Yes____ X X X X ludoviciana phase producta Yes____ X X X ludoviciana phase texana No X X X

^ For additional characters and discussion, see Lassetter {1972).

6. Vicín pulchella Humboldt, Bonpland, Kunth subspecies pulchella (subgenus Vicilla, section Cracca) Showy Vetch (Fig. 7, Map 4)

Vicia pulchella Humboldt, Bonpland, Kunth subsp. throughout. Corolla creamy white to white with pulchcUa, Nov. Gen. et Sp. PI. 6: 499. 1824. Cracca purple veins or pinkish to bluish purple, with pulchella (Humboldt, Bonpland, Kunth) Alefeld, Bonplandia 9 : 119. 1861. TYPE : MEXICO : Hum- standard 5-8-(12) mm long. Androecial sheath holdt é Bonpland, Mescala (HOLOTYPE : P). oblique at apex. Stigma globular. Style apex en- V. suhulata Sessé & Mociño ex Steudel, Nom. Bot. ed. 2: circled by hairs of uniform length. Ovary gla- 765. 1841. (Negative 45453, F) brous. Legume 20-35 by 5-8 mm, straw color, F. melilotoides Wooten & Standley, Contr. U.S. Nati. Herb. obliquely acuminate at both ends, on 1 mm stipe, 16 : 141. 1913. TYPE : UNITED STATES : Standley bearing 5-7 ovules, with inner valve surface 4S64, Winsors Ranch, Pecos River National Forest, N. Mex. (HOLOTYPE: US; ISOTYPE: GH). smooth, twisting tightly during dehiscence. Seed V. canifolia Standley, Field Mus. Nat. Se. Bot. Ser. 22 (2) : 2.2-2.5 mm in diameter, subspherical, smooth dull 83. 1949. TYPE : MEXICO : LeSueur 7U, La Cima, reddish brown mottled with black. Hilum color of Chihuahua (HOLOTYPE: F). seedcoat, oblong to wedge-shaped, 6 times longer than wide, parallel to length of seed, occupying 26 Perennial.—Herbaceous vine, up to 1.5 m long, percent of seed circumference. Lens black, with drying green, pubescent to glabrate. Trichomes raised center 0.7 mm from hilum. white. Leaves pinnately compound, terminating in Distribution.—Vicia pulchella subsp. pulchella a ramified prehensile tendril. Leaflets 8-16, entire, is found in Arizona and New Mexico south to thin to coriaceous, elliptic to linear-oblong, northern Guatemala, exclusive of Baja California oblong-lanceolate or linear, 7-30 by 0.8-11 mm, and the Yucatan Peninsula. apex obtuse to acute and mucronulate, base ta- Selected Mexican specimens examined.— pered. Stipules inconspicuous, eglandular, entire. AGUASCALIENTES : Upper slope over vol- Eacemes elongate, with (6)-8-25 closely spaced canic rocks near Cerro Altamira, elev. 2,200-2,450 flowers, longer to shorter than subtending leaf. m, McVaugh 23767 (MICH). CHIAPAS : Steep Calyx about one-half standard length, teeth sub- slope along highway 190, elev. 2,743 m, Breedlove equal with upper 2 deltoid and lateral 2 and basal U159 (F, MICH, US). CHIHUAHUA: Cool acut^ to acuminate, not gibbose at base, green slopes La Bufa Mountains, Pringle 1231 (F, US). 18 TECHNICAL BULLETIN 1601, U.S. DEPT. OF AGRICULTURE

,^a. â.Mt*^AA4^

FIGURE 7.—Vicia pulchella subsp. pulchella drawn from herbarium specimens: A, Habit ( X 0.75) ; B, leaflets (X 2.5) ; C, inflorescences and subtending leaf ( X 0.75) ; D, flower (X 5) ; ^, androecial sheath and style ( X 10) ; F, stylar hair pattern (X 30) ; G, mature legumes (X l-S) ; H, seeds (X 5). GENUS VICIA WITH NOTES ABOUT TRIBE VICIEAE 19 COAHUILA : Oak forest, Buenos Aires, Kenoyer Vicia pulchella subsp. pvlchella is the most com- & Orum 2698 (MICH). DUEANGO: Sierra mon and widespread native Mexican vetch, and it Madre Occidental, elev. 2,600-2,700 m, Pemiell is one of two vetches endemic to Central America. 18m (MEXU, US). GUANAJUATO: Shaded The other endemic is F. humilis. Vicia pulchella bank near Cerro Zamorano, elev. 2,600 m, Mc- exhibits less variation than most endemic vetches, Vaugh lOm (MICH). HIDALGO : Under cliffs especially considering its wide range. The flowers near Metepec Station, elev. 2,530 m, Pringle 11970 exhibit color variation and the subspecies are based (F, MICH, US). JALISCO : Sierra Madre, Rose on discrete flower lengths. Subspecies mexicana 2961 (US). MEXICO: Amecameca, elev. 2,600 with a flower length of 12 mm or more, usually 13- m, Matuda 18808 (MEXU). MICHOACAN: 18 mm, is the next taxon. Of the approximately Morelia, elev. 2,200 m, Arsène 6636 (US). 250 specimens of subspecies pulchella and mexi- NUEVO LEON : Arroyos and canyon slopes near cana collected in Mexico and Guatemala, only three Galeana, elev. 2,290 m, Mueller & Mueller 665 (F, have an intermediate flower length. These collec- MEXU, MICH). OAXACA: Sierra de San tions were made in Chihuahua. The collection by Felipe, elev. 3,050 m. Smith 568 (F, UC, US). LeSueur 70S has flower lengths from 8.9 to 11 mm. PUEBLA : Esperanza, Purpus 2^76 (POM, UC). Another and more widely distributed collection, SAN LUIS POTOSÍ: Alvarez, Palmer 66-1/2 Townsend & Barher 123, has flower lengths from (F, MEXU, US). SINALOA: Pine forest, elev. 9.5 to 12 mm. The Kruckeherg JfilyS specimens have 1,830-2,130 m. Gentry 6227, 6318 (MICH). flowers approaching 13 mm. Except for these col- SONORA: Burnt over pineland, Puerto de los lections, subspecies pulchella flowers are less than Aserraderos, White 281^8, 3133 (MICH). TA- 10 mm long and usually less than 8 mm. MAULIPAS: Marcella, Stanford, Lauter, & Little is known about the pollination and crop Taylor s M. (RSA, US). potential of F. pulchella. Plants are usually found Guatemalan specimens examined.—HUEHUE- at about 2,000 meters and often associated with TENANGO: Ravines and hillsides below Cala- stony hillsides. Collectors' habitat remarks in- veras, elev. 3,000 m, Williams, Molina R,, & Wil- cluded barren mountain summits, covering vol- liams 21937 (F). QUEZALTENANGO: San canic rocks and shrubs, on steep slopes, shaded ar- Juan Ostuncalco, elev. 3,000 m, Williams, Molina royo bank, under cliffs, crest of ridge, well-drained R., & Williams 2271^0 (F, US) and 25531 (F, NA). soil between rocks, and rocky rhyolitic soil. The SAN MARCOS: Edge of cypress forest, San vines may be abundant. They may grow in shade Sebastian, elev. 3,050 m, Williams, Molina R,, & or full sunlight and in association with or separ- Williams (F). ately from other plants.

7. Vicia pulchella Humboldt, Bonpland, Kunth subspecies mexicana (Hemsley) C. R. Gunn (subgenus Vicilla^ section Cracca) Mexican Showy Vetch (Fig 8, Map 4)

Vicia pulchella Hum'boldt, Bonpland, Kunth subsp. mexi- Hemsley (cited here) used several characters to cana (Hemsley) C. R. Gunn stat. nov. V. mexicana separate F. mexicana from F. pulchella. He said Hemsley, Diag. PI. Nov. 3: 47. 1870. TYPE: MEXICO: Ghiesbrecht, Oaxaca (HOLOTYPE: K). that his species was glabrous, except for the young- V. sessei G. Don, Gen. Hist. Dich. Pis. 2: 318. 1832. est tips of the branches and that it was larger in all TYPE : MEXrOO : V. poliphylla Sessé á Mociño 3636 parts. Two parts were specifically mentioned: (ISOTYPE : MA ; fragment, F). Flowers nearly three times larger and ovary borne Vicia pulchella subsp. mexicana is restricted to on a much longer stalk. Of these characters, only the area of Jalisco east to Hidalgo and south to ñower size is valid in separating subspecies mexi- Colima and Morelos. In addition to the specimens cana from subspecies pulchella. Flowers of the cited here, about 50 others were annotated. In its former are at least 13 mm long, usually 13-18 mm, range, subspecies mexicaria and pulchella are whereas flowers of the latter are usually 10 mm or sympatric. less in length. 20 TECHNICAL BULLETIN 1601, U.S. DEPT. OF AGRICULTURE

FIGURE 8.—Vicia pulchella subsp. mexicana drawn from herbarium specimens: A, Habit (X 0.5) ; B, leaflets (X 1.5) ; (7, flowers (X 1.5) ; D, androecial sheath and style (X 3.5) ; iJ, sty lar hair pattern (X 10). GENUS VICIA WITH NOTES ABOUT TRIBE VICIEAE 21

Pringle 6786 (cited here) is a widely distributed JfS (US). DISTRITO FEDERAL: Wet places, collection of subspecies mexicana^ which has a stiff valley of Mexico, elev. 2,225 m, Pringle 6786 appearance when compared with the average ma- (ENCB, F, MEXU, MICH, NY, UC, US, VT). terial of subspecies pulchella. As a result, all HIDALGO: Luyamaloya, Pringle 13^71 (US). plants of subspecies Tnexicana are often thought of JALISCO: Steep slope of Nevado de Colima, as stiff, but this is generally not true. There are elev. 2,800 m, McVaugh IS 1^11 (MEXU, US). specimens of subspecies rrhexicana that lack this MEXICO : Tlalmanalco, elev. 3,050 m, RzedowsM character, and there are stiff specimens of sub- 191^7 (MEXU, MICH, US). MICHOACAN: species pulchella^ such as Pringle 12S1, Pine-oak forest, elev. 3,150 m, Hinton 131^52 Selected Mexican specimens examined: CO- (MICH, UC, US). MORELOS: Valle do LIMA : Volcano Colima, elev. 2,591 m. Goldsmith Tepeite, Lyonnet <& Elcoro IIH, 1782 (US).

8. Vicia americana Muhlenberg ex Willdenow subspecies americana (subgenus Vicilla^ section Americanae) American Vetch (Fig. 9, Map 5)

Vicia americana Muhlenberg ex Willdenow subsp. ameri- y. copelandii Eastwood, BuU. Torr. Bot. Club 32 (4) : 196. cana, Sp. PI. 3 (2) : 1096. 1802. AMcosa americana 1905. TYPE: UNITED STATES: Copeland 165, (Willdenow) Alefeld, Bonplandia 9:104. 1861. Siskiyou Co., Calif. (HOLOTYPE: CAS). TYPE : UNITED STATES : Muhlenberg 201, Pennsyl- V. pumila Heller, Muhl. 2 (1): 88. 1905. TYPE: vania (HOLOTYPE: B, photo). UNITED STATES : Helle?' 1938, Siskiyou Co., Calif. Orohus diffusus Nuttall, Eras. Oat. 1813. Reprinted in (HOLOTYPE: NY; ISOTYPES: F, GH. ISC, US). Pittonia 2:118. 1890. Lathyrus diffusus (Nuttall) V. waskingtonensis Suksdorf, West Amer. Sei. 15 (130) : Don, Gen. Hist. Dich. Pis. 2:336. 1832. TYPE: 59. 1906. TYPE : UNITED STATES : Suksdorf 26^3, UNITED STATES : No type preserved. Klickitat Co., Wash. (HOLOTYPE: WTU; ISO- V. sylvatica of Nuttall, Gen. 2: 97. 1818. Non L. (Incor- TYPES: F, GH, ISC). rectly attributed to Nuttall by subsequent authors. ) V. perangusta Greene, Leafl. 22: 267. 1912. TYPE: T'. oregana Nuttall in Torrey & Gray, Fl. N. Amer. 1: 270. UNITED STATES: Read, June 16, 1912, Coconino Co., Ariz. (SYNTYPE: Pro parte: Right specimen 1838. F. americana var. oregana (Nuttall) Nelson in Coulter & Nelson, Bot. Rocky Mts. 301. 1909. F. US). americana subsp. oregana (Nutt.) Abrams, Fl. Pac. V. perangusta YSLT. latiuscula Greene, Leafl. 2: 268. 1912. TYPE: UNITED STATES: Read, June 16, 1912, States 2:617. 1944. TYPE: UNITED STATES: Coconino Co., Ariz. (SYNTYPE: Pro parte: Left Nuttall, Oregon plains (HOLOTYPE: BM, photo; ISOTYPES: GH, PH). specimen US). y. hypolasia Greene, Leafl. 2: 268. 1912. TYPE: F. truncata Nuttall in Torrey & Gray, Fl. N. Amer. 1: 270. UNITED STATES: Blumer ISJfS, Chiricahua Mts., 1838. F. americana var. truncata (Nuttall) Brewer Ariz. (SYNTYPES: F, ISC, RM, US). in Brewer and Watson, Bot. Cal. 1:158. 1876. V. acicularis Greene, Leafl. 2: 268. 1912. TYPE: TYPE : UNITED STATES : Nuttall, Columbia plains UNITED STATES: Suksdorf 2013, Klickitat Co., (HOLOTYPE: BM, photo; ISOTYPES: GH, PH). Wash. (SYNTYPES: US, WTU). F. truncata var. villosa Kellogg, Proc. Cal. Acad. Sei. 1: 57. V. vexillaris Greene, Leafl. 2: 269. 1912. TYPE: 1854. F. americana var. villosa (Kellogg) Hermann, UNITED STATES: /. T. Worthley, 1896, Big Horn USDA Agr. Handb. 168: 83. 1960. TYPE : UNITED Co., Wyo. No type located. STATES: Kellogg, El Dorado Co., Calif. No types V. californica Greene var. madrensis Jepson, Fl. Cal. 2 : preserved. 386. 1936. TYPE: UNITED STATES: Noddin 28, F. calif ornica Greene, Fl. Fr. 3. 1891. TYPE : UNITED Madera Co., Calif. (HOLOTYPE: JEPS). STATES: Greene, June 1889, Calaveras Co., Calif. (HOLOTYPE and fragment: ND). Perennial.—Herbaceous vine, up to 2 m long, F. americana var. pallida Suksdorf in Leimbach, Deutsch. drying green, glabrous to pubescent. Trichomes Bot. Monats. 18 (2): 26. 1900. TYPE: UNITED golden or white. Leaves pinnately compound, STATES: Suksdorf 2111, Klickitat Co., Wash. terminating in a simple or ramified prehensile (HOLOTYPE: WTU; ISOTYPES: F, GH). tendril. Leaflets 4-14, entire to toothed, thin to F. durhrowii Eastwood, Bull. Torr. Bot. Club 32 (4) : 196. coriaceous, oval to linear, 5-60 by 1-15 mm, apex 1905. TYPE : UNITED STATES : Eastwood, July 5, 1902, Mariposa Co., Calif. (HOLOTYPE and frag- rounded to truncate to sharply acute and mu- ment: CAS). cronulate, base rounded to sharply acute. Stipules 22 TECHNICAL BULLETIN 1601, U.S. DEPT. OF AGRICULTURE

/ft^û, S/.-^M^^

FIGURE 9.—Ficta americana subsp. americana drawn from herbarium specimens : A, Habit (X 0.5) ; ß, leaflets showing size and shape variation (X 0.75) ; 0, flower ( X 3) ; D, androecial sheath and style (X 7) ; E, stylar hair pattern ( X 21) ; F, mature legumes (X 1) ; Ö, seeds ( X 5). GENUS VICIA WITH NOTES ABOUT TRIBE VICIEAE 23 inconspicuous, eglandular, usually sharply serrate. Mexican specimens examined.—STATE UN- Racemes elongate, with (2)-3-10 widely spaced KNOWN: Wright 9Jt5 (UC). CHIHUAHUA: flowers, shorter than subtending leaf. Calyx less Near Colonia Garcia, elev. 2,280 m, Townsend <& than one-half standard length, teeth unequal with Barter J^ (F,MEXU,MO,NY,POM,TEX,UC). 2 upper shortest poorly developed and curved and DISTKITO FEDERAL : Canada de Contreras, 2 lateral and basal straight deltoid not long at- elev. 3,100 m, Vazquez H, (ENCB). MEXICO : tenuate, gibbous at base, usually partially bluish Municipio de Tlalmanalco, elev. 3,050 m, Moncayo purple toward apex. Corolla varying shades of 25 (US) ; Rafael, elev. 2,700 m, RzedowsU 153 bluish purple to rarely white, with standard 12-25 (ENCB); Rzedowshi 19W (ENCB, ISC); mm long. Androecial sheath oblique at apex. Rzedowshi <è Holguin (ENCB). MORELOS: Stigma globular. Style apex encircled by hairs Z^môdildi, Paray 1280 (ENCB). with abaxial surface bearing elongated hairs form- Although F. americana subsp. amêricana is not ing brush. Ovary bearing soon deciduous golden an obligate self-pollinator, it is polymorphic, ex- hairs. Legume 25-35 by 5-6 mm, light to dark hibiting pronounced variations in leaflet shape, straw color, glabrous (pubescent in small popula- size, and thickness ; pubescence ; stipule shape and tions in Arizona, California, Oregon, and Wash- size ; height ; and flower number. It may be identi- ington), strongly reticulate, oblong, terete to fied easily and consistently by the number of partially flattened, oblique and acute at both ends, flowers per raceme, standard length, shape of on 4-5 mm stipe, bearing 8-14 ovules, with inner stylar hairs, unequal calyx teeth, ovule number, valve surface smooth, twisting loosely to tightly and percentage of seed circumference occupied by during dehiscence. Seed 3-3.5 mm in diameter, the hilum. Its phenotypic variability and wide subspherical to oblong, smooth, dull purplish geographic range have contributed to its taxo- brown to dull light brown, with scattered to dense nomic and nomenclatural confusion. This pheno- dark-brown mottles and points (some seeds so typic variability has been discussed and evaluated densely speckled as to appear almost mono- by Gunn {1968), chrome). Hilum color of seedcoat or silvery (due Vicia americana subsp. minor (Hooker) C. R. to funicular remnants), oblong to slightly wedge- Gunn stat, nov,^ based on V, americana var. minor shaped, 5-6 times longer than wide, parallel to Hooker (Fl. Bor. Am. 1: 157. 1831, syntype K), length of seed, occupying 20-33 percent of seed has not been found in Mexico, though one of the circumference, with lips of hilar groove color of herbarium sheets from Chihuahua {Townsend and hilum. Lens generally darker than seed, with Barber ^, UC) resembles this subspecies. This raised center 1.3-2 mm from hilum. sheet, although superficially similar to subspecies Distribution.—Vicia americana subsp. ameri- minor^ has been identified as subspecies aTnericana cana is found throughout southern Canada and using characters presented by Gunn {1968), Fur- Alaska and the United States essentially north of thermore, there are at least seven sheets of this col- the 74° F isotherm, from New England and ad- lection and the other six do not superficially resem- jacent Canada. The 74° F isotherm separates the ble subspecies minor. microthermal and mesothermal climatic regions One Mexican specimen, Wright 91^5 (UC), pos- (Thornthwaite, 1931), South and west of the sesses unusually narrow leaflets (a length/width terminus of this line in New Mexico (Gunn, 1968^ ratio of 10 or more), which do not exceed 42 by 1 fig, 5), the distribution of subspecies americana is mm. The occurrence of narrow leaflet plants governed by elevation or an ocean-cooled climate. throughout the range of V, americana subsp. amer- In Mexico, this subspecies has not been found be- icana has been discussed (Gunn, 1968,, pp, .197- low 2,200 meters. Sustained heat appears to be det- 198), This phenotype has been assigned specific rimental. The collections in Morelos, Distrito and varietal ranks in the United States. However, Federal, and Mexico represent both the southern I have concluded that these linear leaflet pheno- limit of this subspecies as well as possibly relic types merit no more than forma recognition for populations. There are no discernible differences these reasons: (1) They share all the technical between these central Mexican populations and characters exhibited by broader leaflet plants, (2) comparable populations in the United States. some plants have leaves with both broad and nar- 24 TECHNICAL BULLETIN 1601, U.S. DEPT. OF AGRICULTURE row leaflets on the same stem, and (3) the distribu- subsp. americana,, viz, shape and relative length of tion of the narrow leaflet plants has no geograph- the two upper calyx lobes and length of the style ical significance because they occur throughout covered by stylar hairs (fig. 10). In my study of the range of the broad leaflet plants. There is no the F. americana complex, I have never found a taxonomic advantage to propose formal names for plant that exhibited these two characters singly or either the narrow leaflet form or other forms based collectively. Therefore, two possibilities remain. on such minor characteristics as leaflet shape, pu- The first is the highly unlikely possibility that bescent legumes, or white flowers. Humboldt and Bonpland collected a new Mexican Vicia setifolia Humboldt, Bonpland, Kunth species of Vicia, which has not been collected by resembles the narrow leaflet form of F. americana anyone else. The second and more plausible possi- subsp. americana. The range of F. setifolia,, as re- bility is that they may have seen a narrow leaflet corded by Humboldt, Bonpland, and Kunth plant of F. americana subsp. americana in Mexico, {1821^,, pp, 500-601),, is '^crescit prope Mexico^ but they collected the three F. setifolia sheets in Quito et Santa Fe de Bogota,, alt, 1^170-1.500 Aea?." the Andes Mountains of northwestern South Vicia nigricans is the only vetch native to both America. North and South America. Except for F. humilis During their South American travels from and F. pulchella whose ranges were extended Bogotá, Colombia, to Quito, Ecuador, Humboldt south of the Chiapas border into adjacent Guate- and Bonpland could have seen only one endemic mala, there are no endemic Vicia species in Central Vicia species, F. andícola Humboldt, Bonpland, America. Kunth, which they collected and named (Sand- Two extant sheets of F. setifolia,, one in Paris with, 1926; Sprague, 1926; Steam, 1968), No (P), which is the lectotype (Burkart, 1966),, and other endemic Vicia species has been found in this the other, a fragment, in Berlin (B), were placed part of South America. Vicia andicola is a South in the Willdenow collection and subsequently American vicariate of F. americana subsp. ameri- saved. The main Berlin sheet was destroyed, but cana. It exhibits all the variability of the latter not before it had been photographed, and the nega- taxon and is strikingly similar to it. Only a tive, number 2346, was deposited at the Field Mu- thorough study of the two taxa can determine seum (F). These two specimens and the photo- their taxonomic status. graph are unique among Humboldt and Bonpland One of three technical differences is the relative New World Vicia collections because they contain lengths of the five calyx lobes. The lobes tend to no data about where they were collected. Lacking be more or less equal for F. andicola, whereas they published or unpublished data, one must review are definitely unequal in F. americana. There are the Humboldt-Bonpland pre-1824 New World some specimens of F. andicola that possess unequal itinerary for clues about where these plants were calyx lobes and closely resemble F. americana^ collected. even to being purplish at the apex. Another tech- Humboldt and Bonpland's Mexican travels nical difference is that the length of style covered were primarily in and around the State of Mexico by stylar hairs is longer for F. andicola than for (Humboldt, Bonpland, and Kunth, 182^^ pp. 500- F. americana. The third character is that the an- 501; Sprague, 1926), the southernmost limits of droecial sheath apex is rather truncate in F. andi- F. americana subsp. americana,, a taxon known for cola and oblique in F. americana. its plasticity. As previously noted, F. setifolia and Specimens of F. andicola with the same general this subspecies share several superficial and tech- and technical characters as F. setifolia at the U.S. nical characters that are not shared with other National Herbarium (US) inolviàQ Smith, Idroho, Mexican Vicia species. One may postulate that if <& Jaramillo-Mejie 1029, Usme, Colombia; Idrolo, any of the three sheets of F. setifolia were col- Jaramillo, Mesa-Bernal, <& Mora S22, Cundina- lected in or around the State of Mexico, the col- marca, Colombia; Maguire <& Maguire 61723^ lection either would be part of the F. mmericana Bolivar, Ecuador; and Hitchcock 220Jfi, Chim- complex or would represent a new taxon. borazo, Ecuador. There are two technical characters that may be These reasons would explain why Humboldt used to separate F. setifolia from F. americana and Bonpland did not record finding the narrow GENUS VICIA WITH NOTES ABOUT TRIBE VICIEAE 25 leaflet form of F. andícola. They did ñnd it, but Mexico. they called it V, setifolia. If F. setifolia was col- Burkart {1966) described F. setifolia var. lona- lected in South America and not Mexico, then it riensis Burkart from Argentina as possessing a becomes a synonym of F. andicolcu '^caliz Jtr-7 mm long, campanulado, pubescente- When Kuntze {1898, p, 73) reduced F. setifolia hirtulo, con 5 dientes acutisimos, los 2 superiores to a variety of F. gramínea Linnaeus, some taxon- de long, igual al tubo, inferiores mayores,'''^ The omists stated that F. graminea was an endemic of illustration of the spread calyx shows the two both South and North America. Vicia grandnâ is upper lobes to be no more than 2 mm longer than clearly a separate species and is not endemic to the lower three. The status of these and other F. Mexico. There are no herbarium specimens to in- setifolia-like plants can only be established by a dicate that the species has been introduced to thorough review of South American Vicia species.

9. Vicia americana Muhlenberg ex Willdenow subspecies mexicana C. R. Gunn (subgenus Vicilla, section Americanae) Mexican American Vetch

(Fig. 10, Map 5)

Vicia americana Muhlenberg ex Willdenow subsp. mexi- Mueller é Mueller 510, Nuevo León (HOLOTYPE: cana C. R. Gunn nov. suhsp. TYPE: MEXICO: MEXU ; ISOTYPES : F, TEX).

V. americana subsp. americana V. americana subsp. mexicana

v. andícola (broad leaflet) v. andícola (narrow leaflet)

FIGURE 10.—Vicia americana subsp. americana and subsp. mexicana compared with two phenotypes of V. andicola, a South American species: V. americana subsp. americana inflorescence and leaves (X 1), flower (X 2), androecial sheath and style (X 2), stylar hair pattern ( X 6) ; F. americana subsp. mexicana inflorescence and leaves ( X 1), flower (X 2), androecial sheath and style (X 2), stylar hair pattern (X 4) ; V. andicola (broad leaflet) inflorescence and leaves (XI), flower (X 2), androecial sheath and style (X 2), stylar hair pattern (x 4) ; F. andicola (narrow leaflet) inflorescence and leaves (X 0.5), flower (X 1.5), androecial sheath and style (X 1.5), stylar hair pattern (x 4.5). 26 TECHNICAL BULLETIN 1601, U.S. DEPT. OF AGRICULTURE Ä suhsp, americana calycis dentibus longe at- huila border (US). This subspecies is geographi- tenuatis essentialiter aequalihus differt. Differs cally separated from subspecies americana^ which from subspecies americana by possessing long- is found in the Chihuahua mountains. This is an attenuate calyx teeth of essentially eq,ual length. expected extension of the Arizona and New Mexico Vicia avwricana subsp. mexicana is known only population. In Texas, subspecies americana occurs from Nuevo León in the Sierra Madre Mountains no farther south than the Panhandle. A small from Monterrey to the Coahuila border area. In population of F. americana subsp. americana addition to the collection cited here, this subspecies found in the States of Mexico and Morelos as well has been collected by Mueller and Mueller 511 (F) in Mesa Canyon ; Pringle, 16 June 1888 (MEXU) ; as the Distrito Federal may be a relic of what once and Palmer ^^7, Caracol Mountains at the Coa- was a larger population in Mexico.

10. Vicia leucophaea Greene (subgenus Vicilla, section Mediocinctae) Mogollón Vetch

(Fig. 11, Map 5)

Vicia leucophaea Greene, Bot. Gaz. 6: 217. 1881. TYPE : eter, subspherical to squarish, smooth, monochrome UNITED STATES : Greene, Pinos Altos Mountains, dull brown or greenish ocher and mottled with N. Mex. (HOLOTYPE: NDG; ISOTYPES: F, MO). dark brown. Hilum. brown, oblong, 6 times longer V. mediocincta S. Watson, Proc. Amer. Acad. Arts and Sei. than wide, parallel to length of seed, occupying 21: 425. 1886. V. leucophaea var. mediocincta (S. Watson) Rose, Contr. U.S. Nati. Herb. 1: 119. 1891. about 23 percent of seed circumference, with lips TYPE : MEXICO : Palmer 341, Cumbre, southwestern of hilar groove color of hilum. Lens black, with Chihuahua. Type not located. raised center about 1 mm from hilum. Distribution.—Fíí^ leucophaea is a little Perennial.—Herbaceous vine, up to 8 dm long, studied species. Its range is restricted to south- drying green, sparsely villous. Trichomes golden eastern Arizona and southwestern New Mexico to tawny. Leaves pinnately compound, terminat- and south into Mexico through Sonora and Chi- ing in a simple (seldom ramified) prehensile tendril. Leaflets 6-8, entire, thin to coriaxîeous, huahua into Durango. linear to lanceolate to elliptic-oblong (basal Selected Mexican specimens.—CHIHUAHUA : leaflets occasionally obovate and truncate), 7-25 PHnqle 1230, cool slopes of Sierra Madre (F, by 1-6 mm, apex rounded to acute and mucronu- MEXU, NY, POM, US, VT). DURANGO: late, base tapered. Stipules inconspicuous, egland- Palmer 378, Otinapa (F, GH, MO, NA, NY, UC, ular, entire. Kacemes short, with 1-2 closely spaced US). SONORA : White 3208, burned over pme- flowers, shorter than subtending leaf. Calyx less land (ARIZ, MICH). than one-half to one-half standard length, teeth Vicia leucopJmea is the only North American essentially equal straight attenuate, not gibbous at Vicia species that I have not grown m a field plot base, green throughout. Corolla whitish to cream or a greenhouse. Data about its duration are con- color with standard veins and keel tip purple, with flicting. From studies of herbarium specimens, the standard 7-9 mm long. Androecial sheath obhque species would seem to be a perennial. This con- at apex. Stigma disc-shaped. Style apex glabrous, clusion is based on the appearance of the bases and style encircled by hairs about midway between not on the presence of rhizomes. stigma and ovary. Ovary appressed-villous, with Because of the placement of its stylar hairs, F. white hairs. Legume 25-40 by 4-6 mm, straw color leucophaea is unique among New and Old World to brown, appressed-villous, obscurely reticulate, Vicia species. Its relationship to other New World lanceolate, flattened, obliquely acuminate to almost Vicia species needs further study. This unique beaked at both ends, on 0.5 mm stipe, bearing 8-10 character was incorrectly recorded by Greene ovules, with inner valve surface smooth, twisting {1881), who said "style very villose at apex." He tightly during dehiscence. Seed 2-2.6 mm in diam- differentiated F. leucophaea from other Vicia GENUS VICIA WITH NOTES ABOUT TRIBE VICTEAE 27

FIGURE 11.—Vicia leucophaea drawn from herbarium specimens: A, Habit (x 0.5) ; B, leaflets (X 3) ; (7, flower (X 3) ; D, androecial sheath and style ( X 4.5) ; E, sty lar hair pattern ( X 14) ; F, mature legume (X 1) ; G^, seeds ( X 3). 28 TECHNICAL BULLETIN 1601, U.S. DEPT. OF AGRICULTURE species by its cream-colored standard changing to Some of the widely distributed Palmer 378 purple and its appressed hairs. Watson (1886) sheets bear a notation that is unexpected and not correctly described the placement of stylar hairs verified: '^Garbansea grows among grasses and and recognized its significance by using the char- other plants, it is said to be injurious to animals to acter as the source of his species epithet. eat it, has the same reputation at Alvarez, San Apical stylar hairs hold pollen in juxtaposition Luis Potosi." It is unlikely that this note is re- to the stigma and thus facilitate fertilization. corded on the correct specimen for these reasons : Vicia leucophaea flowers would be expected to be (1) The species is not known from San Luis self-pollinated in the bud stage because of the Potosí, (2) Vicia species are seldom injurious placement of the stylar hairs. They are too low to (poisonous), and (3) it is doubtful that this plant hold pollen around the stigma when the flower is occurs in sufficient numbers to constitute a notice- fully expanded. able range hazard.

11. Vicia sativa Linnaeus (subgenus Vicia^ section Vicia) Common Vetch

(Fig. 12, Map 5)

Vioia sativa Linnaeus, Sp. PI.: 736. 1758. TYPE: or obtuse to truncate or emarginate and mucronu- EUROPE. ( SYNTYPES : LINN, International Docu- late, base tapered to slightly rounded. Stipules in- mentation Centre microfiche 906.20, 906.21, and conspicuous to somewhat conspicuous, bearing 906.23). V, notata Gililbert, Fl. Lituan. 2 : 105. 1781. purplish nectary, usually siharply serrate. Racemes V. Mela Moench, Meth. 148. 1794. subsessile, with l-2-(3) closely spaced flowers, V. alha Moench, Meth. 148. 1794. shorter than subtending leaf. Calyx one-half or V. leucosperma Moeni^h, Meth. 148. 1794. less standard length, teeth subequal linear-subu- F. canadensis Zuccagni, Obs. Bot. Cent. I (39), No. 83. late to lanceolate with lower 3 bearing conspicuous 1806. V. cordata Wulfen ex Hoppe in Sturm, Deutschi. Fl. Heft yellowish-brown nectary, not gibbous at base, 32. 1812. green throughout. Corolla violet purple to rarely V. pimpinelloides Mauri, Pl. Rom. Cent. 13: 35, t. 1. 1818. white, with standard usually 18-30 mm long (8-18 V. incisa M. Bieberstein, Fl. Taur. Cauc. 3: 471. 1819. mm long in subspecies nigra). Androecial sheath V. glalra Schleicher, Cat. PL Helv. ed. 4: 37. 1821. oblique at apex. Stigma globular. Style apex en- V. intermedia Viviani, Fl. Lob. 42, t. 19, f. 1. 1824. circled by hairs with abaxial surface bearing elon- V. cosentini Gussone, Fl. Sie. Prod. 2: 426. 1828. V. melanosperma Reichen'bach, Fl. Germ. Exe. 530. 1832. gated hairs forming a brush. Ovary pubescent. V. erythosperma Reichembach, Fl. Germ. Exe. 530. 1832. Legume (25)-35~80 by 6-12 mm or 4.5-6 mm in F. nemoralis Tenore, Fl. Nap. 5: 118. 1835. subspecies cordata, yellowish to brown, rarely F. macrocarpa Bertoloni, Fl. Ital. 7: 511. 1847. white or black (black or nearly so in subspecies F. euneata Grenier & Godron, Fla. Fr. 1: 459. 1848. nigra), rather pubescent, strongly reticulate, F. nemoralis Boreau, Fl. Centr. Fr. ed. 3, 2: 172. 1857. linear, compressed to nearly terete, torulose or F. morisiana Jordan ex Boreau, Fl. Centr. Fr. ed. 3,2:172. smooth in subspecies macrocarpa, oblique and 1857. acute at both ends, stipe absent, bearing 5-13 F. communis Rouy, Fl. Fr. 5 : 208. 1899. ovules, with inner valve surface bearing fibrous F. maculata Rouy in Rouy & Foucard, Fl. Fr. 5: 211. 1900. cross ribs, twisting loosely to tightly during dehiscence. Seed 3.5-6 mm in diameter, subspherical Annual.—Herbaceous vine, up to 1 m long, dry- to spherical, sublenticular, lenticular, ovoidal, or ing green, glabrescent. Trichomes white. Leaves cuboidal, smooth, dull pale straw to dark greenish pinnately compound, terminating in a ramified ocher to green, lightly to intensely mottled and prehensile tendril. Leañets 4-18, usually entire pointed with light to dark brown, some so densely (toothed or incised in subspecies incisa)^ thin, mottled as to appear monochrome dark brown to linear or lanceolate to oblong or obovate, or blackish. Hilum generally color of seedcoat, elliptic to obcordate, 10-40 by 2-15 mm, apex acute wedge-shaped, 3-4 times longer than wide, parallel GENTJS VICIA WITH NOTES ABOUT TRIBE VICTEAE 29

^0.0. I^ucflu^

FIGURE 12.—Vida sativa L. subsp. sativa drawn from greenhouse-grown plants: A, Habit (X 0.5) ; B, stipule with nectary patch (X 3) ; (7, leaflets showing shape and apex variation (X 2) ; D, flower (X 2) ; JE7, androecial sheath and style (X 2.5) ; 2^, sty lar hair pattern (X 6) ; G, immature legume (X 0.5) ; IT, mature legume (X 1) ; /, seed shape and color variation (from white to nearly black) ( X 3). 30 TECHNICAL BULLETIN 1601, U.S. DEPT. OF AGRICULTURE

to length of seed, occupying 14-15 percent of seed Based on the surveyed herbaria, F. sativa subsp. circumference, with lips of hilar groove color of sativa is the only subspecies found in Mexico and hilum. Lens usually blackish or brown to light Central America. It is unlikely that subspecies straw on seeds with light-colored seedcoats, with amphicarpa (Dorthes) Ascherson & Graebner and raised center usually 1 mm from hilum (occa- subspecies incisa (M. Bieberstein) Arcangeli are sionally about 0.7 mm). in Mexico or Central America. Lack of mature Distribution.—Vieia sativa is a widely planted legumes and seeds often prevents the correct iden- seed and fodder crop in North America. The tification of subspecies cordata (Wulfen ex species has become naturalized in diverse areas of Hoppe) Ascherson & Graebner and subspecies southern Mexico and Guatemala. macrocarpa (Moris) Arcangeli. Of these two, Selected Mexican specimens examined.— cordata is more frequently found in the United CHIAPAS: Grassy slope with Quercv^^ elev. States than mdcrocarpa. Although some of the 1,860 m. Ton lJß7 (MICH, US). MEXICO: Guatemalan specimens are labeled F. angustifolia Comunidad Temascaltepec, elev. 2,600 m, Hinton Linnaeus, now subspecies nigra (Linnaeus) Ehr- 2m (MEXU, US). MICHOACAN : Cultivated, hart, apparently none is labeled correctly accord- Magaña 30 (MEXU). ing to flower length or legume color and shape. Selected Guatemalan specimens examined.— Of the five other subspecies, nigra is the most CHIMALTENANGO: Weed in garden, elev. likely to be found in Mexico and Central America 1,830 m, Stamdley 80^80 (F). QUEZALTE- and is depicted in figure 13. Care should be taken NANGO: Oak forest, elev. 2,400 m, Standley in measuring dried standards to determine 66^69 (F). SAN MAKCOS: Moist woods or whether they are below or above 18 mm in length. sandy field above Rio Tacana, elev. 2,700 m, They are often allowed to wither before pressing Standley 66068, 66085, 66131 (F). and thus their length is shortened. The legume The list of synonyms for this species includes color and tereteness are the best characters to only species epithets, and the list is incomplete. The multitude of synonyms at the species level and separate subspecies sativa from subspecies nigra. below is staggering. Six subspecies are keyed and Narrow leaflet plants are not necessarily the latter. described in "Flora Europaea" (Ball, 1968). This Because V. sativa is a self-pollinating polymor- is the best available treatment even though a phic species and often used as a seed crop, numer- flowering mature fruiting specimen must be avail- ous forms are named based on different colors, able to use the key. Unfortunately most Mexican shapes, and sizes of seeds, as well as flower color and Guatemalan specimens of V, sativa were col- variation. Vicia sativa exhibits more leaflet varia- lected in flower and thus the value of the key is tion than most Vicia species. It is still being limited. studied and bred for special uses. The cultivar The Ball key is as follows : Warrior is designed for use in the Southern

1. At least some leaflets toothed or incised subsp. incisa 1. AU leaflets entire, or at most crenate-dentate at apex. 2. Plant with underground stems bearing apetalous flowers and white, 1- to 2- seeded legumes subsp. amphicarpa 2. Plant without underground stems. 3. Corolla (8)-10-18 mm; calyx teeth shorter than tube; legume black or very dark brown subsp. nigra 3. Corolla 18-30 mm ; calyx teeth as long as or longer than tube ; legume yellow brown to dark brown, rarely almost black. 4. Legume 4.5-6 mm wide subsp. cordata 4. Legume 6-11 mm wide. 5. Legume contracted between the seeds, brown or yellow brown; seeds 3.5-6.5 mm subsp. sativa 5. Legume not contracted between the seeds, dark brown or almost black ; seeds 5.5-8 mm subsp. macrocarpa GENUS VICIA WITH NOTES ABOUT TRIBE VICIEAE 31

FIGURE 13.—Vicia sativa subsp. nigra drawn from greenhouse-grown plants : A, Habit (X 0.5) ; ß, stipule with nectary patch (X 3) ; G, leaflets {X 2) ; D, flowers (X 2) ; E, androecial sheath and style (X 4) ; F, sty lar hair pattern (X 8) ', G, mature legumes (X 1) ; H, 2 mottled and 2 black seeds (X 4). 32 TECHNICAL BULLETIN 1601, U.S. DEPT. OF AGRICULTURE

United States and may be of value in Mexico and brid of Lens culinaris and F. sativa. This claim is Central America. not accepted by most taxonomists. The different subspecies of V, sativa hybridize, In southern Mexico and Guatemala, F. sativa is and crosses made between F. sativa and F. known as alherja and grown as a seed, forage, or angicstifolia are often cited as one of the few ex- green manure crop. Seeds are planted on dry, well- amples of Vicia species hybridization. There is a drained soil and do best in full sunlight. They are remarkable claim by Moritz and vom Berg {1931) often planted in old grainfields or with oats (El that F. leganyi Rapaico & Lengyel arose as a hy- Agricultor Mexicano, 1908),

12. Vicia faba Linnaeus (subgenus Vicia^ section F aha) Horse-Bean, Faba (Fig. 14, Map 5)

Vicia faha Linnaeus, Sp. PI. 1: 737. 1753. Faha vulgaris hairs on adaxial surface. Ovary puberulous. Moench, Meth. 150. 1794. V. esculenta SaUsbury, Legume 30-200 by 10-30 mm, dark brown to black, Prod. Stirp. Hort. Chapel AUerton 339. 1796. glabrous, reticulate, oblong, terete to partially Orolus faha (Linnaeus) Brotero, Fl. Lusit. 2: 147. 1804. V. vulgaris S. F. Gray, Nat. Arr. Br. PL 2: 617. flattened (plump and bulging over seeds), 1821. V. equina Steudel, Norn. ed. 1: 881. 1821 and obliquely acuminate at both ends, on a 4-5 mm Reichenbach, Fl. Germ. Exc. 532. 1832. Faha faha long stipe, bearing 2-5 ovules, with inner valve (Linnaeus) House, N.Y. State Mus. BuU. No. 254: 457. surface bearing fibrous cross ribs between seeds, 1924. TYPE: NOT RECORDED. (HOLOTYPE: twisting loosely or occasionally rather tightly dur- LINN, International Documentation Centre microfiche 306.34). ing dehiscence. Seed variable in size and shape, 7- Faha hona Medikus, Vorles. Churpf. Phys. Ges. 2: 360. 30 by 6-17 mm, oblong to oval, flattened or terete, 1787. smooth, bright reddish brown to light to dark Faha major Desfontaines, Tabl. ed. 1: 196. 1804. greenish brown or light to dark purple and Potamogetón hifolius Lapeyrouse, Hist. Abr. Pyren. Sup. obscurely mottled and pointed with colors similar 27. 1818 {fide Bentham, Cat. PI. Pyren. 113. 1826). to the base colors. Hilum blackish to brown, com- Faha minor Roxburgh, Fl. Ind. 3: 323. 1832. pletely or partially covered by scurfy remnants of Annual or biennial.—Erect herb, up to 2 m tall, funiculus, elliptic on smaller seeds, oblong on drying black, glabrous. Leaves pinnately com- larger seeds, 3-6 times longer than wide, at right pound, terminating in a cusp. Leaflets 2-6, entire angles to length of seed, occupying about 18 percent of seed circumference, with lips of hilar (rarely toothed), fleshy, oval, 3-10 by 1-5 cm, apex grooves often white. Lens conspicuous, brown, rounded and mucronulate, base rounded. Stipules with raised center 1.3-2 mm from hilum. foliaceous, bearing purplish nectary, sharply serrate. Racemes subsessile, with up to 12 closely Distribution.—Vicia faba may not have become spaced flowers, much shorter than subtending leaf. naturalized in North or Central America, and its Calyx less than one-half standard length, teeth occurrence is probably governed by its cultivation. unequal with 2 upper shortest and poorly devel- Mexican specimens examined.—CHIAPAS : oped and 2 lateral and basal deltoid to somewhat Tenejapa, elev. 2,100-2,300 m, Breedlove 6833 attenuate, slight gibbous at base, occasionally (US) and 12503 (TEX, US) ; Tenejapa, elev. partially bluish purple toward apex. Corolla white 2,700 m. Ton 686 (MEXU, US) ; Zinacatán Cen- with faint brown streaks, bearing a large black or ter, elev. 200 m, Laughlin 373 (MEXU) and 2710, deep maroon patch on each wing (wings rarely 2715, 2719 (US). PUEBLA: Puebla, Arsène yellowish brown, reddish, or variegated), with (MO, US) ; Nicolas (US). ZACATECAS: Be- standard 2-3.5 cm long and 15 man wide. An- tween Bolanos and Guadalajara, Rose 3032 (US). droecial öheath oblique at apex. Stigma conical on Guatemalan specimens examined.—ALTA the abaxial side of the style. Style apex with brush VERAPAZ: Rarely planted, elev. 1,300 m, of hairs on abaxial surface and scattered short Standley 92166 (F). QUEZALTENANGO : GENUS VICIA WITH NOTES ABOUT TRIBE VICIEAE 33

FIGURE 14.—Vicia faha L. drawn from greenhouse-grown plants : A, Habit (X 0.5) ; B, stipule with nectary patch ( X 2> ; C, flowers (X 2) ; D, androecial sheath and style (X 4) ; E, sty lar hair pattern (X 8) ; F, dying plant with mature legumes (X 0.5) ; G, seed in legume (X 0.6). 34 TECHNICAL BULLETIN 1601, U.S. DEPT. OF AGRICULTURE Sold in market for food, elev. 2,300 m, Standley separated. Fruiting f)lants may be separated, 86565,86566 (F). though not with the ease that seeds can. Between Linnaeus {1735) recognized the Tournefort these two extremes are many phenotypes that have genus Faha in his first reference to this species. been selected by man and perpetuated by self- However, the binomial Vicia faVa was used by pollination. Many of them were named by Alef eld Linnaeus {1753, p. 737), and this is the legitimate, {1866, pp. 27^32) and Muratova {1931). A key to validly published name for this species in the these taxa using seed and flower characters may be genus Vicia, It is a matter of opinion as to whether found in Guinea {1953, pp. 75-79), who cited Gams F. faha is sufficiently different from other Vida {192Ji) as the source of his data. A complicated species to warant recognition of the genus F aba. codification of subspecific taxa (Hanelt, 1972) may The characters that may be used to circumscribe be biologically sound but will be difficult for lay- this genus are reliable, but according to most men to use. The holotype of F. faha var. faha is taxonomists they are not of a quality to clearly LINN 906.34, and the neotype of F. faha var. establish a separate genus. These characters have minor (Petermann) Beck is Westphal IfiS, col- also been used to place V, faha in its own subgenus lected in Ethiopia (Westphal, 197i, pp. 205-213). (Gray, 1821, pp, 617-618) or section (Ascherson Although I have found no record of variety minor and Graebner, 1909, pp. 987-989). The major ones in Mexico or Central America, it likely has been are (1) hilum at right angles to length of seed, (2) planted as a seed crop. plant without tendrils (erect) and blackening Vicia faha var. faha is a widespread Mediter- when drying, and (3) seed without significant con- ranean seed crop plant, known as Tiaha in Mexico centrations of any of the nonprotein amino acids and Central America, but it has not become nat- or related compounds found in most other tested uralized in these countries. Because haha seldom seeds of Vicia species (Bell, 1966; Bell and Tiri- sets seeds in warm regions, plants are usually cul- manna, 1965). tivated in the Tropics at an elevation of 2,000 The problem in using these characters to estab- meters or more. Most haha seeds are produced in lish the genus Faha is that other Vieia species also southern Mexico and northern Guatemala. Ac- exhibit these characters, except for the basal cording to Berlin, Breedlove, and Raven {197It) ^ hilum, which is unique for F. faha. Vicia narho- the Mayan-speaking people of the Chiapas high- nensis Linnaeus, often cited as a near relative of lands intercrop Vicia faha var. faha and Zea m/iys V. faha, has generally been excluded from the Linnaeus. The seeds are used as green shell and segregate genus Faha. dry beans to feed man and livestock. In Central A simple explanation, though not necessarily America, liaba is commonly cultivated in the correct, of why F. faha has these unusual charac- Guatemalan highlands, where it is an important ters involves these considerations. Like a few other source of food. members of the tribe Vicieae, flowers of F. faha Standley and Steyermark {19Iß) estimated that are self-pollinated in the bud. Therefore, any over 4 million pounds of seed are produced an- viable trait or trend that becomes established may nually in the Departments of Alta Verapaz, be perpetuated. The introduction of these traits Chimaltenango, Quezaltenango, San Marcos, Suchitepequez, and Totonicapan. Most seeds are and trends has been increased by the selective in- roasted and used like seeds of Arachis hypogaea fluence of man, who has had more effect on this Linnaeus (peanut) even though they are much Vicia species than most other species. Parallel ex- harder to chew than the latter. Although seeds are amples, though lacking the extreme phenotypes, eaten by most individuals without any danger, raw include V. ervilia, V. narhonenm Linnaeus, and or cooked seeds may cause severe hemolytic anemia V. sativa (including F. angustifolia). in some individuals 2 to 3 days after consumption. Linnaeus {1753, p. 737) mentioned a variant of This condition, known as "favism," is an inherited F. faha, "ß. Faha minor /. equina. Bauh. pin. 338," trait characterized by a deficiency of glucose-6- which may be recognized by its much smaller phosphate dehydrogenase. About 10-15 percent of terete seeds and legumes. Although the large flat the Black race and about 1 percent of Caucasians seeds of variety faha can be readily separated from of Greek and Italian origin have this deficiency seeds of variety minor, flowering plants cannot be (Hardin and Arena, 1974, pp. 88-^9). GENUS VICIA WITH NOTES ABOUT TRIBE VICIEAE 35 EXCLUDED SPECIES

Vicia pedunculata Peyritsch, Linn. 30: 80. 1859-60. Tex., and Wewoka, Okla., (both localities between TYPE : MEXICO : HOLOTYPE : Heller S74, Mexico 96° and 97" longitude). Even if V. reverchonii is con- (location unknown). Peyritsch has described a Vida sidered an element of the more widespread species, V. or vetchlike plant. His description is too general to be minutiflora Dietrich, the taxon should remain ex- useful. cluded from Mexico. The southernmost collection of Vicia reverchonii S. Watson. A sheet deposited at ARIZ V. minutiflora is around Galveston Bay, Tex. is correctly labeled V. reverchonii, but it bears this Vicia trifoliata Sessé & Mociño, FI. Mex. ed. 2:168. 1894. legend : J. Reverchon, 1881, Mexico. There is no col- lateral evidence that this species is found in Mexico. TYPE : UNKNOWN. Based on their description, the Vicia reverchonii has been collected only from Dallas, authors described a species of Galactia.

LITERATURE CITED

ALEFELD, F. GUINEA, E. 1866. LAND WIRTH SCHAFTLICHE FLORA. 363 PP. 1953. ESTUDIO BOTáNICO DE LAS VEZAS Y ARVEJAS Wiegandt and Hempel, Berlin. ESPAÑOLAS. 227 pp. Typographia Artística, AscHERSON, P., and GRAEBNER, P. Madrid. 1909. SYNOPSIS DER MITTELEUROPAICHEN FLORA. V. GUNN, C R. 6 (2), 1,255 pp. Engelmann, Leipzig. 1968. THE VICIA AMERICANA COMPLEX ( LEGUMINOSAE ). BALL, P. W. Iowa State Jour. Sei. 42: 171-214. 1968. VICIA. In Tutin, T. G., et al.. Flora Europaea, V. 2, pp. 129-136. Cambridge Univ. Press, 1969. GENERA, TYPES, AND LECTOTYPES IN THE TRIBE Cambridge, England. VICIEAE (LEGUMINOSAE). Taxou 18: 725-733. BARULINA, H. I. 1930. LENTILS OF THE U.S.S.R. AND OF OTHER COUN- 1971. SEEDS OF NATIVE AND NATURALIZED VETCHES OF TRIES. Bui. Appl. Bot., Genet, and Plant NORTH AMERICA. U.S. Dept. Agr. Agr. Handb. Breeding Sup. 40, pp. 1-307. 392, 42 pp. BELL, E. A. - and KLUVE, J. 1966. AMINO ACIDS AND RELATED COMPOUNDS. In 1976. ANDROECIUM AND PISTIL CHARACTERS FOR TRIBE Swain, T., Comi>arative Phytochemistry, VICIEAE (FABACEAE). Taxou 25 : 663-675. chap. 12. Academic Press, New York. HANELT, P. • and TiRiMANNA, A. S. L. 1972. DIE INTRASPEXIFISCHE VARIABILITÄT VAN VICIA 1965. ASSOCIATIONS OF AMINO ACIDS AND RELATED FABA L. UND IHRE GLIEDERUNG. KulturpflanZC COMPOUNDS IN THE SEEDS OF FORTY-SEVEN 22: 75-128. SPECIES OF VICIA I THEIR TAXONOMY AND NUTRI- HARDIN, J. W., and ARENA, J. M. TIONAL SIGNIFICANCE. Biochem. Jour. 97: 1974. HUMAN POISONING FROM NATIVE AND CULTI- 104-111. VATED PLANTS. 194 pp. Duke Univ. Press, BERLIN, B., BREEDLOVE, D. E., and RAVEN, P. H. Durham, N.C. 1974. PRINCIPLES OF TZELTAL PLANT CLASSIFICATION. HERMANN, F. J. 660 pp. Academic Press, New York. 1960. VETCHES OF THE U.S.—NATIVE, NATURALIZED, BURKART, A. AND CULTIVATED. U.S. Dept. Agr. Agr. Handb. 1966. NOTAS SOBRE LAS ESPECIES ARGENTINAS DE VICIA 168, 84 pp. (LEGUMINOSAE) DEL AREA MESOPOTAMINCO- PAMPEANA. Darwiniana 14: 161-194. HITCHCOCK, C. L. 1952. A REVISION OF THE NORTH AMERICAN SPECIES EL AGRICULTOR MEXICANO. OF LATHYRUS. Uuiv. Wash. Pub. Biol. 15: 1- 1908. ALBERJA. Agr. Mex. 25 (1) : 13-14. 104. GAMS, H. 1924. LEGUMINOSAE. In Hegi, G., Illustrierte Flora HOLMGREN, P. I., and KEUKEN, W. 1974. INDEX HERBARIORUM. Part 1, ed. 6. Regnum von Mittel-Europa, v. 4 (33), pp. 155^-1562. Lehmanns, München. Veg. 92: 1-397. HUMBOLDT, F. H. A. VON, BONPLAND, A. J. [G.], and GRAY, S. F. 1821. A NATURAL ARRANGEMENT OF BRITISH PLANTS. KUNTH, C. S. V. 2, 757 pp. Baldwin, Cradock, and Joy, 1824. VOYAGE AUX REGIONS EQUINOCTIALES DU London. NOUVEAU CONTINENT, FAIT EN 1799-1804, GREENE, E. L. PARTIE 6, BOTANIQUE, SECT. 3, NOVA GENERA ET 1881. NEW PLANTS OF NEW MEXICO AND ARIZONA. SPECIES PLANTARUM. V. 6, 422 pp. Gidc Son, Bot. Gaz. 6: 217-219. Paris. 36 TECHNICAL BULLETIN 1601, U.S. DEPT. OF AGRICULTURE

HUTCHINSON, J. PLITMANN, U. 1964. THE GENEBA OF FLOWERING PLANTS. 516 pp. 1968. BIOSYSTEMATICAL STUDY IN ANNUAL SPECIES OF Olarendon Press, Oxford, England. VICIA OF THE MIDDLE-EAST. In Zohary, M., KUNTZE, C. E. O. Critical Revision of Native Legumes and 1898. REVISIO GENERUM PLANTARUM. V. 3, 576 pp. Grasses With Special Reference to Economi- Felix, Leipzig. cally Important Varieties. U.S. Dept. Agr. KupicHA, F. K. Project No. CR-11, Final Tech. Rpt., Grant 1973. STUDIES IN THE VICIEAE 1 : THE NEW GENUS No. FG-lS-134. 137 pp. Washington, D.C. RADZHI, A. D. ANATROPOSTYLiA. Roy. Bot. Gard., Edinb. Notes 32: 247-250. 1971. EVOLUTION OF THE GENERA OF THE TRIBE VICIEAE ADANSON. Bot. Zhur., Moscow 56: 978-981. SANDWITH, N. Y. 1975. OBSERVATIONS ON THE VASCULAR ANATOMY OF 1926. HUMBOLDT AND BONPLAND'S ITINERARY IN THE TRIBE VICIEAE ( LEGUMINOSAE) . Bot. JOUr. ECUADOR AND PERU. Kew Roy. Bot. Gard. Bul. Linn. Soc. 70: 231-242. Misc. Inform. 1926, pp. 181-190. SHîNNERS, L. H. 1976. THE INFRAGENERIC STRUCTURE OF VICIA. Roy. 1948. THE VETCHES AND PEA VINES (VICIA AND Bot. Gard., Edinb. Notes 34: 287-326. LATHYRUS) OF TEXAS. Field & Lab. 16: 18-29. SPRAGUE, T. A. 1977. THE DELIMITATION OF THE TRIBE VICIEAE 1926. HUMBOLDT AND BONPLAND'S ITINERARY IN (LEGUMINOSAE) AND THE RELATIONSHIP OF COLOMBIA. Kew Roy. Bot. Gard. Bul. Misc. ciCER L. Linn. Soc. London, Jour. Bot. 74: Inform. 1926, pp. 23-40. 131-162. STANDLEY, P. C, and STEYERMARK, J. A. LASSETTER, J. S. 1946. FLORA OF GUATEMALA. Fieldiana : Bot. 24(5) : 1972. A BIOSYSTEMATIC STUDY OF THE VICIA LUDO- 361-363. VICIANA COMPLEX (LEGUMINOSAE). 73-9458, STEARN, W. T. Univ. Microfilms, Ann Arbor, Mich. 1968. HUMBOLDT, BONPLAND, KUNTH AND TROPICAL AMERICAN BOTANY. 159 pp. -f- map. Cramer, 1975. TAXONOMIC STATUS OF VICIA HASSEI (LEGUMI- Lehre. NOSAE). Madroño 23: 73-78. THORNTHWAITE, C. W. 1931. THE CLIMAXES OF NORTH AMERICA ACCORDING TO LINNAEUS, C. A NEW CLASSIFICATION. Gcog. Rev. 21: 633- 1735. SYSTEMA NATURAE, SIVE REGNA TRIA NATURAE 655. SYSTEMATICE PROPOSITA PER CLASSES, ORDINES, ToRREY, J., and GRAY, A. GENERA, ET SPECIES . . . LUGDUNI BATAVORUM, 1838. A FLORA OF NORTH AMERICA. V. 1, 711 pp. APUD T. HAAK. 11 pp. J. Wilhelm de Groot, Wiley and Putnam, New York. Amsterdam. TURNER, B. L. 1959. THE LEGUMES OF TEXAS. 284 pp. Uuiv. Texas 1753. SPECIES PLANTARUM. V. 2, 1231 pp. Salvie, Press, Austin. Stockholm. WATSON, S. MoRiTZ, O., and VOM BERG, H. 1886. CONTRIBUTIONS TO AMERICAN BOTANY. Amer. 1931. SEROLOGICAL STUDIEN ÜBER DER LINSWECKEN- Acad. Arts and Sei. Proc. W. S. 21, N. S. 8, pp. PROBLEM. Biol. Zentbl. 51: 290-^307. 414-468. MURATOVA, V. S. WESTPHAL, E. 1931. COMMON BEANS (VICIA FABA L.). Bul. Appl. 1974. PULSES IN ETHIOPIA, THEIR TAXONOMY AND Bot., Genet, and Plant Breeding Sup. 50, pp. AGRICULTURAL SIGNIFICANCE. 263 pp. Cent. 1-298. Agr. Publishing and Doc, Wageningen. GENUS VICIA WITH NOTES ABOUT TRIBE VICIEAE 37

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U.S. DEPARTMENT OF AGRICULTURE SCIENCE AND EDUCATION ADMINISTRATION POSTAGE AND FEES PAID WASHINGTON, D.C. 20250 U.S. DEPARTMENT OF OFFICIAL BUSINESS AGRICULTURE PENALTY FOR PRIVATE USE, $300 AGR 101