Pacific Science (1979), vol. 33, nO. 1 © 1980 by The University Press of Hawaii. All rights reserved Vicia menziesii Sprengel (Fabaceae) Rediscovered: Its Taxonomic Relationshipsl J. STUART LASSETTER 2 and CHARLES R. GUNN 3 ABSTRACT: Taxonomic relationships among five taxa of Vicia, traditionally recognized at the species level, were studied. Data used included plant and seed morphology, seed amino acid content, palynology, and cytology. Two Old World species, V. dumetorum Linnaeus and V. pisij'ormis Linnaeus, are shown not to be closely related to the Hawaiian and American taxa. The Hawaiian species, V. menziesii Sprengel, is maintained as a species and is shown to be closely related to V. gigantea Hooker of North America and V. nigricans Hooker & Arnott ofSouth America. The latter two species are treated as subspecies under the name V. nigricans. The karyotype and chromosome number are reported for the first time for the endangered species, V. menziesii. Vicia menziesii Sprengel, which is endemic to first three eophylls each bearing four leaflets the island of Hawaii and was last collected and a tendril. in 1915, has been regarded as extinct. It was A review of·New and Old World vetches rediscovered in 1973 on the same island by located four other species with diagnostic Wayne Gagne and Mae Mull (c. H. Lamou­ characters similar to those of Vicia menziesii: reux and D. Herbst, personal communica­ V. nigricans, V. gigantea, V. pisiformis, and tions). It is now considered an endangered V. dumetorum. Although Kupicha (1976) species (United States Department of the apparently combines V. nigricans Hooker & Interior 1978). This species was not men­ Arnott of Chile and Argentina and V. tioned by Kupicha (1976) when she presented gigantea Hooker of the Pacific Northwest of a general worldwide taxonomic treatment of North America, she neither formally pro­ the genus. posed this combination nor made any tax­ Vicia menziesii exhibits these diagnostic onomic judgment at the subspecific level. characters: (1) plants large, rank and darken­ These three New World taxa were originally ing or blackening with age or on drying; described as species: V. menziesii in 1826, (2) mature flowers rose-purple; (3) oblong V. nigricans in ·1830, and V. gigantea in 1831. standard; (4) compressed styles with encircl­ They traditionally have been· treated as ing apical hairs; (5) stipitate legumes without species in various floras or monographs of inner "woolly" parenchymatous tissue; (6) their respective geographical regions (Degen­ seeds with hilum occupying 75 percent of the er, Degener, and Gunn 1970, Hermann 1960, seed circumference and containing canava­ Reiche 1898, Welsh 1974). Vicia pisiformis nine; (7) six pairs of acrocentric chromo­ Linnaeus of middle Europe and V. dume­ somes, one pair with satellites, and one torum Linnaeus of Eurasia have been treated submetacentric pair; and (8) seedlings with as species in various floras or monographs of the .first eophylls at the fourth node and the their respective geographical regions (Ball 1968, Fedtschenko 1948), and both were studied by Kupicha. 1 Manuscript accepted 15 October 1978. 2 Eastern Kentucky University, Department of Bio­ logical Sciences, Richmond, Kentucky 40475. 3 Plant Taxonomy Laboratory, Science and Education MATERIALS AND METHODS Administration, United States Department of Agricul­ ture, Agricultural Research Center, Beltsville, Maryland Specimens from the following herbaria 20705. were examined: BAFC, BISH, CONC, F, 85 86 PACIFIC SCIENCE, Volume 33, January 1979 15mm V. DUMETORUM V. GIGANTEA V. MENZIESII V. NIGRICANS V. PISIFORMIS FIGURE 1. Styles (tOP row) and standards (bottom row) of five species of Vicia. GH, NA, NY, R, SGO, SP, US [herbarium Pridham 1967). When lengths of flowers, abbreviations throughout this paper defined leaflets, legumes, seeds, hila, or stems are by Holmgren and Keuken (1974)]. Morpho­ considered, these five species are among the logical characters examined include those largest in the genus. used by Gunn (1968), Lassetter (1975,1978), Vicia nigricans and V. gigantea have and Kupicha (1976). External seed characters ochreoleucous to orange flowers, often with were compared, and seed amino acid con­ a tinge of rose-purple, Vicia menziesii flowers tents were evaluated. Pollen was examined, are ochreoleucous when young and rose­ and karyotypes were compared. No artificial purple when mature. Vicia dumetorum is hybridizations of these perennials could be blue-purplish flowered, and V. pisiformis is attempted due to lack of suitable growth yellow flowered. Although the significance of chambers that would provide cool tempera­ flower color may not have been adequately tures and high humidity required by the surveyed for the genus, Kupicha (1976) taxa. Greenhouse and field plot attempts to noted that it is of little value at the section grow Vicia gigantea and V. nigricans at level. Among the native North American Beltsville, Maryland, failed to produce flow­ species, V. gigantea is the only species with ering plants. No attempt was made to grow ochreoleucous to orange flowers tinged with V. menziesii to flower. rose-purple. If Hawaii is considered, V. menziesii is the only native New World­ Oceania species with a distinct rose-purple pigment in the flower. The rose-purplish RESULTS AND DISCUSSION pigment in both species becomes more Vicja menziesii, V. nigricans, and V. gigan­ pronounced as flowers age. Burkart (1966), tea, but not V. pisiformis and V. dumetorum, Moore and Scotter (1976), and Reiche (1898) usually darken or blacken with age or upon listed about 33 native species for South drying due to the presence of L-3, 4-dihy­ America. About 20 of the 33 species are droxyphenylalanine (L-dopa) (Andrews and valid and only 7 of these have ochreoleucous Vida menziesii Sprengel Rediscovered-LAssETTER AND GUNN 87 to orange or reddish flowers. Four of these similar study, Tschiersch and Hanelt (1967) represent forms of V. nigricans and should be also established three groups: (1) canavanine­ regarded as synonyms. Flowers of the other producing species; (2) species without cana­ native North and South American species vanine but with fJ-cyanoalanine and its y­ are bluish, white, lavender, or purple, but glutamyl derivative; and (3) species without never with a definite orangy or rose-purple canavanine but with high levels of arginine. color. Tschiersch and Hanelt (1967) placed V. The shape of the standard in all five dumetorum and V. pisiformis in their group 3. species is oblong (Kupicha 1976 and Figure Of the five species under consideration, only 1). V. dumetorum was included by Bell and The nature of stylar pubescence and com­ Tirimanna (1965) and Bell (1966) when they pression ofthe style is an important morpho­ surveyed the genus for seed amino acid con­ logical character in Vida (Gunn and Kluve tent. This species contains y-hydroxyarginine 1976, Kupicha 1976, Lassetter 1975). Gunn and an additional compound which is and Kluve found that about 40 percent of probably hydroxycitrulline (Bell 1966; Bell, the 106 species surveyed had apical hairs personal communication). Vida pisiformis, encircling the style. Members of several of analyzed at a later time, exhibited no amino Kupicha's sections, including Cassubicae acid pattern typical forthe genus. It contains with V. nigricans and Vicilla with V. pisi­ an unusual neutral amino acid and an unusual formis and V. dumetorum, have stylar pubes­ acidic amino acid (Bell; personal communica­ cence distributed evenly around the style. tion). Based on this biochemical compari­ Vida dumetorum is an exception in its section, son, V. dumetorum and V. pisiformis are because its stylar pubescence is densely con­ different. Tschiersch and Hanelt (1967) found centrated on the abaxial side (Figure 1). The that V. dumetorum and V. pisiformis were other four species iIi this study have evenly similar, differing only in amounts ofthreonine pubescent styles, the predominant condition and arginine and in presence or absence of in the subgenus Vicilla. Compression of alanine. These authors, however, may not styles is dorsal in all five species. Kupicha have analyzed for the neutral and acidic acids discussed the types of stylar compression. Bell found in V. pisiformis. Their data show The legumes ofthe five species are similar, that neither species is a canavanine producer. except that those of Vida menziesii may be Vida menziesii, V. nigricans, and V. gigantea much larger. These general legume characters contain appreciable amounts of canavanine are typical for the subgenus Vidlla (Kupicha with no trace of fJ-cyanoalanine or y­ 1976). glutamyl-fJ-cyanoalanine (Bell and B. V. All five species have a circumlinear hilum Charlwood, personal communications). This that occupies 75 percent of the seed circum­ would place these vetches in the canavanine­ ference. Only 15 percent of 100 species (± 150 containing group of Bell and Tschiersch and species in the genus) surveyed exhibited a Hanelt. Tschiersch and Hanelt (1967) stated circumlinear hilum (Gunn 1970). The seeds that biochemically defined species groupings of the five species have a similar external and taxonomically (they presumably mean morphology, except that Vicia menziesii seeds morphological data) defined groups are never average about 25 percent greater in diameter completely identical. Bell (1971) similarly (Figure 2). noted that his three main groups correspond On the basis of seed amino acid data, Bell in general to accepted subgeneric groups (1971) divided Vida into three groups: (1) based on morphology and cytology. These canavanine-producing species; (2) species not groups predate Kupicha's report, and in this producing canavanine but having high con­ instance the biochemical and morphological centrations ofy-glutamyl-fJ-cyanoalanine and data mesh. Kupicha (1976) placed V. pisi­ lower concentrations of free fJ-cyanoalanine; formis and V. dumetorum in a different and (3) species without canavanine and with­ section from V. nigricans. out compounds in the second group. In a It has been clearly established that karyo- U~I Y.MENZIESII A. .n. ,j :r\5·~ nn un Y.MENZIESII B. Jt j' n;(11 It " I 23'?4 5 6 7 V.