Lotus roudairei Bonnet and taxonomic relation­ ships between African and North American species of the tribe (Papilionaceae)

Tatjana E. KRAMINA Higher Department, Biological Faculty, Moscow State University, 119899 Moscow, Russia. [email protected]

Dmitry D. SOKOLOFF Higher Plants Department, Biological Faculty, Moscow State University, 119899 Moscow, Russia. [email protected]

ABSTRACT A comparative analysis of morphological features in NW African rou­ dairei, N American Lotus sect. Simpeteria, and American Lotus sect. Microlotus (= gen. s. str.) was carried out. According to the data obtained, these three taxa seemed not to form the distinct or subgenus KEYWORDS Lotus roudairei, Acmispon sensu P. LASSEN (1986). A new section Pseudosimpeteria with a Papilionaceae, single species, L. roudairei, is described within the Old World Lotus subgen. Loteae, Lotus. The relationships between Old World and New World Loteae are Africa, N America. briefly discussed.

RÉSUMÉ Une analyse morphologique comparative de Lotus roudairei nord-ouest-afri­ cain, Lotus sect. Simpeteria nord-américain et Lotus sect. Microlotus (= gen. Acmispon s. str.) américain a été réalisée. En partant des données obtenues, il apparaît que ces 3 taxons ne peuvent pas former un genre (ou sous-genre) particulier, Acmispon sensu P. LASSEN (1986). Une section nouvelle MOTS CLÉS Lotus roudairei, Pseudosimpeteria, avec une seule espèce L. roudairei, est décrite dans le genre Papilionaceae, Lotus subgen. Lotus, répandu dans l'Ancien Monde. Les relations entre les Loteae, Loteae de l'Ancien Monde et ceux du Nouveau Monde sont brièvement dis- Afrique, N Amérique.

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INTRODUCTION Generally, the native N American species of Lotus s.l. are distributed from Mexico to S The taxonomic boundaries of the genus Lotus Canada and from the Atlantic to Pacific coasts; remain one of the most complicated issues in the the only native S American species occurs in intetgenetic delimitation of the tribe Loteae. In a Chile. The section Simpeteria contains 2-3 btoad sense, the genus comprises, according to annual and 10 perennial species in the SW part diffetent views, from 100 species (POLHILL 1981) of U.S.A. (Arizona, California, Colorado, to 176 species (KlRKBRlDE 1994) on all conti­ Nevada, New Mexico, Texas, Utah), and in nents except the Antarctic. The majority of spe­ Mexico, south to Veracruz and Puebla. The spe­ cies occut in the Meditettanean tegion, cies tend to be geographically separated from Macaronesia, and in the western patt of North each othet, e.g. two endemic species occur in America, especially in California. All native Mexico, one in Nevada, and one in Arizona. The North-American species of Loteae belong to the center of diversity of sect. Simpeteria is located in genus Lotus in the broadest sense. Mexico and Arizona. The section Microlotus A number of authors disagree with the broad contains about 8 annual species primarily in concept of the genus Lotus and tend to bteak it Western N America (British Columbia, up into several separate genera. In particular the Washington, Oregon, California, Arizona, New taxonomic position of North-American species Mexico, and Mexico). One species, L. subpinna- has been disputed. The review of the discussion tus Lag., is however, restricted to Chile, and one was presented by OTTLEY (1923) and CALLEN species, L. unifoliolatus (Hook.) Benth. has a (1959). OTTLEY (1923) recognized four main wide area of disttibution, extending from approaches to the of N American Mexico, Texas and Arkansas to British Columbia Loteae. and Manitoba, and from the Pacific coast to North and South Carolina. In contrast with sect. 1. All N American species should be included Simpeteria, sect. Microlotus has a center of diver­ in the Old World genus Lotus. sity in California, and all the species reported for 2. All N American species should be treated as a United States occur also in California. Even the distinct genus Dougl. ex Benth. Chilean L. subpinnatus is very close to L. wrange- 3. American species should be excluded from lianus Fisch. & Mey. from California, and some­ Lotus of the Old World and segregated into seve­ times the two species ate merged. The members ral genera: Hosackia, Acmispon Rafl, Syrmatium of subgenera Hosackia and Syrmatium, which are Vogel, and Anisolotus Bernh. not the subject of this papet, occur in Western N 4. The genus Hosackia should include the America, from British Columbia and Idaho to majority of American species, whereas remaining Mexico. species should be left within the Old World genus Lotus. OTTLEY (1923) accepted a broad concept of the According to GlLLETT (1958), only one insigni­ genus Lotus and tecognized three subgenera in ficant trait separates Old Wotld Lotus species America: subgen. Hosackia Ottley (syn. gen. from American Loteae, viz. leaf morphology. Old Hosackia Dougl. ex Benth. s. stt.), subgen. World species have five leaflets, with the lower Acmispon Ottley (syn. gen. Acmispon Rafl, gen. pair (of which one leaflet is occasionally absent) Anisolotus Bernh.), and subgen. Syrmatium Ottley situated at the base of the tachis, simulating (syn. gen. Syrmatium Vogel). Species of the first foliaceous stipules, very close to the true stipules subgenus have membranaceous or foliaceous sti­ which, if present, are reduced to glands. Less pules, while in the remaining two subgenera they often, they have three leaflets, with petiolules but are glandulai. Subgen. Syrmatium differs from without petiole or rachis. American species have subgen. Hosackia and subgen. Acmispon by the three leaflets with a definite petiole or rachis, or indéhiscent fruits. Later OTTLEY (1944) recogni­ 4-19 leaflets, pinnately arranged, often alternate, zed two sections—Microlotus Benth. and the lowest one situated above the base of the Simpeteria Ottley—in subgen. Acmispon. tachis and well separated from the glandulat or

322 ADANSONIA, sér. 3 • 1997 • 19(2) Lotus roudairei (Papilionaceae)

membranaceous stipules. GlLLETT (1958) noted The name "Acmispon roudairef is accepted by that "even this definition fails for L. roudairei LOCK (1989) and by GREUTER et al. (1989). On Bonnet from Morocco which has up to 6 alter­ the other hand, LEBRUN & STORK (1992) and nately pinnate leaflets, the lowest well above the KlRKBRlDE (1994) again place the species in the base of the rachis and separated from the glandu­ genus Lotus; it is not clear however whether these lar stipules just as in Hosackia" (p. 363). COSSON authors include L. roudairei in sect. Simpeteria. named this species "L. bosackioides" (in herb.) thus implying a similarity to American species. BONNET (1893), describing L. roudairei in accor­ MATERIAL AND METHODS dance with the rules of botanical nomenclature, also compared it with North American Hosackia. The work is based on the study of herbarium The species is distributed in the W of N Africa specimens from herbaria LE, MHA, and P. The (Morocco, Algeria, and Tunisia), and in adjacent following American species were studied: sect. parts of Tropical Africa. Simpeteria: L. argyraeus (Greene) Greene, MAIRE et al. (1935) described a new species, L. L. grandiflorus (Benth.) Greene, L. greenei Ottley, simonae Maire, Weiller & Wilczek from the SE L. mearnsii (Britton) Greene, L. oroboides foothills of the Anti-Atlas mountains in Morocco (Humboldt, Bonpland & Kunth) Ottley, L. rigi- and included it, along with L. roudairei, in the dus (Benth.) Greene, L. strigosus (Nuttal ex proposed new section Stipulati Maire, Weiller & Torrey & A. Gray) Greene, L. tomentellus Wilczek. Recognition of sect. Stipulati was based Greene, L. utahensis Ottley, L. wrightii (A. Gray) on a single feature, the presence of true stipules Greene; sect. Microlotus: L. denticulatus (Drew) reduced to small dark glands. MONOD (1980) Greene, L. humistratus Greene, L. micranthus justly noted that not only L. simonae and L. rou­ Benth., L. salsuginosus Greene, L. subpinnatus dairei, but a number of Old World Lotus species, Lag., L. unifoliolatus (Hook.) Benth. [L. purshia- demonstrated glandular stipules, and therefore nus (Benth.) Clements & Clements], L. wrange- rejected the sect. Stipulati. According to LASSEN lianus Fisch. & Mey. (1986), L. simonae and L. roudairei have nothing For the study of floral morphology flowers were in common except the structure of the stipules; placed for two days into a mixture of equal parts in floral and vegetative characters L. simonae is a of glycerin, ethyl alcohol and water, and then true Lotus. LASSEN (1986, 1989) accepted at least dissected. For the study of ovule arrangement two genera of N American Loteae, namely and orientation additional material was used, Hosackia and Acmispon. He recognized sect. namely the herbarium specimens from MW, and Simpeteria within the genus Acmispon in accor­ flowers of L. corniculatus L., L. krylovii Schischk. dance with OTTLEY'S classification, where the & Serg., L. ucrainicus Klok., fixed in 70% ethyl section was included in subgen. Acmispon. alcohol in the field from several locations in LASSEN (1986) transferred L. roudairei to European Russia, Ukraine, and Kazakhstan. Acmispon sect. Simpeteria (Ottley) Lassen making a new combination, Acmispon roudairei (Bonnet) Lassen. Since sect. Stipulati had been described RESULTS AND DISCUSSION earlier than sect. Simpeteria, he selected L. simonae as lectotype of sect. Stipulati so that the latter A comparative morphological study of Lotus name would not interfere with OTTLEY's epithet1. roudairei and New World Loteae does not sup­ port the conclusion of LASSEN (1986), that L. roudairei belongs to the N American sect. Simpeteria. 1. LASSEN also reported, that he had studied the lectotype of L. simonae In MPU. Lectotyplflcatlon Is, however, not needed, OTTLEY (1944) gave a short and clear diagnosis because the type specimen exists ("[Morocco, prov. Tatta.] In of sect. Simpeteria: "Herbae vel suffrutices; carina alveo lapldoso amnls Bouzezza ad septentr. oasis Tatta ad obtusa, quam alis breviore; vexillo sine ungue; radices merldlonales Anti Atlantis. 730 m. 3 apr. 1934. Maire et Wilczek.", PI). stilo sub stigmate circumcirca barbato". Accord-

ADANSONIA, ser. 3 • 1997 • 19(2) 323 Kramina T.E. & Sokoloff D.D.

TABLE 1.—The main differences between Lotus roudairei Bonnet, Lotus sect. Microlotus Benth., and Lotus sect. Simpeteria Ottley.

Characters Microlotus Simpeteria L. roudairei

Can be annuals Yes Yes No Can be perennials No Yes Yes Corolla asymmetrical: wings and keel turned to the one side and obliquely oriented in relation to standard No Yes No Standard blade abruptly clawed Yes No Yes Wings conspicuously longer than the keel No Yes No Ovules*: micropylae... inferae alternantes alternantes Stylodium with a collar of spreading hairs below the stigma No Yes No Pollen grains: number of apertures** 4 4-7 3 Geographical distribution America America Africa

* Mlcropyle infera means that the ovule has the micropyle oriented towards the proximal end of the ovary; micropyle supera means that micropyle is oriented towards the distal end of the ovary. Micropylae alternantes means that the ovules are alternately oriented in the ovary, i.e. the micropylae of two nearest ovules are oriented in the opposite directions (see TIKHOMIROV & SOKOLOFF 1997). This important character was first used in the taxonomy of Loteae by LASSEN (1989). He demonstrated that the genus differed in this feature from and . All Old World Lotus species seemed to have micropylae alternantes (TIKHOMIROV & SOKOLOFF 1997). ** According to CROMPTON & GRANT (1993) and DÍEZ & FERGUSON (1994).

ing to our data, Lotus roudairei has neither an tic charactet of a group formed by these 3 sec­ obtuse keel, nor long wings, nor a standard blade tions. In particular, leaf morphology cannot be with indistinct claw, nor a stylodium with a col­ used as such a diagnostic featute. The definition lar of spreading hairs below the stigma. L. rou­ by GlLLETT (1958, see above) appears deficient dairei is indeed a perennial herb but this fact not only for L. roudairei but also for some othet alone does not seem to be a sufficient teason to species. There are several species in sect. transfer this species to sect. Simpeteria, because Simpeteria with sessile palmately compound this section, as well as the Old World Lotus spe­ leaves {L, utabensis Ottley, L. wrightii (A. Gtay) cies, comprises both annual and perennial plants. Greene). Their leaves do not display any signifi­ The main differences between sect. Simpeteria, cant difference from those of some species of sect. Microlotus and L. roudairei are summarized Lotus and Dorycnium (such as Dorycnium penta- in Table 1. Lotus roudairei differs as much from phyllum Scop, or Lotus polyphyllus Clarke) (see both sect. Simpeteria and sect. Microlotus as these OTTLEY 1924, 1944). On the other hand, sections diffet from each other, therefore, we LASSEN (1986) himself transfered the N Africian believe that L. roudairei should be accepted as a species Benedictiella benoistii Maire with 7-9- member of a monotypic new section, pinnately compound leaves into the genus Lotus Pseudosimpeteria. The name "Stipulat? cannot be and considered it a member of sect. Heinekenia used, because Lotus sect. Stipulati is lectotypified Webb. & Berth., which comprises several species by L. simonae (see above). with leaves typical for Old World Lotus species. We believe that L. roudairei demonsttates the It is difficult to include all 3 sections under dis­ charactets of Old World Lotus subgen. Lotus cussion {Simpeteria, Microlotus, and Pseudos­ (= subgen. Edentolotus Brand). When BRAND impeteria) in the genus, ot subgenus, Acmispon. (1898) recognized sections in this subgenus he Moreover, we are unable to indicate any diagnos­ attached a significant importance to leaf sttuctu-

324 ADANSONIA. ser. 3 • 1997 • 19(2) Lotus roudairei (Papilionaceae)

re to distinguish sect. Ononidium Boiss. (leaves 2. The foliage leaf has in the axil a shoot with a sessile, 3-foliolate) and sect. Quadrifolium Brand single leaf, namely that subtending the flower. (leaves sessile, of 4 leaflets, 3 of them attached to This subtending leaf of the flower is represented the top of rachis) from sect. Xantholotus Brand by two free stipules (Fig. IB). This structure can and sect. Erythrolotus Brand (leaves sessile, 5- arise from the case typical for the genus Coronilla foliolate with distinct rachis). Thus, we include and its closest allies where an axillary head occurs the section Pseudosimpeteria with leaves pinnately without any foliage leaf on the peduncle. The compound and distinct petiole, in Lotus subgen. number of flowers then becomes reduced to 1. Lotus. 3. The flower with two bracteoles is situated in In addition to unusual leaf structure, L. roudai­ the axil of a foliage leaf (Fig. 1C). Bracteoles are rei differs from the majority of species of subgen. found in a few Loteae species unrelated to each Lotus (including L. simonae) by the arrangement other (e.g. lotoides Fenzl.- of flowers. Tikhomirov, SOKOLOFF 1996, Lotus strictus Fisch. & Mey). Flowers situated in the axils of foliage Lotus roudairei has very short peduncles in the leaves were found in tribe Loteae only in Dory­ axils of foliage leaves (i.e. leaves with distinct cnium sanguineum (VURAL & KIT TAN 1983). blades). Each peduncle bears two dark glands, The listed types of flower arrangement may be each close by other, and a single flower. These distinguished through the study of relative orien­ dark glands are very similar to the stipules of tation of organs (see Fig. 1). A detailed study foliage leaves. The subtending leaves of flowers2 shows the inflorescence of L. roudairei to be in represented in Lotus (as well as in a number of accordance with the first of the 3 types listed 3 other Loteae ) by the leaves without a blade, cor­ above. Therefore, the inflorescence of L. roudairei, responding in morphology to the stipules of being strongly reduced, possesses a typical flower foliage leaves. Sometimes subtending leaves of arrangement for the genus Lotus. The unusual flowers consisting of two almost free stipules may structute of the inflorescence does not prevent the be observed. The structure of the inflorescence in inclusion of sect. Pseudosimpeteria into the subgen. L. roudairei may be treated in 3 different ways Lotus, because the SW Asian and NE African sect. according to the discussed data. Ononidium is characterized by similar (and even 1. The foliage leaf has in the axil a shoot with more reduced) 1-flowered heads. two aggregated (subopposite) leaves, each redu­ The only character distinguishing L. roudairei ced to one dark gland. One of the leaves (redu­ from all Old World species of Lotus s.l. [except ced to glands) subtends a flower (Fig. 1A). This for Himalayan Podolotus bosackioides Benth. = structure can arise from the typical case for the Lotus bosackioides (Benth.) Ali] is revealed by genus Lotus where an axillary head occurs posses­ DfEZ & FERGUSON (1994). Lotus roudairei has sing at the base a foliage leaf without any flower pollen grains with endoapertures 5-9 X 10- in the axil. The number of flowers then becomes 14 mm, while in remaining species they are 1- reduced to 1, and the blade of the foliage leaf is 5 X 3-10 mm. We believe however, that this cha­ lost. racter is insufficient evidence for recognizing L. roudairei as a member of a separate monotypic subgenus. On the other hand, an important dif­ ference was found between L. roudairei and the 2. i.e. the leaves bearing flowers in their axils. We do not use American species of sections Simpeteria and the term "bract" because the foliage leaf on the peduncle of Microlotus in the number of apertures (Table 1). Lotus and Is often Incorrectly regarded as a bract. Really, the foliage leaf on the peduncle bears no flower in the Lotus roudairei, as well as all Old World Lotus axil and cannot be treated as a bract. species, has 3 apertures (DlEZ & FERGUSON 3. The exceptions are pseudocytisus (Boiss.) Fertlg 1994), while in sect. Microlotus and sect. and Dorycnium sanguineum Vural. In both species the distinct Simpeteria pollen gains are tetra- or stephanocol- stipules are absent. The subtending leaves of flowers are broad scales in C. pseudocytisus, and leaves with entire green blade porate (CROMPTON & GRANT 1993; DIEZ & In D. sanguineum (VURAL & KIT TAN 1983; SOKOLOFF 1997). FERGUSON 1994).

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Lotus sect. Pseudosimpeteria Kramina & D.D. Perennial herbs without creeping rhizome. Sokoloff, sect. nov. (subgen. Lotus) Leaves shortly petiolate, with 3-6 leaflets. Rachis well developed, with more or less alternately atta­ Lotus sect. Eulotus auct. non Set., p.p.: Bonnet, J. Bot. ched leaflets. Stipules small, but conspicuous, fle­ (Morot) 7: 232 (1893). shy, dark. Heads 1-flowered, with peduncles Lotus sect. Stipulati Maire, Weiller & Wilczek, Bull. Soc. Hist. Nat. Afrique N. 26: 121 (1935), p.p., much shotter than foliage leaves. Peduncle bea­ excl. lectotypo. ring two small subopposite fleshy datk leaves, Acmispon Raf. sect. Simpeteria (Ottley) Lassen, one of which subtends the flower. Bracteoles Willdenowia 16: 107 (1986), p. min. p., excl. typo, absent. Calyx campanulate. Petals yellow; stan­ non Lotus sect. Simpeteria Ottley, Brittonia 5: 81 dard glabrous, abruptly clawed; keel roundly cur­ (1944). Acmispon auct. non Raf.: Greuter, Burdet & Long, ved, acute, slightly shorter than the wings. Med-Checklist, 4: 4 (1989); Lock, Legumes of Stylodium below the stigma glabrous. Ovules Africa. A Check-list: 339 (1989). with alternate micropylae. Fruit many-seeded, dehiscent, glabrous. Seeds smooth. Pollen grains Plantae perennes rhizomatis repentibus nullis, foliis with 3 apertutes. breviter petiolatis 3-6-foliolatis, Rachis distincta foliolis plus minusve alternatim ajfixis. Stipulae parvae, sed bene conspicuae, carnosae, nigrae. Capitula uniflora Sect. Pseudosimpeteria differs from sect. pedunculis quam folia laminas ferentes multo breviori- Ononidium Boiss. by the well developed rachis bus. Pedunculus apice par foliorum suboppositorum and from sections Lotus, Quadrifolium Brand, squamuliformium, carnosorum, nigrorum babens, quo­ Lotea (Medik.) Ser., Stipulati Maire, Weiller & rum unum in axilla sua florem ferens. Bracteolae nullae. Wilczek, Krokeria (Moench) Ser. and Erythrolotus Calyx campanulatus. Petala lutea, vexillum glabrum unguiculatum, carina rotundato-curvata, acuta, vix bre- Brand by the foliage leaf morphology and by the vior quam alae. Stylodium sub stigmate glabrum. Ovula single flowered peduncle always without a foliage micropylis alternantibus. Eructus polyspermi, dehiscentes, leaf, but with two small fleshy subopposite leaves glabri. Semina levia. Pollina 3-colporata. lacking a blade. Sect. Pseudosimpeteria can also be A sect. Ononidio Boiss. rachidi distincta, a sect. Loto, distinguished from L. benoistii (Maire) Lassen by Quadrifolio Brand, Lotea (Medik.) Ser., Stipulatis the dehiscent fruit; from sect. Microlotus Benth. Maire, Weiller & Wilczek, Krokeria (Moench) Ser. et Erythroloto Brand pedunculo folio laminam habente by alternate micropylae, pollen grains with 3 semper nullo, sed par foliorum squamuliformium flo- apettures, and the perennial habit; and from sect. remque unicum ferente atque structura foliorum reliquo- Simpeteria Ottley by the stylodium without a rum, a L. benoistii (Maire) Lassen fructu dehiscente collat of hairs below the stigma, the structure of differt. A sect. Microloto Benth. ovulis micropylis alter­ nantibus, pollinibus 3-colporatis habituque perenni, a the keel, wings, and standard, and pollen grains sect. Simpeteria Ottley stylodio sub stigmate glabro, nec with 3 apertutes. circumcirca barbato, carinae structura atque pollinibus 3-colporatis bene differt. TYPUS (et species unica).—Lotus roudairei Bonnet.

Fig. 1.—Diagrams of possible types of flower arrangement in Lotus roudairei. Each diagram demonstrates structures situated in the axil of a single foliage leaf. A detailed study shows the inflorescence of L. roudairei \o be In accordance with the diagram A. See also explanation in the text.

326 ADANSONIA, sér. 3 • 1997 • 19(2) Lotus roudairei (Papilionaceae)

Lotus roudairei Bonnet include in Lotus a number of taxa that are now accepted as distinct, e.g. Hammatolobium Fenzl. J. Bot. (Morot) 7: 232 (1893).—Acmispon roudairei and Medik. (Bonnet) Lassen, Willdenowia 16: 108 (1986).— Lectotype (hie designatus): Letourneux s.n. [Regnum Tunetanum] "In lapidosis inter Aïn Kebirita et oued Acknowledgments Chaba. lOJunio 1884" (P!). We are thankful to Prof. Dr. V.N. TlKHOMIROV and Lotus fruticulosus Coss., Bull. Soc. Bot. France 22: 57 Dr. S.B. YAZVENKO for helpful discussion, to Prof. Dr. Ph. MORAT, Prof. G. AYMONIN, J.-C. JOLINON, (1875), nom. nud., non Desf. Lotus hosackioides Coss., nom. in sched. and A.-M. BOUYER for their hospitality and assistance in D.D. SOKOLOFF'S work in herbarium of Laboratoite de Phanérogamie, Museum National GEOGRAPHICAL DISTRIBUTION.—North Africa d'Histoire Naturelle, Paris, and to Dr. I.K. (Tunisia, Algeria, Morocco) and North-Western FERGUSON, Dr. W.F. GRANT, Dr. J.H. KIRKBRIDE, Jr. who sent us the copies of their papers. part of Tropical Africa (Sahara).

Thus L. roudairei seems to be a true Old World Lotus and should not be considered as a "connec­ REFERENCES ting link" between the N American and African Loteae. There is a second African species often BONNET E. 1893.—Notes sut quelques plantes rares, nouvelles ou critiques de Tunisie (suite). /. Bot. thought to be closely related to N American spe­ {Morot) 7: 229-237. cies of Lotus. OTTLEY (1944) reported that "if it BRAND A. 1898.—Monographie der Gattung Lotus. should be desirable to segregate the American Bot. Jahrb. Syst. 25:166-232. species [of Lotus] that have indéhiscent from CALLEN E.O. 1959.—Studies in the genus Lotus those with dehiscent fruits, the logical procedure (Leguminosae). I. Limits and subdivisions of the genus. Canad. J. Bot. 37: 157-165. would be to unite them with the genus CROMPTON C.W. & GRANT W.F. 1993.—Pollen Helminthocarpon, and not to set them off in a morphology in Loteae (Leguminosae) with particular genus by themselves". The correct name for reference to the genus Lotus L. Grana 32: 129-153. Helminthocarpon A. Rich, is Vermifrux Gillett. DIEZ M.J. & FERGUSON I.K. 1994.—The pollen The single species of Vermifrux, V. abyssinica (A. morphology of the tribes Loteae and Coronilleae (Papilionoideae: Leguminosae). 2, Lotus L. and rela­ Rich.) Gillett, is restricted to E Africa and ted genera. Rev. Palaeobot. Palynol. 81: 233-255. Yemen. Vermifux is often considered as closely GILLETT J.B. 1958.—Lotus in Africa south of the related to Lotus s.l. and is sometimes included in Sahara (excluding the Cape Verde islands and this genus (see for example POLHILL 1981). Socotra) and its distinction from Dorycnium. Kew Detailed studies showed that the genus Bull. 13:362-381. Vermifrux could neither be treated as being rela­ GREUTER W., BÜRDET H.M. & LONG G. 1989.— Med-Checklist. A critical inventory of vascular plants of ted to American Lotus species, nor to any other the circum-mediterranean countries. 4. Genève, Berlin. species of Lotus and should be merged with the KlRKBRIDE J.H. Jr. 1994.—Taxonomic Circum­ monotypic W Mediterranean genus Dorycnopsis scription of the Genus Lotus Linnaeus (, (TlKHOMIROV & SOKOLOFF 1997). Thus, we Loteae), Its Tribal Position, and Its Species: 11-15, would conclude, that close relatives of N in Proc. 1 Intern. Lotus Symposium. St. Louis. LASSEN P. 1986.—Acmispon roudairei, Lotus benoistii, American taxa of Loteae are absent in African in GREUTER W. & RAUS Th. (eds.), Med-Checklist flora; they are absent also in Europe. We suggest, Notulae, 13. Willdenowia 16: 108, 111, 112. that neither subgenera nor sections comprising LASSEN P. 1989.—A new delimitation of the genera both native American and Mediterranean species Coronilla, Hippocrepis, and Securigera (Fabaceae). can be distinguished within the genus Lotus. The Willdenowia 19: 49-62. New World Lotus species form several distinct LEBRUN J.-P. & STORK A.L. 1992.—Enumeration des Plantes à Fleurs dAfrique Tropicale. 2. Lotus: 101- groups that strongly differ from each other. They 102, Conservatoire et Jardin Botaniques de la Ville could be joined only within a very broad concept de Genève. of the genus Lotus as POLHILL does (1981). In LOCK J.M. 1989.—Legumes of Africa. A checklist. this case, it would perhaps be necessary to also Royal Botanic Gardens, Kew. MAIRE R., WEILLER M. & WILCZEK E. 1935.—

ADANSONIA, sér. 3 • 1997 • 19 (2) 327 Kramina T.E. & SokoloffD.D.

Sertulum austro-maroccanum. Bull. Soc. Hist. Nat. Cytisopsis pseudocytisus (Boiss.) Fertig and taxonomic Afrique N. 26: 120-122. position of this species in the tribe Loteae MONOD Th. 1980.—Contribution a l'étude des Lotus (Papilionaceae): 14. in Abstr. 6th Conf. Young bot. in (Papilionaceae) ouest-sahariens et macaronésiens. St. Petersburg. St. Petersburg [in Russian]. Adansonia, sér. 2, 19: 367-402. TIKHOMIROV V.N. & SOKOLOFF D.D. 1996.—On OTTLEY A.M. 1923.—A revision of the Californian the genera Hammatolobium Fenzl and Tripodion species of Lotus. Univ. Calif. Publ. Bot. 10: 189-305. Medik. (Papilionaceae, Loteae s.l.). Feddes Repert. OTTLEY A.M. 1944.—The American Loti with special 107: 209-217. consideration of a proposed new section, Simpeteria. TIKHOMIROV V.N. & SOKOLOFF D.D. 1997.— Brittonia 5: 81-123. Taxonomic position of Vermifrux abyssinica (A. POLHILL R.M. 1981.—Loteae DC, Coronilleae Rich.) Gillett and taxonomy of the tribe Loteae s.l. (Adans.) Boiss: 371-375, in POLHILL R.M. & RAVEN (Papilionaceae). Feddes Repert. 108: 335-344. P.H. (eds.), Advances in Legume Systematics 1. Kew. VURAL M. & KIT TAN 1983.—New taxa and records SOKOLOFF D.D. 1997.—On the morphology of from Turkey. Not. Roy. Bot. Card. Edinb. 41: 65-76.

Manuscript received 23 May 1997; revised version accepted 25 July 1997.

328 ADANSONIA, sér. 3 • 1997 • 19(2)