Desmodium subsecundum Meibomia subsecunda (Vogel) Kuntze, Revis. Gen. Pl. 1: 198. 1891 Meibomia leiocarpa var. subsecunda (Vogel) Schindl., Rep. Spec. Nov. Beiheft 49: 370. 1928 Desmodium discolor Vogel, Linnaea 12: 103. 1838 Meibomia discolor (Vogel) Kuntze, Revis. Gen. Pl. I : 198. 1891 Desmodium leiocarpum Vogel, Linnaea 12: 102. 1838 Meibomia discolor var. villosa Hoehne, Mem. Inst. Butantan 1(1): 23. 1921 Desmodium discolor var. villosum (Hoehne) Malme, Ark. Bot. 18(7): 17. 1922 Meibomia discolor var. pohlii Schindl., Repert. Spec. Nov. Regni Veg. 20: 151. 1924 A Taxonomic Revision of Desmodium (Leguminosae, Papilionoideae) in Brazil Lima, Laura Cristina Pires Queiroz, Luciano Paganucci De Tozzi, Ana Maria Goulart De Azevedo Lewis, Gwilym Peter Phytotaxa 2014 2014-05-16 169 1 1 119 353J5 Vogel, Linnaea Vogel, Linnaea [194,683,1087,1112] Magnoliopsida Fabaceae Desmodium Plantae Fabales 80 81 Tracheophyta species subsecundum     Desmodium discolor Vogel, Linnaea12: 103. 1838.   Meibomia discolor(Vogel) Kuntze, Revis. Gen. Pl. I: 198. 1891. Lectotype(designated here):— BRAZIL. Brasil Meridional. F. Sellow4955( holotypeB†, lectotype FOBN002259!).    Desmodium leiocarpumvar. β  Vogel, Linnaea 12: 102. 1838, nom nud. Based on:— BRAZIL. Brasil Meridional, entre Campos e Vitória, F. Sellow s.n.(E!, K!).    Meibomia discolor var. villosaHoehne, Mem. Inst. Butantan1(1): 23. 1921.   Desmodium discolorvar. villosum(Hoehne) Malme, Ark. Bot.18(7): 17. 1922. Lectotype(designated here):— BRAZIL. São Paulo: Cantareira,  01 March 1918,  Horto Oswaldo Cruz1570 (SP!). Remaining syntypes: Brazil. São Paulo: Horto Oswaldo Cruz 2234(SP!), syn. nov.    Meibomia discolor var. pohliiSchindl., Repert. Spec. Nov.Regni Veg. 20: 151. 1924. Lectotype(designated here):— BRAZIL. Goiás: Santa Cruzde Goiás, J.B.E. Pohl2603( lectotype W0027268!, isolectotypes M!, K!). Remaining syntype: Brazil. Minas Gerais: Lagoa Santa, E. Warming 2970(C*!, P!), syn. nov. Erect, branched shrub, 1–3 mtall, without a xylopodium; stems not virgate, not slender, cylindrical, striate, densely velutinous or villose or uncinate or sparsely tomentose, uncinate and puberulous-uncinate, rarely puberulousuncinate, not glaucous; internodes 2.6–5.9 cmlong. Stipules 7–11 × 2–4 mm, ovate-lanceolate, auriculate, semiamplexicaul, inserted perpendicularly at the base of the leaf petiole, free from each other, apex caudate, margin ciliate, densely tomentose or sparsely tomentose and uncinate, rarely densely puberulous-uncinate, veins conspicuous, caducous or generally persistent at the base of the inflorescence; auricle 2–4 mmlong. Leaves trifoliolate; petiole 5–36 mmlong, cylindrical, canaliculate, densely velutinous or villose or sparsely tomentose and uncinate; rachis 5–16 mmlong; stipels 2–6.5 mmlong, lanceolate or subulate, margin straight, tomentose or puberulous-uncinate or glabrescent on the outer surface, caducous or persistent; leaflets discolorous, chartaceous, membranaceous, papyraceous or coriaceous, venation eucamptodromous, primary, and secondary veins prominent, another veins flush with the abaxial surface, indumentum densely or sparsely tomentose or velutinous or puberulous on the primary, secondary and tertiary veins on the abaxial surface, adaxial surface puberulous and    DESMODIUM(LEGUMINOSAE, PAPILIONOIDEAE) IN BRAZIL  Phytotaxa169 (1) © 2014 Magnolia Press • 81 82• Phytotaxa169 (1) © 2014 Magnolia Press   FIGURE 29.  Desmodium subsecundum.A. Flowering and fruiting branch. B. Stipules inner surface. C. Calyx inner surface (opened). D. Parts of the corolla from the bottom to top: keel petals, wing petals, and standard. E. Androecium monadelphous (opened). F. Gynoecium. G. Fruits, above loment with the first article aborted, below, loment without the first article aborted. H. Seed. (A: Cunha s.n.; B, G–H: Kummrow 1094; C–F: Salimena 996).  LIMA ET AL. sparsely velutinous, uncinate hairs covering both surfaces, terminal leaflet 4.3–8.9 × 1.9–4.8 cm, elliptic, lanceolate, oblong, ovate or ovate-rhombic, base oblique, obtuse or truncate, apex acute, obtuse or retuse, apiculate, lateral leaflets 2.8–5.1 × 1.6–2.7 cm, the same shape as the terminal leaflet. Inflorescence a terminal or axillary pseudoraceme; the main axis longer than the adjacent leaves, 22–36.5 cmlong, densely or sparsely hispid and uncinate, rarely velutinous, 2 flowers per node; primary bract 1.5–2 mmlong, ovate-lanceolate, margin ciliate, hispid or tomentose and puberulous-uncinate on the outer surface, caducous, veins conspicuous on the outer surface; secondary bract ca. 0.5 mmlong, subulate, margin ciliate hispid and puberulous-uncinate on the outer surface, caducous, veins inconspicuous on the outer surface; pedicel 7–8 mmlong, densely or sparsely hispid and puberulous-uncinate or densely uncinate. Flowers 7–10 mmlong; calyx bilabiate, tube campanulate, 1–1.2 mmlong, tomentose and puberulous-uncinate on the outer surface; upper lip bifid, the 2 teeth joined for ca. ¾ of their length, acute, ca. 0.1 mmlong; lower lip trifid, lateral tooth triangular, 1–1.5 mmlong, central tooth triangular, 1.5–2 mmlong; corolla lilac, standard 8–8.5 × 5–7.5 mm, broadly obovate, apex obcordate, maculate at the base, claw 0.5–1 mmlong; wing petals 8–10 × 1.5–2.5 mm, oblong, apex obtuse, without callosities, claw 0.8–1 mmlong; keel petals 8–10 × 2.5–3 mm, narrowly obovate, apex subacute or obtuse, with callosities, claw 2.5–3 mmlong; androecium monadelphous, 7–10 mmlong, vexillary stamen partially fused with the other from the base 6–7 mmlong; ovary 3–4 mmlong, glabrous, stipe 2–3 mmlong, glabrous. Loment 1.5–2.2 cmlong, stipe 2–3 mmlong, glabrous, isthmus central, both margins sinuate; articles uniform 3–7, 2–3.5 × 2–3 mm, orbicular, not tortuous, indehiscent, membranaceous or subcoriaceous, veins conspicuous, sparsely puberulous-uncinate becoming glabrous at maturity. Seed 2–2.5 × 1–1.5 mm, oblong, hilum subcentral.   Selected specimens examined:—  BRAZIL. Acre: Rio Branco: s. loc.,  September 1975, fl.,  V .   Patiño82( PAMG).  Bahia: Senhor do Bonfim: Serra da Jacobina, oeste de Estiva, ca. de 12 km N. de Senhor do Bonfim,  1 March 1974, fl., fr.,  R. M .   Harleyet al. 16608( K, RB). Distrito Federal: Brasília: Baciado rio São Bartolomeu,  17 March 1980, fr.,  E. P .   Heringeret al. 3997( K, LISC*).  Espírito Santo: Santa Teresa: Penha,  15 May 1984, fl., fr.,  W .   Boone152( RB).  Goiás: Hidrolândia: Morro Feio, ca. de 5 Km  NdeHidrolândia,  8 April 1988, fl.,  R. R .   Brookset al. 17( E, K).  Mato Grosso do Sul: Rio Verde: s. loc., 1975, fl.,  V .   Patiño116( PAMG).  Minas Gerais: Belo Horizonte: Parque das Mangabeiras,  13 March 1997, fl.,  J .   Damasceno147( BHCB, HUEFS); Lima Duarte: Hotel Serra do Ibitipoca,  23 February 2002, fl.,  F. R .   Salimenaet al. 996( ESA). Pará: Belém: s. loc.,  January 1963, fl., fr.,  J. M .   Pires8112( UB).  Paraná: Tibagi: fazenda Monte Alegre,  15 January 1954,  G .   Hatschbachet al. 3495( PAMG, MBM).  Rio de Janeiro: Teresópolis: s. loc.,  8 April 1959, fl., fr.,  A. P .  Duarte& E. Pereira4709( B, PAMG).  Rio Grande do Sul: Bom Jesus: fazenda do Cilho,  12 February 2007, fr.,  R. B .   Setubalet al. 924( HUEFS).  Santa Catarina: Florianópolis: Vargem do BomJesus- Ingleses,  4 April 1991, fr., D. A .   Machado388( FLOR). São Paulo:  São Paulo: Horto Florestal,  26 March 1952, fl., fr.,  M. A .   Cunhas.n. ( HUEFS169174).  Tocantins: Arrayas: s. loc.,  April 1840, fl.,  G .   Gardner3680( BM).   Distribution and Ecology:—restricted to Brazil, in the Distrito Federal and the States of Acre, Bahia, Espírito Santo, Goiás, Mato Grosso do Sul, Minas Gerais, Pará, Paraná, Rio de Janeiro, Rio Grande do Sul, Santa Catarina, São Paulo, and Tocantins.  D. subsecundumoccurs in gallery and riparian forests, pastures, and grasslands within Cerradoand Rain Forest biomes.  Conservation Assessment:—Least Concern (LC), not endangered ( IUCN 2001).  Phenology:—flowering and fruiting from January to May.   Etymology:—from the Latin sub(= somewhat) + secundus(= secund), in a reference to the subsecund flowers.  Common Name:—amores-do-campo; beiço-de-boi; carrapichinho; carrapicho; marmelada-de-cavalo; marmelada; pega-pega; trevo-dos-pampas.  Taxonomic notes:—the shrub or subshrub habit, auriculate, semi-amplexicaul stipules persistent at the base of the young inflorescences, discolorous leaves, flowers 7–10 mmlong, the long-stipitate ( 1–2 mm), glabrous or puberulentous ovary, and orbicular fruit articles (that are glabrous when mature) constitute a diagnostic suite of characters for  Desmodium subsecundum. The species is morphologically similar to  D. album,  D. leiocarpum, and  D. venosumdue to similar habit and paniculate inflorescences (see notes under each of those species). Most specimens of  D. subsecundumexamined had the first fruit article aborted (see Fig. 29). The specimens cited by Azevedo (1981)as  D. subsecundumwere examined in this study. Because they possess trifoliolate or unifoliolate leaves next to the base of their inflorescences, abaxial surfaces of their leaflets with conspicuous primary, secondary, and tertiary veins, tomentose ovaries with a stipe to 0.5–1 mmlong, and elliptic   DESMODIUM(LEGUMINOSAE, PAPILIONOIDEAE) IN BRAZIL  Phytotaxa169 (1) © 2014 Magnolia Press • 83fruit articles, they better correspond morphologically with  D. venosum, a species recognised as distinct in this treatment. The collection Hatschbach 3495was cited by Azevedo (1981)as a new species,  D. hatschbachii, but this name was never validly published. In our study the specimen is recognised as  D. subsecundum. The difficulties associated with the morphological delimitation and nomenclature of  Desmodium discolor,  D. subsecundum,  D. leiocarpum, and  D. venosumhas existed since Bentham (1859), who cited  D. venosumas a possible variety of  D. leiocarpumbased only on vegetative characters such as leaflet consistency and venation and petiole length. In our study other differences between  D. venosumand  D. leiocarpumbecame evident, such as the indumentum of the ovary and the fruit article length.  Schindler (1928)established  Desmodium subsecundumas a variety of  D. leiocarpumbecause, besides the vegetative differences, these taxa can be distinguished on the basis of flower length, indumentum of the ovary, fruit stipe length, and article shape. The identity of  D. discolorand  D. subsecundumis obscured by our failure to trace the typespecimen of  D. subsecundumThe typeof D. discolorwas analysed based on a photograph at F (negative 2259) of a specimen originally at B. The photograph F2259 bears a label confirming it as  D. discolorVogel, apparently fide Schindler, August 1920. Amorphological comparison was made between  D. subsecundumand  D. discolorbased on the original publication by Vogel (1838). Uncinate-pubescent stems in  D. subsecundum(vs. hirsute-tomentose in  D. discolor), oblong-lanceolate leaves (vs. ovate or ovate-oblong in  D. discolor), flowers organized in large panicles (in both species), puberulous pedicels 0.8 cmlong (vs. 0.6–0.8 cmlong in  D. discolor), flowers not described (vs. flowers ca. 0.6 cmlong in  D. discolor), fruit stipe 2 mmlong, and fruits subsecund with the first article aborted (vs. without indication in  D. discolor) together suggest that the two species are closely related. The two were were differentiated by Bentham (1859)and Hoehne (1921)only on the indumentum density and leaflet shape. In this study we consider the two names to be conspecific,  D. discoloris proposed as a new synonym of  D. subsecundum.  Vogel (1838)also described  Desmodium leiocarpumvar. βfor “ Brasilmeridional, entre Campos e Vitória” without giving it a name. Schindler (1928)considered this taxon to be a synonym of  Meibomia leiocarpa(Spreng.) O. Kuntze. Inthis study it is identified as  D. subsecundumdue to article shape and fruit stipe length.   Meibomia discolorvar. pohliipublished by Schindler (1924)has the same inflorescence, and fruit and leaflet shape as  D. subsecundum, differing only in the petiole length (more than 40 mmlong), a variable vegetative characteristic that was not used to treat these specimens as varieties. [367,520,1269,1291] F, B Brazil Sellow 80 81 FOBN002259 3 lectotype [357,792,1392,1414] 1918-03-01 SP Brazil Horto Oswaldo Cruz Cantareira 80 81 2 Sao Paulo lectotype [349,629,1484,1506] J, B, E, M, K, C, P Brazil Pohl Santa Cruz 80 81 W0027268 3 Goias lectotype [595,1305,836,861] 1975-09 V Brazil Rio Branco 82 83 1 Acre PAMG Brazil Patino 82 83 1 [263,1151,871,896] N de Estiva Brazil Serra da Jacobina Senhor do Bonfim 82 83 1 Bahia 1974-03-01 R, M Brazil Senhor do Bonfim 82 83 1 1980-03-17 K, RB, E, P Distrito Federal & Brasilia & Bacia Brazil rio Sao Bartolomeu Harley 82 83 1 [388,751,943,968] K, LISC Brazil Heringer 82 83 1 [762,1435,943,968] 1984-05-15 W Brazil Penha Santa Teresa 82 83 1 Espirito Santo [151,340,980,1005] RB Brazil Boone 82 83 1 [351,1317,980,1005] 1988-04-08 N, R Km Brazil Morro Feio Hidrolandia 82 83 1 Goias E, K Brazil Brooks 82 83 1 [319,936,1015,1040] V Brazil Rio Verde 82 83 1 Mato Grosso do Sul [948,1192,1015,1040] PAMG Brazil Patino 82 83 1 1997-03-13 J Parque das Mangabeiras Brazil Belo Horizonte 82 83 1 Minas Gerais 2002-02-23 BHCB, HUEFS, F, R Lima Duarte Brazil Hotel Serra do Ibitipoca Damasceno 82 83 1 1963-01 ESA, J, M Para & Belem Brazil Salimena 82 83 1 [208,395,1124,1149] UB Brazil Pires 82 83 1 [405,1067,1124,1149] 1954-01-15 G Monte Alegre Brazil Tibagi 82 83 1 Parana PAMG, MBM Brazil Hatschbach 82 83 1 [244,932,1159,1184] 1959-04-08 A, P Brazil Teresopolis 82 83 1 Rio de Janeiro [943,1081,1160,1184] E Brazil Duarte 82 83 1 2007-02-12 R, B Cilho Brazil Bom Jesus 82 83 1 Rio Grande do Sul [1027,1358,1196,1221] HUEFS Brazil Setubal 82 83 1 1991-04-04 A Ingleses Brazil Vargem do Bom Florianopolis 82 83 1 Santa Catarina [1049,1305,1231,1256] FLOR Brazil Machado 82 83 1 [151,784,1268,1293] 1952-03-26 M, A Brazil Horto Florestal 82 83 1 Sao Paulo [796,1121,1268,1293] HUEFS Brazil Cunha 82 83 HUEFS169174 1 1840-04 G Brazil Arrayas 82 83 1 Tocantins [355,586,1303,1328] BM Brazil Gardner 82 83 1