Meibomia leiocarpa var. subsecunda (Vogel) Schindl., Rep., 2014

Lima, Laura Cristina Pires, Queiroz, Luciano Paganucci De, Tozzi, Ana Maria Goulart De Azevedo & Lewis, Gwilym Peter, 2014, A Taxonomic Revision of Desmodium (Leguminosae, Papilionoideae) in Brazil, Phytotaxa 169 (1), pp. 1-119 : 81-84

27. Desmodium subsecundum Vogel, Linnaea 12: 99. 1838. Meibomia subsecunda (Vogel) Kuntze, Revis. Gen. Pl. 1: 198. 1891. Meibomia leiocarpa var. subsecunda (Vogel) Schindl., Rep. Spec. Nov. Beiheft 49: 370. 1928. Type:— BRAZIL. Brasil meridional, F. Sellow s.n. (holotype B†). Neotype (designated here):— BRAZIL. Paraná: Ponta Grossa, Parque Vila Velha, 17 March 1976, R. Kumrow & W. Anderson 1094 (neotype MBM!). Fig. 29 .

Desmodium discolor Vogel, Linnaea 12: 103. 1838. Meibomia discolor (Vogel) Kuntze, Revis. Gen. Pl. I : 198. 1891. Lectotype (designated here):— BRAZIL. Brasil Meridional. F. Sellow 4955 (holotype B†, lectotype FOBN002259 !).

Desmodium leiocarpum var. 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. villosa Hoehne, Mem. Inst. Butantan 1(1): 23. 1921. Desmodium discolor var. villosum (Hoehne) Malme, Ark. Bot. 18(7): 17. 1922. Lectotype (designated here):— BRAZIL. São Paulo: Cantareira, 01 March 1918, Horto Oswaldo Cruz 1570 (SP!). Remaining syntypes: Brazil. São Paulo: Horto Oswaldo Cruz 2234 (SP!), syn. nov.

Meibomia discolor var. pohlii Schindl., Repert. Spec. Nov. Regni Veg. 20: 151. 1924. Lectotype (designated here):— BRAZIL. Goiás: Santa Cruz de Goiás, J.B.E. Pohl 2603 (lectotype W0027268 !, isolectotypes M!, K!). Remaining syntype: Brazil. Minas Gerais: Lagoa Santa, E. Warming 2970 (C*!, P!), syn. nov.

Erect, branched shrub, 1–3 m tall, 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 mm long. Leaves trifoliolate; petiole 5–36 mm long, cylindrical, canaliculate, densely velutinous or villose or sparsely tomentose and uncinate; rachis 5–16 mm long; stipels 2–6.5 mm long, 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

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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 cm long, densely or sparsely hispid and uncinate, rarely velutinous, 2 flowers per node; primary bract 1.5–2 mm long, 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 mm long, subulate, margin ciliate hispid and puberulous-uncinate on the outer surface, caducous, veins inconspicuous on the outer surface; pedicel 7–8 mm long, densely or sparsely hispid and puberulous-uncinate or densely uncinate. Flowers 7–10 mm long; calyx bilabiate, tube campanulate, 1–1.2 mm long, 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 mm long; lower lip trifid, lateral tooth triangular, 1–1.5 mm long, central tooth triangular, 1.5–2 mm long; corolla lilac, standard 8–8.5 × 5–7.5 mm, broadly obovate, apex obcordate, maculate at the base, claw 0.5–1 mm long; wing petals 8–10 × 1.5–2.5 mm, oblong, apex obtuse, without callosities, claw 0.8–1 mm long; keel petals 8–10 × 2.5–3 mm, narrowly obovate, apex subacute or obtuse, with callosities, claw 2.5–3 mm long; androecium monadelphous, 7–10 mm long, vexillary stamen partially fused with the other from the base 6–7 mm long; ovary 3–4 mm long, glabrous, stipe 2–3 mm long, glabrous. Loment 1.5–2.2 cm long, stipe 2–3 mm long, 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 . Patiño 82 ( 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 . Harley et al. 16608 ( K, RB). Distrito Federal: Brasília: Bacia do rio São Bartolomeu , 17 March 1980, fr., E. P . Heringer et al. 3997 ( K, LISC *) . Espírito Santo: Santa Teresa: Penha , 15 May 1984, fl., fr., W . Boone 152 ( RB) . Goiás: Hidrolândia: Morro Feio , ca. de 5 Km Nde Hidrolândia, 8 April 1988, fl., R. R . Brooks et al. 17 ( E, K) . Mato Grosso do Sul: Rio Verde : s. loc., 1975, fl., V . Patiño 116 ( PAMG) . Minas Gerais: Belo Horizonte : Parque das Mangabeiras, 13 March 1997, fl., J . Damasceno 147 ( BHCB, HUEFS); Lima Duarte: Hotel Serra do Ibitipoca , 23 February 2002, fl., F. R . Salimena et al. 996 ( ESA). Pará: Belém: s. loc., January 1963, fl., fr., J. M . Pires 8112 ( UB) . Paraná: Tibagi : fazenda Monte Alegre, 15 January 1954, G . Hatschbach et al. 3495 ( PAMG, MBM) . Rio de Janeiro: Teresópolis : s. loc., 8 April 1959, fl., fr., A. P . Duarte & E . Pereira 4709 ( B, PAMG). Rio Grande do Sul: Bom Jesus : fazenda do Cilho, 12 February 2007, fr., R. B . Setubal et al. 924 ( HUEFS) . Santa Catarina: Florianópolis: Vargem do Bom Jesus- Ingleses, 4 April 1991, fr., D. A . Machado 388 ( FLOR) . São Paulo: São Paulo: Horto Florestal , 26 March 1952, fl., fr., M. A . Cunha s.n. ( HUEFS169174 ) . Tocantins: Arrayas : s. loc., April 1840, fl., G . Gardner 3680 ( 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. subsecundum occurs in gallery and riparian forests, pastures, and grasslands within Cerrado and 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 mm long, 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. venosum due to similar habit and paniculate inflorescences (see notes under each of those species). Most specimens of D. subsecundum examined had the first fruit article aborted (see Fig. 29 ).

The specimens cited by Azevedo (1981) as D. subsecundum were 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 mm long, and elliptic

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The collection Hatschbach 3495 was 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. venosum has existed since Bentham (1859), who cited D. venosum as a possible variety of D. leiocarpum based only on vegetative characters such as leaflet consistency and venation and petiole length. In our study other differences between D. venosum and D. leiocarpum became evident, such as the indumentum of the ovary and the fruit article length.

Schindler (1928) established Desmodium subsecundum as a variety of D. leiocarpum because, 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. discolor and D. subsecundum is obscured by our failure to trace the type specimen of D. subsecundum The type of D. discolor was 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. discolor Vogel , apparently fide Schindler, August 1920.

Amorphological comparison was made between D. subsecundum and D. discolor based 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 cm long (vs. 0.6–0.8 cm long in D. discolor ), flowers not described (vs. flowers ca. 0.6 cm long in D. discolor ), fruit stipe 2 mm long, 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. discolor is proposed as a new synonym of D. subsecundum .

Vogel (1838) also described Desmodium leiocarpum var. β for “ Brasil meridional, entre Campos e Vitória” without giving it a name. Schindler (1928) considered this taxon to be a synonym of Meibomia leiocarpa (Spreng.) O. Kuntze. In this study it is identified as D. subsecundum due to article shape and fruit stipe length.

Meibomia discolor var. pohlii published by Schindler (1924) has the same inflorescence, and fruit and leaflet shape as D. subsecundum , differing only in the petiole length (more than 40 mm long), a variable vegetative characteristic that was not used to treat these specimens as varieties.

Figures

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).