Desmodium wydlerianum Urb., Symb. Antill.

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 : 99-105

34. Desmodium wydlerianum Urb., Symb. Antill. 2(2): 302. 1900. Meibomia wydleriana (Urb.) Britton, Sci. Surv. Porto Rico e Virgin Islands. 5: 403. 1924. Lectotype (designated by Schubert 1980):— PORTO RICO. Schwaneeke, June 1827, H. Wydler 140 (lectotype G!, isolectotype FI-W!). Remaining syntypes: Nicaragua. Granada in sylvis interioribus montis Felix 400m. alt. H.H.F.A.B. von Eggers 6009 (B†, not seen); Tobago in convalli fluminis superioris Great Dog River, H.H.F.A.B. von Eggers 5794 (B†, not seen). Fig. 36 .

Desmodium lunatum Huber, Bol. Mus. Goeldi Hist. Nat. Ethnogr. 4: 568. 1906. Meibomia lunata (Huber) Hoehne, Anexos Mem. Inst. Butantan, Secc. Bot. , 1(1): 39. 1921. Type:— PERU. Chinganilla, Pampa del Sacramento, 24 November 1898, J.E. Huber 1504 (holotype MG!).

Prostrate, branched shrub, lacking stolons rooting at the nodes, without a xylopodium; stems virgate, slender, cylindrical, striate, glabrescent, not glaucous; internodes 5.8–6.2 cmlong. Stipules 3–4 × 1–1.5 mm, ovate, not auriculate, semi-amplexicaul, inserted perpendicularly at the base of leaf petiole, free from each other, apex caudate, margin straight, sparsely uncinate, striate on the outer surface, veins conspicuous, caducous. Leaves trifoliolate; petiole (29–) 41–52 mm long, cylindrical, sulcate, sparsely puberulous-uncinate; rachis 9–14 mm long; stipels 2–4 mm long, subulate, margin straight, glabrous on the outer surface, persistent; leaflets concolorous, membranaceous, venation craspedodromous, primary and secondary veins prominent, other veins inconspicuous on the abaxial surface, indumentum sparsely puberulous-uncinate or glabrous on the primary veins on the abaxial

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surface, adaxial surface puberulous or glabrous, uncinate hairs covering both surfaces, terminal leaflet (5.5–)6.5–8.6 × 3.3–5.6 cm, lanceolate or trullate, base truncate, obtuse or attenuate, apex acuminate, subacute, obtuse or slightly retuse, lateral leaflets 5.4–6 × 2.4–3.2 cm, the same shape as the terminal leaflet. Inflorescence a terminal or axillary pseudoraceme; the main axis longer than the adjacent leaves, 15–29.5 cm long, sparsely or densely hirsutous–uncinate, 2 flowers per node; primary bract ca. 2 mm long, lanceolate, margin straight, glabrescent on the outer surface, caducous, veins inconspicuous on the outer surface; secondary bract 1–1.5 mm long, linear, margin straight, glabrescent on the outer surface, caducous, veins inconspicuous on the outer surface; pedicel 11–21 mm long, densely hirsutous–uncinate. Flower 3–3,5 mm long; calyx 5–laciniate, tube cylindrical, 1–1.5 mm long, tomentose and uncinate on the outer surface; teeth subequal, triangular, upper lip ca. 1 mm compr; lower lip 1–1.5 mm long; corolla lilac, standard 2.5–3.5 × 2–2.5 mm, obovate, apex obtuse, maculate at the base, claw ca. 0.5 mm long; wing petals ca. 2 × 0.5 mm, narrowly obovate, apex obtuse, without callosities, claw ca. 0.5 mm long; keel petals 2.5–3 × 1–1,5 mm, oblong, apex subacute, without callosities, claw ca. 1 mm long; androecium diadelphous, 3–3.5 mm long, vexillary stamen free; ovary 2–2,5 mm long, sparsely tomentose, stipe ca. 0,5 mm long, glabrous. Loment 0.8–1.8 cm long, stipe ca. 1 mm long, uncinate, isthmus marginal, upper suture slightly invaginated, lower suture crenate; articles uniform 1–3, 8–9 × 3–4 mm, obovate-depressed, slightly tortuous, indehiscent, membranaceous, veins insconspicuous, densely uncinate. Seed ca. 2 × 1 mm, ovate, hilum subcentral.

Specimens examined:— BRAZIL. Acre: Nova Olinda: de São Luiz ao Acre, 26 October 1923, fl., fr., J. G . Kuhlmann 730 ( RB, U, US) . Amazonas: Presidente Figueiredo: Rebio Uatumã , s.d., fl., fr., J. G . de Carvalho- Sobrinho et al. 1431 ( INPA). Mato Grosso: Chapada dos Guimarães: Agropecuária do Cachimbo , 23 August 1977, fr., L. A . Moretti 31 ( RB). Pará: Óbidos: Rio Branco de Óbidos , 14 September 1927, fl., A . Ducke s.n. ( RB20376 ). Rondônia: Alta Floresta do Oeste : Serra dos Parecis , ca. 27 Km de Alta Floresta doeste, 29 November 1982, fl., fr., P . Lisboa et al. 2497 ( MG). Roraima: Alto Alegre : s. loc., 9 June 1986, fl., fr., M. J. G . Hopkins et al. 627 ( K) .

Additional specimens examined:— PERU. Arequipa: s. mun.: Entre Sarayaçu e Catalina , 24 November 1898, fr., J. E . Huber s.n. ( RB10626 ) .

Distribution and Ecology: — West Indies, Central America, and South America in Brazil, Colombia, Guyana, French Guiana, Peru, and Venezuela ( Schubert 1980, Capdevila 1994, Aymard et al. 2007, Lima et al. 2010). In Brazil D. wydlerianum occurs in the Amazon Forest in the States of Acre, Amazonas, Mato Grosso, Pará, Rondônia, and Roraima.

Conservation Assessment:— Least Concern (LC), not endangered ( IUCN 2001).

Phenology:— flowering and fruiting from June to November.

Etymology:— the specific epithet honours the naturalist Heinrich Wydler, who travelled in the West Indies and collected the type material of this species.

Common Name:— unknown.

Taxonomic notes:— membranaceous leaflets with craspedodromous venation, loments with 2–3 articles, and an invaginate upper suture help in the identification of D. wydlerianum . The prostrate habit and flowers 3–3.5 mm long are characters that are also found in D. axillare , but the two species otherwise differ morphologically (see the notes under D. axillare ).

Conflicting information exists about the country where the type material of Desmodium lunatum was collected: “Chinganilla, Pampa del Sacramento, 24 XI 1898.” Schubert (1980) referred to this locality as being in Brazil, whereas Capdevila (1994) referred to it as in Peru. In concordance with Huber (1906), Peru is correct country of origin of the type material.

Schindler (1928) and Schubert (1963) cited D. spirale var. stoloniferum (Rich ex Poir.) DC. in the synonymy of D. wydlerianum . As the name published by De Candolle was established as part of the publication by Poiret (1805), it is treated as a synonym of Hedysarum stoloniferum (see article 33 of the ICBN). However, the circumscription of De Candolle for D. spirale var. stoloniferum corresponds to the morphology of D. wydlerianum .

EXCLUDED NAME

Desmodium bracteatum Micheli, Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn. 73. 1875. Meibomia bracteata (Micheli) Hoehne, Anex. Mem. Inst. Butantan I , 1: 30.1921. Lectotype (designated

DESMODIUM (LEGUMINOSAE, PAPILIONOIDEAE ) IN BRAZIL

Phytotaxa 169 (1) © 2014 Magnolia Press • 101 here):— BRAZIL. ad Quinta prope Rio de Janeiro, Jun , A. F. M. Glaziou 4784 (lectotype C *!, isolectotype P![2 sheets]) .

Azevedo (1981) mentioned the specimen Glaziou 4787 deposited in the Rherbarium as an isotype; moreover the

protologue of Desmodium bracteatum cited Glaziou 4784. The specimen deposited in the Cherbarium is designated

here as the lectotype of this taxon. The correct placement of this species is in genus Codariocalyx Hassk. , as a

synonym of Codariocalyx gyroides (Roxb.) X.Y. Zh.

NAMES NOT CONFIRMED FOR BRAZIL

Here are included names cited for Brazil, or synonyms linked to these names, that have not been confirmed to occur in Brazil.

Desmodium aparines (Link) DC., Prodr. 2: 330. 1825. Hedysarum aparines Link, Enum. Pl. 2: 247.1822. Meibomia aparines (Link) Schindl., Repert. Spec. Nov. Regni Veg. 23: 274.1926. Type:— MEXICO. C. Alaman s.n. (holotype G-DC!).

Reported by Bentham (1859) and Oliveira (1983) as a synonym of Desmodium uncinatum .

Desmodium bridgesii (Schindl.) Burkart, Darwiniana 3(2): 194. 1939. Meibomia bridgesii Schindl., Repert. Spec.

Nov. Regni Veg. 23: 280. 1926. Lectotype (designated here):— BOLIVIA. ohne Ort , Bridges s.n. (lectotype K!,

isolectotypes BM!, US *!).

Reported for Brazil by Azevedo (1981) and Lima et al. (2010). In the protologue of Meibomia bridgesii, Schindler (1926) cited only one collection by Bridges in Bolivia as definitely belonging to his new taxon: " Bolivia, ohne Ort". He cited three other incomplete collections as: " Meibomi a minisincompleta forsan M. bridgesii Schindl. vel affinis". Two of these were examined by us: Pohl 1689, from Brazil (W!), is recognised here as D. glabrescens , and Ule 6539, from Peru (HBG!) as D. cajanifolium .

Desmodium hirsutum M. Martens & Galeotti, Bul l. Acad. Roy. Sci. Bruxelles 10(2): 186. 1843. Meibomia

hirsuta ( M. Martens & Galeotti) Kuntze, Revis. Gen. Pl. 1: 198. 1891. Lectotype (designated

here):— MEXICO. dans les savanes et bois de Zacuapan et Mirador, 2500-3500 pieds, October, H.G. Galeotti

3275 ( lectotype BR 989407 !, isolectotypes BR 989411 !, K!, P!) .

This species was cited by McVaugh (1987) as a synonym of Desmodium distortum , but this is not accepted in our study, based on material seen from Central America and Mexico including types and species protologues. These species shared characters as the uni or trifliolate leaves, paniculate inflorescence and tortuous articles, but there are morphological diferences that helps to recognize them as a distinct taxa. Desmodium distortum is diferenciate of D. hirsutum by tomentose, uncinate and puberulous or puberulous-uncinate, hispid or glabrescent not ferrugineous stems (vs. hirsute ferrugineous in D. hirsutum ), leaflet elliptic, lanceolate, broadly ovate, ovate, ovate-rhombic or rhombic margin straight (vs. elliptic–rhombic or ovate-oblong, margin ciliate), pedicel 2–5 mm long, hispid and puberulous-uncinate or glabrescent (vs. hirsute–ferrugineous 4–6 mm long), flowers 3–5 mm long (vs. 6–8 mm long) and articles ovate-elliptic or orbicular (vs. rhombic).

Among the syntypes of Desmodium hirsutum analysed in the BR herbarium, the material BR 989407 is designated here as lectotype .

Desmodium incanum var. angustifolium (Kunth) Griseb., Fl. Brit. W. I. 187. 1859. Hedysarum angustifolium Kunth , Nov. Gen. Sp. (quarto ed.) 6: 517. 1823. Desmodium angustifolium (Kunth) DC., Prodr. 2: 328. 1825. Meibomia angustifolia (Kunth) Kuntze, Revis. Gen. Pl. 1: 197. 1891. Meibomia incana var. angustifolia (Kunth) Cook & Collins, Contr. U.S. Natl. Herb. 8: 189. 1903. Desmodium supinum var. angustifolium (Kunth) Urb. Symb. Antill. 4: 491. 1905. Desmodiumfrutescens var. angustifolium (Kunth) Schindl., Repert. Spec. Nov. Regni Veg. 21: 10. 1925. Desmodium canum var. angustifolium (Kunth) León & Alain, Contr. Ocas. Mus. Hist. Nat. Colegio "De La Salle" 9: 18. 1950. Type:— MEXICO. "Cresciti in regno Mexicano" (holotype P- HUMB!).

Reportedby Schindler (1925) as avarietyof Desmodiumfrutescens (= D. incanum ).

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Desmodium intermedium Burkart, Darwiniana 3(2): 208. 1939, nom. illeg, non Bom ex Baill. 1884. Type:— ARGENTINA. Formosa: Guayulec , February 1919, P. Joergensen 2721 (holotype SI!) .

Reported by Vanni (2001) as occurring in Brazil.

Desmodium lupulinum Schltdl., Linnaea 12: 317. 1838. Meibomia lupulina (Schltdl.) Kuntze, Revis. Gen. Pl. 1: 198. 1891. Type :— MEXICO. Jalapam, 29 May 1829, C.J.W. Schiede s.n. (holotype HAL 51037!) .

Reported by Bentham (1859) and Oliveira (1983) as a synonym of Desmodium uncinatum .

Desmodium procumbens f. quinquefolium Standl. & L. O. Wilians , Ceiba 3(3): 224. 1953. Type:— HONDURAS. Morazán, campo de la Escuela Agrícola Panamericana um la orilla de un dormitorio, alt. 800 m, 24 October 1949, P. C. Standley 24252 (holotype US!) .

Reportedby Standey & Williams (1953) as a form of Desmodium procumbens .

Desmodium sonorae A. Gray Smithsonian Contr. Knowl. 5(6): 47. 1853. Meibomia sonorae (A. Gray) Kuntze, Revis. Gen. Pl. 1: 198. 1891. Type:— MEXICO. Sonora: Low Valley , on the Sonoite, near deserted Rancho, September, C. Wright 1014 (holotype GH!, isotype K!, PH *!, US!) .

Reported by Bentham (1859) and Oliveira (1983) as a synonym of Desmodium uncinatum .

Desmodium supinum var. amblyophyllum Urb., Symb. Antill. 7: 229. 1911. Desmodium frutescens var. amblyophyllum (Urb.) Schindl., Repert. Spec. Nov. Regni Veg. 21: 10. 1925. Lectotype (designated by Schubert 1980):— DOMINICAN REPUBLIC. Santo Domingo: prope Constanza , H. von Türckheim 3183 (lectotype NY!, isolectotype BR!).

Reported by Schubert (1980) as a synonym of Desmodium incanum .

Desmodium tenellum (Kunth) DC., Prodr. 2: 333. 1825. Hedysarum tenellum Kunth , Nov. Gen. Sp. Pl. 6: 522 (4 ed.). 1824. Type:— VENEZUELA. "Caracas, 440 hex., crescit in umbrosis humidis" (holotype P-HUMB!, isotype B-WILLD!).

Reported by Schubert (1980) as a synonym of Desmodium procumbens var. longipes .

Desmodium trigonum ( Sw.) DC., Prodr. 2: 332. 1825. Hedysarum trigonum Sw., Prodr. 107. 1788. Lectotype (designated here):— JAMAICA. "Jamaica Hispaniola" (lectotype S!, isolectotype LE!).

Schindler (1928) proposed this name as a synonym of Desmodium intortum , a placement confirmed by analysis of

type material. Nevertheless , Desmodium intortum is recorded for Brazil .

Meibomia distorta var. michelii Schindl., Repert. Spec. Nov. Regni Veg. 23: 282. 1926. Lectotype (designated here):— GUATEMALA. Santa Rosa: Cerro Redondo, E.T. Heyde & E. Lux 6106 (M!). Remaining syntypes: Guatemala. Chimaltenango: Volcan del Fuejo, E.G. Seler 2421 (B†), Santa Rosa: Cerro Redondo , E.T. Heyde & E. Lux 6107 (B†)

From analysis of the specimen Heyde & Lux 6106 (M!), cited in the protologue of Meibomia distorta var. michelii ; as well as other materials analysed in the G herbarium, which were determined by Schindler as Desmodium asperum var. michelii , we conclude that these specimens equate to the circumscription of Desmodium hirsutum (see notes under that species), a taxon not recorded from Brazil.

The name Desmodium asperum var. michelii Schindl. was cited only in a checklist of Plantae Selerianae by Loesener (1923), without description, diagnosis, or reference to a protologue, and is thus an invalid name in accordance with the ICBN ( McNeill et al. 2006).

DOUBTFUL NAMES

Cyclomorium caracasanum Walp., Repert 2: 890. 1843. Type:— VENEZUELA. Caracas , J.W.K. Moritz 25 (B†). Referred to by Azevedo (1981) as a synonym of Desmodium procumbens , although from the diagnosis it could be

either D. procumbens or D. tortuosum .

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Desmodium ancistrocarpum (Ledeb.) DC., Prodr. 2: 331. 1825. Hedysarum ancistrocarpum Ledeb., Index horti Dorpat Suppl. 1823. Type:— BRAZIL. Brasilia. C.F. von Ledebour s.n. (not seen)

Schindler (1928) cited Desmodium ancistrocarpum as a synonym of D. frutescens , a name related to D. incanum .

Nicolson (1978) treated D. ancistrocarpum as a doubtful name. The type material was not seen and the diagnosis of

this species is incomplete. We agree with Nicolson (1978) and recocognise the taxon as doubtful.

Desmodium bullamense G. Don, Gen. hist. 2: 294. 1832. Type:—NEW GUINEA. Serra of Leone , Costa of Bullam (not seen).

Reported by Schindler (1928), Schubert (1980), and Azevedo (1981) as a synonym of Desmodium triflorum . There

is morphological character overlap between D. bullamense , D. triflorum and D. heterophyllum .

Desmodium chamissonis Vogel, Linnaea 10: 588. 1836. Meibomia chamissonis (Vogel) Kuntze, Revis. Gen. Pl. 1: 198. 1891. Type:—Locality unknown (not seen).

Schindler (1928) and Schubert (1940) reconised Desmodium chamissonis as a synonym of D. procumbens . The

type collection has not been seen and the protologue does not include enough information to accept this synonymy.

Desmodium physocarpos Vogel, Linnaea 12: 104. 1838. Meibomia physocarpa (Vogel) Kuntze, Revis. Gen. Pl. 1: 198. 1891. Meibomia physocarpa (Vogel) Hoehne, Anexos Mem. Inst. Butantan, Secc. Bot. 1(1): 44. 1921. Type:— BRAZIL. Brasil meridional, F. Sellow s.n. (B†).

Bentham (1859) reported Desmodium physocarpos as being morphologically similar to D. tortuosum . Schindler (1928) considered D. physocarpos , as a synonym of Meibomia distorta (= Desmodium distortum ). As the type collection was not found and the diagnosis of D. physocarpos includes morphological characters that overlap between D. distortum and D. tortuosum , we chose to treat D. physocarpos as a doubtful name.

Desmodium portoricense (Spreng.) G. Don, Gen. Hist. 2: 294. 1832. Hedysarum portoricense Spreng., Syst. Veg. 3: 314. 1826. Type:— PORTO RICO. C.G. Bertero s.n. (not seen).

Reported by Schindler (1928) as a synonym of Desmodium incanum . Comparing the diagnosis of Hedysarum portoricense "stipulis subulatis abbreviatis, panicula terminali ebracteata, lomenti articulis reticulatis pubescentibus" with the circumscription of D. incanum , there exists overlap in the characters of these two taxa but as the type collection of Hedysarum portoricense was not found, this name is considered as doubtful.

Desmodium sclerophyllum var. tortuosa Pilg., Bot. Jahrb. Syst. 30: 161. 1901. Type:— BRAZIL. Mato Grosso, R. K .F. Pilger 615 ( B †) .

Malme (1931) referred to Desmodium sclerophyllum var. tortuosa as a doubtful name. Based on Pilgerʼs (1901)

original diagnosis this variety could be considered a synonym of either Desmodium sclerophyllum or D. distortum .

Desmodium wade (Vand.) DC., Prodr. 2: 331. 1825. Hedysarum wade Vand. Fasciculus plantarum: 17.1771.

Meibomia wade (Vand.) Kuntze, Revis. Gen. Pl. : 198. 1891. Type:— BRAZIL. Para (not seen).

Reported by Dietrich (1847), Bentham (1859), Hoehne (1921), and Azevedo (1981) as a doubtful species. Schindler (1928) did not recognise Hedysarum wade under the genera Desmodium , Nephromeria , Nicolsonia or Meibomia . According to Stafleu & Cowan (1985), the type materials of Vandelli are deposited in BM, Pand LISU herbaria; two of those herbaria were consulted in this study, but the type collection of Hedysarum wade was not encountered.

Comparing the diagnosis of Hedysarum wade "leguminibus membranaceis, scabris, trarticulatis" with the morphology of Desmodium species recognised in this study, strongly suggests that this species is not a Desmodium , but more probably belongs to the genus Cranocarpus Benth.

Hedysarum erectum Vell., Fl. Flumin. : 318. 1829. Type:—Tabula 149.

Reported by Bentham (1859) and Azevedo (1981) as a synonym of Desmodium leiocarpum , but plate 149 and the diagnosis published by Vellozo (1829, 1831) were analysed by us, and we find that the inflorescence position, and the shape and venation of leaflets do not agree with the circumscription of D. leiocarpum as recognised by us here.

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Uraria lagocephala (Link) DC., Prodr. 2: 324. 1825. Hedysarum lagocephalum Link., Enum. Pl. 2: 248.1822. Type:— BRAZIL. (B†).

Hedysarum lagocephalum was reported by Schindler (1928), Bentham (1859) and Azevedo (1981) as a synonym of Desmodium barbatum . Part of the diagnosis of Hedysarum lagocephalum : "estipulis lanceolatis, panicula terminali contracta, legumen triarticulatum pilosum" includes morphological characteristics seen in Desmodium barbatum and D. juruenense . As a type collection of Hedysarum lagocephalum was not found, and the diagnosis is superficial, this name is considered doubtful.

Figures

FIGURE 36. Desmodium wydlerianum. A. Habit. B. Stipules inner surface. C. adaxial surface of leaflet. D. Calyx inner surface (opened). E. Androecium diadelphous (opened). F. Gynoecium. G. Fruit. H. Seed. (A: Moretti 31; B, E–F, H: Ducke s.n. RB20376; C, G: Huber s.n. RB10626).