Leandra melastomoides Raddi (1820: 386)

Reginato, Marcelo, 2016, Taxonomic revision of Leandra sect. Leandra (Melastomataceae, Miconieae), Phytotaxa 262 (1), pp. 448-450 : 448-450

11. Leandra melastomoides Raddi (1820: 386) . Leandra involucrata Raddi (1829: 145) . nom. superfl. Type:— BRAZIL. Rio de Janeiro: “ Boschi di Mandiocca e del Corcovado", G. Raddi s.n. (holotype PI!, isotypes FI!, G, PI!). ( Figs. 52 , 53 , 54 , 55 , 56 , 57 )

Leandra scabra de Candolle (1828: 154) . Lectotype (designated here):— BRAZIL. Minas Gerais: C.F.P von Martius s.n. ( M 173421 !, isolectotype G image!). Additional syntype: BRAZIL: Martii Herbarium Florae 4 (GH!, M!, MO, NY!, P).

Leandra villosa de Candolle (1828: 154) . Type:— BRAZIL. Rio de Janeiro: C. Gaudichaud-Beaupré 734 (holotype G image!, isotypes P (×3)).

Leandra asperifolia Chamisso (1835: 33) . Type:— BRAZIL: F. Sellow s.n. (not located).

Leandra fragilis Cogn. in Martius et al. (1888: 88). Lectotype (designated here):— BRAZIL. Santa Catarina: L.A. von Chamissoo s.n., ( P 117371 image!). Additional syntype: BRAZIL: C. Gaudichaud-Beupré 24 (G image!).

Leandra melastomoides var. major Cogn. in Martius et al. (1888: 607). Type:— BRAZIL. Rio de Janeiro: Serra dos Órgãos, P. Schwacke 4389 (holotype BR image!, isotype RB).

Shrubs or treelets, 0.5–4(5) m tall. Branches, petioles, leaves, inflorescences and bracteoles covered by appressed, unbranched trichomes (0.4–2 mm), these sparser leaf blade surfaces, denser on branches, inflorescences and hypanthia, seldom unbranched glandular trichomes (ca. 1 mm) mixed on the hypanthia. Leaves slightly anisophyllous in each pair (8:10 ratio); petioles 0.2–3 cm long; blades 4–25 × 1.5–8 cm, lanceolate to elliptic, apex acute to slightly acuminate, base obtuse or cuneate, margin crenulate or entire, ciliate (0.7–1.5 mm), chartaceous; 3 acrodromous nerves, plus 2 additional pair of faint veins, basally nerved or plinerved, distant 0–35 mm above the base, nerves slightly conspicuous on adaxial surface and acrodromous nerves prominent, transversal flat on abaxial, reticulation conspicuous or not. Inflorescences terminal, 1–3(5) per node, (1.5) 4–12.5 cm long, 2–4 pairs of opposite paraclades, accessory branches absent or seldom present,> (10)20 flowers per inflorescence, these in glomerules; bracteoles 3–11 × 2–5 mm, ovate, elliptic, oblong or obovate, seldom lanceolate, glabrous, glabrous towards the margins, or covered by unbranched trichomes in the whole surface, involucral, persistent. Flowers (5-)6-merous, sessile. Hypanthium 3.7–5 × 2.4–3 mm,

TAXONOMIC REVISION OF LEANDRA SECT. LEANDRA

Phytotaxa 262 (1) © 2016 Magnolia Press • 61 tubular or slightly campanulate, torus indumentum absent. Calyx tube 0.4–0.5 mm long; inner lobes 1.5–2.7 × 0.6–1.5 mm, triangular; external teeth 2–3 mm long. Petals white, 4–6.5 × 1–1.5 mm, linear or lanceolate, apex acute to acuminate, glabrous or seldom with an apical glandular trichome (ca. 0.5), spreading or reflexed at anthesis. Stamens (10-)12, opposite to the style; filaments geniculation present, the larger 5–7 mm long, the smaller 4–5.5 mm long; anthers pink, the larger 3.4–4.5 mm long, the smaller 2.7–3.5 mm long, linear-subulate, dorsally curved or straight, pore 0.10–0.15 mm wide, connectives dorsally produced below the anthers 0.3–0.8 mm, appendage absent or present, in the larger anthers as a basal-dorsal bifurcation. Ovary (3-)4-celled, 2.5–3.5 × 1.2–2 mm, 30–40 % inferior, apex covered by unbranched trichomes (0.6–1 mm). Style 10–13 mm long, sigmoid, opposite of the stamens, stigma 0.2–0.3 mm diam. Berries dark purple or black, 5–10 × 4–6 mm. Seeds 0.8–1 × 0.4–0.6 mm, long-obpyramidal, hilum covering 9/10 of the seed length, anticlinal cell walls flat.

Notes:— Leandra melastomoides is the most widespread and morphologically variable species recognized in this treatment. Among the species with persistent bracteoles and up to 5 acrodromous nerves may be distinguished from L. amplexicaulis by the non-amplexicaulous leaf base and from L. glazioviana by the smaller flower structures and fruits (see notes under that species). Wurdack (1970) noticed that de Candolle (1828) described L. scabra with doubt regarding whether it would be a potential synonym of L. melastomoides , and concluded that L. scabra should be treated as such. Cogniaux (1888) used the number of petals to distinguish L. fragilis (5-merous) from L. scabra (6-merous). That author also mentioned in his key that L. fragilis has bracts with trichomes restricted to the area along the midvein (i.e., glabrous towards the margin), but he did not mention this character for L. scabra . Leandra fragilis has been recognized in recent local floras ( Wurdack 1962, Camargo & Goldenberg 2007, Baumgratz & Souza 2011), where it was exclusively differentiated from L. melastomoides by the bracts glabrous towards the margin (the number of petals was suggested to be variable). Nonetheless, all authors highlighted the similarities to L. melastomoides , with a more incisive suggestion made by Wurdack (1962), where

62 • Phytotaxa 262 (1) © 2016 Magnolia Press

REGINATO he stated that L. fragilis should be treated as a less pubescent variety of L. scabra (he later suggested L. scabra as synonym of L. melastomoides, Wurdack 1970 ). In summary, if it does appear that the types of L. melastomoides , L. scabra and L. fragilis might belong to distinct taxa, but it also appears that they would be difficult to diagnose, given the continuous variation observed in the complex.

In an attempt to understand the status and possible differences among L. melastomoides , L. scabra and L. fragilis , I further explored the morphological variability of leaves, bracts and inflorescences in the complex. The specimens were a priori divided in four morphotypes, defined as follows: (1) "melastomoides "—specimens with bract pubescence on the whole surface, mainly from Rio de Janeiro and Espírito Santo; (2) "minifolia"—specimens with small leaves, with a very faint external most nerve, mainly from Santa Maria Madalena region (Rio de Janeiro); (3) "seminervia"—specimens with basally nerved or slightly plinerved leaves, from southern Bahia; (4) "scabra "—specimens with bracts glabrous towards the margin (or totally glabrous), from several regions. The last morphotype includes both L. scabra and L. fragilis types, since in the specimens annotated by A. Cogniaux as L. scabra , the bracts are glabrous towards the margin. In general, it is possible to code the bracts regarding the pubescence, but in several instances this task becomes highly arbitrary, and for these cases geographical distribution was taken into account.

The measurements taken for the morphotypes are depicted as box plots in Fig. 55 and were summarized in the PCA ( Fig. 54-B ). The geographical distribution of the specimens measured is presented in Fig. 54-A , color coded by morphotype (same coding in the morphospace). Overall, the "scabra " morphotype present larger leaves, higher leaf length to width ratio (more lanceolate), acrodromous nerves diverging farther up the leaf, and larger inflorescences. The "melastomoides " morphotype is mainly characterized by larger leaf width, smaller leaf ratio (more elliptic), and longer bracts. The "seminervia" morphotype is distinguished by the basally nerved to slightly plinerved leaves, especially when the plinervy ratio is taken into account (i.e. plinervy / leaf length), while the "minifolia" presents small leaves. The first two principal components explained 60 % of the variation. The morphospace present some structure regarding the a priori groups, although overlapping, with the "minifolia" and "seminervia" morphotypes

TAXONOMIC REVISION OF LEANDRA SECT. LEANDRA

Phytotaxa 262 (1) © 2016 Magnolia Press • 63 64 • Phytotaxa 262 (1) © 2016 Magnolia Press

REGINATO TAXONOMIC REVISION OF LEANDRA SECT. LEANDRA

Phytotaxa 262 (1) © 2016 Magnolia Press • 65 66 • Phytotaxa 262 (1) © 2016 Magnolia Press

REGINATO more isolated from the others. Nonetheless, under a taxonomic perspective, any of the morphological characters measured here can undoubtedly identify the recognized morphotypes. For instance, the "scabra " morphotype includes the leaf size range of "minifolia", and there are several "scabra " morphotypes in southern Bahia sympatric to "seminervia" (not included in the analysis). It is not clear whether those sympatric specimens would be closely related, which would increase the plasticity of plinervy in "seminervia", making it less diagnosable.

Although, only L. melastomoides is recognized as a species here, it seems that there is some structured diversity inside this taxon, even with potential additional morphotypes not included in the analysis due to very small sampling (i.e., a glabrous form from one locality in Minas Gerais). A populational genetic study is very desirable to test whether or not L. melastomoides and/or these morphotypes constitute natural groups. Such a study would also help to better establish its circumscription in relation to the two close relatives L. amplexicaulis and L. glazioviana . It is noteworthy that some specimens treated here as L. melastomoides look more similar to these two species than to some conspecifics of a different morphotype.

Selected specimens examined:— BRAZIL. Bahia: Abaíra, Harley 50863 ( CEPEC); Almadina, Borges 429 ( UPCB); Amargosa, Cardoso 1502 ( CEPEC); Arataca, Thomas 14553 ( CEPEC); Barra do Choça, Mori 9408 ( CEPEC); Barro Preto, Thomas 14317 ( NY); Camacan, Thomas 13767 ( NY); Eunápolis, Almeida 75 ( RB); Ilhéus, Silva 1554 ( RB); Itamaraju, Borges 791 ( CEPEC); Porto Seguro, Santos 869 ( CEPEC); Rio de Contas, Harley 24529 ( SPF); Una, Silva 90 ( CEPEC); Uruçuca, Mori 11769 ( NY); Wenceslau Guimarães, Thomas 9355 ( CEPEC). Distrito Federal: Brasília, Sevilha 1750 ( SP). Espírito Santo: Alfredo Chaves, Hatschbach 61421 ( CEPEC); Cariacica, Kollmann 10659 ( RB); Castelo, Amorim 7829 ( RB); Domingos Martins, Peixoto 431 ( RB); Ibitirama, Colletta 410 ( RB); Santa Leopoldina, Fontana 3026 ( RB); Santa Maria de Jetibá, Goldenberg 1025 ( NY); Santa Teresa, Reginato 1205 ( NY); São Roque do Canaã, Reginato 1188 ( UPCB); Vargem Alta, Sakuragui 873 ( UPCB); Viana, Goldenberg 1232 ( NY); Vitória, Santos 974 ( CEPEC). Goiás: Corumbá de Goiás, Irwin 34289 ( NY). Minas Gerais: Alpinópolis, Goldenberg 462 ( MBM); Alto Caparaó, Souza 111 ( RB); Antônio Carlos, Krieger 1272 ( RB); Araponga, Silva 2269 ( UPCB); Barão de Cocais, Kollmann 6025 ( UPCB); Barroso, Assis 13 ( MBM); Belo Horizonte, Barreto 6722 ( F, NY); Brumadinho, Martens 530 ( NY); Caldas, Regnell 28 ( M); Carandaí, Duarte 628 ( RB); Carangola, Mexia 4267 ( F, NY); Catas Altas, Mello-Silva 2541 ( NY); Conceição, Barreto 10807 ( UPCB); Congonhas do Norte, Riina 1327 ( NY); Cruzília, Andrade 10732 ( MBM); Delfinópolis, Pacheco 565 ( MBM); Diamantina, Romariz 103 ( RB); Felício dos Santos, Viana 3688 ( UPCB); Fervedouro, Leoni 5025 ( NY); Gouveia, Mello-Silva 2446 ( SPF); Jaboticatubas, Semir 4353 ( NY); Lima Duarte, Valente 70 ( UPCB); Monte Belo, Vieira 133 ( RB); Nova Lima, Williams 6184 ( NY); Ouro Branco, Schembri 12907 ( MBM); Ouro Preto, Reginato 1164 ( NY); Patrocínio, Pereira Neto 219 ( SP); Perdizes, Mendes 814 ( MBM); Poços de Caldas, Santos 5903 ( R); Rio Preto, Matozinhos 303 ( UPCB); Santa Bárbara, Pirani 720 ( SPF); Santa Luzia, Barreto 8958 ( UPCB); Santa Maria do Salto, Lombardi 5871 ( UPCB); Santana do Riacho, Muniz 7873 ( SP); Santo Antônio do Itambé, Anderson 35719 ( NY); São Gonçalo do Rio Preto, Foresto 264 ( SPF); São Roque de Minas, Romero 2305 ( NY); Tiradentes, Rutter 144 ( R). Paraná: Adrianópolis, Ribas 3037 ( MBM); Antonina, Reginato 684 ( MBM); Campo Largo, Hatschbach 3216 ( MBM); Guaraqueçaba, Ziller 118 ( MBM); Guaratuba, Reginato 1239 ( NY); Jaguariaíva, Uhlmann 116 ( MBM); Londrina, Silva 205 ( MBM); Matinhos, Straube 38 ( MBM); Morretes, Hatschbach 766 ( RB); Paranaguá, Ziller 629 ( MBM); São José dos Pinhais, Motta 270 ( MBM); Sengés, Souza 26 ( MBM); Tibagi, Silva 1844 ( RB); Tunas do Paraná, Camargo 102 ( UPCB); Ventania, Estevan 198 ( SPF). Rio de Janeiro: Itatiaia, Lanstyak 175 ( NY); Macaé, Farney 250 ( RB); Magé, Vidal 5445 ( R); Nova Friburgo, Ule 4559 ( R); Nova Iguaçu, Fromm 1277 ( R); Parati, Ribeiro 1 ( CEPEC); Petrópolis, Wawra 424 ( NY); Porciúncula, Ribeiro 65 ( CAS); Resende, Martinelli 10775 ( RB); Rio de Janeiro, Sucre 1718 ( NY); Santa Maria Madalena, Reginato 1210 ( UPCB); Silva Jardim, Calvente 126 ( NY); Teresópolis, Vidal 32 ( R). Santa Catarina: Águas Mornas, Krapovickas 44750 ( MBM); Blumenau, Verdi 5103 ( MBM); Brusque, Klein 36 ( RB); Florianópolis, Klein 7125 ( MBM); Garuva, Cervi 8804 ( MBM); Ibirama, Gevieski 24 ( M); Ilhota, Reginato 1454 ( UPCB); Itajaí, Landrum 2483 ( MBM); Jaraguá do Sul, Melo Jr. 689 ( MBM); Palhoça, Reitz 2462 ( MBM); Rio do Campo, Sobral 8386 ( MBM); São Bento do Sul, Meyer 571 ( UPCB); São Francisco do Sul, Vieira 663 ( UPCB); São Pedro de Alcântara, Falkenberg 6071 ( MBM). São Paulo: Apiaí, França 2548 ( RB); Campos do Jordão, Porto 3172 ( NY); Cananéia, Toledo 47 ( NY); Caraguatatuba, Cordeiro 2376 ( NY); Cubatão, Toledo 432 ( F); Cunha, Baitello 493 ( SPF); Guarulhos, Arzolla 1142 ( MBM); Ibiúna, Toledo 351 ( NY); Iguape, Brade 8175 ( GH); Iporanga, Souza 5968 ( SPF); Itaberá, Chiea 679 ( NY); Itararé, Dusén 9641 ( GH); Jundiaí, Goldenberg 767 ( MBM); Miracatu, Pirani 3087 ( SPF); Paranapiacaba, Pereira 5931 ( RB); Parelheiros, Godoy 364 ( RB); Pariquera-Açu, Bernacci 982 ( NY);

TAXONOMIC REVISION OF LEANDRA SECT. LEANDRA

Phytotaxa 262 (1) © 2016 Magnolia Press • 67 Peruíbe, Sobral 6626 ( MBM); Registro, Eiten 6064 ( NY); Santo André, Smith 1915 ( F, NY); São Bernardo do Campo, Kirizawa 173 ( SP); São José dos Campos, Tamashiro 911 ( SP); São Miguel Arcanjo, Dias 512 ( MBM); São Paulo, Smith 1809 ( F, NY); Serra Negra, Chiea 712 ( NY); Sete Barras, Benson 10885 ( SP); Ubatuba, Kirizawa 2446 ( CEPEC).

Figures

FIGURE 52. Distribution of Leandra melastomoides.

FIGURE 53. Leandra melastomoides. A. Habit. B–C. Branch with inflorescence. C. Leaf on adaxial view. E–F. Detail of the inflorescences. G. Flower. H. Branch with young infrustecences.

FIGURE 54. Distribution of the specimens measured (A) and PCA of these measurements (B). of Leandra melastomoides. In B first component in the y axis and second component in the x axis. The morphotypes are colorcoded following the legend, and representative specimens are labeled in both graphs.

FIGURE 55. Boxplot of leaf and inflorescence measurements of Leandra melastomoides morphotypes. B = larger leaf; S = smaller leaf.

FIGURE 56. Leandra melastomoides. A–E. Branch with inflorescences (at the same scale). F–K. Leaf on abaxial view (at the same scale). L–N. Detail of the glomerules showing the bracts and bracteoles. O–S. Detail of the leaf on adaxial view (at the same scale). Vouchers:Goldenberg 1232 (A, H, R), Gardner 382 (B, J), Caddah 867 (C, G, Q), Reginato 1239 (D, S), Reginato 1205 (E), Reginato 1092 (F), Reginato 1145 (I, M, P), Fiaschi 1898 (K, L, O), Reginato 1237 (N).

FIGURE 57. Leandra melastomoides. A–B. Flower. C. Bracteole. D. Petal. E. Stamens. F. Flower bud. G. Hypanthium and style. H. Detail of the calyx. I. Anthers. J. Detail of anther apex. K. Hypanthium and calyx longitudinal section. Scales: A–G, I, K = 1 mm; H, J = 0.25 mm. Voucher: Reginato 1318.