Lavoisiera glandulifera Naudin (1844: 149)

Martins, Angela B. & Almeda, Frank, 2017, A Monograph of the Brazilian endemic genus Lavoisiera (Melastomataceae: Microlicieae), Phytotaxa 315 (1), pp. 448-450 : 448-450

17. Lavoisiera glandulifera Naudin (1844: 149) . Lavoisiera glandulifera var. genuina Cogniaux (1883: 149) . Type:— BRAZIL. Minas Gerais: “In montibus vulgo Serro Frio ( Diamantina ) frequens praesertim in locis humidioribus et sabulosis,” A.-C. Vauthier 17 (lectotype, designated by Martin and Cremers (2007): P-00481693!; isolectotypes: F!, G-DC!).

= Lavoisiera glandulifera var. brevifolia Cogniaux in Martius (1883: 149). syn. nov. Type:— BRAZIL. Minas Gerais: “ad Caxoeira do Campo,” P. Claussen 24 or 191 (holotype: G!; isotype: BR-518569, n.v., online image!, BR-522741, n.v., online image, W!). The labels on the holotype and isotype at W give the collector’s number as both “24” and “191”. One of the isotypes at BR has only “24” on the label and the other one at BR has only “191” on the label. We interpret both of these numbers as one collection and not syntypes as might be surmised from the protologue.

= Lavoisiera glandulifera var. longifolia Cogniaux in Martius (1883: 149). syn. nov. Type:— BRAZIL. Minas Gerais: “loco haud indicato,” H. L. Sello s.n. (holotype: B, destroyed).

Erect, much-branched shrubs 0.4–2 m tall. Branches and branchelets subrounded to obscurely quadrangular, furrowed longitudinally on opposite faces, glabrous, frequently defoliating and decorticating basally with age but sometimes leafy to the base; internodes 2–5 mm long, with knobby thickenings that persist where a leaf has fallen away, older branches with a thick, caducous, cork-like bark, nodes with short vinaceous glandular trichomes. Leaves sessile to subsessile or with a flattened petiole ca. 1 mm long, spreading to subimbricate on the uppermost branchlets; blade 10–35 × 6–14 mm, flat, chartaceous, oblong-elliptic to elliptic or elliptic-ovate, base attenuate, apex broadly acute to subobtuse or occasionally subrounded, margin entire to obscurely subserrulate, glandular-ciliate, sparsely to moderately hirsute with glandular trichomes 1–2 mm long on both surfaces, the trichomes surrounded at the base by minute sessile glands, flat, 3-nerved. Flowers 5-merous, solitary or paired, terminal (but sometimes pseudoaxillary with the elongation of lateral shoots), sessile or on pedicels up to 2 mm long. Bracts or the uppermost leaves subtending the flowers eventually becoming broader and suborbicular, rounded at the apex, but otherwise similar to the principal leaves. Hypanthium (at anthesis) 8–10 × 3–4 mm, oblong-subcylindrical, constricted above the ovary, 7–8 mm distally, moderately covered with spreading glandular trichomes from the base to the middle, apex glabrous, or totally glandular. Calyx tube ca. 0.5 mm long, inconspicuous; calyx lobes (at anthesis) 7–11 × 1–1.5 mm, membranaceous, caducous, linear-triangular to triangular-oblong, acute at the apex tapering to an eglandular trichome 0.5–1 mm long, abaxially glandular-hirsute, adaxially glabrous, margins obscurely serrulate, glandular-ciliolate. Petals 15–30 × 10–20 mm, pink with a yellow-white base, rarely pale yellow throughout, obovate, apex rounded, asymmetrically and shortly apiculate with a glandular trichome 1 mm long, base attenuate, margins eciliate. Stamens 10, dimorphic: large (antesepalous) stamens 5, filaments 8–10 mm long, anther thecae 3.5–4 × 1–1.2 mm, oblong, yellow, rostrum ca. 0.5 mm long, yellowish-white, pedoconnective 4–5 mm long, appendage 1.5 mm long, bilobed, yellow; small (antepetalous) stamens 5, filaments 7–9 mm long, anther thecae 3.5 × 1.1 mm, oblong, yellow, rostrum 0.5 mm long, pedoconnective 2 mm long, appendage 0.7 mm long, obscurely lobed. Ovary 5-locular, 4/5 to almost totally inferior, style 8–9 mm long, curved at the apex, glabrous, stigma punctiform. Fruiting hypanthium (including calyx lobes) 11–13 mm long. Capsule (at maturity) 9–10 × 5 mm long, oblong, dehiscing from the base to the apex, vascular strands occasionally persisting long after

94 • Phytotaxa 315 (1) © 2017 Magnolia Press

MARTINS & ALMEDA capsule walls have fallen away. Seeds 0.66–1.01 × 0.32–0.41 mm, reniform to somewhat L-shaped or oblong, dark brown, periclinal cell walls of the testa convex and low domed (colliculate) becoming concave (foveolate), raphal zone about 20–50% the length of the seed. Chromosome number: unknown.

Illustrations:— Figure 38 ; Cogniaux (1883: t. 33).

Photographic images:— Figures 2J–L ; 8C, D .

Phenology:—Flowering January through October; fruiting April, June, August, and October.

Distribution and habitat:—Endemic to the Cadeia do Espinhaço of Minas Gerais where it is largely restricted to the Serra do Cipó and the Serra do Caraça in campo rupestre, gallery forest margins, campo limpo and campo arenoso at 840–1950 m elev. Figure 39 .

LAVOISIERA ( MELASTOMATACEAE )

Phytotaxa 315 (1) © 2017 Magnolia Press • 95

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MARTINS & ALMEDA

Conservation status:—This species has been collected repeatedly over many decades on the Serra do Cipó where it can be locally common. We have examined some 60 collections. The EOO is 2,529 km ² and the AOO is 40 km ². Some populations are protected in Parque Nacional Serra do Cipó and the Serra do Caraça population is protected in a Private Natural Heritage Reserve. Periodic fires pose an ongoing threat. In view of its limited AOO and fragmented population structure we recommend a classification of Endangered (EN): B1ab(iii).

Discussion:— Lavoisiera glandulifera is readily recognized by its oblong-elliptic to elliptic or elliptic-ovate leaf blades that are sparsely to moderately covered with spreading gland-tipped trichomes on both surfaces. These trichomes are unusual in that each one is beset at the base with minute sessile glands ( Figure 38B ). Other distinguishing features of L. glandulifera include its 5-merous flowers, glandular-hirsute hypanthia and calyx lobes, and 5-locular ovary.

Morphologically, Lavoisiera glandulifera is uniform throughout its geographic range. It frequently grows with L. caryophyllea , L. firmula , L. imbricata , and L. sampaioana on the Serra do Cipó. Two anomalous collections (Almeda et al. 9201 and 9202) from Serra do Cipó combine characters of L. glandulifera and L. imbricata in a way that can only be explained as an example of interspecific hybridization. Almeda et al. 9201 has ovate-oblong leaves that lack a callose-thickened margin (like L. glandulifera ) but they are adaxially glabrous (like L. imbricata ) and copiously beset with smooth gland-tipped (in part) trichomes abaxially that have the minute glands at the base of each trichome (like L. glandulifera ). The leaves are intermediate in size (8–10 × 6–10 mm) between the putative parents at that particular Serra do Cipó locality, as is the length of the fruiting hypanthia (10 mm). The ovary is 6-locular (like L. imbricata ), the calyx lobes are scarious (like L. imbricata ) and tardily caducous. In L. imbricata the calyx lobes are scarious but persistent and in L. glandulifera they are not scarious but are caducous. Almeda et al. 9202 also has ovate-oblong leaves of intermediate size (10–12 × 5–7 mm) that are not callose-thickened (like L. glandulifera ) with scattered gland-tipped trichomes along the distal margins adaxially, an intermediate character state. These trichomes also have basal sessile glands at the base (like L. glandulifera ). The abaxial surface of the leaves in this collection have scattered gland-tipped trichomes along the whole length of the blade and each trichome also has sessile glands at the base (like L. glandulifera ). The fruiting hypanthia are intermediate in length (8 mm) as opposed to 6 mm in L. imbricata and 11–13 mm in L. glandulifera . In this collection the flowers are 5-merous, the ovary is 5-locular, and the calyx lobes are caducous (like L. glandulifera ).

The closest relative of Lavoisiera glandulifera is clearly L. harleyi of Bahia. These two species share similar foliar morphology and indumentum details on leaves, hypanthia, and calyx lobes. The latter differs by its 6-merous flowers, 6-locular ovary, shorter calyx lobes (3–4 mm vs. 7–11 mm), and larger anther thecae (6–6.5 mm vs. 3.5 mm) in both the antesepalous and antepetalous whorls.

Additional specimens examined:— MINAS GERAIS: Mpio. Santana do Riacho , Serra do Cipó , ca. Km 127 na estrada para Conceição do Mato Dentro , 19°10'S, 43°30'W, Almeda et al. 7780 ( CAS!, MO!, NY!, UEC!); Serra do Cipó. Between Kms 111 and 112 along the road from Lagoa Santa to Conceição do Mato Dentro , 19°17'S, 43°34'W, Almeda et al. 8562 ( CAS!, UEC!); Mpio. Serra do Cipó , Km 128, Serra do Cipó. Açude de Cima ca. 40 km from Distrito Cardeal Mota along the road to Fazenda Cachoeira and Pico do Breu , 19°7'S, 43°41'W, Almeda et al. 8592 ( CAS!, MO!, UEC!); Serra do Caraça, Parque do Caraça on the trail to Pico Inficionado, 20˚8’2” S, 43˚27’54.7” W, Almeda et al. 8884 ( CAS!, NY!, UEC!); Mpio. Santana do Riacho , Serra do Cipó , about 23–24 kms from the bridge over the Rio Cipó on the road from Serra do Cipó (formerly Cardeal Mota ), about 38 km S of Conceição do Mato Dentro on a temporary detour road while the main Serra do Cipó / Conceição road was being paved, 19˚14’49.3” S, 43˚30’42.7” W, Almeda et al. 9199 ( BHCB!, BM!, CAS!, COL!, RB!, UEC!); Mpio. Conceição do Mato Dentro , Parque Natural do Riberão do Campo (Tabuleiro), 22 km from the center of Conceição do Mato Dentro, upslope from the visitor center, 19˚4’30.2” S, 43˚32’51.9” W, Almeda et al. 9209 ( CAS!, UEC!); Mpio. Santana do Riacho , Serra do Cipó , 27 km beyond the bridge over the Rio Cipó enroute to Conceição do Mato Dentro , -19.25918, -43.53125, Almeda et al. 9700 ( CAS!, RB!, UEC!); Mpio. Santana do Riacho , Serra do Cipó, Retiro do Alto do Palácio, 25 km NE de Cardeal Mota, Arbo et al. 4254 ( SPF!) ; Mpio. Santana do Riacho , Serra do Cipó , Retiro do Alto do Palácio, 25 km NE de Cardeal Mota, Arbo et al. 4942 ( SPF!) ; Claussen 24 ( P!) ; Mpio. Santana do Riacho , Serra do Cipó, Damazio s.n. ( RB!) ; Mpio. Santana do Riacho , Serra do Cipó, Damazio 1960 (G-2!, RB!) ; Mpio. Santana do Riacho , vicinity of Alto do Palácio along road from Chapéu do Sol to Conceição do Mato Dentro , Daniel & Hensold 2304 ( CAS!, US!); Mpio. Santana do Riacho , Serra do Cipó , Km 131 & Km 137, Duarte 2470 ( BM!, G!, M!, MO!, RB!, US!); Mpio. Santana do Riacho, Serra do Cipó, Km 131 &

LAVOISIERA ( MELASTOMATACEAE )

Phytotaxa 315 (1) © 2017 Magnolia Press • 97 Km 137, Duarte 2693 (B!, NY-2!, RB!, S!); Mpio. Santana do Riacho, Serra do Cipó, Km 137, Duarte 6479 (B!, F- 2!, RB!, UB!); Mpio. Santana do Riacho, Serra do Cipó, Km 138, antiga estrada do Serro, Duarte 9699 (B!, F!, G!, M-2!, RB!); Mpio. Santa Bárbara, Caraça, Morro Carapuça, Glaziou 14757 (C!, K!, LE!, P!, R!); Mpio. Santa Bárbara, Caraça, Morro Carapuça, Glaziou 14758 (C!, G!, K!, LE!, R!, US!); Mpio. Santana do Riacho, Serra do Cipó, Glaziou 19269 (C!, G!, K-2!, LE!, P!, R!, US!); Mpio. Santana do Riacho, Serra do Cipó, Fazenda Palácio, Hatschbach 30055 (C!, MBM!); Mpio. Santana do Riacho, Serra do Cipó, Fazenda Palácio, Hatschbach 31596 (F!, NY!); Mpio. Santana do Riacho, Serra do Cipó, Heringer & Castellanos 6186 (NY!, R!, UB!, US!); Mpio. Santana do Riacho, Serra da Salitreira, Palácio, Serra do Cipó, Horta & Andrade 149 (BHCB!); Mpio. Santana do Riacho, Vacaria, summit of Serra do Cipó, ca. 170 km N of Belo Horizonte, Irwin et al. 20637 (F!, MO!, NY!, RB!); Mpio. Santana do Riacho, Vacaria, summit of Serra do Cipó, ca. 170 km N of Belo Horizonte, Irwin et al. 20646 (NY!); Summit of Serra do Caraça, Irwin et al. 29052 (C!, F!, NY!); Mpio. Santana do Riacho, Km 132 ao longo da Rodovia Lagoa Santa-Conceição do Mato Dentro-Diamantina, Joly et al. 244 (UEC!); Mpio. Santana do Riacho, Serra do Cipó, próximo ao Palácio, Km 135, Lima 359 (RB!); Mpio. Santana do Riacho, Serra do Cipó, Km 140, próximo a ponte Preta, Mattos Fo. & Rizzini 105 (RB!); Mpio. Santana do Riacho, Km 126 da estrada da Serra do Cipó-MG, Medeiros s.n. (MBM!); Mpio. Santana do Riacho, Serra do Cipó Km 134-Estrada Pilar, Mello Barreto 311 (BHCB ex BHMH!, F!); Mpio. Santana do Riacho, Serra do Cipó Km 137, Estrada Pilar, Mello Barreto 1302 (NY!, RB!); Mpio. Santana do Riacho, Serra do Cipó Km 124, Mello Barreto 7036 (F!, NY!); Mpio. Santa Bárbara, Caraça, Mello Filho, L. Emygdio et al. 3516 (R!); Mpio. Santa Bárbara, Caraça, Mendonça 26 ( US!); Mpio. Santa Bárbara, Serra do Caraça, Pereira 2584 (F!, G!, NY-2!, RB!); Mpio. Diamantina, Pereira 2788 (RB!); Mpio. Santana do Riacho, Serra do Cipó, Pereira 2885 (NY!, RB!); Mpio. Congonhas do Norte, estrada para Santana do Riacho, Serra da Carapina ( Serra Talhada na folha IBGE), 18°56'S, 43°41'W, Pirani et al. 4175 (CAS!, SPF!); Mpio. Santana do Riacho, Fazenda Inhame. Serra Mineira , 18°55'S, 43°54'W, Pirani et al. CFSC 8090 (SPF!, US!); Mpio. Santana do Riacho, Serra do Cipó. Fazenda Cachoeira da Capivara, Sakuragui & V. C. Souza 120 (ESA!); Sampaio 6586 (R!); Sampaio 6914 (R!); Mpio. Santana do Riacho, Serra do Cipó, Schwacke 7796 (BHCB!, RB!); Mpio. Santana do Riacho, Serra do Cipó, Schwacke 12286 (W!); Mpio. Santa Bárbara, Caraça, trilha para a Carapuça, Semir et al. 28879 (HUFU!, UEC!); Mpio. Santana do Riacho, Serra do Cipó, entrada para Cachoeira da Anta, J. P. Souza et al. 2199 (ESA!); Mpio. Santana do Riacho, Serra do Cipó. Rodovia Belo Horizonte-Conceição do Mato Dentro, ca. 18 km após Chapéu do Sol, 19°14'S, 43°32'W, V. C. Souza et al. 8197 (ESA!, SPF!); Mpio. Santana do Riacho, Serra do Cipó. Rodovia Belo Horizonte-Conceição do Mato Dentro, ca. 18 km após Chapéu do Sol, 19°14'S, 43°32'W, V. C. Souza et al. 8215 (ESA!); Mpio. Santana do Riacho, Serra do Cipó. Fazenda Cachoeira da Capivara, V. C. Souza et al. 11634 (CAS!); Mpio. Santa Bárbara, Caraça-Cascata Padre Rabelo, Tales 216 (BHCB!); Mpio. Santa Bárbara, Serra do Caraça, Ule 2535 (R-2!, US!); Mpio. Santa Bárbara, Caraça, Vainio s.n. ( US!); Mpio. Santana do Riacho, Serra do Cipó, Vidal s.n. (R!); Mpio. Santana do Riacho, Serra do Cipó. Entre o Palácio e o Morro do Pilar, Vidal II-6064 (F!, R!); Mpio. Santa Bárbara, Serra do Caraça. Caminho para o Mirante, Zappi & Scatena 10905 (CAS!, US!).

Putative hybrids between L. glandulifera and L. imbricata :— MINAS GERAIS: Mpio. Santana do Riacho, Serra do Cipó, about 23–24 kms from bridge over Rio Cipó about 38 km S of Conceição do Mato Dentro on temporary detour road, 19°14’49.3”S, 43°30’42.7”W, Almeda et al. 9201 (CAS!, NY!, UEC!); Ibid., Almeda et al. 9202 (CAS!, UEC!).

Figures

FIGURE 2. Images of Lavoisiera species. A. L. cordata (Almeda et al. 7798). B−C. L. cordata (Almeda et al. 9696). D. L. crassifolia (Almeda et al. 8442). E. L. crassifolia (Almeda et al. 7807). F. L. crassifolia (Almeda et al. 9051). G−H. L. firmula (Almeda et al. 9197). I. L. gentianoides (Almeda et al. 8314). J. L. glandulifera (Almeda et al. 9700). K. L. glandulifera (Almeda et al. 9199). L. L. glandulifera (unvouchered image). Image credits: A−K. Frank Almeda; L. Livia Echternacht.

FIGURE 8. SEM images of seeds for representative Lavoisiera species showing lateral symmetrical plane (left) and detail of testa cells and cell wall microrelief (right). A–B. L. gentianoides (Mori et al. 12491). C–D. L. glandulifera (Glaziou 14758). E–F. L. grandiflora (Macedo 3578). Scale bars: A=300 µm, B=30 µm; C & E=430 µm; D=43 µm, F=150 µm.

FIGURE 38. Lavoisiera glandulifera. A. Habit. B. Adaxial leaf surface with enlarged surface detail (lower right). C. Flower. D. Hypanthium and calyx lobes. E. Ovary cross-section. F. Petal. G. Ventral view of antesepalous stamen. H. Ventral view of antepetalous stamen. I. Seed. (Based on: A–I, Almeda et al. 8592, CAS, UEC.)

FIGURE 39. Distributions of Lavoisiera glandulifera, L. itambana, L. macrocarpa, and L. mucorifera.