Lavoisiera grandiflora A. Saint-Hilaire ex Naudin (1844: 148–149)

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

18. Lavoisiera grandiflora A. Saint-Hilaire ex Naudin (1844: 148–149) . Type:— BRAZIL. Minas Gerais: “In paludosis prope praedium Taipa ad limites provinciarum Minas Geraes et Goyas,” A. Saint-Hilaire 629, Cat. C¹ (holotype: P!; isotypes: F!, P-00723377-n.v., online image!, P-00723378-n.v., online image!).

= Lavoisiera riedeliana Cogniaux in Martius (1883: 141). syn. nov. Type :— BRAZIL. Minas Gerais: “Habitat in uliginosis prov. Minas Geraes ad Villa Franca,” L. Riedel & G. H . Langsdorff 2340 (Lectotype, here designated: LE!, photos: MICH!, NY!, RB!, US!; isolectotypes: LE!, M!, P!; probable isolectotype: W!) .

Erect, dichotomously much-branched virgate shrubs 1–2.5 m tall. Branches and branchlets subquadrangular or quadrangular to gradually subrounded toward the base, distally compressed, slightly furrowed longitudinally on opposite faces, essentially glabrous or sparsely and randomly beset with reddish glandular trichomes 1.5–2 mm long like the hypanthium and calyx lobes, frequently leafy to the base or defoliating basally with age; internodes 6–20 mm long, with knobby thickenings that persist where a leaf has fallen away, nodes with inconspicuous

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MARTINS & ALMEDA yellowish glandular trichomes. Leaves sessile, semi-amplexicaul, spreading or subimbricate and congested at the ends of the branchlets; blade 20–45(–80) × 10–20(–40) mm, subcoriaceous, oblong-lanceolate to ovate-oblong, base rounded to shortly subauriculate, apex subrounded or acute, margins entire, sometimes slightly revolute, obscurely and narrowly subcallose, adaxial surface bright green and glabrous, abaxially pale green to yellowish-green, glandular-punctate or resinous-punctate, basically flat, 7–9-nerved (only the inframarginal pair extending to the apex), with laxly reticulate inconspicuous venules evident on the abaxial surface. Flowers typically 6-merous (sometimes 5-merous), in dichasia or congested pseudodichasia, frequently solitary and at first terminal but becoming pseudolateral because of elongation of lateral shoots, pedicel 3–4 mm long. Bracts 28–40 × 18–20 mm, sessile, ovate, glabrous, faintly 7-nerved, bracteoles 15 × 5 mm, sessile, oblong to obovate, glabrous, 1–3-nerved. Hypanthium (at anthesis) 12–18 mm long, cylindric-oblong and suburceolate, 5 mm wide to 8–9 mm wide distally, glabrous and minutely granulose-furfuraceous or inconspicuously beset with sessile minute glands, occasionally sparsely to moderately glandular-setose with trichomes ca. 1.5 mm long, green. Calyx tube 1.5–2 mm long; calyx lobes 4–6 × 2–3.5 mm, chartaceous, ± subulate to triangular-acute at the apex and sometimes with a gland-tipped trichome, typically glabrous or sparsely glandular-pubescent on both surfaces, the glands sessile, margins entire or inconspicuously ciliolate, persistent. Petals 25–30 × 10–15 mm, pink with a yellow base or white to yellowish-white with a yellow base, obovate-oblong, apex subrounded to truncate, base attenuate, margins entire, eciliate. Stamens 12, dimorphic: large (antesepalous) stamens 6, filaments 12 mm long, yellow, anther thecae 8 × 2 mm, yellow, linear-oblong, rostrum 0.7 mm long, yellow, pedoconnective 10–12 mm long, appendage 3–4 mm long, yellow but often becoming brownish following pollination, deeply emarginate at the apex or truncate; small (antepetalous) stamens 6, filaments 8 mm long, yellow, anther thecae 7 × 1.5 mm, yellow, linear-oblong, pedoconnective 2–3 mm long, appendage 1–1.5 mm long, yellow, subrounded to obscurely bilobed. Ovary 6- locular, 4/5 inferior, style 12–15 mm long, glabrous, curved apically, stigma punctiform. Fruiting hypanthium (including calyx lobes) 25–30 mm long, pedicel 5 mm long, oblong, constricted distally above the ovary summit but below the calyx lobes. Capsule (at maturity) 12–13 × 8–10 mm, oblong, coriaceous, grayish-brown, enveloped by the persistent and enlarged hypanthium and calyx lobes, dehiscing from the apex to the base, the vascular strands often persisting long after capsule walls have fallen away. Seeds 1.20–2 × 0.43–0.62 mm, oblong, abruptly tapering to a spur-like extension on the chalazal end of the seed, brownish or orangish-red, periclinal cell walls of the testa concave (foveolate), the raphal zone about 50% the length of the seed. Chromosome number: n=12.

Illustrations:— Figure 40 ; Cogniaux (1883: t. 31, as L. riedeliana ).

Photographic images:— Figures 3A–C ; 8E, F .

Phenology:—Flowering and fruiting all year.

Distribution and habitat:—Locally common from the Distrito Federal and Goiás south to Minas Gerais where it is local and mostly centered in the Uberlândia and Serra do Caraça areas with a disjunct population extending to Município Franca in São Paulo where it is known only from old collections in cerrado, campo rupestre, brejo (often with standing water), campo úmido, gallery forest margins, rocky stream banks, and vereda borders at 975–1600 m elev. Figure 33 .

Conservation status:—This locally common species is known from some 75 collections gathered over several decades from a comparatively widespread range. The EOO is 31,954 km ² and the AOO is 44 km ². Several populations are protected in Parque Nacional Chapada dos Veadeiros, Parque Nacional de Brasília, Parque Estadual da Serra dos Pireneus, Reserva Ecológico do IBGE and the Fazenda Experimental do Glória (Minas Gerais, Mpio. Uberlândia). In view of its broad EOO and presence in several protected sites we recommend a classification of Least Concern (LC).

Discussion:— Lavoisiera grandiflora is an erect virgate shrub that can form locally dense stands especially when it is growing in brejo. The profuse shoots are evidently short-lived because many dead ones can be found among the living ones on any one individual. It has oblong-lanceolate to ovate-oblong leaves that are 7–9-nerved, prevailingly 6-merous flowers, a 6-locular ovary, pedicels 3–4 mm long, cylindric-oblong hypanthia that are 21–28 mm long and conspicuously constricted distally in fruit ( Figure 3B ), persistent calyx lobes, and unique seeds that are abruptly tapered into a spur-like extension at the chalazal end ( Figure 40J ).

Lavoisiera grandiflora is variable in floral merosity, flower color, and the indumentum on hypanthia and calyx lobes. The populations we studied in Goiás were prevailingly 6-merous but we noted some flowers that were 5- merous and the label on the type notes that the flowers are 6-merous but occasionally 7-merous. However, we have seen no specimens with 7-merous flowers. Two petal color forms can be found in this species. One has intense pink

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MARTINS & ALMEDA petals with an inverted V-shaped bright yellow patch at the base ( Figure 3A ). In the other form the petals are white or yellowish-white (often with a faint flush of pink) with a diffuse yellow patch at the base ( Figure 3C ). Flower color is often homogeneous within a population. The majority of the collections we have studied have pink petals but populations in the Distrito Federal and Minas Gerais can have both color forms. Different individuals in the population at Fazenda Experimental do Glória (Minas Gerais) have pink petals (Romero & Nakajima 593) and white petals (Romero & Nakajima 594) so flower color can vary within a single population. This Minas Gerais population (both pink and white-flowered forms) also has hypanthia and calyx lobes that are moderately beset with spreading gland-tipped trichomes. Judging from the protologue, it appears that Cogniaux (1893) circumscribed L. riedeliana based, in part, on the glandular-setose hypanthia and calyx lobes coupled with 5-nerved (rarely 3- nerved) leaf blades, and triangular calyx lobes (vs. subulate calyx lobes in L. grandiflora ). With respect to foliar venation, it appears that Cogniaux examined only upper immature blades because mature blades on all the specimens he examined and all others we have seen have the typical 7–9-nerved leaf blades of the species. Cogniaux’s calyx lobe description for L. riedeliana also appears to be part of the variation found in L. grandiflora since the lobes can be subulate at anthesis and then become distinctly triangular in post-anthesis and on fruiting hypanthia. The petal color variation and indumentum on hypanthia and calyx lobes do not correlate with other characters in any meaningful way so we treat all of this variation as part of a single species and relegate L. riedeliana to synonymy.

In the protologue, Naudin (1844) suggested that L. grandiflora is closely related to L. gentianoides . Both species have large flowers and leaves but the latter, which is largely centered in Bahia, has leaf blades that are held perpendicular to somewhat reflexed on the stems when fresh, the 5-merous flowers are sessile and borne in congested mostly 6–10-flowered inflorescences ( Figure 37 ), the bracts and bracteoles are glandular-puberulous, the petals are consistently white, the hypanthia and tardily caducous calyx lobes are always covered to some degree with spreading gland-tipped trichomes, the pedoconnective of the larger set of stamens is only 5 mm long, and the ovary is 5-locular.

Perhaps the closest relative is L. alba , which is mostly centered in Minas Gerais. It differs in its overall glabrosity, quadrangular upper branchlets that are prominently angled and carinate to narrowly winged, bluish-green or pale green glaucous leaves, 5-merous flowers, uniformly white petals, and 5-locular ovary.

Additional specimens examined:— DISTRITO FEDERAL: Fazenda Agua Limpa ( FAL) , Campo Experimental da UnB. Córrego da Onça, 15°57'S, 47°54'W, S. P. Almeida et al. 129 ( HUFU!, UB!) ; Mpio. Brasília , Reserva Ecológica do IBGE, 15°56'S, 47°54'W, Azevedo 80 ( RB!) ; Mpio. Brasília , Granja do Riacho Fundo , Cobra 596 ( US!); Mpio. Brasília , Margem do Lago, Duarte 10579 ( RB!) ; Mpio. Brasília , Fercal, L. Duarte & Silva Santos 139 ( NY!) ; Mpio. Brasília , Reserva Ecológica do IBGE.DF, 15°55'S, Furtado 317 ( UEC!) ; Fazenda Agua Limpa , Olho D'água da Onça, 15°56'S, 47°54'W, Guimarães & C. F. R. Cardoso s.n. ( CAS!) ; Mpio. Brasília , Km 8 da estrada Brasília-Anápolis , Heringer 7940-133 ( NY!, US-2!); Cabeça do Veado , Heringer 8307/501 ( US!); Mpio. Brasília , Regiao da Papuda, Heringer 18656 ( UEC!) ; Mpio. Brasília , Estação Florestal Cabeça do Veado , Heringer et al. 5886 ( CAS!, K!, MO!, US!); Mpio. Brasília , Bacia do Rio São Bartolomeu, Heringer et al. 6839 ( MO!) ; Mpio. Brasília , Lagoa Paranoá , confluência do Córrego Canjerana , Heringer et al. 7437 ( US!); Mpio. Brasília, ca. 20 km S of Brasília, Irwin et al. 8593 ( F!, MO!, NY!) ; Mpio. Brasília , ca. 20 km S of Brasília, Irwin et al. 8596 ( UB!) ; Near Sobradinho , Irwin et al. 8727 ( NY!, US!); E of Lagoa Paranoá, Irwin et al. 11222 ( LE!, MO!, NY!, S!) ; Perto da Cachoeira da Forquilha do Ribeirão Sobradinho , 15°44'S, 47°44'W, Kirkbride Jr. 5179 ( F!, K!, NY!, RB!, US!); Mpio. Brasília, Reserva Ecológica do IBGE, Mendonça 225 ( CEPEC!, COL!) ; Mpio. Brasília , Pelanda s.n. ( US!); Mpio. Brasília, Sobradinho, Pereira 9067 ( RB!) ; Mpio. Brasília , Parque Nacional de Brasília, Philcox & Onishi 4297 ( K!) ; Chapada da Contagem , a 6 km da BR 010 , na Rodovia para Brasilândia BR 080 , 15°35'S, 47°57'W, Pirani et al. 1634 ( CAS!, SPF!, US!); Mpio. Brasília , Pires et al. 9771 ( US!); Mpio. Brasília , Chapada da Contagem, NE of Brasília, along highway DF 5, about 25 km W of intersection with BR 020 , 15°36'S, 48°3'W, Plowman 9952 ( CAS!, F!, G!, K!, NY!, RB!) ; Mpio. Brasília , Fazenda Agua Limpa , near Vargem Bonita , ca. 18 km SSW of Brasília TV tower, Ratter & Fonseca 2789 ( K!, NY!, US!). GOIÁS: Mpio. Alto Paraíso de Goiás , off of Hwy GO 118 at Km 184 N of Alto Paraíso de Goiás enroute to Terezina de Goiás, -14.00334, -47.52111, Almeda et al. 9459 ( BHCB!, CAS!, CEN, K!, NY!, RB!, SP!, UEC!) ; Mpio. Pirenópolis , Parque Estadual da Serra dos Pireneus , trail to Cachoeira Sonrizal and beyond the small waterfalls to top of distant ridge, -15.81485, -48.85849, Almeda et al. 9522 ( CAS!, UEC!) ; Chapada dos Veadeiros , 20 km by road N of Alto do Paraíso, Anderson 6389 ( NY!) ; Serra dos Pireneus , ca. 15 km (straight line) N of Corumbá de Goiás , Anderson et al. 10457 ( C!, COL!, F!,

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MO!, NY!, RB!, US!); Mpio. Alto Paraíso, estrada norte do Parque Nacional Chapada dos Veadeiros, Castro & Barbosa 237 ( CAS!, HUFU) ; Près de Cocal , Glaziou 21304 (BR-2!, C!, G!, K!, LE!, NY!, S!) ; Serra dos Veadeiros ao Pouso Alto, Glaziou 21305 (BR-2!, C!, F!, G!, K!, LE!, NY!, R!, RB!, S!, US!); Glaziou 21306 (BR-2!, G!, K!) ; Rod. GO 118 , Kms 5–10 ao Norte de Alto Paraíso, Hatschbach & Nicolack 53922 ( S!) ; Mpio. Alto Paraíso, Chapada dos Veadeiros, Hatschbach et al. 36352 ( MBM!, US!); Mpio. Cristalina, Rio das Perdizes, Hatschbach et al. 53789 ( CAS!, US!); Mpio. Ipameri, Ponte Alta, 5 km oeste de Ipameri, Hatschbach 43801 ( BR!, C!, MICH!, NY!, US!); Rod . BR 040 , 4 km oeste de Cristalina, Hatschbach 44578 ( MBM!) ; Rod. BR 040 , 4 km oeste de Cristalina, Hatschbach 44758 ( COL!, F!) ; Samambaia , afluente do Corumbá, Heringer 9571/1751 ( NY!, UB!, US!); ca. 30 km NW of Veadeiros, 14°S, 47°W, Irwin et al. 12946 ( F!, LE!, M!, NY!, S!) ; Mpio. Cristalina , ca. 3 km N of Cristalina, 17°S, 48°W, Irwin et al. 13249 ( LE!, MO!, NY-2!, S!); Mpio. Corumbá de Goiás, ca. 20 km NW of Corumbá de Goiás, near Pico dos Pireneus, Irwin et al. 19249 ( CAS!, MO!, NY!, UEC!, US!); Mpio. Alto do Paraíso, ca. 20 km N of Alto do Paraíso, Irwin et al. 32155 ( C!, F!, NY!) ; Mpio. Alto do Paraíso , 12–20 km N of Alto do Paraíso ; W of road to Monte Alegre , King & Bishop 8833 ( MO!, UB!, US!); Mpio. Cristalina , BR 050 , ca. 10 km S de Cristalina, Kral et al. 75302 ( CAS!, NY!) ; Mpio. Anápolis, Macedo 3578 ( RB!, S!) ; Mpio. Corumbá de Goiás , Pireneus , Macedo 4324 ( US!); Mpio. Alto Paraíso , ca. 16 km de Alto Paraíso em direção a Teresina de Goiás, 13°46'S, 47°30'W, Munhoz et al. 151 ( UB!) ; Mpio. Alto Paraíso , Chapada dos Veadeiros, 16°0'S, 67°35'W, Ratter et al. 2613 ( K!, NY!, UB!) ; Mpio. Pirenópolis, estrada para a Serra dos Pireneus , próximo ao Morro do Cabeludo , Romero et al. 5607 (CAS-2!); Mpio. Sobradinho , E. Santos 1724 ( NY!, R!) ; Chapada dos Veadeiros ; Rio dos Brancos, próximo à estrada entre S. Gabriel de Goiás e São João da Aliança, Semir s.n. ( UEC!) . MINAS GERAIS: Mpio. Patrocínio , Fazendas da Terra, Farah et al. 978 ( ESA!, UEC!) ; Mpio. Santa Bárbara, Serra do Caraça , Glaziou 19275c ( R!) ; Mpio. Uberlândia , Fazenda Experimental do Glória , Liberali s.n. ( CAS!, HUFU, US!); Mpio. Estrela do Sul , Lagoa da Estiva , Mello Barreto 11616 ( BHCB ex BHMH!); Mpio. Uberlândia, Fazenda Experimental do Glória, Nakajima et al. 248 ( CAS!, HUFU) ; Mpio. Paracatú, E. T. Neto 1835 (BHCB!); Mpio. Uberlândia, Fazenda Experimental do Glória, Nakajima & Romero 251 ( CAS!, HUFU!) ; Mpio. Uberlândia , Fazenda Experimental do Glória , Romero & Nakajima 593 ( CAS!, HUFU, US!); Mpio. Uberlândia , Fazenda Experimental do Glória , Romero & Nakajima 594 ( CAS!, HUFU, US!); Mpio. Uberlândia , Fazenda Experimental do Glória, Romero & Nakajima 596 ( CAS!, HUFU) ; Mpio. Uberlândia, Clube Caça e Pesca Itororó, Romero & Nakajima 2329 ( CAS!, HUFU) ; Mpio. Uberlândia, Clube Caça e Pesca Itororó, Romero 2101 (CAS-2!, HUFU) . SÃO PAULO: Mpio. Franca, Löfgren & Edwall 2078 ( BR!, C!, SP!) ; Mpio. Franca, Lund s.n. (C-2!); Mpio. Franca, Riedel s.n. ( C!) ; exact locality not specified, Riedel s.n. ( BM!) ; Mpio. Franca, Riedel & Lund 48 ( C!) ; Mpio. Villa Franca?, Riedel s.n. ( F!) ; Mpio. Franca, Wacket s.n. ( W!) .

Figures

FIGURE 3. Images of Lavoisiera species. A. L. grandiflora (unvouchered image). B−C. L. grandiflora (Almeda et al. 9459). D. L. harleyi (unvouchered image). E−F. L. humilis (Almeda et al. 8497). G. L. imbricata (Almeda et al. 8497). H. L. imbricata (Almeda et al. 9032). I−K. L. itambana (Almeda et al. 9663). L. L. macrocarpa (Almeda et al. 8545). Image credits: A. Pinterest at www.pictopin.com; B, C, E−L. Frank Almeda; D. Fernando B. Matos.

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 33. Distributions of Lavoisiera cordata, L. gentianoides, L. grandiflora, L. pohliana, and L. setosa.

FIGURE 37. Lavoisiera gentianoides. A. Habit. B. Adaxial leaf surface with enlarged detail of of abaxial surface (upper right). C. Abaxial bract surface. D. Abaxial bracteole surface. E. Flower. F. Hypanthium and calyx lobes. G. Ovary cross-section. H. Petal. I. Ventral view of antesepalous stamen. J. Ventral view of antepetalous stamen. K. Seed. (Based on: A–K, Almeda et al. 8318, CAS, UEC.)

FIGURE 40. Lavoisiera grandiflora. A. Habit. B. Adaxial leaf with surface detail (upper left). C. Flower. D. Hypanthium, calyx lobes, and style with detail of adaxial calyx lobe base (upper right) and hypanthial surface (lower right). E. Ovary cross-section. F. Ovule. G. Petal. H. Ventral view of antesepalous stamen. I. Ventral view of antepetalous stamen. J. Seed. (Based on: A–J, Almeda et al. 9459, CAS, UEC.)