Lavoisiera vestita Almeda & A.B. Martins, 2017

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

41. Lavoisiera vestita Almeda & A.B. Martins , sp. nov.

Shrubs or small trees, densely branched, 1–2.5 m tall. Branches densely covered with smooth eglandular trichomes, intermixed with glandular trichomes. Leaves typically with short flattened petioles 1 mm long with some leaves appearing subsessile, membranaceous, spreading to laxly imbricate apically, flat, densely to moderately beset with appressed to somewhat spreading smooth eglandular trichomes on both surfaces, 1–3(–5)-nerved. Flowers 6-merous, solitary. Hypanthium cylindrical to campanulate, densely covered with appressed to ascending-spreading gland-tipped trichomes. Petals pink with a greenish-yellow base. Ovary 6-locular, 4/5 inferior.

Type :— BRAZIL. Minas Gerais: “Mpio. Santo Antônio do Itambé, Parque Estadual do Pico do Itambé. Encostas de barranco no campo, elev. ca. 1700 m,” 16 November 2010, J. L. M . Aranha 164, R . Belinello , G . Shimizu & D. J. P . Gonçalves (holotype: UEC!; isotype: CAS!) .

Shrub or small tree with densely branched upright habit 1–2.5 m tall. Branches and branchlets rounded to subrounded and somewhat compressed distally, densely covered with ascending to spreading smooth eglandular trichomes (intermixed with glandular trichomes) 0.5–1 mm long on young branchlets, pale green, defoliating and

LAVOISIERA ( MELASTOMATACEAE )

Phytotaxa 315 (1) © 2017 Magnolia Press • 171 tardily decorticating at base with age; internodes (0.3–) 0.5–0.8 mm long, nodes with discrete leaf scars. Leaves with short flattened petioles 1 mm long (some appearing subsessile), spreading to laxly subimbricate apically; blade (13–)15–25(–30) × 6–10 mm, membranaceous, oblong to elliptic-lanceolate, base slightly rounded to attenuate, apex broadly acute with a short pungent trichome, the margins entire, greenish-gray, inconspicuously pulverulent, densely to moderately beset with appressed to somewhat spreading smooth eglandular trichomes on both surfaces, the central vein abaxially and leaf margins densely covered with smooth appressed or ascending trichomes 0.75–1 mm long, flat, 1–3(–5)-nerved, the lateral pairs sometimes inconspicuous. Flowers 6-merous, solitary, terminal on primary and secondary branches, becoming central with elongation of lateral shoots, subsessile, pedicel 1 mm long. Bracts 2.4–5 × 1–2 mm, subsessile, oblong, base slightly rounded, apex broadly acute, glandular-hirtellous, 1-nerved, similar to the principal leaves. Hypanthium (at anthesis) 7–8 mm long, cylindrical to campanulate, densely covered with elongate, mostly gland-tipped trichomes ca. 0.3 mm long, constricted above the ovary, 4 mm wide distally. Calyx tube 1.5 mm long; calyx lobes (at anthesis) 1–1.5 × 1 mm, triangular, acute with a pungent apical trichome 0.4 mm long, margins ciliate, abaxially covered with elongate, mostly gland-tipped trichomes, adaxially glabrous, tardily caducous in post-anthesis. Petals 20–25 × 15 mm, pink with a greenish-yellow base, oblong to ± obdeltoid, apex rounded and bluntly cuspidate, base attenuate, margin entire, sparsely and minutely glandular-ciliolate. Stamens 12, dimorphic: large (antesepalous) stamens 6, filaments 8–10 mm long, glabrous, anther thecae 3.5–4 × 1.5 mm, yellow, oblong, rostrum 1 mm long, pedoconnective 6–8 mm long, appendage 1–1.5 mm long, obscurely bilobed; small (antepetalous) stamens 6, filaments 8–9 mm long, glabrous, anther thecae 3.5 × 1 mm, yellow, oblong, rostrum ca. 1 mm long, pedoconnective 2–3 mm long, appendage ca. 1 mm long, rounded. Ovary 6-locular, 4/5 inferior, globose. Style 8–9 mm long, glabrous, slightly curved distally, stigma punctiform. Fruiting hypanthium (including calyx lobes) 8–10 mm long. Capsule (at maturity) 8–10 × 7–9 mm, globose, enveloped by the persistent hypanthium, dehiscing from the apex to the base. Seeds 1.08–1.17 × 0.52–0.60 mm, oblong to subreniform, dull yellow-brown, periclinal cell walls of the testa concave (foveolate), raphal zone about 35–50% the length of the seed. Chromosome number unknown.

Illustration:— Figure 64 .

Photographic images:— Figures 5K, L .

Phenology:—Flowering in November, fruiting in March.

Distribution and habitat:—Endemic to Minas Gerais where it is restricted to Pico do Itambé, Parque Estadual do Pico do Itambé, growing in campo rupestre at cliff edges among rocks from 1692–1950 m along the trail to the summit of the Pico. Figure 28 .

Conservation status:—We have searched for this species on more than one excursion to Pico do Itambé where we have encountered only four individuals on the upper slopes of Pico do Itambé. The EOO is 4 km ² and AOO is 4 km ². The entire population of this species occurs within Parque Estadual do Pico do Itambé so it is afforded some protection. In view of its limited range, extremely small population size, and threat of habitat degradation by periodic fires we assign a classification of Critically Endangered (CR): B2ab(v).

Discussion:—This rare species is distinguished by its prevailingly short-petiolate (1 mm long) leaf blades that have both surfaces copiously covered with ± appressed smooth eglandular trichomes, hypanthia and uppermost cauline internodes covered with a mixture of gland-tipped and eglandular antrorsely spreading trichomes, 6-merous flowers, calyx lobes that are acute apically and terminating in a pungent trichome, and 6-locular ovary.

Among congeners with 6-merous flowers, a 6-locular ovary, and copiously glandular-pubescent hypanthia, L. vestita most closely resembles L. harleyi and L. mucorifera but neither of these two species have consistently petiolate leaves. Lavoisiera harleyi , a Bahia endemic, has wider leaf blades (9–22 mm) that are crowded in pseudowhorls at the ends of branches and each foliar trichome on both adaxial and abaxial surfaces is surrounded by sessile glands at its base. Lavoisiera harleyi also differs in having hypanthia that are uniformly glabrous distally below the torus.

Lavoisiera mucorifera is also endemic to Minas Gerais where it is restricted to the Diamantina plateau but it has never been collected on Pico do Itambé. It differs from L. vestita in having glandular-punctate and viscid abaxial foliar surfaces, serrulate foliar margins, much longer calyx lobes (7–9 mm) that are nearly twice as long as the hypanthia at anthesis and lack a pungent apical trichome, and shorter petals (12–18 mm) that are pale pink, pale lavender, white, or white flushed with pink.

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Etymology:— The specific epithet is derived from the Latin word vestitus, meaning clothed or covered, alluding to the copious indumentum of appressed or ascending smooth eglandular and glandular trichomes on all vegetative parts, hypanthia, and abaxial surfaces of calyx lobes.

Additional specimens examined:— MINAS GERAIS: Mpio. Santo Antônio do Itambé, Parque Estadual do Pico do Itambé, trail from the Fazenda at 1367 m to the summit of the Pico at 2038 m, -18.39868, -43.34816, Almeda et al. 9672 ( BHCB!, CAS!, NY!, RB!, UEC!) .

DOUBTFUL AND EXCLUDED NAMES

Lavoisiera blanchetiana Naudin (1844: 150) . TYPE:— BRAZIL. Bahia: 1844, J. Blanchet 213 (holotype: P-n.v., online image!; isotype: G-n.v., online image!) = Microlicia blanchetiana (Naudin) Cogniaux (1883: 79) .

Lavoisiera candolleana Barreto (1936: 65) . Type:— Brazil. Minas Gerais: “Habitat in civitate Minas Geraes, ad Serra do Cipó,” 24 May 1892, A. F. M. Glaziou 19273 (holotype: R-n.v., online image!; isotypes: B, destroyed, C!, F!-frag., K!, P!; photo of B isotype: F!, NY!). We are unable to place this name with certainty because the isotypes we examined lack flowers and fruits. Barreto’s illustration in the protologue depicts the habit, a representative leaf and a hypanthium that terminates in what he describes as a double calyx. We are at a loss to explain what he interpreted as a double calyx because we have never seen anything like this in a species of Lavoisiera . The holotype was not among the material received on loan from the Museu Nacional in Rio de Janeiro (R) but we have examined an image of it online that has one detached hypanthium (in bud) that shows the 12-merous condition described by Barreto. This funnel-shaped budding hypanthium is highly reminiscent of the six calyx lobes + six epicalyx lobes produced by species of Diplusodon Pohl (1827: 150) , at least four species of which grow on Serra do Cipó (Calvalcanti 1990). Barreto noted that L. candolleana bears a vegetative resemblance to L. cordata which is locally common on the Serra do Cipó and to L. alba which occurs sporadically along the Cadeia do Espinhaço. Vegetatively, L. candolleana is a close match for L. alba because it has the quadrate uppermost cauline internodes and foliar morphology of that species. Based on the strength of these vegetative similarities, we tentatively annotated the sterile isotypes as L. alba . Both L. cordata and L. alba have fairly large flowers with white petals and narrowly campanulate hypanthia. Barreto described the flowers of L. candolleana as “flores parvi rosei” (small pink flowers). This lends some credence to the possibility that the holotype may be a mixed collection of Lavoisiera and Diplusodon since all the species of the latter that occur on Serra do Cipo have pink or magenta-pink petals.

Lavoisiera centiformis var. viminalis (Martius & Schrank ex Candolle) Naudin (1849: 224–225) . TYPE:— BRAZIL. Bahia: “alpinis campestribus ad Serra do Sincorá alt. 4500 ped.,” C. F. P. Martius s.n. (holotype: M-n.v, online image!; isotypes: BM, G-n.v., online image!, P) = Rhexia viminalis Martius & Schrank mss = Lavoisiera viminalis Martius & Schrank ex Candolle (1828: 103) . = Microlicia viminalis (Martius & Schrank ex Candolle) Triana (1871: 28) .

Lavoisiera cerifera Gardner (1844 : tab. 63). TYPE:— BRAZIL. Minas Gerais: elevated mountain tracts NW of Diamantina, G. Gardner 4581 (holotype: BM!; isotypes: F-frag.!, GH-00072340, n.v., online image!, K-2!, P-00245273!, P-00723380, n.v., online image!, US!, W!) = Microlicia cerifera (Gardner) A. B. Martins & Almeda in Almeda & Martins (2001: 3).

Lavoisiera crassinervia Cogniaux ex Glaziou (1908: 253) . nom. nud. = Lavoisiera aff. nervulosa Naudin.

Lavoisiera dichotoma Cogniaux ex Glaziou (1908: 225) . nom. nud. based on BRAZIL. Minas Gerais: A. F. M. Glaziou 19248. = Lavoisiera adamantium Barreto ex Pedersoli.

Lavoisiera dorifolia Cogniaux ex Glaziou (1908: 256) . nom. nud. = Microlicia giuliettiana A. B. Martins & Almeda in Almeda & Martins (2001: 3).

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Lavoisiera elegans Cogniaux in Martius (1883: 160–161). TYPE:— BRAZIL. Goiás: A. F. M. Glaziou 3758 (holotype: BR!; isotypes: C, P-00245290!, P-00723374-n.v., online image!) = Trembleya elegans (Cogniaux) Almeda & A. B. Martins (2001: 6) .

Lavoisiera glandulifolia Naudin (1845b: 192 , tab. 6, fig. 3). nom. nud. = Lavoisiera glandulifera Naudin. Lavoisiera glandulifolia appears to have been a typographical error that inadvertently created a nomen nudum. The epithet was synonymized under L. glandulifera by Cogniaux (1883: 149; 1891:86).

Lavoisiera glutinosa Cogniaux in Martius (1883: 145). TYPE:— BRAZIL. Minas Gerais: “prope Tejuco ( Diamantina ),” Dec. 1824, L. Riedel 1224 (holotype: LE!; isotypes: BR-2!, C, K-275870!, LE!, M-9854!, MO, NY, P-245305!, P-723375-n.v., online image!, P-723376-n.v., online image!, US, W) = Microlicia longipedicellata Almeda & A. B. Martins (2001: 3) .

Lavoisiera goyazensis Cogniaux (1896: 447) . TYPE:— BRAZIL. Goiás: Serra da Balisa , elev. 1500–1600 m, Sept. 1892, Ule 2903 (holotype: BR!; probable isotypes: R!, RB!, US!) = Microlicia crebropunctata Pilger (1903: 17) .

Lavoisiera linifolia Martius & Schrank ex Candolle (1828: 104) . TYPE:— BRAZIL. Minas Gerais: “pratis subuliginosis prov. Minarum Gen. ad Serro Frio” ( Diamantina ), Martius s.n. (holotype: M; isotype: G-00343753- n.v., online image!) = Rhexia linifolia Martius & Schrank , mss = Microlicia linifolia (Martius & Schrank ex Candolle) Chamisso (1834: 395) . Also see Romero (2013).

Lavoisiera luetzelburgii Markgraf (1927: 47) . TYPE:— BRAZIL. Bahia: P. von Luetzelburg 286 (holotype: M!) = Microlicia giuliettiana A. B. Martins & Almeda in Almeda & Martins (2001: 3).

Lavoisiera marqueslisboana Barreto (1935c: 9) . Type:— BRAZIL. Minas Gerais: “In campis siccis arenosisque in Serra do Cipó, secus margines viae ad vicum Morro do Pilar, ubi rara,” A. J. Sampaio 6567a (holotype: R!). This taxon appears to be a hybrid between L. caryophyllea and L. confertiflora . See the discussion under the latter species for details.

Lavoisiera microphylla Cogniaux ex Glaziou (1908: 254) . nom. nud. The specimens bearing this name at BR! and P! ( BRAZIL:— Minas Gerais, Diamantina a Formação, Glaziou 19254a) have no good flowers. We suspect they are a Microlicia .

Lavoisiera microphylla Naudin (1844: 150) . TYPE:— BRAZIL. Minas Gerais: “in campis prope Tijuco adamantium,” 1816–1821, Cat. B1 n 2081, A. Saint-Hilaire s.n. (holotype: P; isotype: P) = Microlicia decipiens Naudin (1849: 232) . Also see Martin and Cremers (2007).

Lavoisiera mucugensis Wurdack (1988: 294) . TYPE:— BRAZIL. Bahia: Mucugê, Corrego Moreira , 22 Jan. 1984, G. Hatschbach 47502 (holotype: MBM!; isotypes: C!, CEPEC!, HUEFS!, US!) = Microlicia mucugensis (Wurdack) Almeda & A. B. Martins (2001: 4) .

Lavoisiera noblickii Wurdack (1995: 821) . TYPE:— BRAZIL. Bahia: Palmeiras, 19 Nov. 1983, L. R. Noblick & A. Pinto 2769 (holotype: HUEFS!; isotypes: CAS!, CEPEC!, US!) = Microlicia noblickii (Wurdack) A. B. Martins & Almeda in Almeda & Martins (2001: 4).

Lavoisiera novemnervia Cogniaux ex Glaziou (1908: 253) . nom. nud. = Lavoisiera alba Martius & Schrank.

Lavoisiera ordinata Wurdack (1974: 135) . TYPE:— BRAZIL. Goiás: Chapada dos Veadeiros, rocky slopes, 12 Feb. 1966, H. S. Irwin et al. 12695 (holotype: US!; isotypes: F!, MO!, NY!, RB!, S!, W!) = Microlicia ordinata (Wurdack) Almeda & A. B. Martins (2001: 5) .

Lavoisiera parviflora Cogniaux ex Glaziou (1908: 254) . This is a nomem superfluum, based on the same type as Rhexia imbricata Thunberg [TYPE:— BRAZIL. Minas Gerais: “prope Villam Riccam ,” G. W. Freyreiss s.n. (UPS-

LAVOISIERA ( MELASTOMATACEAE )

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THUNB-9051)] = Lavoisiera imbricata (Thunberg) Candolle.

Lavoisiera patulifera Cogniaux ex Glaziou (1908: 252) . nom. nud. = Lavoisiera aff. nervulosa Naudin.

Lavoisiera pulchella Triana (1883: 150) . This name was attributed to Triana and listed as a synonym of L. australis (which is a synonym of L. pulchella Cham. ) by Cogniaux (1883). We see no evidence that Triana (1871) intended to describe a new species with this name because he clearly attributed the epithet “pulchella ” to Chamisso. Thus L. pulchella Triana is illegitimate.

Lavoisiera pulcherrima Martius & Schrank ex Candolle var. a ngustifolia Barreto (1935b: 188). nom. nud. = Lavoisiera pulcherrima Martius & Schrank ex Candolle.

Lavoisiera pulcherrima var. cipoana Barreto (1935b: 189) . nom. nud. = Lavoisiera pulcherrima Martius & Schrank ex Candolle.

Lavoisiera pulcherrima var. latifolia Barreto (1935b: 190) . nom. nud. = Lavoisiera pulcherrima Martius & Schrank ex Candolle.

Lavoisiera pulcherrima var. major Barreto (1935b: 189) . nom. nud. = Lavoisiera pulcherrima Martius & Schrank ex Candolle.

Lavoisiera pulcherrima var. obtusifolia Barreto (1935b: 190) . nom. nud. = Lavoisiera pulcherrima Martius & Schrank ex Candolle.

Lavoisiera pulcherrima var. ovalifolia Barreto (1935b: 189) . nom. nud. = Lavoisiera pulcherrima Martius & Schrank ex Candolle.

Lavoisiera pulcherrima var. venusta Barreto (1935b: 189) . nom. nud. = Lavoisiera pulcherrima Martius & Schrank ex Candolle.

Lavoisiera reticulata Moricand (1847 : t. 99). According to Art. 52 of the I.C.N. (McNeill et al. 2012) this species is illegitimate because it was superfluous when published. It is to be rejected because it was based on the same type as Lavoisiera nervulosa Naudin , an epithet that should have been adopted.

Lavoisiera scabrella Naudin (1849: 226) . nom. nud. = Lavoisiera scaberula Naudin. This epithet was apparently a typographical error that inadvertently created a nude name. Cogniaux (1883: 226) cited it as a synonym of L. scaberula .

Lavoisiera suberosa Cogniaux (1896: 447) . TYPE:— BRAZIL. Goiás: Serra da Balisa, Sept. 1892 , E. G. H. Ule 766 (holotype: R-14787!; isotypes: BR-550887-n.v., online image!, R-14787a!). = Stenodon suberosus Naudin, (1844: 146) . TYPE:— BRAZIL. “In arenosis loci dicti vulgo Chapadaó, in Brasilia australi.”, [Cat. C1, n°546], A. Saint-Hilaire s.n. (holotype: P; isotype: P). See Martin & Cremers (2007). These two names are based on different types, thus the Cogniaux epithet is a heterotypic synonym of S. suberosus Naudin.

Lavoisiera vernicosa Barreto ex Pedersoli (1979 /1980: 25). TYPE:— BRAZIL. Minas Gerais: Serra do Cipó, 13 Aug. 1933, H. L. de Mello Barreto 325 (holotype: BHCB ex BHMH!) = Microlicia vernicosa (Barreto ex Pedersoli) A. B. Martins & Almeda in Almeda & Martins (2001: 6).

ACKNOWLEDGMENTS

We have benefitted from the assistance and support of many colleagues, friends, and organizations during this study. For help with many aspects of one or more field expeditions in Brazil we thank Renato Belinello, Rosana Romero, Renato Goldenberg, Jimi N. Nakajima, Peter W. Fritsch, Mary Beth Almeda, João L. M. Aranha, Paulo

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J.F. Guimarães, Orbélia R. Robinson, the late Donald Robinson, and Catherine Bush. We thank Darrell Ubick and Scott Serata for help in the Scanning Electron Microscope Laboratory at CAS; Fei-Mei Chuang for laboratory assistance with cytological preparations; Camille Nowell for preparing the distribution maps; Orbélia R. Robinson for assembling the initial database of collections from various herbaria; Gustavo Shimizu for help with collectors’ names; Cristiana Koschnitzke for providing bibliographic references and photographs of type specimens from the Museu Nacional, Rio de Janeiro (R); João Renato Stehmann for help with various bibliographic references; Renato Goldenberg for bibliographic and editorial assistance; Kathryn Jee for technical assistance with the SEM images; Becca Berezuk for technical help with the figures; Mary Beth Almeda for editorial assistance, and assembling Figures 1–13 and Appendix III; Thomas F. Daniel for assistance with the assembly of Figures 1–13 ; Roberta Brett for facilitating identification of the buprestid beetles that feed on the stamens of Lavoisiera macrocarpa ; Beth Guy for help with preliminary sketches for some of the line drawings; Ricardo Pacifico for new collections of rare species; Elizabeth Woodgyer for checking collections at K; João L. M. Aranha Renato Belinello, Gustavo Shimizu, and Deise Josely Pereira Gonçalves for special assistance on an expedition to Parque Estadual Pico do Itambé, and Don Mahoney and staff at the San Francisco Botanical Garden nursery for attempts to grow seeds of various species of Lavoisiera . Our special thanks go to Rogério Lupo who prepared the line drawings in Figures 15 , 17 , 19 , 21 , 23 , 25 27 , 29 , 31 , 32 , 34 , 36–38 , 40–54 , and 56–64; Sean V. Edgerton for Figures 30 and 35 ; and Jenny Speckels for Figure 55 . For selected digital images we are grateful to the following: Karina Fidanza for the images of Lavoisiera arachnoidea ( Figures 1D, E ); Mats Hjertson for an image of the holotype of L. imbricata ; Pin Flor do Brejo on Pinterest (www.picstopin.com) for the pink-flowered form of L. grandiflora ( Figure 3A ); Fernando B. Matos for the image of L. harleyi ( Figure 3D ); Gustavo Shimizu for images of L. vestita on Pico do Itambé ( Figures 5K, L ); Livia Echternacht for the image of the yellow-flowered form of L. glandulifera ( Figure 2L ) taken at Cachoeira Braúna (south of Serra do Cipó) and Ernesto Lemes for the two images of L. sampaioana ( Figures 5D, F ) taken at Lapinha da Serra (Serra do Cipó). We also thank the Instituto Brasileiro do Meio Ambiente e dos Recursos Naturais Renováveis (IBAMA) for granting permission to collect in protected areas of Brazil, the Universidade Estadual de Campinas, the California Academy of Sciences and their respective herbaria (UEC and CAS) for facilities and logistical support. Initial field support for this study was provided by the National Geographic Society (NGS Grant 6173–98 to Almeda). The M. Stanley Rundel Charitable Trust provided several subsequent grants to Almeda for field expeditions and production of all the line drawings. Martins’ postdoctoral year at CAS (19992000) was made possible by a grant from the Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP grant No. 99/0694-3 to Martins) as was a portion of her 2002 visit (FAPESP Grant No. 02/03676- 0 to Martins). Her 2002 and 2013 collaborative study visits to CAS were also supported by fellowship grants from the Lakeside Foundation to Almeda. We thank all of these funding agencies and Benjamin and Ruth Hammett for their outstanding support. Martins also extends thanks to the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq grant No. 301346/86-0) for supporting her research on Brazilian Melastomataceae . Lastly, we express our grateful appreciation to Rosana Romero and an anonymous reviewer for helpful reviews of the manuscript and the curators and staffs of the following herbaria for specimen loans, gifts, and/or special assistance during study visits: ALCB, B, BHCB, BHMH, BM, BR, C, CAS, CEPEC, COL, DS, ESA, F, FUEL, G, HUEFS, HUFU, IAC, K, LE, M, MBM, MICH, MO, NY, P, R, RB, S, SP, SPF, UB, UEC, US, W.

Figures

FIGURE 1. Images of Lavoisiera species. A. L. adamantium (Almeda et al. 8504). B–C. L. alba (Almeda et al. 9624). D–E. L. arachnoidea (unvouchered image). F. L. belinelloi (Almeda et al. 8532). G. L. bradeana (Almeda et al. 9718). H. L. canastrensis (Almeda et al. 7878). I. L. caryophyllea (Almeda et al. 9172). J–K. L. chamaepitys (Almeda et al. 8500). L. L. confertiflora (Almeda et al. 9725). Image credits: A–C, F–L. Frank Almeda; D–E. Karina Fidanza.

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 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 4. Images of Lavoisiera species. A−C. L. macrocarpa (Almeda et al. 9173). D. L. mucorifera (Almeda et al. 9607). E. L. pulchella (Almeda et al. 9830). F−G. L. pulcherrima (Almeda et al. 9666). H−I. L. punctata (Almeda 9709). J. L. rigida (Almeda et al. 9089). K. L. rigida (Almeda et al. 9604). L. L. rigida (Almeda et al. 9635). Image credits: A−L. Frank Almeda.

FIGURE 5. Images of Lavoisiera species. A−C. L. rundeliana (Almeda et al. 9667). D, F. L. sampaioana (unvouchered images). E. L. sampaioana (Almeda et al. 9198). G−H. L. senae (Almeda et al. 9186). I. L. subulata (Almeda et al. 8571). J. L. tetragona (Almeda et al. 7808). K−L. L. vestita (Aranha et al. 164). Image credits: A−C, E, G−J. Frank Almeda; D, F. Ernesto Lemes; K, L. Gustavo Shimizu.

FIGURE 6. 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. alba (Hatschbach 27282). C−D. L. caryophyllea (Tamashiro 25639). E−F. L. chamaepitys (Markgraf et al. 3437). Scale bars: A=150 µm, B=38 µm; C=380 µm, D=38 µm; E=231 µm, F=10 µm.

FIGURE 7. 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. confertiflora (Occhioni 4932). C−D. L. cordata (Glassman & Gomes 8059). E−F. L. crassifolia (Almeda et al. 7807). Scale bars: A=250 µm, B=60 µm; C=430 µm, D=100 µm; E=380 µm, F=38 µm.

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 9. 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. harleyi (Mori et al. 12455). C–F. L. imbricata (Irwin & Soderstrom 5769). Scale bars: A & C=380 µm; B, E & F=38 µm; D=30 µm.

FIGURE 10. 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. imbricata (Joly et al. 2257). C–F. L. mucorifera (Hatschbach 27445). Scale bars: A=380 µm, B=38 µm; C & E=176 µm; D & F=43 µm.

FIGURE 11. 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. nervulosa (Mori & Benton 13167). C–D. L. pulchella (Rossi & Catharino 1540). E–F. L. pulchella (Reitz & Klein 17495). Scale bars: A & C=300 µm; B=75 µm; D=30 µm; E=380 µm; F=38 µm.

FIGURE 12. SEM images of seeds for representative Lavoisiera species showing lateral symmetrical plane (left) and detail of testa cells and cell wall microrelief (right). A–D. L. pulcherrima (Arbo et al. 4339). E–F. L. quinquenervis (Heringer et al. 4061). Scale bars: A & C=380 µm; B & D=38 µm; E=300 µm, F=75 µm.

FIGURE 13. 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. rundeliana (Kameyama et al. 11255). C–D. L. senae (Anderson et al. 36197). E–F. L. subulata (Figuereido & Kubitski 83-22). Scale bars: A & C=250 µm; B=60 µm; D=30 µm; E=200 µm; F=50 µm.

FIGURE 15. Lavoisiera adamantium. A. Habit. B. Abaxial leaf surface. C. Adaxial bract surface. D. Flower. E. Hypanthium, calyx lobes and style with enlarged detail of adaxial calyx lobe apex (upper left) and detail of adaxial intercalycine sinus (upper right). F. Ovary cross-section. G. Petal with margin detail (right). H. Ventral view of antesepalous stamen. I. Ventral view of antepetalous stamen. J. Seed. (Based on: A–I, Araújo et al. 198, HUFU, UEC; J, Almeda et al. 8504, CAS, UEC.)

FIGURE 17. Lavoisiera alba. A. Habit. B. Adaxial leaf surface. C. Bud and subtending bracts. D. Flower. E. Hypanthium and calyx lobes. F. Ovary cross-section. G. Petal with enlarged margin detail (lower right). H. Ventral view of antesepalous stamen. I. Ventral view of antepetalous stamen. J. Seed. (Based on: A–J, Almeda et al. 8388, CAS, UEC.)

FIGURE 19. Lavoisiera angustifolia. A. Habit. B. Branch with terminal flower. C. Abaxial leaf surface. D. Adaxial leaf surface. E. Abaxial bract surface. F. Hypanthium and calyx lobes. G. Ovary cross section. H. Petal with margin detail (right). I. Ventral view of antesepalous stamen. J. Ventral view of antepetalous stamen. K. Seed. (Based on: A–K, Libon s.n., C.)

FIGURE 21. Lavoisiera arachnoidea. A. Habit. B. Abaxial leaf surface. C. Abaxial bract surface. D. Hypanthium and calyx lobes with enlarged surface detail (left) and enlarged adaxial intercalycine detail (right). E. Ovary cross-section. F. Petal. G. Ventralview of antesepalous stamens. H. Lateral view of antepetalous stamen. I. Seed. (Based on: A, Kirkbride 4588, US; B–I, Irwin et al. 15382, NY.)

FIGURE 23. Lavoisiera belinelloi. A. Habit. B. Adaxial leaf surface. C. Adaxial bract surface with enlarged detail of base (lower right). D. Flower. E. Hypanthium (at maturity) and calyx lobes with enlarged adaxial surface detail (left) and enlarged detail of calyx lobe apex (lower right). F. Ovary cross-section. G. Petal with margin detail (lower left). H. Lateral view of antesepalous stamen. I. Lateral view of antepetalous stamen. J. Seed. (Based on: A–J, Almeda et al. 8532, CAS, UEC.)

FIGURE 25. Lavoisiera canastrensis. A. Habit. B. Abaxial leaf surface. C. Abaxial bract surface. D. Flower. E. Hypanthium, calyx lobes and style with enlarged detail of intercalycine sinus (above). F. Ovary cross-section. G. Petal with enlarged detail of apex (above). H. Ventral view of antesepalous stamen. I. Ventral view of antepetalous stamen. J. Fruiting hypanthium. K. Capsule, dehiscing from the base. L. Seed. (Based on: A–L, Almeda et al. 7878, CAS, UEC.)

FIGURE 26. Distributions of Lavoisiera canastrensis, L. cogniauxana, L. crassifolia, L. pulchella, and L. rigida.

FIGURE 27. Lavoisiera caryophyllea. A. Habit. B. Adaxial surface of leaf-like flower-subtending bract. C. Flower. D. Hypanthium and calyx lobes. E. Ovary cross-section. F. Petal with enlarged detail of lower margin (left). G.Ventral view of antesepalous stamen. H. Ventral view of antepetalous stamen. I. Seed. (Based on: A–I, Almeda et al. 8567, CAS, UEC.)

FIGURE 28. Distributions of Lavoisiera caryophyllea, L. chamaepitys, L. minima, L. senae, and L. vestita.

FIGURE 29. Lavoisiera chamaepitys. A. Habit. B. Detached branchlet. C. Adaxial leaf surface. D. Abaxial bract surface. E. Flower. F. Hypanthium and calyx lobes. G. Ovary cross-section. H. Petal with enlarged detail of margin (lower right). I. Ventral view of antesepalous stamen. J. Ventral view of antepetalous stamen. K. Post-mature fruit. L. Persisting columella and placenta. M. Seed. (Based on: A–M, Almeda et al. 8500, CAS, UEC.)

FIGURE 30. Lavoisiera cogniauxana. A. Habit. B. Cauline nodes and internodes. C. Abaxial leaf surface. D. Flower. E. Petal and detail of margin (upper left). F. Stamens (ventral view), antepetalous (left) and antesepalous (right). G. Ovary cross-section. H. Hypanthium in fruit. I. Hypanthium at anthesis (petals removed) with enlargement of adaxial calyx lobe sinus (above). J. Seed. (Based on: A, C, E–G, I, J, Mello Barreto 8943, F, US; B, H, Glaziou 19270, LE; D, Furlan et al. CFSC 8459, SPF, US.)

FIGURE 31. Lavoisiera confertiflora. A. Habit. B. Abaxial (right) and adaxial (left) surfaces of the bract subtending the bud. C. Abaxial (right) and adaxial (left) surfaces of the bract subtending the fruit. D. Flower. E. Hypanthium and calyx lobes with enlarged detail of adaxial lobe sinus (above). F. Ovary cross-section. G. Petal. H. Ventral view of antesepalous fertile stamen. I. Antepetalous staminode. J. Seed. (Based on: A, B, D–J, Almeda et al. 8568, CAS, UEC; C, Almeda et al. 8585, CAS, UEC.)

FIGURE 32. Lavoisiera cordata. A. Habit. B. Adaxial leaf surface. C. Flower. D. Hypanthium and calyx lobes. E. Ovary crosssection. F. Petal with enlarged detail of lower margin (lower left). G. Lateral view of antesepalous stamen. H. Ventral view of antepetalous stamen. I. Seed. (Based on: A–I, Almeda et al. 8572, CAS, UEC.)

FIGURE 34. Lavoisiera crassifolia. A. Habit. B. Adaxial leaf surface. C. Flower. D. Hypanthium and calyx lobes. E. Ovary crosssection. F. Petal with enlarged detail of margin (lower left). G. Ventral view of antesepalous stamen. H. Ventral view of antepetalous stamen. I. Seed. (Based on: A, Almeda et al. 8442, CAS, UEC; B–I, Almeda et al. 8477, CAS, UEC.)

FIGURE 35. Lavoisiera daviesiana. A. Habit. B. Abaxial leaf surface with detail of revolute margin (left). C. Abaxial bract surface. D. Hypanthium and calyx lobes at anthesis (petals removed) with enlarged surface detail (lower right). E. Stamens (ventral view), antesepalous (left) and antepatalous (right). F. Flower. (A–F, Hatschbach et al. 66393, MBM.)

FIGURE 36. Lavoisiera firmula. A. Habit. B. Adaxial leaf surface. C. Hypanthium and calyx lobes with enlarged detail of adaxial lobe sinus (right). D. Ovary cross-section. E. Petal with enlarged detail of margin (lower left). F. Ventral view of antesepalous stamen. G. Ventral view of antepetalous stamen. H. Seed. (Based on: A–H, Stehmann & Sobral 1137, UEC.)

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 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 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.)

FIGURE 41. Lavoisiera harleyi. A. Habit. B. Abaxial leaf with enlarged surface detail (right). C. Adaxial bract surface. D. Hypanthium and calyx lobes. E. Ovary cross-section. F. Petal and margin detail (lower right). G. Ventral view of antesepalous stamen. H. Ventral view of antepetalous stamen. I. Seed. (Based on: A–I, Harley et al. 24484, UEC.)

FIGURE 42. Lavoisiera humilis. A. Habit. B. Adaxial leaf surface. C. Adaxial bract surface. D. Flower. E. Hypanthium and calyx lobes with enlarged calyx lobe margin detail (right). F. Ovary cross-section. G. Petal with enlarged margin detail (lower left). H. Ventral view of antesepalous stamen. I. Ventral view of antepetalous stamen. J. Seed. (Based on: A–J, Almeda et al. 8497, CAS, UEC.)

FIGURE 43. Lavoisiera imbricata. A. Habit. B. Abaxial leaf surface. C. Abaxial bract surface. D. Flower bud. E. Flower. F. Hypanthium, calyx lobes and bracts. G. Ovary cross-section. H. Petal with margin detail (lower right). I. Ventral view of antesepalous stamen. J. Ventral view of antepetalous stamen. K. Seed. (Based on: A–K, Almeda et al. 8401, CAS, UEC.)

FIGURE 44. Some of the named variants of Lavoisiera imbricata. A. Hypanthium and calyx lobes (above) and abaxial leaf surface (below); B. Hypanthium and calyx lobes (above) and abaxial leaf surface (below); C. Hypanthium and calyx lobes (above) and abaxial leaf surface (below); D. Hypanthium and calyx lobes (above) and abaxial leaf surface (below); E. Hypanthium and calyx lobes (above) and abaxial leaf surface (below). (Based on: A – L. pectinata form, Almeda et al. 8599, CAS, UEC; B – L. francavillana form, A.B. Joly et al. 2447, UEC; C – L. insignis form, Mello Barreto 318, UEC; D – L. bergii form, Carvalho et al. s.n., FUEL, UEC; E – L. compta form, Sciamarelli et al. 26540, UEC.)

FIGURE 45. Lavoisiera itambana. A. Habit with detail of abaxial leaf apex (right). B. Adaxial bract surface. C. Flower. D. Hypanthium and calyx lobes with enlargement of adaxial sinus (above). E. Ovary cross-section. F. Petal. G. Ventral view of antesepalous stamen. H. Ventral view of antepetalous stamen. I. Seed. (Based on: A–I, Zappi et al. CFCR 11267, UEC.)

FIGURE 46. Lavoisiera macrocarpa. A. Habit. B. Adaxial leaf surface. C. Flower. D. Hypanthium, calyx lobes, and ovary apex with style. E. Ovary cross-section. F. Petal with enlarged margin detail (right). G. Ventral view of antesepalous stamen. H. Ventral view of antepetalous stamen. I. Seed. (Based on: A–D, F–H, Almeda et al. 8545, CAS, UEC; E, I, Almeda et al. 8597, CAS, UEC.)

FIGURE 47. Lavoisiera mellobarretoi. A. Habit. B. Abaxial leaf surface. C. Abaxial flower bract surface. D. Abaxial bud bract surface. E. Flower. F. Hypanthium and calyx lobes. G. Ovary cross-section. H. Ventral view of antesepalous stamen. I. Ventralviewof antepetalous stamen. J. Seed. K. Petal, with enlargement of margin detail (right). (Based on: B–H, I, Pirani & Mello-Silva CFCR 10764, SPF; A, J, Cordeiro et al. CFCR 961, SPF.)

FIGURE 48. Lavoisiera minima. A. Habit. B. Abaxial leaf surface. C. Adaxial leaf surface. D. Abaxial bract surface. E. Hypanthium and calyx lobes. F. Ovary cross-section. G. Petal with enlargement of margin (lower left). H. Lateral view of antesepalous stamen. I. Ventral view of antepetalous stamen. J. Seed. (Based on: A–J, Menezes et al. CFCR 195, UEC, US.).

FIGURE 49. Lavoisiera mucorifera. A. Habit. B. Flower. C. Adaxial leaf surface. D. Adaxial bract with enlargement of surface detail (upper left). E. Hypanthium and calyx lobes. F. Ovary cross-section. G. Petal with enlargement of margin detail (right). H. Ventral view of antesepalous stamen. I. Ventral view of antepetalous stamen. J. Seed. (Based on: A–J, Almeda et al. 8464, CAS, UEC.)

FIGURE 50. Lavoisiera nervulosa. A. Habit. B. Abaxial leaf surface (from young branch) with enlargement of surface detail (upper right). C. Abaxial bract surface. D. Flower. E. Hypanthium and calyx lobes. F. Ovary cross-section. G. Petal. H. Ventral view of antesepalous stamen. I. Ventral view of antepetalous stamen. J. Seed. (Based on: A–J, Giulietti et al. CFCR 1553, CAS, UEC.)

FIGURE 51. Lavoisiera pohliana. A. Habit. B. Abaxial bract surface. C. Flower. D. Hypanthium and calyx lobes with enlargement of calyx lobe margin (upper right) and hypanthial surface (lower right). E. Ovary cross-section. F. Petal with enlargement of margin detail (lower right). G. Ventral view of antesepalous stamen. H. Ventral view of antepetalous stamen. I. Seed. (Based on: A–B, Koschnitzke & Matsumoto 35624, UEC; C–H, Irwin et al.13296, UEC; I, Romero et al. 5509, UEC.)

FIGURE 52. Lavoisiera pulchella. A. Habit. B. Abaxial leaf surface. C. Abaxial bract surface. D. Flower. E. Hypanthium and calyx lobes. F. Ovary cross-section. G. Petal with apex detail (above) and margin detail (lower left). H. Ventral view of antesepalous stamen. I. Ventral view of antepetalous stamen. J. Seed. (Based on: A–J, Shepherd & Andrade 6158, UEC.)

FIGURE 53. Lavoisiera pulcherrima. A. Habit. B. Adaxial leaf surface. C. Flower. D. Hypanthium and calyx lobes. E. Ovary crosssection. F. Ventral view of antesepalous stamen. G. Ventral view of antepetalous stamen. H. Old disintegrating hypanthia and capsule with persisting vascular strands. I. Seed. J. Petal with enlarged detail of apical margin (upper left) and lateral margin detail (lower right). (Based on: A, Romero & Nakajima 5948, UEC; B–J, Pirani et al. CFSC 10270, UEC.)

FIGURE 54. Lavoisiera punctata. A. Habit. B. Abaxial leaf surface with enlarged surface detail (upper left). C. Hypanthium, calyx lobes, and style with enlarged detail of calyx lobe base and intercalycine sinus (above). D. Ovary cross-section. E. Petal with enlarged detail of margin (right). F. Ventral view of antesepalous stamen (left), lateral view of antesepalous stamen (right). G. Ventral view of antepetalous stamen (left), lateral view of antepetalous stamen (right). H. Disintegrating mature hypanthium and enclosed capsule. I. Seed. (Based on: A, B, H, I, Semir 28670, UEC; C–G, Almeda et al. 9709, CAS, UEC.)

FIGURE 55. Lavoisiera quinquenervis. A. Habit. B. Abaxial leaf surface. C. Hypanthium, calyx lobes and bracteoles. D. Ovary cross-section. E. Petal. F. Ventral view of antesepalous stamen (left); lateral view of antepetalous stamen (right). G. Seeds. (Based on: A–G, Heringer et al. 4061, MO, NY, US.)

FIGURE 64. Lavoisiera vestita. A. Habit. B. Abaxial leaf surface (left), adaxial leaf surface (right) with surface detail enlargement (lower left). C. Flower. D. Hypanthium and calyx lobes with enlarged hypanthial surface detail (right) and enlarged adaxial surface of calyx lobe (above). E. Ovary cross-section. F. Petal with enlarged detail of apex (left) and lateral margin (right). G. Ventral view of antesepalous stamen. H. Ventral view of antepetalous stamen. I. Style (right) and stigma (left). (Based on: A–I, Aranha et al. 164, CAS, UEC.)