Malanea bahiensis Malanea megalantha Malanea macrophylla f. bahiensis Malanea bahiensis Malanea macrophylla f. cuneata Malanea macrophylla var. megalantha Malanea megalantha The genus Malanea Aubl. (Rubiaceae) in the Atlantic Forest of Brazil Roxo, Rafaela Sales Pereira Barbosa, Maria Regina de Vasconcellos Plant Ecology and Evolution 2023 2023-08-17 156 3 257 275 6R54S 33FF4757-6EB3-5CEE-9BC6-24B0B23B289B Bartl. ex Griseb. (Grisebach 1861: 337) Bartl. ex Griseb. (Grisebach 1861 Magnoliopsida Rubiaceae Malanea treatment-meta Plantae Malanea macrophylla Gentianales 0 257 Tracheophyta species macrophylla   Figs 2L-P , 3A-D   Malanea bahiensis Muell.Arg. (  Mueller1875: 453, 457) - Type: BRAZIL - Bahia• "Brasilia intratropica circa Bahia"; 1831; Blanchet 586; lectotype (designated by Steyermark 1965: 250): P n.v.; isolectotypes: G [G00237733], MO [MO 3152039], NY [132139].  Malanea megalanthaWernham ( Wernham 1913: 221) - Type: TOBAGO • s.l.; 21 Oct. 1910; Broadway 4024; holotype: BM [001008922]; isotypes: MO [MO 2532742], NY [132153], K [K000975676], GH [00096445], US [00138485].  Malanea macrophylla f. bahiensis( Muell.Arg.) Steyerm. ( Steyermark 1965: 250) - Type: same as for Malanea bahiensis.  Malanea macrophylla f. cuneataSteyerm. ( Steyermark 1965: 250) - Type: GUYANA [as British Guiana] • basin of Kuyuwini river (Essequibo tributary), about 150 miles from mouth; 21-26 Nov. 1937; Smith 2618; holotype: NY [132151]; isotypes: MO [MO 3150990], US [US00138484].  Malanea macrophylla var. megalantha(Wernham) Steyerm. ( Steyermark 1965: 250) - Type: same as for Malanea megalantha.   Type.   GUYANA• Guiana, "Gujana Anglica ad fl. Morocco"; Sep. 1843; Schomburgk 1484; first-step lectotype(designated by Steyermark 1965: 250), second-step lectotype( designated here): GOET [GOET008943]; isolectotype: F [V0296275F].   Figure 3. A- D.  Malanea macrophylla. A. Habit. B- C. Stipule. D. Adventitious roots. E- F.  Malanea martiana. E. Stipule. F. Fruit. Photos by Rafaela Roxo, 2022.  Description. Liana or scandent shrub 2-15 m tall; branchlets reddish-brown with circular lenticels, glabrous or glabrescent to pubescent when young, occasionally with adventitious roots. Stipules oblong to ovate or obovate, 7-20 x3-10 mm, round at apex, pubescent in central portion, glabrous near margins outside. Leaves with petioles 0.7-2.0 cm long, glabrescent; blades ovate, oblong-ovate or broadly elliptic, 6.2-18.1 x(3.3-)4.2-10 cm, acute to obtuse at base, acute to round at apex, margin flat, chartaceous to leathery, discolorous, glabrous above, occasionally lustrous, glabrous to pubescent below, trichomes adpressed; secondary veins 6-8 on each side of the midrib, impressed above, prominent below; tertiary veins reticulate. Inflorescence paniculate, 2.8-7 cm long, 1-2 per axil, glomerules 3-7, with (3-)5-9(-18) flowers each; peduncle (0.4-)0.5-1.7(-2.3) cm long, sericeous; bracts ovate-triangular or elliptic, 2-5 mm long, sericeous; bracteoles inconspicuous, sericeous. Flower buds sericeous. Calyx truncate or subtruncate, ca 2 mm long, glabrous or glabrescent. Corolla white, cream or yellowish, tube 2-4 mm long, externally strigose, internally lanate; lobes rounded to acute, 1.5-2 mm long, externally sericeous, internally villous. Stamens with filaments 0.8-1 mm long; anthers 0.8-1 mm long. Style 2.5-4.3 mm long. Fruits oblong-ellipsoid or cylindrical, 5-12 x2-5 mm, purple, glabrous; persistent calyx lobes glabrescent.  Distribution.   Malanea macrophyllais the most widely distributed species of  Malanea, ranging from Venezuela and the Guianas into the Brazilian Amazon, and along the coast to the Atlantic Forest, where it occurs from Rio Grande do Norte to Rio de Janeiro (Fig. 1B).  Habitat and ecology. It is usually found on the edge of lowland seasonal or moist forests, or in montane or submontane forests.  Phenology. Flowers in January to April, June, October to December; fruits from December to May.  Vernacular name.  Cipo-de-brejo(Alagoas, Rodrigues 1443), Arco-de-barri (Bahia, Cardoso 1591).  Etymology. The epithet macrophyllarefers to the large size of the leaves.  Preliminary IUCN conservation assessment. Least Concern (LC). The extent of occurrence (EOO) of this species in the Atlantic Forest domain was calculated as 586,645 km2, qualifying it for the category Least Concern (LC). The area of occupancy (AOO) was estimated as 336 km2, falling into the category Endangered (EN) ( Bachman et al. 2011; IUCN 2012, 2022), however, no data for material outside Brazil was included. Due to its wide distribution in the Atlantic Forest, and its occurrence in several other vegetation types and different phytogeographic domains and countries, we consider this species as Least Concern.  Notes.  Schomburgk (1848: 947) accepted  Malanea macrophyllaas a new species recognized by Bartling in mss (manuscript). However, the only information in Schomburgk'sFlora about the species is "An den ufern des Morocco. Bluehtim September. Strauch" (On the banks of the Morocco. Flowers in September. Shrub) and it is a nomen nudum. Grisebach (1861: 337) validly published the name and cited the collections by Guilding (St. Vincent), Crueger (Trinidad), and Schomburgk (Guyana), without citing the herbaria. Steyermark (1965)cited the Schomburgk collection from Guyana as the type, but did not mention the herbarium where it is deposited, providing a first-step lectotypification. The second-step lectotype designated here, Schomburgk 1484at GOET (GOET008943), has well-preserved leaves, inflorescences, and flowers and has handwritten annotations similar to those in the original publication.  Comment.   Malanea macrophyllais widely distributed and has leaf blades with variable shape, apex, base, and indument. Individuals collected under tall trees have much larger leaf blades, with a more acuminate apex and denser indumentum. Individuals collected at the edge of forests or in more open areas have smaller leaves, with a rounded apex and sparse indumentum. Steyermark (1965)recognized two varieties. Malanea macrophylla var. macrophyllaand M. macrophylla var. megalantha, distinguished by leaf blade type and indumentum, flower size, and distribution. In the typical variety, which occurs in Brazil, he considered three forms, distinguished by the density of the indumentum and the shape of the leaves. However, these three forms overlap and, in our opinion, may reflect different stages of vegetative development and/or specific environmental conditions. Taylor and Steyermark (2004)considered these differences not taxonomically informative. Delprete (2010)also treats the three forms as synonyms. We agree and did not consider as distinct the different forms proposed by Steyermark (1965).  Malanea macrophyllais distinguished from other species of the genus in the Atlantic Forest by the pubescent stipules round at apex, sericeous peduncle, and truncate or sub-truncate calyx.  Material examined.  BRAZIL-  Alagoas• ColoniaLeopoldina, Serra do Livramento; 7 Sep. 2010; fb., fr.; Chagas-Mota 8293; MAC • Flexeiras, Estacao Ecologicade Murici; 24 Mar. 2011; fb.; Chagas-Mota 10504; MAC • Ibateguara, Petropolis; 11 Dec. 2010; fb.; Chagas-Mota 9726; MAC • Ibateguara, Usina Grande; 14 Mar. 2010; fb., fr.; Cavalcante 418; MAC • Ibateguara, Usina Grande; 8°58 '32"S, 35°55 '48"W; 24 Jan. 2010; fr.; Lyra-Lemos 12837; MAC • Ibateguara, Usina Grande, Mata de Coimbra; 8°58 '32"S, 35°55 '46"W; 23 Jan. 2010; fr.; Lyra-Lemos 12811; MAC • Marechal Deodoro; 23 Feb. 1999; fr.; Bayman & Barros 211; JPB, MAC • Murici, Serra da Bananeira; 20 Jan. 2005; fr.;  Mendonca409; MAC • Murici, Serra do Ouro; 7 Sep. 2003; fb.;  Falcao42; MAC • Passo de Camaragibe, Fazenda Santa Justina; 9°12 '05"S, 36°08 '03"W; fb.; Lyra-Lemos 8186; MAC • Penedo, Fazenda Gameleira; 10°04 '00"S, 36°29 '17"W; 26 Dec. 1998; fb., fl.; Rodrigues 1443; JPB, MAC • Quebrangulo, REBIO Pedra Talhada; 9°23'°S, 36°44'W; 770 m; 14 May 2014; fb., fr.; Nusbaumer & Ammann 4054; JPB, UFP • Quebrangulo, REBIO Pedra Talhada; 9°24'S, 36°44'W; 740 m; 2 Dec. 2014; fb.; Nusbaumer & Cailliau 4475; MAC, JPB, UFP • Quebrangulo, REBIO Pedra Talhada; 9°23'S, 36°44'W; 770 m; 14 May 2014; fb., fr.; Nusbaumer & Ammann 4050; MAC, JPB, UFP • Sao Joseda Lage, BR 104; 11 Nov. 2010; fb.; Chagas-Mota9282; MAC • Sem municipio; 28 Oct. 1980; fb., fl.; Andrade-Lima 80-9762; IPA • Uniaodos Palmares, Fazenda Santo Antonio; 26 Nov. 1966; fb.; Andrade-Lima 66-4751; IPA. -  Bahia• Almadina, Serra do Corcovado/ SaoDomingos, trilha que daacesso a Areado Carvoeira; 14°41'S, 39°35'W; 25 Jan. 2014; fb.; Marinho 601; HUEFS, CEPEC • Amargosa, Serra do Timbo, Fazendo Mucuri, Riacho do Meio; 13°10 '20"S, 39°09 '34"W; 531 m; 25 Jan. 2007; fb.; Cardoso 1591; HUEFS, CEPEC • Amargosa, Serra do Timbo; 13°10 '20"S, 39°09 '34"W; 531 m; 26 Jan. 2007; fr.; Cardoso 1631; HUEFS, CEPEC • Amargosa, Serra do Timbo; 13°10'S, 39°09'W; 531 m; 17 Mar. 2007; fr.;  Paixao1093; HUEFS • Amargosa, Serra do Timbo; 13°10'S, 39°09'W, 28 Apr. 2007; fr.;  Paixao1160; HUEFS • Andarai, Cachoeira dos Funis; 2 Jan. 2017; fl.; Oliveira 126; HUEFS • Andarai, Chapada Diamantina; 12°48'S, 41°23'W; 2 Nov. 1999; fb.; Guedes 6969; ALCB • Apuarema, Concessaoda Rio Tinto; 13°53 '46"S, 39°41 '10"W; 680 m; 20 Nov. 2013; fb.; Aona 3276; RB • Belmonte, a 25kmSW da cidade; 6 Jan. 1981; fb.; Carvalho 441; CEPEC • Belmonte, a 30 kmSW da cidade; 8 Jan. 1981; fb.; Carvalho 484; CEPEC • Cabralia, Reserva BiologicaPau-Brasil; 5 Jan. 1972; fb., fl.; Eupunino 159; CEPEC • Camamu, Estrada para Cajaiba; 14°00 '49"S, 39°00 '43"W; 30 Nov. 2015; fb.; Marinho 1209; CEPEC • ElisioMedrado, ca 1 kmdo povoado de Monte Cruzeiro; 12°52 '10"S, 39°28 '18"W; 600 m; 2 Dec. 2004; fb.; Jardim 4283; HUEFS • ElisioMedrado, Serra da Jiboia; 12°51 '10"S, 39°28 '18"W; 650 m; 2 Dec. 2004; fb., fl.; Jardim 4283; HUEFS • Entre Rios, Areial; 12°15'S, 37°53'W; 7 Nov. 2012; fb.; Popovkin 1269; HUEFS • Entre Rios, Areial; 12°15'S, 37°53'W; 20 Jun. 2012; st.; Popovkin 1130; HUEFS • Entre Rios, Areial; 12°15'S, 37°53'W; 31 Jul. 2012; st.; Popovkin 1196; HUEFS • Eunapolis, trecho de Mata Atlanticaentre Eunapolise Porto Seguro; 16°26'S, 39°03'W; 29 Mar. 1995; fr.; Ferreira 83; ALCB • Igrapiuna, assentamento mirante; 13°54 '20"S, 39°20 '59"W; 18 Dec. 2001; fb., fr.; Loureiro 393; ALCB, CEPEC • Ibirapitanga, Serra do Papua, APA do Pratigi, descida da parcela 3 antes do corrego1; 13°54 '58"S, 39°27 '57"W; 13 Mar. 2013; fb.; Aona 2400; RB • Ilheus, Parque Municipal Mata da Esperanca, acesso pelo Banco da Vitoria; 14°46'S, 39°05'W; 8 Jan. 2003; fb., fl.; Jardim 4098; HUEFS, CEPEC • Itacare, Fazenda Capitao, 7.9 kma partir da juncaoentre BA 001 e a rodovia Itacare-Ubaitaba; 31 Mar. 2004; fr.; Fiaschi 2187; NY, RB • Itacare, proximoa boca do Rio de Contas; 14°17'S, 38°59'W; 28 Jan. 1977; fb.; Harley 18317; CEPEC, IPA, NY, RB • Itamaraju, Assentamento Pedra Bonita, aproximadamente 20 kmda rodovia vicinal de Itamaraju sentido Jucurucu; 16°50 '19"S, 39°37 '53"W; 500 m; 13 Feb. 2000; fl.; Borges 785; CEPEC, NY • Itamaraju, Assentamento Pedra Bonita, aproximadamente 20 kmda rodovia vicinal de Itamaraju sentido Jucurucu; 16°50 '19"S, 39°37 '53"W; 500 m; 13 Feb. 2000; fb.; Borges 808; CEPEC, NY, RB • Itanagra, Fazenda Brejo Verde; 9 Aug. 1975; fb.;  Gusmao145; ALCB • Itubera, 11 kmda estrada Itubera/Valenca; 5 Feb. 1983; fr.; Carvalho 1459; HUEFS, CEPEC • Itubera, km 11 da estrada Itubera/ Valenca; 5 Feb. 1983; fb.; Carvalho 1459; CEPEC • Jacobina, Cachoeira Itaitu; 11°20 '04"S, 40°30 '11"W; 610 m; 30 Mar. 1996; fb., fl.; Guedes 2647; HUEFS, ALCB • Jacobina, Piemonte da Diamantina, Barro Bananeira; 11°10 '49"S, 40°29 '53"W; 23 Nov. 2011; fb.; Loureiro 457; ALCB, CEPEC • Jaguaripe; 28 Feb. 2011; fr.; Ferreira 57; ALCB • Lencois; 12°34'S, 41°23'W; 30 Jul. 2006; 600 m; fr.; Couto 201; HUEFS • Lencois, Serra da Chapadinha; 12°27 '03"S, 41°25 '07"W; 600 m; 6 Feb. 1995; fr.; Guillietti 1614; ALCB, CEPEC • Lencois, Chapada Diamantina, Foz da Capivara; 9 Mar. 2000; fb.; Stradmann 0706; ALCB • Lencois, Chapada Diamantina; 26 Jan. 2000; fr.; Jardim 2526; CEPEC, NY • Lencoisde Una, proximo aponte sob rio Acuipe; 21 Feb. 2022; fr.; Roxo 43; JPB • Marau, Estrada que liga Ponta de Mutaa Marau; 6 Feb. 1979; fr.; Mori 11435; CEPEC, RB • Mucuri, Km 4 na estrada Mucuri/BR101; 3 Jan. 1991; fr.; Farney 2639; CEPEC, RB • Palmeiras, Morro do Pai Inacio; 12°28'S, 41°27'W; 1000 m; 12 Mar. 1997; fr.; Gasson 6193; HUEFS, ALCB, CEPEC • Palmeiras, Pai Inacio; 12°26 '37"S, 41°29 '16"W; 690 m; 1 Jul. 1995; fr.; Guedes 2116; ALCB, CEPEC • Palmeiras, Morro do Pai Inacio; 12°27 '20"S, 41°28 '15"W; 1080 m; 4 Feb. 1995; fr.; Guilietti 1531; ALCB, CEPEC, HUEFS • Palmeiras, Morro do Pai Inacio; 12°27 '16"S, 41°28 '17"W; 1040 m; 30 Dec. 1994; fb., fl.; Guilietti 1506; CEPEC • Piraido Norte; 13°46 '12"S, 39°19 '52"W; 6 Nov. 2014; fb.; Braga 41; JPB • Pontal dos Ilheus, saidapara Buererema; 17 May 1986; fr.;  Belem3571; CEPEC • Presidente Tancredo Neves, ponto 63; 13°23 '24"S, 39°18 '57"W; 1 Feb. 2011; fr.; Ferreira 45; ALCB • Porto Seguro, Parque Nacional Monte Pascoal, along trail leading to summit of Monte Pascoal; 150-400 m; 15°15 '53"S, 40°34 '29"W; 25 Mar. 1996; fr.; Thomas 11155; CEPEC, NY • Porto Seguro, Parque Nacional do Pau-Brasil, trilha do Mucununga, areaaberta proximo amata; 16°26 '06"S, 39°19 '30"W; 91 m; 19 Feb. 2022; fr.; Roxo 36; JPB • Porto Seguro, Parque Nacional do Pau-Brasil, trilha do Mucununga, proximo amargem de rio; 16°26 '06"S, 39°19 '30"W; 91 m; 19 Feb. 2022; fr.; Roxo 34; JPB • Porto Seguro, RPPN Manona; 20 Dec. 2005; fr.; Alves 102; ALCB, CEPEC • Porto Seguro, Reserva Biologicado Pau-Brasil, 17 kmao W da cidade, Rodovia para Eunapolis; 16°04 '00"S, 39°18 '33"W; 20 Jan. 1997; fb.; Harley 18110; CEPEC, RB • Porto Seguro, Reserva Florestal de Porto Seguro - CVRD; 16°26 '58"S, 39°03 '51"W; 11 Jan. 90; st.; Folli 1044; NY • Salvador, Biribeira-Paralela; 8 Nov. 1994; fb.; Bandeira s.n.; ALCB • Salvador, Jardim Botanicode Salvador; 12°55 '37"S, 39°03 '51"W; 6 Feb. 2004; fr.; Queiroz 353; HUEFS, ALCB • Salvador, regiaometropolitana, barragem do Cobre; 12°58'S, 38°30'W; 16 Apr. 2012; fr.; Queiroz 5312; ALCB • Santa Cruz Cabralia, Santo Andre, proximoao estuariodo Rio Acuba; 16°14'S, 39°02'W; 2 Nov. 2003; fb.; Guedes 10624; ALCB • SaoMiguel das Matas, Fazenda Engenho da Lama, ca 4,5 km ao S da cidade; 13°02 '49"S, 39°25 '56"W; 400 m; 24 Feb. 2000; fb., fr.; Jardim 2884; ALCB, CEPEC, HUEFS, NY • Serra das Lontras, ca de 7 kmno ramal que liga o distrito de Itatingui aSerra; 15°12 '09"S, 39°24 '28"W; 31 Mar. 2006; fb.;  Paixao927; CEPEC, NY • Ubaira, arredores de Tres Bracos; Jan. 1991; fb., fl.; Sobral 6702; SP • Una, Mata litoranea; 19 May 1965; fb.;  Belem1062; CEPEC • Una, proximo aReserva BiologicaUna; 16°28 '10"S, 39°16 '46"W; 21 Feb. 2022; fr.; Roxo 41; JPB • Una, Reserva Biologicado Mico-leao(IBAMA); 8-12 Mar. 1993; fb.; Amorim 1130; CEPEC, NY • Una, Reserva Biologicado Mico-leao(IBAMA); 26 Apr. 1994; fl.; Carvalho 4515; CEPEC, NY • Una, Reserva Biologicado Mico-leao(IBAMA); 3 Mar. 1998; fr.;  Sant'Ana640; CEPEC • Una, Reserva Biologicado Mico-leao(IBAMA), 15°06 '58"S, 39°02 '11"W; 20 Mar. 1999; fr.; Jardim 2068; CEPEC • Urucuca, estrada de Itacarepara Ubaitaba; 14°29 '59"S, 39°06 '54"W; 380 m; Apr. 2004; st.; Martini sn.; CEPEC • Urucuca, Parque Estadual Serra do Conduru ca 11.5 kmde Serra Grande em direcaoa Urucuca, com 1kmdo ramal a esquerda; 18 May 2000; fr.;  Sant'Ana855; CEPEC. -   EspiritoSanto• Linhares, Reserva Natural Vale do Rio Doce (Particular). Reserva de Linhares; 31 Jan. 1972; Sucre 8338; RB • Santa Teresa; 19°58'S, 40°32'W; 600-900 m; 18 Nov. 1991; fr.; Pizziolo 363; JPB • Santa Teresa; 4 Mar. 1997; fb., fl.; Boudet Fernandes 3195; JPB • Santa Teresa; 10 Mar. 1999; fl.; Kollmann 2109; JPB • Santa Teresa; 30 Mar. 1999; fr.; Kollmann 2320; JPB. -  Minas Gerais• Uberlandia, Parque do Sabia; 22 Jun. 1995; fr.; Nakagima 1644; HUFU • Uberlandia, TangaraCountry Club; 18°55 '07"S, 48°16 '37"W; 28 Nov. 1995; fl.; Schiavini 410; HUFU. -   Paraiba• Bananeiras, Reserva Estadual Goiamunduba; 6°44 '28"S, 35°36 '22"W; 800 m; 30 Jun 2021; st.; Roxo 03; JPB • Conde, APA Tambaba, Loteamento Enseada de Jacuma; 7°19 '24"S, 34°48 '47"W; 4 Dec. 2009; fr.;  Araujo219; JPB • EspiritoSanto, Engenho SaoPaulo; 25 Nov. 1968; fl.; Andrade-Lima 68-5475; IPA • JoaoPessoa, Praia da Penha, Rio Aratu; 8 Nov. 1992; fb., fl., fr.; Moura 892; JPB • Mamanguape; 6°07 '22"S, 35°18 '20"W; 10 Feb. 2012; fb., fr.; Thomas 15655; JPB • Mamanguape, Estacao EcologicaMamanguape; 17 Aug. 1989; fb.; Felix 30; JPB. -  Pernambuco• Barreiros, Mata do Porto; 4 Jan. 2000; fb., fl.; Lucena 784; PEUFR • Bonito; s.d.; fb., fr.; Silva 444; PEUFR • Bonito, Reserva Municipal de Bonito; 08°29 '40"S, 35°41 '45"W; 444 m; 6 Mar. 1996; fr.; Campelo 92; PEUFR • Bonito, Reserva Municipal de Bonito; 08°29 '40"S, 35°41 '45"W; 444 m; 6 Mar. 1996; fr.; Marcon 127; PEUFR, IPA, UFP • Bonito, Reserva Municipal de Bonito; 15 Mar. 1995; fr; Sales de Melo 29; PEUFR, IPA, UFP • Escada, Engenho Conceicao; 22 Nov. 1967; fb.; Costa 143-67; IPA • Gravata, Fazenda Harmonia; 10 Oct. 1970; fb., fl.; Andrade-Lima 70-6022; IPA • Igarassu, Mata de Piedade; 7°50 '11"S, 35°00 '28"W; 17 Dec. 2009; fb.;  Garcia-Gonzalez1374; CEPEC, JPB, HUEFS, NY UFP • Igarassu, Mata de Piedade; 7°49 '48"S, 35°59 '55"W; 2 Mar. 2010; fr.;  Garcia-Gonzalez1445; JPB, IPA; CEPEC, UFP • Igarassu, Mata de Piedade, areaprotegida propriedade da Usina Sao Jose; 7°49 '38.5"S, 35°0 '16.1"W; 91 m; 12 Aug. 2009; fr.;  Garcia-Gonzalez1070; UFP • Lagoa dos Gatos, RPPN Pedra D'Anta; 7 Jun. 2011; fb.; Pessoa 647; JPB • Maraial, Serra do Urubu; 20 Apr. 1994; fr.; Miranda 1609; IPA • Recife, Jardim Botanicado Curado; 13 Sep. 1994; fb.; Rita Pereira s.n.; IPA • Rio Formoso, Mata do Ribeira; 6 Mar. 2002; fr; Lira 314; PEUFR • Rio Formoso, Mata do Ribeira, 6 Mar. 2002; fb.; Lira 306; PEUFR • Rio Formoso, Mata do Ribeira; 7 Mar. 2002; fb.; Lira 458; PEUFR • SaoVicente Ferrer, Mata do Estado; 7°35'S, 35°30'W; 600 m; 12 Nov. 1995; fb., fl.;  Laurencio237; NY, PEUFR • SaoVicente Ferrer, Mata do Estado; 07°35'S, 35°30'W; 600 m 13 Jan. 2000; fb.; Ferraz 810; PEUFR • Serinhaem, Usina Trapiche; 8°33 '14"S, 35°08 '48"W; 110 m; 21 Dec. 2010; fb.; Melo 713; JPB, UFP • Tamandare, Mata do conde, Fazenda estivas; 21 Jan. 2000; fb., fr.; Oliveira 541; PEUFR. -  Rio de Janeiro• Carapebus, Fazenda Lazaro; 23 Jun. 1994; fb.; Bovini 367; JPB, RB • Carapebus, Restinga de Carapebus. Fazenda Sao Lazaro; 23 Apr. 1994; fb.; Bovini 367; RB • Rio das Ostras, Mar do Norte, no final da Rua Ivan Lins; 19 May 2005; st.; Marquete 3632; RB • Nova Friburgo, Serra de Friburgo; 22°28 '19"S, 42°53 '11"W; 15 Jul. 1975; fr.; Occhioni 7620; RFA • Silva Jardim, Reserva Biologicade Pocodas Antes, trilha do Morro do Calcario; 22°30'S, 42°15'W; 5 Mar. 1993; fr.; Barreto 29; CEPEC, RB • Silva Jardim, Reserva Biologicade Pocodas Antas, trilha do Morro do Calcario, ponto 1000; 22°00'S, 42°00'W; 9 Jan. 1993; fb.; Lima 4585; NY, RB • Silva Jardim, Reserva Biologica Pocodas Antas, Trilha do Morro do Calcario; 22°30'S, 42°15'W; 8 Jan. 1993; fb.; Sylvestre 828; RB. -  Rio Grande do Norte• Canguaretama; 6°31 '52"S, 35°10 '33"W; 69 m; 20 Sep. 2014; inf.; Jardim 6728; UFRN • Macaiba, Escola Agricolade Jundiai; 5°53 '35"S, 35°21 '01"W; 54 m; 10 Feb. 2012; fr.; Costa-Lima 628; UFRN, UFP • Porto Velho, Areade ProtecaoAmbiental Piquiri-Una; 6°23 '36"S, 35°15 '22"W; 26 Feb. 2019; fr; Roque 2576; CEPEC. -  Sergipe• Campo do Brito, Povoado Tapera da Serra; 14 Apr. 1982; fr.; Viana 408; IPA • Campo do Brito, Tapera da Serra; 11 Nov. 1981; fb., fl.; Carneiro 158; IPA • Capela, RVS Mata do Junco; 10°31 '10"S, 37°03 '05"W; 20 Apr. 2012; fr.; Gomes 440; JPB • Capela, RVS Mata do Junco; 10°32'S, 37°03'W; 15 Feb. 2012; fr.; Chagas 154; JPB • Capela, RVS Mata do Junco, bica do estreito Mata Atlantica; 18 Oct. 2012; fb.; Costa-Silva 60; UFP • Santa Luzia de Itanhy; 11°22 '31"S, 37°25 '24"W; 9 Sep. 2011; fb., fl.; Gomes 192; JPB, HUEFS, RB • Santa Luiza do Itanhy, ca 2.5 kmdo Distrito do Crato, na estrada para Santa Luiza do Itanhi; 27 Nov. 1993; fb.; Amorim 1473; CEPEC • Santa Luzia do Itanhy, RPPN Mata do Crasto; 11°23 '36"S, 37°25 '15"W; 3 Mar. 2011; fr.; Gomes 67; JPB • Santa Luzia do Itanhy, RPPN Mata do Crasto; 11°22 '01"S, 37°25 '01"W; 2-20 m; 2 Mar. 2011; fb., fr.; Thomas 15276; JPB • Santa Luzia do Itanhy, RPPN Mata do Crasto; 11°23 '33"S, 37°25 '07"W; 26 Jan. 2011; fr.; Prata 2576; JPB, NY.  Additional material examined from outside the Atlantic Forest domain.   BRAZIL-   Amapa•   Calcoene, BR156 entre  Calcoenee Rio AmapaGrande, ca  5 kmS de Calcoene; 2°12'N, 50°55'W; 12 Dec. 1984; fb.; Rabelo 3005; NY • Rio Araguari, at mouth of Anicahy, above Camp 14; 8 Oct. 1961; fb.; Pires 51555; NY. -   Amazonas• Santa Isabel do Rio Negro; 0°25'S, 64°40'W; 9 Oct. 1987; fb.; Cid Ferreira 9314; RB, NY. -   Distrito Federal•  Brasilia, ca 10 kmof  Brasilia, proximo de Sobradinho; 15°43 '09"S, 47°51 '25"W; 1 Oct. 1965; fb.; Irwin 8854; NY • s.l.; 5 Oct. 1965; fl.; Irwin 8970; RB. -    Goias• Mineiros, Nascente do RibeiraoGloria,  200-300 mafora da cerca do Parque Nacional das Emas; 17°54'S, 52°55'W; 2 Oct. 1998; fr.; Delprete 6809; NY • Mineiros, Parque Nacional das Emas, corrego Cabecerao; 18°2'S, 52°52 '12"W; 1 Oct. 1998; fb.; Delprete 6780; NY • Pirenopolis, Cachoeira Santa Maria- RPPM; 15°49 '23"S, 48°54 '25"W;  1150 m; 23 Jun. 2001; fr.;  Marquete3157; RB •  Pirenopolis, Chapada dos Veadeiros. Fazenda Pai Jose; 27 Oct. 1994; fb.;   Mendonca2229; RB • Planaltina; 20 Jul. 1966; fb.;  Irwin18296; RB • Serra do Caiapo,  40 kmSouth of Caiaponia; 26 Oct. 1964; fr.;  Prance59673; NY. -    Maranhao•  SaoLuiz, Ilha de SaoLuiz; 2°57'S, 44°20'W; Feb.-Mar. 1939; fb.; Lemos Froes11737; NY. -   Mato Grosso• Northca 2 km. E. of base camp; 19 Oct. 1968; fr.;  Harley10728; NY, RB • Sinop, Rd.to Fazenda Atlantica, 10-14 kmW of BR 163 and 8 kmN of rd to  Portodos Gauchos; 11°06'S, 55°57'W; 25 Sep. 1985; fr.; Thomas 4061; NY. -    Para•  Belem; 24 Aug. 1922; fb.;  Duckes.n.; RB • Ilha do Marajo, Rio Anajas; 0°59'S, 49°55'W; 2 Nov. 1987; fr.; Prance 30227; NY • Rio Mocooes,  1 kmabove Anajas; 0°58'S, 49°56'W; 7 Nov. 1987; fr.; Beck 366; NY. -   Roraima•  Caracarai, PARNA do  Virua, Gradedo PPBio, trilha L01, segment 1400-1450; 1°29 '12"N, 61°0 '15"W; 19 Sep. 2012; fb.; Perdiz 1600; RB. 4178097311 1843-09 GUYANA Guiana 1 lectotype 4178097312 1961-10-08 1984-12-12 1961-10-08 Calcoene, BR BRAZIL Calcoene 2.2 Rio Amapa Grande 1307 -50.916668 Calcoene 1 Amapa 4178097303 1987-10-09 Brazil -0.41666666 Santa Isabel do Rio Negro 1308 -64.666664 1 Amazonas 4178097317 1965-10-01 1965-10-05 1965-10-01 de Sobradinho Brazil -15.719166 Brasilia 21 -47.85694 Brasilia 1 Distrito Federal 4178097319 1964-10-26 2001-06-23 1964-10-26 Parque Nacional das Emas, Marquete, Pirenopolis, Chapada dos Veadeiros. Fazenda Pai Jose, Mendonca, Planaltina, Irwin, Prance Brazil Mineiros 250 -15.823055 Serra do Caiapo 21 -48.906944 Nascente do Ribeirao Gloria 1 Goias 4178097306 1939-02 1939-03-31 1939-02 Brazil -2.95 Ilha de Sao Luiz 1307 -44.333332 Sao Luiz 1 Maranhao 4178097307 1968-10-19 1985-09-25 1968-10-19 Porto dos Gauchos Brazil North -11.1 Sinop 1296 -55.95 Harley 1 Mato Grosso 4178097322 1922-08-24 1987-11-07 1922-08-24 Brazil Belem -0.96666664 Ilha do Marajo 1308 -49.933334 Ducke 1 Para 4178097321 2012-09-19 Brazil Caracarai 1.4866667 Grade 21 -61.004166 Virua 1 Roraima