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Research Article
The genus Malanea Aubl. (Rubiaceae) in the Atlantic Forest of Brazil
expand article infoRafaela Sales Pereira Roxo‡, Maria Regina de Vasconcellos Barbosa§
‡ Federal University of Pernambuco, Recife, Brazil
§ Federal University of Paraíba, João Pessoa, Brazil

Corresponding author: Rafaela Sales Pereira Roxo ( rafaela.roxo95@gmail.com )

Academic editor: Brecht Verstraete
© 2023 Rafaela Sales Pereira Roxo, Maria Regina de Vasconcellos Barbosa.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Roxo RSP, Barbosa MRV (2023) The genus Malanea Aubl. (Rubiaceae) in the Atlantic Forest of Brazil. Plant Ecology and Evolution 156(3): 257-275. https://doi.org/10.5091/plecevo.103395
Open Access

Abstract

Background and aims – Malanea (Rubiaceae) is a Neotropical genus of ca 40 species, 15 of which occur in Brazil, with nine found in the Atlantic Forest domain. This study aims to present a taxonomic treatment of the species occurring in the Atlantic Forest domain.

Material and methods – We studied collections deposited in 15 Brazilian herbaria, ALCB, BHCB, BHZB, CEPEC, HUEFS, HUFU, IPA, JPB, MAC, OUPR, PEUFR, RFA, RB, UFP, and UFRN, and collections in NY were studied online. Specimens were identified using protologues and digital images of types available online.

Key results – Of the nine species of Malanea previously recognized in the Atlantic Forest domain, we accepted seven, six of which are endemic. Malanea macrophylla has a wide distribution also occurring in other Brazilian phytogeographic domains, and adjacent countries. We expanded the distribution of four species and identified Bahia as the possible centre of diversity for the group in the domain. In addition, an identification key to the species, detailed descriptions, taxonomic notes, typifications, illustrations, maps of geographic distribution, and assessments of conservation status are provided. The names Malanea evenosa, M. forsteronioides, M. macrophylla, M. martiana, and M. spicata are here typified. The calculated conservation status revealed three Endangered species, two Vulnerable ones, M. forsteronioides as Near Threatened, and M. macrophylla with a status of Least Concern.

Keywords

diversity, flora, Guettardeae, neotropics, taxonomy

Introduction

Malanea Aubl. includes ca 40 Neotropical species that occur from eastern Brazil to northern Bolivia (Taylor 2022). In Brazil, 15 species are currently recognized, present in the Amazon, Cerrado, and Atlantic Forest phytogeographic domains (Barbosa 2020). These species mainly occur in moist or seasonal forests, in areas close to bodies of water. In the Atlantic Forest, nine species are recognized, six of which are endemic to this domain, representing 40% of Brazilian species (Barbosa 2020).

According to molecular phylogenies (Bremer and Eriksson 2009; Pessoa 2016), Malanea is a genus in the tribe Guettardeae whose monophyly is not confirmed. Its representatives are lianas or climbing shrubs with a conspicuous terminal pair of stipules on each branchlet, axillary, paniculate or spiciform inflorescences, sessile flowers, turbinate hypanthium, corolla pubescent in the mouth, and ellipsoid, fusiform or cylindrical drupaceous fruits (Barbosa 2020).

The taxonomic treatment of Müller (1881) is the most comprehensive one of Malanea species present in Brazil. In a previous paper, Müller (1875), based on inflorescence type, divided the genus into two sections: Eumalanea (= Malanea), with paniculate inflorescence (6 spp.), and Pseudomalanea, with cymose inflorescence (2 spp.). Later, he transferred the species of section Pseudomalanea to Chomelia Jacq (Müller 1881).

In the 20th and 21st centuries, the Brazilian species of Malanea were treated only in parts, with focus on other South American countries (Hochreutiner 1910; Standley 1940; Steyermark 1965; Taylor and Steyermark 2004; Amaya 2008; Amaya and Delprete 2010), or in regional treatments in Brazil (Delprete et al. 2005; Barbosa 2007; Taylor et al. 2007; Delprete 2010). Worthy of mentioning are Kirkbride (1997) and Amaya and Popovkin (2012) for the description of new species for the Atlantic Forest in north-eastern Brazil, and Barbosa (2020) for the monograph of the genus for the online Flora of Brazil.

Despite the high number of Malanea species endemic to the Atlantic Forest domain, there is no detailed taxonomic treatment of the species that occur there. This paper presents a modern taxonomic treatment of Malanea in the Atlantic Forest, with an identification key to the species, detailed descriptions, taxonomic notes, typifications, illustrations, maps of geographic distribution, and conservation status assessments for each species.

Material And Methods

The Atlantic Forest domain extends more than 3,300 km along the Brazilian coast (Morellato and Haddad 2000), from the state of Rio Grande do Norte to the state of Rio Grande do Sul. It is home to native forest formations and associated ecosystems that vary with altitude, longitude, precipitation, and temperature (Moura 2006; Ribeiro et al. 2009; Stehmann 2009). Because of its latitudinal extension and climate variation the Atlantic Forest presents high richness and endemism (Morellato and Haddad 2000).

We define all types of naturally occurring forests of the coastal zone and all restinga and tabuleiro formations, according to Thomas and Barbosa (2008). The forest types were classified as Tropical Moist Forest, Mixed Moist Forest (a subtropical forest, also known as Araucaria Forest), and Seasonal Forest (IBGE 2012). We also included enclaves of submontane and montane forests in the north-east (Brasil 2006).

The present study is based on the analysis of 269 Malanea specimens collected in the Atlantic Forest and deposited in ALCB, BHCB, BHZB, CEPEC, HUFU, HUEFS, IPA, JPB, MAC, NY*, OUPR, PEUFR, RFA, RB, UFP, and UFRN (Thiers 2023). The collections at NY (marked with an asterisk) were consulted online. In addition, collecting expeditions took place between August 2021 and February 2022, in areas that had no records or were poorly collected. The material was processed following the usual herbarium techniques (Mori et al. 1985) and deposited in the JPB Herbarium.

The species were identified using the protologues, the analysis of photos of nomenclatural types, and regional taxonomic treatments for Brazil (Delprete et al. 2005; Barbosa 2007, 2020; Taylor et al. 2007; Delprete 2010). Additional types and specimens were analysed virtually through JSTOR (https://plants.jstor.org), JACQ (https://www.jacq.org), Natural History Museum (https://data.nhm.ac.uk), REFLORA (http://floradobrasil.jbrj.gov.br/reflora), and SpeciesLink (https://specieslink.net/search).

The morphological terms used in the descriptions followed Radford et al. (1974) and Harris and Harris (2001). The specimens examined are listed in alphabetical order by state and location and, when necessary, were supplemented with additional relevant samples from locations outside the Atlantic Forest domain.

Geographical distribution and phenological data in the Atlantic Forest domain were obtained from specialized literature and from specimen labels. Georeferenced data were imported into GeoCat (Bachman et al. 2011) to estimate the Extent of Occurrence (EOO) and Area of Occupancy (AOO) for each species in order to generate an assessment of the risk of extinction following the Red List categories and criteria of the IUCN (2012, 2022). Distribution maps were created using QGIS v.3.14 (QGIS Development Team 2020). All the images were edited using Photofilter v.11.4.1 (http://www.photofiltre.com).

Results

Seven species of Malanea are recognized in this study as occurring in the Atlantic Forest domain: M. evenosa Müll.Arg., M. forsteronioides Müll.Arg., M. harleyi J.H.Kirkbr., M. macrophylla Bartl. ex Griseb., M. martiana Müll.Arg., M. revolutifolia A.Amaya & Popovkin, and M. spicata Müll.Arg. They are found mainly along the edges of moist or seasonal forests, or in areas with more sunlight close to bodies of water.

Malanea sarmentosa Aubl. and M. obovata Hochr., also cited for the Atlantic Forest by Barbosa (2020), occur only in the Amazon. Malanea sarmentosa registered as occurring in Bahia, has no voucher collected in that state. The collection Thomas et al. 13443 (NY, CEPEC), from Bahia, identified as M. sarmentosa in 2005, had its identification corrected by us to Malanea revolutifolia, a species described in 2012. A similar misidentification occurred with M. obovata. The collection Belém 3117 (CEPEC) was misidentified and is actually M. spicata. The distinction between these two species is discussed in the comments under M. spicata.

The geographic distribution (Fig. 1) of four species is being extended in this treatment: M. evenosa also occurs in Minas Gerais, M. martiana in Pernambuco and Alagoas, M. revolutifolia in Minas Gerais, and M. spicata in Bahia. Except for M. macrophylla, the other six species studied are recognized as endemic to the Atlantic Forest. Malanea macrophylla has a wide distribution and occurs in other Brazilian phytogeographic domains and adjacent countries. Bahia is the state with the greatest richness, with six species of Malanea. It is the centre of diversity for the genus in the Atlantic Forest. At the extremes of the domain, a gradual decrease in richness was observed. In the northern portion, where the climate is seasonal and dry, only Malanea macrophylla and M. martiana were registered. In the southern part of the range, where the climate is subtropical, only M. forsteronioides occurs.

Figure 1. 

Geographical distribution of the species of Malanea in Atlantic Forest based on georeferenced localities of herbarium specimens. A. M. evenosa (yellow circles) and M. forsteronioides (pink diamonds). B. M. macrophylla (blue circles). C. M. harleyi (turquoise diamonds) and M. martiana (orange circles). D. M. revolutifolia (red diamonds) and M. spicata (green circles).

The calculated conservation status revealed three species that deserve Endangered status (M. evenosa, M. harleyi, and M. spicata), two that should be considered Vulnerable (M. martiana and M. revolutifolia), one that deserves to be Near Threatened (M. forsteronioides), and one that should be considered Least Concern (M. macrophylla). These results reflect the importance and need of floristic and taxonomic studies to support the development of conservation strategies in one of the world’s biodiversity hotspots.

Taxonomic Treatment

Malanea Aubl. (Aublet 1775: 106, t. 41)

Cunninghamia Schreb. (Schreber 1791: 789) – Type species: Cunninghamia sarmentosa (Aubl.) Forsyth f.

Type species

Malanea sarmentosa Aubl.

Description

Lianas or scandent shrubs, unarmed; branchlets cylindrical, with lenticels or not, the young ones compressed or coplanar. Stipules entire, interpetiolar, free, caducous after 1 to 2 distalmost nodes, membranaceous, apex round, acute or acuminate. Leaves simple, opposite, petiolate; blade oblong, ovate, obovate, or elliptical to narrow-elliptic, margin entire, flat or revolute, membranaceous to leathery, glabrous, sparsely pubescent to adpressed pubescent or tomentose to hirsute; venation brochidodromous, secondary veins impressed or sulcate above, prominent or impressed below, tertiary veins reticulate or inconspicuous, with or without axillary hairy domatia. Inflorescence axillary, pedunculate, paniculate or spiciform, bracts 2 at base. Flowers 4-merous, bisexual, heterostylous, sessile, actinomorphic; subtended by 3 bracteoles; hypanthium turbinate. Calyx lobed or dentate, or truncate to subtruncate, glabrous to glabrescent or tomentose. Corolla funnelform, valvate in bud, externally glabrous or sericeous, internally villous or lanate at mouth, white, cream or yellowish. Stamens free at base, exserted, inserted near the corolla mouth; anthers linear, dorsifixed, opening by introrse longitudinal splits. Style cylindrical, glabrous, 2-lobed; nectariferous disc present. Ovary 2-locular, ovule 1 per locule, apically attached, axile, pendulous. Fruits drupaceous, cylindrical, ellipsoid, fusiform or oblong, glabrous to glabrescent or hirsute to tomentose, reddish, purple or black when ripe, with persistent calyx lobes; pyrenes 1–2.

Key to identify the Malanea species present in the Atlantic Forest

1. Stipules round at the apex 2
– Stipules acute to acuminate at the apex 3
2. Stipules glabrous outside; leaves concolorous, glabrous, tertiary veins inconspicuous; calyx lobed; corolla externally glabrous M. harleyi
– Stipules pubescent on the central portion outside; leaves discolorous, glabrous to pubescent, tertiary veins reticulate and evident; calyx truncate or subtruncate; corolla externally strigose M. macrophylla
3. Stipules, petiole, and leaves glabrous; leaves membranaceous to chartaceous, concolorous; calyx subtruncate M. spicata
– Stipules, petiole, and leaves not glabrous; leaves chartaceous to leathery, discolorous; calyx dentate or lobed 4
4. Leaves strongly revolute, tomentose to hirsute below; secondary veins 7–11 on each side of the midrib, grooved adaxially; inflorescence tomentose; calyx deeply lobed M. revolutifolia
– Leaves slightly revolute or flat, sparsely pubescent or tomentose below, secondary veins 4–6 on each side of the midrib, impressed or inconspicuous adaxially; inflorescence hirsute, sericeous, or strigose; calyx dentate or slightly lobed 5
5. Petiole glabrescent; leaf blades cuneate to attenuate at the base, lustrous above; inflorescence paniculate, longer than 5 cm; inflorescence bracts triangular; calyx dentate; corolla yellow, lobes villous M. martiana
– Petiole pubescent to sericeous or hirsute-hispid; leaf blades obtuse to rounded at the base, not lustrous above; inflorescence spiciform, up to 5 cm long; inflorescence bracts lanceolate to narrow-elliptic; calyx slightly lobed; corolla white to cream, lobes lanate 6
6. Stipules sericeous outside; petiole pubescent to sericeous; leaves chartaceous to leathery, margin slightly revolute, secondary veins inconspicuous above and impressed below; calyx lobes equal; corolla tube up to 2 mm long; fruits oblong M. evenosa
– Stipules tomentose outside; petiole hirsute-hispid; leaves chartaceous, margin flat, secondary veins impressed above and prominent below; calyx lobes unequal; corolla tube longer than 2 mm; fruits fusiform M. forsteronioides

Malanea evenosa Müll.Arg. (Müller 1875: 453, 457)

Fig. 2A–B

Type

BRAZIL – Bahia • “partie méridionale de la prov.”; 1840; Blanchet 3201a; lectotype (designated here): G [G00237735]; isolectotypes: F [F0069701F], LD [LD1817538], W [W0028441].

Description

Scandent shrub, 2–4 m tall; branchlets greyish brown, striated, glabrous to pubescent when young. Stipules narrow-triangular to triangular, 3.5–6 × 0.8 mm, acute, sericeous outside. Leaves with petioles 2–6(–9) mm long, pubescent to sericeous when young; blades elliptic, (2–)4–6.5 × 2–3.8 cm, obtuse to round at base, acute to acuminate at apex, margin slightly revolute, chartaceous to leathery, discolorous, glabrous above, pubescent with short golden trichomes adpressed below; secondary veins 5–6 on each side of the midrib, inconspicuous above, impressed below; tertiary veins inconspicuous on both sides; domatia hairy below. Inflorescence spiciform, 1.4–3 cm long, 1 per axil, glomerules 2–3 with 5–7 flowers each, peduncle 0.6–1.5 cm long, strigose; bracts lanceolate to narrow-elliptic, 2–2.9 mm long, externally pubescent to strigose, internally glabrous; bracteoles triangular, 0.7–0.9 mm long, pubescent. Flower buds sericeous. Calyx shallowly lobed, lobes ca 1 × 1 mm, externally puberulent. Corolla white to cream, tube 2 mm long, externally glabrous to sparse sericeous, internally lanate near the mouth; lobes acute 1.1 × 0.3–1.1 mm, externally sparsely sericeous, internally lanate. Stamens with inconspicuous filaments; anthers ca 0.6 mm long. Style ca 3 mm long. Fruits oblong, 7–8 × 4–5 mm, purple, glabrous; persistent calyx lobes glabrescent.

Figure 2. 

Species of Malanea from the Atlantic Forest. A–B. M. evenosa (Cardoso 1417). A. Lower surface of leaf with domatia. B. Inflorescence. C–F. M. forsteronioides. C. Lower surface of leaf (Hoehne 3547). D. Stipule (Furlan 1238). E. Inflorescence (Furlan 1238). F. Fruits (Bernacci 1137). G–K. M. harleyi. G. Habit (Carvalho 1364). H. Lower surface of leaf (Thomas 10378). I. Stipule (Mattos-Silva 3929). J. Inflorescence (Thomas 9494). K. Fruits (Thomas 10378). L–P. M. macrophylla. L. Habit (Silva 218). M. Lower surface of leaf (Kollmann 2109). N. Stipule (Bayman 211). O. Flower (Cardoso 1591). P. Fruits (Viana 408).

Distribution

Endemic to Brazil, it occurs in Bahia, mostly in forest remnants in the Chapada Diamantina National Park and in the mountains of Minas Gerais (Fig. 1A).

Phenology

Flowers in May and November; fruits in March.

Etymology

The epithet evenosa refers to the inconspicuous tertiary veins (Müller 1875, 1881).

Preliminary IUCN conservation assessment

Endangered (EN) B1ab(iii)+2ab(iii). The extent of occurrence (EOO) of this species was calculated as 990 km2, qualifying it for the category Endangered (EN). Its area of occupancy (AOO) was estimated as 16 km2, falling into the category Endangered (EN) (Bachman et al. 2011; IUCN 2012, 2022). Malanea evenosa is known by six collections in only two locations in the Chapada Diamantina Nacional Park, in the central portion of the state of Bahia where the Atlantic Forest domain mixed with other domains, such as Cerrado and Caatinga, and a single location in the state of Minas Gerais. Since 1701, the economy of Chapada Diamantina region was largely dominated by mining of gold and diamonds that was responsible for the economic development and urban expansion in the area (Giudice and Souza 2009; Giudice 2022). Only in 1985, the Brazilian government created the Chapada Diamantina National Park, aiming to preserve the natural resources and promote the tourism. Due to this, approval for all mining activities needs previous studies to measure the impacts on flora and fauna. However, currently the permitted mining exceeds the limits established by law, destroying the surrounding vegetation in the area. Therefore, we considered this species as Endangered (EN) according to IUCN criteria (IUCN 2012, 2022).

Notes

Müller (1875: 457) described Malanea evenosa in a brief diagnosis and cited the gatherings Blanchet 3201 and Blanchet 3201a. Five specimens of the two collections were found in F, G, LD, and W. One of them, Blanchet 3201a at G, with barcode G00237735, is designated here as the lectotype. It has handwritten notes by the author of the species, as well as leaves, buds, and fruits in good condition, which allows a correct application of the name.

Comment

Although often misidentified as Malanea revolutifolia, M. evenosa is easily distinguished from the former by its glabrous to pubescent indumentum on young branches (vs hispid to ferruginous-tomentose), the pubescent to sericeous petiole when young (vs strigose), leaves that are acute to acuminate at the apex (vs acute, in young leaves, to rounded), margins slightly revolute (vs strongly revolute), pubescent with short golden trichomes adpressed beneath (vs tomentose to hirsute), hairy domatia in the axils of secondary veins (vs domatia absent), 5–6 secondary veins on each side of the midrib (vs 7–11), inconspicuous on the upper side (vs grooved), and calyx slightly lobed (vs deeply lobed). Few specimens of M. evenosa have been collected so far. Most of the specimens identified as M. evenosa in herbaria are, in fact, M. revolutifolia. These identifications, however, were made before the publication of M. revolutifolia.

Material examined

BRAZIL – Bahia • Ibiquera; 13°31’09”S, 41°18’33”W; 22 Jun. 1978; fr.; Vaillant 61; RB • Palmeiras, Parque Nacional da Chapada Diamantina; 12°27’S, 41°28”W; 1100 m; 18 Nov. 2006; fb., fl.; Cardoso 1417; HUEFS • Serra dos Lençóis, ca 2 km NE de Caeté-Açu (Capão Grande); 12°36’S, 41°29’W; 25 May 1980; infl.; Harley 22644; HUEFS, CEPEC • Serra do Sincorá, Barra da Estiva on the Capão da Volta road; 13°35’S, 41°27’W; 1300 m; 22 Mar. 1980; fr.; Harley 20710; CEPEC, K [K00015533] • s.l.; s.d.; Blanchet 3201; F [F0069701F], LD [LD1817538], W [W0028441]. – Minas Gerais • Serra do Espinhaço, 17°41’37”S, 43°47’35”W; 950–1150 m; 15 Feb. 1972; fr.; Anderson 36045; H, NY [NY00950457], US [US02506579].

Malanea forsteronioides Müll.Arg. (Müller 1875: 453, 458)

Fig. 2C–F

Malanea forsteronioides var. pilosa Standl. (Standley 1931: 365) – Type: BRAZIL – Paraná • Paraty, “ad rivulam scandens”; 30 Dec. 1911; Dusen 13769; holotype: S [S16-54156].

Type

BRAZIL – São Paulo • s.l.; s.d.; Burchell 3770; lectotype (designated here): BR [BR0000005314782]; isolectotypes: G frag. [G00237736], K [K000432743].

Description

Scandent shrub 2–3 m tall; branchlets greyish brown, glabrous to hirsute when young. Stipules triangular-lanceolate, 4–8 × 1.5–2 mm, acute to acuminate, tomentose outside, trichomes long, stiff. Leaves with petioles 5–8 mm long, densely hirsute-hispid; blades oblong-lanceolate, narrow lanceolate to narrow elliptic, 4.5–7 × 2.2–3 cm, obtuse at base, acuminate at apex, margin flat, chartaceous, discolour, glabrous or with hirsute trichomes on primary vein above, tomentose below; secondary veins 4–6 on each side of the midrib, impressed above, prominent below, tertiary veins inconspicuous. Inflorescence spiciform, 2.5–3 cm long, 1 per axil, glomerules 4–5(–7) with 3–4 flowers each, peduncle 0.5–1 mm long, hirsute; bracts lanceolate, ca 2 mm long, externally hairy, ciliated; bracteoles narrowly triangular, 0.2–0.27(–4) mm long, sericeous. Flower buds sericeous. Calyx lobed, lobes unequal, the two larger 2–3 mm long, the two smaller 1–2.5 mm, externally hirsute. Corolla white, tube 2.6–3.5 mm long, externally glabrescent, internally lanate; lobes rounded to acute, 2.1–3.1 mm long, externally tomentose, internally puberulent or lanate close to the mouth. Stamens with filaments 1.8–2 mm long; anthers 1–1.2 mm long. Style 2–2.3 mm long. Fruits fusiform, 8.6–9.5 × 3.8–5.2 mm long, reddish to purple, glabrous; persistent calyx lobes hirsute.

Distribution

Endemic to Brazil, it occurs from Espírito Santo to Santa Catarina, along the eastern coast of Brazil (Fig. 1A).

Habitat and ecology

Known only from submontane moist forests in the Brazilian Atlantic Forest.

Phenology

Flowers from October to December; fruits in January, April, and from July to September.

Etymology

The epithet forsteronioides refers to the similarity of this species with species of Forsteronia G.Mey. (Apocynaceae).

Preliminary IUCN conservation assessment

Near Threatened (NT). The extent of occurrence (EOO) was calculated as 135,845 km² qualifying the species as Least Concern (LC). The area of occupancy (AOO) was evaluated as 68 km², falling into the Endangered (EN) category (Bachman et al. 2011; IUCN 2012, 2022). Malanea forsteronioides occurs mainly on the coast of the Southeast portion of the Atlantic Forest. Although this area had an increase in tourism and an expansion of private houses for occasional use (Sabino 2012), with impacts on the surrounding biodiversity, the species occupies more than ten locations. For this reason, we consider it as Near Threatened (NT).

Notes

Müller (1875: 453) described Malanea forsteronioides and cited the gathering Burchell 3770. Three specimens of this gathering were found in BR, G, K. The lectotype here designated is the specimen in BR, with barcode BR0000005314782, because it has an annotation handwritten by Müller, as well as leaves, buds, and flowers in good condition, which allows a precise application of the name.

Comment

Barbosa (2007) indicated heterostyly in Malanea forsteronioides, which was corroborated here. However, we emphasize that long-styled individuals are more abundant than short-styled individuals in the specimens analysed. However, the size of the corolla, androecium, and style do not show great variation between the two flower forms. Only the size of the bracteoles and calyx lobes vary, being slightly larger in long-styled individuals.

Material examined

BRAZIL – Paraná • Antiga estrada para Superaqui-Guaraqueçaba; 23 Mar. 1989; fr.; Caetano Filho s.n.; RB • Jacarezinho, Jacarehy; 2 Sep. 1914; fb., fl.; Standley s.n; NY • Paranaguá, Caiobá, Morro do Bio; 25°31’13”S, 48°30’33”W; 100 m; 1 Nov. 1965; fb.;. Hatschbach 13070; NY. – Rio de Janeiro • Arraial do Cabo, Ilha do Farol; 30 Oct. 2013; fb.; Uller s.n.; RB • Paraty, Pico do cairuçú - acesso pela praia negra; 25 Nov. 1990; fl.; Farney 2548; RB • Rio de Janeiro, Serra da Tijuca; Jan. 1932; fb.; Brade 11319; NY. – Santa Catarina • Florianópolis, Ilha de Santa Catarina, Jureré; 23 Feb. 1970; fr.; Klein 8606; NY • Itapoá, Reserva Volta Velha; 26°15’S, 48°65’W; 9 Jan. 1998; fb.; Canha s.n.; NY • Vargem do Macário, Governador Celso Ramos; 18 Nov. 1971; fb.; Klein 9955; NY. – São Paulo • Alto da Serra, Estação Ecológica; 8 Dec. 1919; fb., fl.; Hoehne 3547; SP • Antiga São Paulo – Santos, Alto da Serra; 14 Oct. 1961; fb., fl.; Pereira 5932; RB • Biritiba Mirim, Estação Ecológica de Boracéia; 23°38’S, 45°53’W; 950 m; 14 Nov. 1983; fb.; Custodio-Filho 1839; NY, SP • Parelheiros, Sítio à esquerda na Estrada Eng. Marsilac; 23°53’46”S, 46°43’40”W; 14 Feb. 1995; fb.; Godoy 373; SP [SP300813] • Pariquera-Açu, Propriedade de Antônio Povinski; 29 Sep. 1995; fb.; Ivanauskas 419; RB • Pariquera-Açu, Rodovia para Cananeia, Fazenda Esteiro do Morro de F. Capobianco; 12 Jan. 1995; fr.; Bernacci 1137; IAC • Peruibe; 23 Oct. 1990; fl.; Furlan 1238; HRCB • Peruíbe, Estação Ecológica da Juréia; Oct. 1990; fb.; Sobral 6868; HRCB • Praia Grande; 26 Oct. 1998; fl.; Loefgren 4146; SP.

Malanea harleyi J.H.Kirkbr. (Kirkbride 1997: 366)

Fig. 2G–K

Type

BRAZIL – Bahia • Ca 26 km SW of Belmonte along road to Itapebi, and 4 km alongside road towards the sea; 16°03’S, 39°02’W; 25 Mar. 1974; fr.; Harley, Renvoize, Eskine, Brighton & Pinheiro 17413; holotype: CEPEC [11800]; isotypes: K [K000015545, K000015546, K000852369, K000852370], MO [MO 100477410], RB [RB00344453], US [US02367811, US00512890], VEN [VEN176883].

Description

Liana or scandent shrub 2–3 m tall; branchlets dark brown with sparse subcircular lenticels, glabrous. Stipules elliptic to ovate, (5–)7–15 × 3–9 mm, round at apex, glabrous. Leaves with petioles 0.4–1.8(–2.3) cm long, glabrous; blades elliptic to ovate, (6.4–)8–14.5(–16) × (3–)4–7(–8.1) cm, obtuse to round at base, acuminate at apex, margin slightly revolute, subcoriaceous to leathery, lustrous, glabrous, concolorous; secondary veins (5–)6–7(–8) on each side of the midrib, impressed or slightly sulcate above, prominent below; tertiary veins inconspicuous. Inflorescence spiciform, 4–11 cm long, 1–2 per axil, glomerules 7–8, with 5–7 flowers each, the basal pair of glomerules with branches 0.4–1.4 cm long, the others sessile; peduncle 0.5–3.1 cm long, sparse strigose; bracts triangular, 0.14–0.2(–3.3) mm long, pubescent; bracteoles two smaller and one larger, triangular to deltoid, 0.1–0.5 × 0.1–0.3 mm, glabrescent. Flower buds pubescent. Calyx lobed, lobes 0.7–1.3 mm long, glabrous. Corolla cream-yellow, tube 1.4–2.5(–3.0) mm long, externally glabrous, internally lanate; lobes rounded, 1.2–1.8 mm long, externally glabrous, internally lanate. Stamens with filaments 0.6 mm long; anthers 0.5–0.8 mm long. Style 1.2–3 mm long. Fruits oblong, 7–12 × 3–6.5 mm, purple, glabrous; persistent calyx lobes glabrous.

Distribution

Endemic to the state of Bahia (Fig. 1C).

Habitat and ecology

Known only from forests on the coastal white sands (restinga forests) or from vegetation bordering grassy open areas on damp white sand at sea level.

Phenology

Flowers from January to March, and in November; fruits from February to April, and from September to November.

Vernacular name

Cravo-do-campo (Bahia, Mattos-Silva 3929).

Etymology

The epithet harleyi honours Raymond Mervyn Harley, a botanist and honorary researcher at the Royal Botanic Gardens (Kew), and a specialist in neotropical biodiversity and one of the greatest scholars of the flora of Bahia, where he continues to collect and study the flora.

Preliminary IUCN conservation assessment

Endangered (EN) B1ab(ii,iii)+2ab(ii,iii). The extent of occurrence (EOO) of this species was calculated as 4,394 km2, qualifying it as Endangered (EN). The area of occupancy (AOO) was estimated as 88 km2, also falling into the Endangered (EN) category (Bachman et al. 2011; IUCN 2012, 2022). Malanea harleyi is an endemic species found in only five locations of Restinga forests in southern Bahia. That forest was severely degraded due to human activities such as housing construction, agriculture, and selective logging that threaten the local diversity (Delprete and Jardim 2019).

Notes

In the protologue of the species, Kirkbride (1997) comments that the flowers have fallen off after anthesis and he did not describe their characters. In this treatment, we present the first information about the floral structures of M. harleyi. However, this species is clearly distinct from the others due to its yellowish stipules, leaves, peduncle, and calyx when dried, subcoriaceous to leathery leaves, and glabrous vegetative structures.

Material examined

BRAZIL – Bahia • Belmonte, ca 26 km SW de Belmonte ao longo da Rod. para Itapebí, e 4 km ao longo da Rod. para o mar; 16°05’S, 39°03’W; 25 Mar. 1974; fr.; Harley 17413; CEPEC, RB • Belmonte; 6 Jul. 1966; fr.; Belém 2498; CEPEC, RB • Camamu, povoado de Barcelos do Sul; 16 Jul. 2005; fr.; Miranda 5083; HUEFS, ALCB • Camamu, povoado de Barcelos do Sul; 16 Jul. 2005; fr.; Miranda 5186; UFRN • Ilhéus, 19.8 km W de Olivença na rodovia para Vila Brasil; 14°57’51”S, 39°06’48”W; 14 Feb. 1994; fr.; Thomas 10378; JPB, CEPEC • Ilhéus, 8.9 Km ao SW de Olivença, Rod. Maruim/Vila Brasil; 14°59’36”S, 39°3’W; 21 May 1995; fb., fl., fr.; Thomas 10956; CEPEC, NY • Ilhéus, ca 5 km, estrada que liga Olivença ao povoado do Maruim; 14°47’20”S, 39°04’57”W; 20 May 2002; fl.; Paixão 185; CEPEC • Ilhéus, Fazenda Barra do Manguinho. Ramal com entrada no Km 10 Rod. Pontal/Olivença, lado direito. 3 km a W da Rod.; 14°47’20”S, 39°04’57”W; 5 Feb. 1982; fr.; Mattos-Silva 1431; CEPEC • Ilhéus, Faz. Barra do Manguinho. Ramal c/ entr. km 10 Rod. Pontal/Olivença. Coletas 3.5 km ao W da praia; 14°47’20”S, 39°04’57”W; 11 Feb. 1993; fr.; Mattos-Silva 2911; CEPEC • Ilhéus, Faz. Guanabara. Ramal c/ entrada no km 10 a direita da Rod. Pontal/Olivença, 3 km ao W da entrada; 14°47’20”S, 39°04’57”W; 16 Oct. 1980; fr.; Mattos-Silva 1161; CEPEC • Ilhéus, Fazenda Guanabara (junto a Fazenda Barra do Manguinho). Ramal com entrada no km 10 da Rod. Ilhéus/Olivença, á direita, 4 km a Oeste da Rodovia; 14°47’20”S, 39°04’57”W; 9 Mar. 1997; fr.; Mattos-Silva 1851; CEPEC • Ilhéus, road from Olivença to Maruim, 6–8 km W of Olivença; 50 m; 10 May 1981; infl.; Mori 13935; CEPEC, RB • Itacaré, Fazenda Campo Cheiroso; 14°27’S, 38°99’W; 150 m; 31 Mar. 1993; fr.; Mattos-Silva 3929; HUEFS, ALCB, CEPEC • Itacaré, Campo Cheiroso; 14°22’50”S, 39°02’23”W; 125 m; 9 Feb. 2011; fr.; Lacerda 47; CEPEC • Itacaré, Campo Cheiroso; 14°22’50”S, 39°02’23”W; 100 m; 5 Apr. 2011; st.; Lacerda 127; CEPEC • Itacaré, Campo Cheiroso; 14°22’50”S, 39°02’23”W; 100 m; 22 Sep. 2011; st.; Lacerda 211; CEPEC • Itacaré, Campo Cheiroso, 14 km N da Serra Grande; 14°22’S, 39°04’W; 15 Nov. 1992; fl., fr; Thomas 9494; ALCB, CEPEC, HUEFS, NY • Itacaré, Estr. liga Serra Grande ramal 13 que leva ao Campinho Cheiroso. Coletas entre Km 15–16 a partir de Serra Grande; 14°16’39”S, 38°59’47”W; 26 Aug. 1992; fr.; Amorim 730; CEPEC • Ituberá, região da praia do Pratigi; 13°42’47”S, 39°09’64”W; 20 m; 27 Mar. 2003; fl.; Paixão 220; CEPEC • Maraú, a 3.9 km da balsa Itacaré/Maraú; 14°14’41”S, 39°00’59”W; 19 m; 22 May 2008; fr.; Vignoli-Silva 284; CEPEC • Maraú, beira da estrada, mata aberta, pouco antes de chegar à BR030; 14°14’39”S, 39°00’56”W; 28 Feb. 2007; fr.; Souza 148; CEPEC, NY • Maraú, ca 3 km a E de Maraú, na Rod. para Saquaíra; 14°06’20”S, 39°0’24”W; 2 Mar. 1971; fl.; Jardim 2648; CEPEC, NY • Maraú, Rod. BR-030, a 24 km trecho Maraú/Porto de Campinhos. 19 km a E do entroncamento; 14°04’S, 38°57’W; 13 Jun. 1979; fb., fr.; Mattos-Silva 451; CEPEC, IPA, RB • Una, ca 3 km SW da sede do município; 15°29’32”S, 39°07’52”W; 22 Feb. 1992; fr.; Carvalho 3821; CEPEC, HUEFS, NY • Una, Fazenda Bolandeira, ramal com entrada no km 10 da rodovia Una/Canaveiras; 15°24’35”S, 39°05’49”W; 14 Apr. 2000; fr.; Mattos-Silva 4058; HUEFS, ALCB, CEPEC • Una, próximo a Reserva Biológica de Una, beira de estrada; 15°11’16”S, 39°01’36”W; 434 m; 7 Jan. 2002; fl.; Nunes 732; HUEFS • Una, ramal a 5 km da estrada Una/Canavieras, 15°29’32”S, 39°07’52”W; 8 Jan. 1983; fr.; Carvalho 1364; CEPEC.

Malanea macrophylla Bartl. ex Griseb. (Grisebach 1861: 337)

Figs 2L–P, 3A–D

Malanea bahiensis Müll.Arg. (Müller 1875: 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 megalantha Wernham (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 (Müll.Arg.) Steyerm. (Steyermark 1965: 250) – Type: same as for Malanea bahiensis.

Malanea macrophylla f. cuneata Steyerm. (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 × 3–10 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 × (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 × 2–5 mm, purple, glabrous; persistent calyx lobes glabrescent.

Distribution

Malanea macrophylla is 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

Cipó-de-brejo (Alagoas, Rodrigues 1443), Arco-de-barri (Bahia, Cardoso 1591).

Etymology

The epithet macrophylla refers 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 macrophylla as a new species recognized by Bartling in mss (manuscript). However, the only information in Schomburgk’s Flora about the species is “An den ufern des Morocco. Blüht im 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 1484 at GOET (GOET008943), has well-preserved leaves, inflorescences, and flowers and has handwritten annotations similar to those in the original publication.

Comment

Malanea macrophylla is 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. macrophylla and 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 macrophylla is 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 • Colônia Leopoldina, Serra do Livramento; 7 Sep. 2010; fb., fr.; Chagas-Mota 8293; MAC • Flexeiras, Estação Ecológica de Murici; 24 Mar. 2011; fb.; Chagas-Mota 10504; MAC • Ibateguara, Petrópolis; 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.; Mendonça 409; MAC • Murici, Serra do Ouro; 7 Sep. 2003; fb.; Falcão 42; 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 • São José da Lage, BR 104; 11 Nov. 2010; fb.; Chagas-Mota 9282; MAC • Sem município; 28 Oct. 1980; fb., fl.; Andrade-Lima 80-9762; IPA • União dos Palmares, Fazenda Santo Antônio; 26 Nov. 1966; fb.; Andrade-Lima 66-4751; IPA. – Bahia • Almadina, Serra do Corcovado/São Domingos, trilha que dá acesso a Área do Carvoeira; 14°41’S, 39°35’W; 25 Jan. 2014; fb.; Marinho 601; HUEFS, CEPEC • Amargosa, Serra do Timbó, 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 Timbó; 13°10’20”S, 39°09’34”W; 531 m; 26 Jan. 2007; fr.; Cardoso 1631; HUEFS, CEPEC • Amargosa, Serra do Timbó; 13°10’S, 39°09’W; 531 m; 17 Mar. 2007; fr.; Paixão 1093; HUEFS • Amargosa, Serra do Timbó; 13°10’S, 39°09’W, 28 Apr. 2007; fr.; Paixão 1160; HUEFS • Andaraí, Cachoeira dos Funis; 2 Jan. 2017; fl.; Oliveira 126; HUEFS • Andaraí, Chapada Diamantina; 12°48’S, 41°23’W; 2 Nov. 1999; fb.; Guedes 6969; ALCB • Apuarema, Concessão da Rio Tinto; 13°53’46”S, 39°41’10”W; 680 m; 20 Nov. 2013; fb.; Aona 3276; RB • Belmonte, a 25km SW da cidade; 6 Jan. 1981; fb.; Carvalho 441; CEPEC • Belmonte, a 30 km SW da cidade; 8 Jan. 1981; fb.; Carvalho 484; CEPEC • Cabrália, Reserva Biológica Pau-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 • Elísio Medrado, ca 1 km do povoado de Monte Cruzeiro; 12°52’10”S, 39°28’18”W; 600 m; 2 Dec. 2004; fb.; Jardim 4283; HUEFS • Elísio Medrado, Serra da Jibóia; 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 • Eunápolis, trecho de Mata Atlântica entre Eunápolis e Porto Seguro; 16°26’S, 39°03’W; 29 Mar. 1995; fr.; Ferreira 83; ALCB • Igrapiúna, 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 córrego 1; 13°54’58”S, 39°27’57”W; 13 Mar. 2013; fb.; Aona 2400; RB • Ilhéus, Parque Municipal Mata da Esperança, acesso pelo Banco da Vitória; 14°46’S, 39°05’W; 8 Jan. 2003; fb., fl.; Jardim 4098; HUEFS, CEPEC • Itacaré, Fazenda Capitão, 7.9 km a partir da junção entre BA 001 e a rodovia Itacaré-Ubaitaba; 31 Mar. 2004; fr.; Fiaschi 2187; NY, RB • Itacaré, próximo a 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 km da rodovia vicinal de Itamaraju sentido Jucuruçu; 16°50’19”S, 39°37’53”W; 500 m; 13 Feb. 2000; fl.; Borges 785; CEPEC, NY • Itamaraju, Assentamento Pedra Bonita, aproximadamente 20 km da rodovia vicinal de Itamaraju sentido Jucuruçu; 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.; Gusmão 145; ALCB • Ituberá, 11 km da estrada Ituberá/Valença; 5 Feb. 1983; fr.; Carvalho 1459; HUEFS, CEPEC • Ituberá, km 11 da estrada Itubera/Valença; 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 • Lençóis; 12°34’S, 41°23’W; 30 Jul. 2006; 600 m; fr.; Couto 201; HUEFS • Lençóis, Serra da Chapadinha; 12°27’03”S, 41°25’07”W; 600 m; 6 Feb. 1995; fr.; Guillietti 1614; ALCB, CEPEC • Lençóis, Chapada Diamantina, Foz da Capivara; 9 Mar. 2000; fb.; Stradmann 0706; ALCB • Lençóis, Chapada Diamantina; 26 Jan. 2000; fr.; Jardim 2526; CEPEC, NY • Lençóis de Una, próximo à ponte sob rio Acuípe; 21 Feb. 2022; fr.; Roxo 43; JPB • Maraú, Estrada que liga Ponta de Mutá a Maraú; 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 Inácio; 12°28’S, 41°27’W; 1000 m; 12 Mar. 1997; fr.; Gasson 6193; HUEFS, ALCB, CEPEC • Palmeiras, Pai Inácio; 12°26’37”S, 41°29’16”W; 690 m; 1 Jul. 1995; fr.; Guedes 2116; ALCB, CEPEC • Palmeiras, Morro do Pai Inácio; 12°27’20”S, 41°28’15”W; 1080 m; 4 Feb. 1995; fr.; Guilietti 1531; ALCB, CEPEC, HUEFS • Palmeiras, Morro do Pai Inácio; 12°27’16”S, 41°28’17”W; 1040 m; 30 Dec. 1994; fb., fl.; Guilietti 1506; CEPEC • Piraí do Norte; 13°46’12”S, 39°19’52”W; 6 Nov. 2014; fb.; Braga 41; JPB • Pontal dos Ilhéus, saída para Buererema; 17 May 1986; fr.; Belém 3571; 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 Muçununga, área aberta próximo à mata; 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 Muçununga, próximo à margem 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 Biológica do Pau-Brasil, 17 km ao W da cidade, Rodovia para Eunápolis; 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 Botânico de Salvador; 12°55’37”S, 39°03’51”W; 6 Feb. 2004; fr.; Queiroz 353; HUEFS, ALCB • Salvador, região metropolitana, barragem do Cobre; 12°58’S, 38°30’W; 16 Apr. 2012; fr.; Queiroz 5312; ALCB • Santa Cruz Cabrália, Santo André, próximo ao estuário do Rio Acuba; 16°14’S, 39°02’W; 2 Nov. 2003; fb.; Guedes 10624; ALCB • São Miguel 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 km no ramal que liga o distrito de Itatinguí à Serra; 15°12’09”S, 39°24’28”W; 31 Mar. 2006; fb.; Paixão 927; CEPEC, NY • Ubaira, arredores de Três Braços; Jan. 1991; fb., fl.; Sobral 6702; SP • Una, Mata litorânea; 19 May 1965; fb.; Belém 1062; CEPEC • Una, próximo à Reserva Biológica Una; 16°28’10”S, 39°16’46”W; 21 Feb. 2022; fr.; Roxo 41; JPB • Una, Reserva Biológica do Mico-leão (IBAMA); 8–12 Mar. 1993; fb.; Amorim 1130; CEPEC, NY • Una, Reserva Biológica do Mico-leão (IBAMA); 26 Apr. 1994; fl.; Carvalho 4515; CEPEC, NY • Una, Reserva Biológica do Mico-leão (IBAMA); 3 Mar. 1998; fr.; Sant’Ana 640; CEPEC • Una, Reserva Biológica do Mico-leão (IBAMA), 15°06’58”S, 39°02’11”W; 20 Mar. 1999; fr.; Jardim 2068; CEPEC • Uruçuca, estrada de Itacaré para Ubaitaba; 14°29’59”S, 39°06’54”W; 380 m; Apr. 2004; st.; Martini sn.; CEPEC • Uruçuca, Parque Estadual Serra do Conduru ca 11.5 km de Serra Grande em direção a Uruçuca, com 1km do ramal a esquerda; 18 May 2000; fr.; Sant’Ana 855; CEPEC. – Espírito Santo • 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 • Uberlândia, Parque do Sabiá; 22 Jun. 1995; fr.; Nakagima 1644; HUFU • Uberlândia, Tangará Country Club; 18°55’07”S, 48°16’37”W; 28 Nov. 1995; fl.; Schiavini 410; HUFU. – Paraíba • 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 Jacumã; 7°19’24”S, 34°48’47”W; 4 Dec. 2009; fr.; Araújo 219; JPB • Espírito Santo, Engenho São Paulo; 25 Nov. 1968; fl.; Andrade-Lima 68-5475; IPA • João Pessoa, Praia da Penha, Rio Aratú; 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, Estação Ecológica Mamanguape; 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 Conceição; 22 Nov. 1967; fb.; Costa 143-67; IPA • Gravatá, 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.; García-González 1374; CEPEC, JPB, HUEFS, NY UFP • Igarassu, Mata de Piedade; 7°49’48”S, 35°59’55”W; 2 Mar. 2010; fr.; García-González 1445; JPB, IPA; CEPEC, UFP • Igarassu, Mata de Piedade, área protegida propriedade da Usina São José; 7°49’38.5”S, 35°0’16.1”W; 91 m; 12 Aug. 2009; fr.; García-González 1070; 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 Botânica do 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 • São Vicente Ferrer, Mata do Estado; 7°35’S, 35°30’W; 600 m; 12 Nov. 1995; fb., fl.; Laurêncio 237; NY, PEUFR • São Vicente Ferrer, Mata do Estado; 07°35’S, 35°30’W; 600 m 13 Jan. 2000; fb.; Ferraz 810; PEUFR • Serinhaém, Usina Trapiche; 8°33’14”S, 35°08’48”W; 110 m; 21 Dec. 2010; fb.; Melo 713; JPB, UFP • Tamandaré, Mata do conde, Fazenda estivas; 21 Jan. 2000; fb., fr.; Oliveira 541; PEUFR. – Rio de Janeiro • Carapebus, Fazenda Lázaro; 23 Jun. 1994; fb.; Bovini 367; JPB, RB • Carapebus, Restinga de Carapebus. Fazenda São Lázaro; 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 Biológica de Poço das Antes, trilha do Morro do Calcário; 22°30’S, 42°15’W; 5 Mar. 1993; fr.; Barreto 29; CEPEC, RB • Silva Jardim, Reserva Biológica de Poço das Antas, trilha do Morro do Calcário, ponto 1000; 22°00’S, 42°00’W; 9 Jan. 1993; fb.; Lima 4585; NY, RB • Silva Jardim, Reserva Biológica Poço das Antas, Trilha do Morro do Calcário; 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 • Macaíba, Escola Agrícola de Jundiaí; 5°53’35”S, 35°21’01”W; 54 m; 10 Feb. 2012; fr.; Costa-Lima 628; UFRN, UFP • Porto Velho, Área de Proteção Ambiental 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 Atlântica; 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 km do 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 – Amapá • Calçoene, BR156 entre Calçoene e Rio Amapá Grande, ca 5 km S de Calçoene; 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 • Brasília, ca 10 km of Brasília, próximo 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. – Goiás • Mineiros, Nascente do Ribeirão Gloria, 200–300 m afora 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, córrego Cabecerão; 18°2’S, 52°52’12”W; 1 Oct. 1998; fb.; Delprete 6780; NY • Pirenópolis, Cachoeira Santa Maria- RPPM; 15°49’23”S, 48°54’25”W; 1150 m; 23 Jun. 2001; fr.; Marquete 3157; RB • Pirenópolis, Chapada dos Veadeiros. Fazenda Pai José; 27 Oct. 1994; fb.; Mendonça 2229; RB • Planaltina; 20 Jul. 1966; fb.; Irwin 18296; RB • Serra do Caiapó, 40 km South of Caiaponia; 26 Oct. 1964; fr.; Prance 59673; NY. – Maranhão • São Luiz, Ilha de São Luiz; 2°57’S, 44°20’W; Feb.–Mar. 1939; fb.; Lemos Fróes 11737; NY. – Mato Grosso • North ca 2 km. E. of base camp; 19 Oct. 1968; fr.; Harley 10728; NY, RB • Sinop, Rd. to Fazenda Atlântica, 10–14 km W of BR 163 and 8 km N of rd to Pôrto dos Gaúchos; 11°06’S, 55°57’W; 25 Sep. 1985; fr.; Thomas 4061; NY. – Pará • Belém; 24 Aug. 1922; fb.; Ducke s.n.; RB • Ilha do Marajó, Rio Anajás; 0°59’S, 49°55’W; 2 Nov. 1987; fr.; Prance 30227; NY • Rio Mocoões, 1 km above Anajás; 0°58’S, 49°56’W; 7 Nov. 1987; fr.; Beck 366; NY. – Roraima • Caracaraí, PARNA do Viruá, Grade do PPBio, trilha L01, segment 1400–1450; 1°29’12”N, 61°0’15”W; 19 Sep. 2012; fb.; Perdiz 1600; RB.

Malanea martiana Müll.Arg. (Müller 1875: 453)

Figs 3E–F, 4A–E

Type

BRAZIL • Bahia, Ilhéus; s.d.; Martius 394; lectotype (designated here): M [M0198135]; isolectotypes: BR [BR0000005314775, BR0000005315109, BR0000005315437], F [F0BN000418], F frag. [V0069707F], G [G00237739, G00237740], GH [GH00096449], K [K000015350], L [L0057647], NY [NY00132152], W [W 0028443, W 1889-0287298].

Description

Liana or scandent shrub up to 15 m tall; branchlets greyish with elliptical to circular lenticels, glabrous. Stipules triangular-lanceolate, 6–8.5(–10) × 1–3 mm, acute, strigose. Leaves with petioles (0.2–)0.3–0.5 cm long, glabrescent at base; blades obovate to elliptic, 5.5–13 × 3.4–5.5 cm, cuneate to attenuate at base, acute to cuspidate at apex, margin flat, chartaceous, discolorous, glabrous above, lustrous, sparse pubescent below, trichomes adpressed; secondary veins 4–5 on each side of the midrib, impressed above, prominent below; tertiary veins inconspicuous. Inflorescence paniculate, 5–12.5 cm long, 1 per axil, glomerules 3–5(–7), with 13–18 flowers each; peduncle 2.5–6.5 cm long, sericeous; bracts triangular, 2–2.9 mm long, glabrescent; bracteoles inconspicuous, triangular, puberulent. Flower buds strigose. Calyx dentate, teeth triangular, 0.4–0.8 mm long, glabrescent. Corolla yellowish, tube 1.4–4 mm long, externally glabrescent to strigose, internally lanate; lobes rounded, 1.5–2.9(–3.5) mm long, externally strigose, internally villous. Stamens with filaments 1–1.5 mm long; anthers 0.7–1(–2) mm long, curved. Style 2.5–3.3(–4) mm long. Fruits fusiform, 8–14 × 2–3 mm, purple or black, glabrous; persistent calyx lobes glabrescent.

Figure 4. 

Species of Malanea from the Atlantic Forest. A–E. M. martiana. A. Habit (Guedes 24971). B. Lower surface of leaf (Sant'Ana 439). C. Stipule (Guedes 24971). D. Flower (Guedes 24971). E. Fruits (Bautista 1748). F–J. M. revolutifolia. F. Habit (Popovkin 25). G. Lower surface of leaf (Ferreira 721). H. Stipule (Ferreira 721). I. Inflorescence (Ferreira 721). J. Fruits (Gusmão 469). K–N. M. spicata. K. Lower surface of leaf (Nunes 751). L. Stipule (Thomas 9885). M. Flower (Carvalho 214). N. Fruits (Fiaschi 1106).

Distribution

It occurs in the states of Alagoas, Bahia, Pernambuco, and Sergipe (Fig. 1C).

Habitat and ecology

It is found in moist forests or forests bordering restinga areas.

Phenology

Flowers in June, and from September to October; fruits in July to August, and from December to April.

Etymology

The epithet martiana honours Carl Friedrich Philipp von Martius (1794–1868), the collector of the type material.

Preliminary IUCN conservation assessment

Vulnerable (VU) B2ab(iii). The extent of occurrence (EOO) of this species was calculated as 104,959 km2, qualifying it for the category Least Concern (LC). The area of ​​occupancy (AOO) was estimated as 100 km2, falling into the category Endangered (EN) (Bachman et al. 2011; IUCN 2012, 2022). Malanea martiana is known from eight locations that occupy a considerable area, however only few populations were sampled in the northern portion of the Atlantic Forest domain (states of Alagoas, Pernambuco, and Sergipe), where forests were largely replaced by plantations of sugarcane (Targino et al. 2010), leaving only small forest fragments located in areas unsuitable for cultivation (Barbosa and Rios 2006). However, considering that extraction of wood persists in the area (Machado 2008), threatening the local biodiversity, we categorize it as Vulnerable (VU) according to the IUCN criteria (IUCN 2012, 2022).

Notes

Müller (1875) described Malanea martiana refering to Malanea sarmentosa Mart. in Herbarium Florae Brasiliensis as a synonym. Consulting Martius (1839), we found that he had identified his collection 394 as Malanea sarmentosa Aubl., not intending to describe a new species. We found specimens of Martius 394 in several herbaria. We designate here M0198135 at M as the lectotype because it shows Müller’s determination handwritten by him and is the best preserved and complete specimen.

Comment

Malanea martiana is heterostylous, but the size of the calyx, corolla, and androecium do not vary between the two forms in the analysed specimens. Only the style length varies. Malanea martiana is distinguished from the other studied species by its acute stipules, the paniculate inflorescence larger than 5 cm, and the dentate calyx.

Material examined

BRAZIL – Alagoas • Maceió, Serra da Saudinha; 18 Sep. 2009; fb.; Chagas-Mota 5529; MAC. – Bahia • Cairu, Litoral Sul; 13°29’S, 39°2’W; 28 Sep. 2001; fl; Guedes 9229; ALCB • Conde, Mata da Maré; 12°02’27”S, 37°43’30”W; 14 Dec. 1995; fr.; Bautista 1748; ALCB, CEPEC, IPA, HUEFS, RB • Cravolândia, povoado Três Braços, Ilha Formosa; 13°30’S, 39°42’W; 15 Jan. 1994; fr.; França 913; HUEFS • Feira de Santana, BR 234, km 35; 5 Jan. 1981; fr.; Gusmão 504; ALCB • Feira de Santana, BR 234, km 35; 17 Sep. 1980; Gusmão 471; ALCB, UB [111794] • Igrapiúna, Reserva Ecológica da Michelin: Cachoeira da Pancada Grande; 13°46’57”S, 39°10’00”W; 4 Dec. 2015; fb., fl.; Silva 668; JPB • Ilhéus, Mata da Esperança. 3 km da Rodoviária, mata ao N do reservatório; 14°46’54”S, 39°04’08”W; 17 Jan. 1995; fr.; Thomas 10799; CEPEC, NY • Ilhéus, Rodovia Uma, próximo à Olivença; 14°33’S, 39°56’W; 17 Jan. 1999; Argôlo 02; ALCB • Itacaré, ao SW da Rod. Itacaré/Ubaitaba, ao S da foz do Rio de Contas; 14°11’24”S, 39°00’59”W; 29 Jan. 1977; fr.; Harley 18383; CEPEC, RB • Itacaré, trilha em direção a Faz. Prainha; 19 Aug. 2010; st.; García-González 1496; RB • Itacaré, Serra Grande, trilha dentro da Fazenda Miranda; 22 Jan. 2022; fr.; Roxo 44; JPB • Itanagra, Fazenda Brejo Verde; 9 Aug. 1975; fr.; Gusmão 145; ALCB • Ituberá/Grapiuna, Michelin, Faz. Biriba, na margem da estrada; 13°43’S, 39°8’W; 23 Sep. 2006; fb.; Guedes 12602; ALCB • Maraú; 14°14’42”S, 39°00’56”W; 27 Jul. 2000; fr.; Mattos-Silva 4376; HUEFS • Olivença, Mata do Balneário Tararomba; 15°26’S, 39°06’W; Sep. 2016; fl.; Guedes 24971; ALCB • Porto Seguro, Parque Nacional do Pau-Brasil, trilha do Muçununga, próximo à margem de rio; 16°26’06”S, 39°19’30”W; 91 m; 19 Jan. 2022; st.; Roxo 35; JPB • Porto Seguro, Parque Nacional do Pau-Brasil, trilha do Muçununga; 16°26’06”S, 39°19’30”W; 91 m; 19 Jan. 2022; fr.; Roxo 37; JPB • Porto Seguro, Trancoso, 16°23’S, 39°08’W; 2 Oct. 1997; fb., fl.; Guedes 5333; ALCB, IPA • Salvador, Região Metropolitana de Salvador; 12°58’S, 38°30’W; 19 Apr. 2011; fr.; Guedes 18773; ALCB, PEUFR • Taperoá, Ramal da Faz. São Braz, c/ entrada no km 6 Rod; 13°32’17”S, 39°05’54”W; 21 Sep. 1988; fb.; Mattos-Silva 2566; CEPEC • Una, Reserva Biológica do Mico-leão (IBAMA), entrada no km 46 da Rod. BA-001 Ilhéus/Una; 15°09’S, 39°05’W; 8–12 Mar. 1993; fr.; Amorim 1095; CEPEC, NY • Una, rodovia Una-Comandatuba, fazenda de Antônio Pimenta; 26 Oct. 1971; fl.; Pinheiro 1658; CEPEC • Uruçuca; 14°35’54”S, 39°17’44”W; 12 Feb. 2004; fr.; Amorim 3919; CEPEC, RB • Uruçuca, 7.4 km N de Serra Grande Rod. P/ Itacaré. Faz. Lagoa do Conj. Sta. Cruz. Inventário; 14°42’33”S, 39°06’05”W; 8 May 1995; infl.; Thomas 10823; CEPEC, NY • Uruçuca, ca 5 km de Serra Grande; 14°47’22”S, 39°02’17”W; 135 m; 1 Oct. 2008; fl.; Queiroz 13848; CEPEC, HUEFS • Uruçuca, distrito de Serra Grande, 7.3 km na estrada Serra Grande/Itacaré, Fazenda Lagoa do Conjunto Fazenda Santa Cruz; 14°25’S, 39°01’W; 7 Sep. 1991; fb., fl.; Carvalho 3644; CEPEC • Uruçuca, Serra Grande, caminho do mirante II; 14°17’S, 39°03’W; 2 Mar. 2018; fr.; Guedes 30341; ALCB • Valença, RPPN Água Branca, estrada de Valença para Guaibim, ca 2 km; 13°19’44”S, 39°05’25”W; 26 Jun. 2004; fl.; Fiaschi 2345; CEPEC, NY. – Pernambuco • Igarassu, Mata de Piedade; 7°49’48”S, 34°59’55”W; 2 Mar. 2010; fr.; García-González 1448; JPB, UFP • Igarassu, Usina São José; 7°42’12”S, 34°58’11”W; 22 Dec. 2007; fr.; Araújo 556; IPA, UFP. – Sergipe • Santa Luzia do Itanhi, Entrada a 1 km a esquerda da estrada Sta. Luzia/Crasto, ca 1 km adentro; 11°35’07”S, 37°44’82”W; 29 Nov. 1993; fr.; Sant’Ana 439; CEPEC, NY.

Malanea revolutifolia A.Amaya & Popovkin (Amaya and Popovkin 2012: 930)

Fig. 4F–J

Type

BRAZIL • Bahia, Entre Rios, Fazenda Rio do Negro; 12°01’S, 38°02’W; 150 m; 14 Aug. 2008; fl.; Popovkin 349; holotype: HUEFS [143271]; isotype NY [02714491].

Description

Liana ca 2 m tall; branchlets greyish with elliptic lenticels, glabrous, hispid to ferruginous-tomentose and without lenticels when young. Stipules deltate to triangular, 4–6 × 2–4 mm, acute, externally tomentose to sericeous. Leaves with petioles 0.5–1 cm long, strigose; blades elliptic, 2.8–10.1 × 2–6.7 cm, obtuse to round at base, acute at apex (round in young leaves), strongly revolute, leathery, discolorous, glabrous above, tomentose to hirsute below; secondary veins 7–11 on each side of the midrib, impressed to sulcate, tomentose up to 3rd pair above, prominent below; tertiary veins inconspicuous. Inflorescence spiciform, 4.5–6 cm long, 1 per axil, glomerules 2–3 with 4–7 flowers each; peduncle 2.2–3.3 cm long, densely tomentose; bracts ovate-triangular, 0.26–0.6 mm long, tomentose; bracteoles triangular, 2–3.5 × 0.9 mm, externally tomentose, revolute. Flower buds tomentose. Calyx deeply lobed, lobes 1.1–3 × 0.6–2 mm long, tomentose. Corolla white or cream, tube 1.5–3 mm long, externally glabrescent, internally lanate; lobes rounded, 1.3–2 mm long, externally tomentose, internally lanate. Stamens with filaments ca 0.3 mm long; anthers 0.4–09 mm long. Style 1.6–2 mm long in short-styled flowers; 2–3 mm long in long-styled flowers. Fruits fusiform to elliptical, 7 × 4 mm, purple, glabrous; persistent calyx lobes tomentose.

Distribution

Previously known only from open moist forests in Bahia and Sergipe, it is being registered here as occurring in the state of Minas Gerais, in the municipality of Santa Maria do Salto, close to the border with Bahia (Fig. 1D).

Habitat and ecology

Known only from open moist forests.

Phenology

Flowers from March to October; fruits in January, April, and July to September.

Vernacular name

Cipó-prego (Bahia, Ferreira 721).

Etymology

The epithet revolutifolia refers to the strongly revolute leaf margins.

Preliminary IUCN conservation assessment

Vulnerable (VU) B2ab(iii). The extent of occurrence (EOO) of this species was calculated as 136,339 km2, qualifying it as Least Concern (LC). The area of occupancy (AOO) was estimated as 80 km2, falling into the category Endangered (EN) (Bachman et al. 2011; IUCN 2012, 2022). Malanea revolutifolia is known from 27 collections from only ten locations. The locality of Entre Rios and adjacent municipalities have 14 collections, including the type. This specific area is protected by law (Governo do Estado da Bahia, Decreto 1046) since 1992. However, the extraction of the piaçava palm (Attalea funifera Mart.) destroys the surrounding forest. The other collections are all from isolated remaining forest patches in the states of Bahia, Minas Gerais, and Sergipe. Therefore, we consider this species as Vulnerable according to IUCN criteria (IUCN 2012, 2022).

Comment

Although often misidentified as Malanea evenosa, M. revolutifolia is distinguished from it by the hispid to ferruginous-tomentose indumentum in the young branchlets (vs glabrous to pubescent in M. evenosa), leaves with strigose petiole (vs pubescent to sericeous), acute, when young, to rounded apex (vs acute to acuminate), strongly revolute margins (vs slightly revolute), tomentose to hirsute below (vs pubescent with short golden trichomes adpressed), hairy domatia absent (vs present), 7–11 pairs of secondary veins (vs 5–6), secondary veins conspicuous above (vs inconspicuous), and deeply lobed calyx (vs slightly). Before the description of M. revolutifolia (Amaya and Popovkin 2012), specimens with strongly revolute leathery leaves from Bahia were mostly identified as M. sarmentosa, a species restricted to the Amazon. However, M. revolutifolia can be distinguished from that by its flat to inconspicuous tertiary veins on the upper side of the leaves, inflorescences 4.5–6 cm long, deeply lobed calyx, and fusiform to elliptical fruits. Malanea sarmentosa presents reticulate tertiary veins, inflorescences 1.5–2.5 cm long, dentate calyx, and cylindrical fruits (Taylor et al. 2007).

Material examined

BRAZIL – Bahia • Almadina, Serra do Corcovado, acesso pela vertente oposta à de Seu Francisco; 14°42’06”S, 39°36’12”W; 8 Jun. 2012; fr.; Coelho 830; CEPEC • Cardeal da Silva, próximo a BA-400; 16 Feb. 1978; fb.; Orlandi s.n.; RB • Conde, Fazenda do Bu; 12°01’33”S, 37°42’14”W; 1 Jun. 1995; fb., fl.; Ferreira & Silva 721; CEPEC, HUEFS, IPA, JPB • Entre Rios, Areial; 12°15’S, 37°53’W; 60 m; 9 Jan. 2012; fl.; Popovkin 1027; HUEFS • Entre Rios, Areial; 12°15’S, 37°53’W; 60 m; 13 Apr. 2012; fl.; Popovkin 1102; HUEFS • Entre Rios, Areial; 12°15’S, 37°53’W; 60 m; 13 Apr. 2012; fr.; Popovkin 1103; HUEFS • Entre Rios, Areial;12°01’S, 38°02’W; 153 m; 13 Feb. 2012; st.; Popovkin 1060; HUEFS • Entre Rios, Fazenda Experimental da Escola de Medicina Veterinária (UFBA); 11°56’30”S, 38°05’03”W; 24 Oct. 2009; fb.; Roque 2244; ALCB • Entre Rios, Fazenda Rio do Negro;12°01’S, 38°02’W; 150 m; 14 Aug. 2008; fl.; Popovkin 349; HUEFS • Entre Rios, Fazenda Rio do Negro; 12°01’S, 38°02’W; 150 m; 30 Apr. 2011; fl.; Popovkin 877; HUEFS • Entre Rios, Fazenda Rio do Negro; 12°01’S, 38°02’W; 150 m; 11 Jul. 2011; fl., fr.; Popovkin 899, HUEFS • Entre Rios, Fazenda Rio do Negro; 12°53’S, 37°57’W; 153 m; 26 May 2007; fl.; Popovkin 25; HUEFS • Entre Rios, Imbé; 12°04’S, 38°00’W; 15 Mar. 2012; fl.; Popovkin 1075; HUEFS • Entre Rios, rodovia BA 099, km 98 à direita; 12°13’34”S, 37°50’13”W; 50 m; 14 Nov. 2000; fl.; Alves 2030, CEPEC, RB • Ibirapitanga, 22 km N of Itamarati on BR 101, then 6.8 km E on the road to Embratel Tower, Reserva Municipal Cachoeira do Pau; 13°53’27”S, 39°27’33”W; 690 m; 19 May 2003; fl.; Thomas 13443; CEPEC, NY • Itanagra; 12°26’30”S, 38°04’16”W; 14 Sep. 1980; fr.; Gusmão 469; ALCB, PEUFR • Itanagra, Fazenda Brejo Verde; 12°25’S, 38°03’W; 18 Apr. 1976; fl.; Gusmão 397; ALCB, RB • Santa Teresinha, Serra da Jibóia ca 4 km de Pedra Branca; 12°51’10”S, 39°28’32”W; 27 Sep. 2000; fr.; Queiroz 6388; HUEFS • Santa Teresinha, Serra da Jibóia; 12°51’17”S, 41°48’53”W; 15 Apr. 2000; fr.; Silva 55; HUEFS • Santa Teresinha, Serra da Jibóia; 800 m; 12°51’S, 39°28’W, 8 Jul. 1999; fr.; França 3198; HUEFS • Santa Terezinha, Vila da Pedra Branca, Serra da Jibóia, entrada para a torre de tv; 850 m; 11 Nov. 2000; fr.; Alves 2001; CEPEC, RB • Simões Filho; 12°47’04”S, 38°24’14”W; 10 Nov. 1980; fl.; Gusmão 501; ALCB • Uruçuca, estrada de Serra Grande para Uruçuca, área do inventário do plano de manejo do Parque Estadual da Serra do Conduru; 14°29’59”S, 39°06’54”W; 380 m; 2 Apr. 2004; fb., fl.; Fiaschi 2208; CEPEC. – Minas Gerais • Santa Maria do Salto, Povoado de Talismã, Fazenda Duas Barras, Reserva Alto Cariri, trilha do sul que sai em Santa Maria do Salto; 16°24’13”S, 40°03’16”W; 950 m; 22 Apr. 2006; fl.; Amorim 5853; CEPEC. – Sergipe • Linha Verde, km 74 ca 5 km da divisa com o estado da Bahia; 11°47’23”S, 37°55’25”W; 12 Oct. 2000; fr.; Silva 55; HUEFS • Santa Luzia do Itanhi, entrada 2 km à esquerda na estrada Santa Luzia/Crasto; 11°16’06”S, 37°26’19”W; 14 Jun. 1994; fb.; Jardim 449; CEPEC, NY.

Malanea spicata Müll.Arg. (Müller 1881: 457)

Fig. 4K–N

Type

BRAZIL • Rio de Janeiro, Mauá; 8 Apr. 1875; fl., fr.; Glaziou 8166; lectotype (designated here): P [P00507166]; isolectotype: G frag. [G00074054].

Description

Scandent shrub ca 6 m tall; branchlets greyish brown, glabrous. Stipules obovate, 5–7 × 2.5–3 mm, acute, glabrous. Leaves with petioles 0.4–0.84 cm long, glabrous; blades elliptic-lanceolate to obovate, 8–10.4 × 3.1–4.15 cm, acute to cuneate at base, acute to cuneate at apex, margin flat, membranaceous to chartaceous, concolorous, glabrous on both sides; secondary veins 5–7 on each side of the midrib, impressed above, prominent below; tertiary veins inconspicuous. Inflorescence paniculate, 4.5–7 cm long, 1–2 per axil, glomerules (4–)5–7(–8) with 7–10 flowers each; peduncle 1.2–2.1 cm long, strigose; bracts ovate to triangular, 2.2–6.6 mm long, pubescent; bracteoles triangular, 0.1 × 0.2 mm, externally tomentose. Flower buds glabrous to glabrescent. Calyx subtruncate, ca 0.7 mm long, glabrous. Corolla white or cream, tube 1.2–1.5 mm long, externally glabrescent, internally villous; lobes rounded, 1.1–1.5 × 0.5–0.9 mm, externally glabrous to glabrescent, internally villous. Stamens with filaments ca 0.3 mm long; anthers ca 0.5 mm long. Style on short-styled flowers ca 2 mm beneath the mouth of the corolla, on long-styled flowers 3–3.5 mm long above. Fruits fusiform to ellipsoid, 5.2–6 × 3.5 mm, purple, glabrous; persistent calyx lobes tiny.

Distribution

Previously known only from Rio de Janeiro. It is registered here for the first time in Bahia (Fig. 1D).

Phenology

Flowers from December to February; fruits in January, May, and August.

Etymology

The epithet spicata refers to the spiciform inflorescence.

Preliminary IUCN conservation assessment

Endangered (EN) B2ab(iii). The extent of occurrence (EOO) of this species was calculated as 57,711 km2, qualifying it for the category Least Concern (LC). The area of ​​occupancy (AOO) was estimated as 28 km2, falling into the category Endangered (EN) (Bachman et al. 2011; IUCN 2012, 2022). The species has only the type specimen collected in the state of Rio de Janeiro, in 1875. All the other eight collections are from Bahia, from only three locations in a very restricted and fragmented area in southern Bahia (Delprete and Jardim 2019). For this reason, we consider this species as Endangered (EN).

Notes

Müller (1881) described Malanea spicata and cited the gathering Glaziou 8166. One complete specimen of Glaziou 8166 was found in P, a fragment in G, and a photo of the destroyed specimen from B in F. Only the fragment at G has a label handwritten by Müller. Despite that, the specimen at P (P00507166) is being chosen as the lectotype because it is the only one with leaves, flower buds, and flowers in anthesis in good condition, which well characterize this species.

Comment

Malanea spicata is frequently misidentified as M. obovata Hochr. (Hochreutiner 1910). However, it can be distinguished by its membranaceous to chartaceous leaves that are acute to cuneate at the base (vs leathery, attenuate at the base in M. obovata), 5–7 secondary veins on each side (vs 7–9), and the externally glabrous corolla (vs hairy). Furthermore, M. obovata occurs only in the Amazon domain, while M. spicata is endemic to the Atlantic Forest.

Material examined

BRAZIL – Bahia • Belmonte, Estação Ecológica Gregório Bondar; 16°06’11”S, 39°12’26”W; 120 m; 8 Jan. 2002; fr.; Nunes 791; HUEFS • Belmonte, Estação Ecológica Gregório Bondar; 12 Aug. 1981; fr.; Brito 57; CEPEC • Belmonte; 16°08’S, 39°15’W; 12 May 1993; fr.; Thomas 9885; CEPEC, NY [416392] • Itacaré, estrada de Itacaré para Maraú, pouco após a desembocadura do Rio de Contas; 14°15’S, 39°00’W; 12 Dec. 2002; fr.; Fiaschi 1106; CEPEC, NY • Maraú, Rod. BR-030, trecho Porto de Campinhos-Maraú, km 11; 26 Feb. 1980; fb., fl.; Carvalho 214; HUEFS, CEPEC • Maraú, mata costeira; 13 Jan. 1967; fl.; Belém 3117; CEPEC, RB • Ubaitaba-Maraú; 13 Dec. 1967; fb.; Vinha 31; CEPEC. – Rio de Janeiro • Rio de Janeiro, Mauá; 8 Apr. 1875; fr.; Glaziou 8166; RB.

Acknowledgements

We would like to thank all the staff and curators of the herbaria visited; the Graduate Program in Plant Biology (PPGBV) at the Federal University of Pernambuco for financial support for field expeditions; the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES - finance code 001) for funding the RSPR scholarship. We also thank Maria do Céo Rodrigues Pessoa, Jomar Jardim, William Wayt Thomas, and the anonymous reviewers and editor of Plant Ecology and Evolution for their valuable comments and suggestions to the manuscript.

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