Flora of Ceará, Brazil: Vochysiaceae

Abstract

As part of the “Flora of Ceará: Knowing to Conserve” project, this study aimed to contribute to the knowledge of the diversity and distribution of Vochysiaceae species in the state of Ceará. The identifications, descriptions, comments on ecology and distribution, illustrations and distribution maps of the taxa were based on the analysis of samples obtained in the field, in collections deposited in herbaria, and specialized bibliographies. Eight species of Vochysiaceae belonging to four genera occur in Ceará: Callisthene (2), Qualea (2), Salvertia (1), and Vochysia (3). The species were recorded in 41 municipalities. Callisthene fasciculata and Qualea parviflora were the species with the most collection records. The species inhabit different phytoecological units in Ceará and were recorded mainly in the Dense Ombrophilous Forest (Mata úmida), Steppic Savanna (Caatinga/Carrasco), and Neotropical Savanna and Neotropical Savanna (Cerrado). Seven species occur in five different Conservation Areas. Vochysia oppugnata is a new occurrence for the state of Ceará and was recorded in Dense Ombrophilous Forest.

Key words:
diversity; Myrtales; northeastern Brazil; semiarid

Resumo

Vinculado ao projeto “Flora do Ceará: conhecer para conservar”, o estudo teve como objetivo contribuir para o conhecimento da diversidade e distribuição das espécies de Vochysiaceae no estado do Ceará. Com base na análise de amostras obtidas em campo, em coleções depositadas em herbários e bibliografias especializadas foram feitas as identificações, descrições, comentários sobre a ecologia e distribuição, ilustrações e mapas de distribuição dos táxons. No território cearense ocorrem oito espécies de Vochysiaceae pertencentes a quatro gêneros: Callisthene (2), Qualea (2), Salvertia (1) e Vochysia (3). As espécies foram registradas em 41 municípios. Callisthene fasciculata e Qualea parviflora foram as espécies mais coletadas. As espécies habitam diferentes unidades fitoecológicas no Ceará e foram coletadas principalmente em Floresta Ombrófila Densa (Mata úmida), Savana Estépica (Caatinga) e Savana (Cerrado). Sete espécies ocorrem em cinco diferentes Unidades de Conservação. Vochysia oppugnata é uma nova ocorrência para o estado do Ceará e foi registrada na Floresta Ombrófila Densa.

Palavras-chave:
diversidade; Myrtales; nordeste do Brasil; semiárido

Introduction

Vochysiaceae comprises nine genera and ca. 250 species presenting a predominantly neotropical distribution, with the genera Erismadelphus Mildbr. and Korupodendron Litt & Cheek endemic to Africa and the others (Callisthene Mart., Erisma Rudge, Mahechadendron W.Ariza, Cortés-Ballén & Fern.Alonso, Qualea Aubl., Ruizterania Marc.-Berti, Salvertia A.St.-Hil. and Vochysia Aubl.) to the American continent (Litt & Stevenson 2003; Shimizu & Yamamoto 2012a; Souza & Lorenzi 2019). Six genera and 166 species occur in Brazil and 61% (101 spp.) are endemic (Shimizu et al. 2024).

Representatives of the Vochysiaceae are trees, shrubs or rarely subshrubs with leaves simple, opposite or verticillate, petiolate or sessile; inflorescence usually in thyrses, terminal or terminal and axillary or cymes axillary; flowers showy, strongly monosymmetric, with a spurred calyx, most often a reduced number of petals (one, three or five), a single fertile stamen; superior trilocular ovary that develops into a loculicidal capsule or inferior unilocular ovary which matures into an indehiscent winged fruit (Erismadelphus and Erisma), seed winged or non-winged (Litt & Stevenson 2003; Shimizu et al. 2024).

The monophyly of all genera has been supported in recent phylogenetic studies, except Qualea, which has emerged paraphyletic with Ruizterania nested within it (Gonçalves et al. 2020). These same authors suggested that Vochysiaceae originated at approximately 100 m.y., splitting into an Afrotropical lineage (sister to the rest of the family) and a Neotropical one ~50 m.y. Vicariance is currently the most likely explanation for the family’s amphi-Atlantic disjunction pattern (Gonçalves et al. 2020).

Several species of Vochysiaceae, especially those belonging to Qualea and Vochysia, have medicinal properties and are used in folk medicine to treat different diseases (Rodrigues & Carvalho 2001). Qualea parviflora Mart. has antiulcer, anti-diarrheal, antioxidant, and anti-hemorrhagic activity (Mazzolin et al. 2010). Qualea grandiflora Mart. also has antiulcer activity (Hiruma-Lima et al. 2006) and its leaves and bark are used to treat inflammation and wounds (Almeida et al. 1998).

Several taxonomic studies have been carried out on representatives of the Vochysiaceae in Brazil. Among these, we highlight those carried out in the states of São Paulo (Barbosa 1999), Rio de Janeiro (Vianna 2006), Paraná (Negrelle et al. 2007; Negrelle 2011), Minas Gerais (Yamamoto 2009; Shimizu & Yamamoto 2012a, 2012b; Gonçalves et al. 2013, 2017), Goiás (Souza 2014; Sampaio 2017; Teixeira & Teles 2022); Pará (Shimizu & Gonçalves 2017), and in the Distrito Federal (França 1998).

In the Northeastern region of Brazil, representatives of this family have only been studied for Bahia (Kawasaki 1995; França 1996; Passos & França 2003; Azevedo et al. 2015) and Sergipe (Shimizu & Gonçalves 2015). In the state of Ceará, Vochysiaceae species are only mentioned in floristic surveys (Silva et al. 2012; Loiola et al. 2015, 2020; Silveira et al. 2020a, 2020b). This study, as part of the “Flora of Ceará: Knowing to Conserve” project, aims to survey the species of Vochysiaceae occurring in the territory of Ceará.

Material and Methods

Comparative morphological analyses were carried out using collections deposited in the herbaria ASE, CEN, EAC, IPA, HUEFS, HUVA, HVSAF, MAC, PEUFR, R, SPF, and UB (acronyms according to Thiers, continuously updated) and samples obtained in the field. For a detailed analysis of the morphological structures, a Nikon SMZ 1500 stereomicroscope was used.

The authorship of the species was based on IPNI - The International Plant Names Index (2024) and Flora e Funga do Brasil (Shimizu et al. 2024). Species identifications were made according to specialized literature (Stafleu 1948, 1952, 1953; Shimizu & Yamamoto 2012b) and confirmed with exsiccates and images from type collections available on REFLORA (2023) and speciesLink (CRIA 2023a).

Samples collected in the state of Ceará were used to describe the family and species. Additional specimens from other Brazilian states were included to complement the descriptions of some taxa when the samples collected in Ceará were incomplete in regarding floral or fruit characters. For each genus and its respective taxa, specific terminology and description standards were adopted. Morphological characters were described based on Harris & Harris (2001).

Information on habit, species distribution, vegetation type, phenology, and popular names was taken from the labels on the specimens. When there was a lack of information on the location coordinates on the labels of the specimens, georeferencing was carried out using the “geoLoc” tool (CRIA 2023b).

The vegetation types where the species were recorded are in accordance with Figueiredo (1997) and the Technical Manual of Brazilian Vegetation (IBGE 2012): Vegetation Complex of the Coastal Zone (comprising Pioneer Psammophilous Vegetation, Forest behind the Dunes and Lowland Semideciduous Seasonal Forest = Mata de Tabuleiro), Semideciduous Seasonal Forest (Mata seca), Dense Ombrophilous Forest (Mata úmida), Neotropical Savanna (Cerrado), Forested Savanna (Cerradão), Steppic Savanna (Caatinga/Carrasco), and Vegetation under Fluvial and/or Lacustrine Influence (Mata de galeria).

Results and Discussion

Eight species of Vochysiaceae were recognized in the state of Ceará: Callisthene fasciculata Mart., C. microphylla Warm., Qualea grandiflora, Q. parviflora, Salvertia convallariodora A.St.-Hil., Vochysia oppugnata (Vell.) Warm., V. pyramidalis Mart. and V. thyrsoidea Pohl. The species were recorded in 41 municipalities in different phytoecological units, although they were most commonly found in Steppic Savanna (Caatinga) and Neotropical Savanna (Cerrado).

Callisthene fasciculata and Qualea parviflora are the species with the collection records in the state, with 16 and 19 records, respectively. Dense Ombrophilous Forest, Steppic Savanna, and Neotropical Savanna were the phytogeographic units with the most species records, seven, four, and four, respectively. Seven species (88 %) were recorded in five different Conservation Areas: Pedra da Andorinha Wildlife Refuge (C. fasciculata); Aiuaba Ecological Station (C. microphylla); Ubajara National Park (Q. grandiflora); Araripe-Apodi National Forest (Q. parviflora and V. thyrsoidea) and Serra da Ibiapaba Environmental Protection Area (V. oppugnata and V. pyramidalis).

Taxonomic treatment

Vochysiaceae A.St.-Hil., Mém. Mus. Hist. Nat. 6: 265. 1820.

Trees, shrubs, rarely subshrubs; branches cylindrical or quadrangular, suberose or not. Leaves simple, opposite or verticillate, petiolate; stipules deciduous or persistent. Terminal or terminal and axillary thyrse or axillary cincinni (Callisthene). Flower buds fusiform, turbinate or ovoid. Flowers dichlamydeous, monoclinous, zygomorphic; calyx 5-merous, 1 sepal with spur or non-spurred; petals 1, 3 or 5, white, yellow or lilac; stamen 1, anther rimose; ovary superior, 3-carpellate, 3-locular; stigma 1, terminal, subterminal or lateral. Capsule ellipsoid, globose, ovoid or oblong, glabrous or glabrescent.

Identification key of the species of Vochysiaceae occurring in Ceará state

1. Leaves opposite.

2. Cincinni 1-florous; bracts persistent 1.2. Callisthene microphylla

2’. Cincinni 2−3-florous; deciduous bracts.

3. Leaf blade chartaceous to coriaceous; petals lilac 2.2. Qualea parviflora

3’. Leaf blade chartaceous; petals yellow.

4. Stipules persistent; capsule oblong 4.2. Vochysia pyramidalis

4’. Stipules deciduous; capsule ellipsoid.

5. Leaf abaxially ferruginous-tomentose; inflorescence sericeous 2.1. Qualea grandiflora

5’. Leaf abaxially glabrous or pubescent; inflorescence glabrous 1.1. Callisthene fasciculata

1’. Leaves verticillate.

6. Whorl with 6−7 leaves; petals 5, white; ovary tomentose 3. Salvertia convallariodora

6’. Whorl with 4 leaves; petals 3, yellow; ovary glabrous.

7. Leaf coriaceous, margin revolute; style straight; capsule ovoid 4.3. Vochysia thyrsoidea

7’. Leaf chartaceous, margin subrevolute; style curved; capsule oblong 4.1. Vochysia oppugnata

1. Callisthene Mart., Nov. Gen. Sp. Pl. (Martius) 1(4): 123 (1826).

Trees or shrubs; branches cylindrical, straight. Stipules deciduous. Petiole without extrafloral nectaries. Leaves opposite, arranged in phyllomorphic branchlets (similar to a pinnate compound leaf); blade elliptic, lanceolate, oblong or obovate, base rounded, apex acute, rounded, attenuate, retrorse, truncate or emarginate, margin entire, flat, ciliate or not, adaxial and abaxial surfaces glabrous to pubescent, chartaceous to coriaceous, brochidodromous venation. Inflorescence in axillary cincinni, glabrous, 1−3-florous; flowers with one petal; bracts persistent or deciduous; flower buds fusiform or turbinate; spur straight, incurved or subincurved; petals yellow or white; ovary globose, glabrous; style cylindrical, terminal, glabrous; stigma terminal, capitate. Capsule ellipsoid or globose, glabrous.

1.1. Callisthene fasciculata Mart., Nov. Gen. Sp. Pl. (Martius) 1(4): 126 (1826). Figs. 1; 2a-d

Tree or shrub, 2.5−15 m tall. Branchlets with 1−5 pairs of leaves. Petiole 2−4 mm long, pubescent; leaf 4.8−13.5 × 2.2−6 cm, elliptic, lanceolate, oblong or obovate, apex rounded, attenuate or emarginate, adaxial and abaxial surface glabrous or pubescent, chartaceous. Cincinni 2−3-florous; bracts deciduous; pedicels ca. 4 mm long; flower buds fusiform; spur ca. 5 mm long, straight or subincurved; spurred sepal 8−13 × 4−7 mm, ovate, acute apex; non-spurred sepals 2.4−3 × 1.7−2 mm; petal 10−17 × 8−12 mm, yellow with reddish patches and stripes in the central region; stamen 1.5−2 mm long, filament 0.8−1 mm long, anther 0.7−1 mm long; ovary globose; style 10−12 mm long. Capsules 18−25 × 10−14 mm, ellipsoid.

Examined material: Aurora, proximidades do Sítio Santo Domingos, 21.VIII.2014, fr., L.B. Pimentel 268 (HUEFS). Brejo Santo, reservatório Cana Brava, 20.VIII.2013, fr., D.G. Oliveira 1025 (HVASF). Caririaçu, 1.X.1958, fr., R. Schnell (P 4776106). Caucaia, Catarina, 12.VIII.2007, fr., A.S.F. Castro 1977 (EAC); Croatá, à margem da BR-222, 10.XII.1978, fl. and fr., A. Fernandes (EAC 5108). Granjeiro, Rio Granjeiro, 7.XII.1933, fl., P. Luetzelburg 26167 (EAC, IPA). Ibiapina, 4.VI.2012, fr., M.I.B. Loiola 1770 (EAC). Iguatu, Reservatório em formação do AHE Serra da Mesa, 5.V.1998, fr., B.M.T. Walter 4116 (CEN). Lavras da Mangabeira, 06°50’24”S, 39°03’36”W, 16.V.2014, fr., W. Batista 398 (EAC). Milagres, 22.X.1978, fl., A. Fernandes et al. (EAC 5199). Moraújo, Sítio Boa Vista, 10.VI.1988, M.A. Figueiredo & Margarete (EAC 15505). Mucambo, pé da Serra Grande, 29.VII.1976, fl. and fr., A. Fernandes (EAC 2905). Nova Olinda, 4.XII.1981, fl., D. Andrade-Lima et al. 1175 (MAC, IPA). Santa Quitéria, Serra Pau Fincado, São Cosme, 04°13’21”S, 40°1’26”W, 15.VII. 2005, fr., J.R. Lemos 418 (HUEFS). São Luiz do Curu, 27.I.1968, fl., A. Lima 5181 (IPA, PEUFR). Sobral, distrito de Taperuaba, Refúgio de Vida Silvestre Pedra da Andorinha, 04°03’51”S, 39°59’51”W, 6.I.2017, fl., F.F. Araújo 93 (EAC, HUEFS, HUVA).

Figure 1
Geographic distribution of the Vochysiaceae species occurring in the state of Ceará.

Callisthene fasciculata differ from the other species occurring in Ceará by its 2−3-florous cincinni. It differs from C. microphylla by presenting branchlets with 1−5 pairs of leaves (vs. 4−18 pairs of leaves), leaf blade measuring 4.8−13.5 × 2.2−6 cm (vs. 0.6−1.4 × 0.4−0.6 cm), and petals yellow with reddish patches and stripes in the central region (vs. petals white with yellow patches in the central region).

Callisthene fasciculata in Brazil is widely distributed but has not been recorded in the southern region, and occurs in the phytogeographic domains of the Amazon, Caatinga, Cerrado, and Pantanal (Shimizu et al. 2024). It has been recorded in 16 municipalities in Ceará in Neotropical Savanna, Steppic Savanna, Semideciduous Seasonal Forest and in the Vegetation Complex of the Coastal Zone (Lowland Semideciduous Forest). It has only been recorded in the Pedra da Andorinha Wildlife Refuge.

The species was collected with flowers in October, December, and January and with fruits in January, May, June, July, August, October, and December.

Uses for this species are not recorded.

The vernacular name for this species are cravo, cravoeiro.

1.2. Callisthene microphylla Warm., Fl. bras. (Martius) 13(2): 28 (1875). Figs. 1; 2e-f

Tree or shrub, 1−5 m tall. Branchlets with 4−18 pairs of leaves. Petiole ca. 2 mm long, pubescent; leaf 0.6−1.4 × 0.4−0.6 cm, ovate, sometimes oblong, apex straight retuse, truncate or acute, margin flat and ciliate, adaxial surface slightly pubescent to glabrous, abaxial surface pubescent or glabrescent, chartaceous to coriaceous. Cincinni 1-florous; bracts persistent; pedicels 1−2 mm long; floral buds turbinate; spur ca. 1 mm long, incurved or subincurved; sepal spurred 7−8 × 2 mm, ovate, apex acute, sepals non-spurred ca. 3 × 2 mm, ovate, apex acuminate; petal 6−8 × 5−8 mm, withe with yellow patches in the central region; stamen ca. 1.5 mm long, filament ca. 0.5 mm long, anther ca. 1 mm long; ovary globose; style ca. 4−6 mm long. Capsules 8−10 × 5−7 mm, globose.

Examined material: Aiuaba, Estação Ecológica de Aiuaba, 9.VI.2005, fr., J.R. Lemos & P. Matias (EAC 38373). Guaramiranga, Sítio Arábia, 19.III.1989, fl. and fr., M.A. Figueiredo et al. (EAC 16393). Maranguape, 26.XI.1955, fl. and fr., A. Lima & A. Fernandes (EAC 1529). Meruoca, Serra da Meruoca, Sítio Lages, 15.II.1957, fr., A. Fernandes (EAC 1698). Mucambo, encosta da Serra da Ibiapaba, 8.IX.2001, fr., A.S.F. Castro 1054 (EAC). Parambu, 19.X.2009, fr., R.G. Ferreira (EAC 45498). Pereiro, Fazenda Campos, 6.VII.1988, fl., A. Fernandes & M.A. Figueiredo (EAC 15893). Sobral, Fazenda Andorinha, 25.IV.1980, fr., A. Fernandes & E. Nunes (EAC 8549). Viçosa do Ceará, Chapada da Ibiapaba, Cocalzinho, 5.I.1987, fl., A. Fernandes & M.A. Figueiredo (EAC 14874).

Additional material: BRAZIL. PIAUÍ: Gilbués, estrada Bom Jesus, 27.XI.1979, fl., A. Fernandes & Matos (EAC 7236).

Callisthene microphylla is a well-defined species and its main characteristics are branchlets with 4−18 pairs of leaves; reduced leaf blade size (0.6−1.4 × 0.4−0.6 cm); cincinni 1-florous and petals white with yellow patches in the central region. These characteristics differentiate it from C. fasciculata which has branchlets with 1−5 pairs of leaves glabrous; leaves 4.8−13.5 × 2.2−6 cm; cincinni 2−3-florous, and petals yellow with reddish patches and stripes in the central region.

Callisthene microphylla is endemic to Brazil, with records in the Amazon, Caatinga, and Cerrado phytogeographic domains in different regions, except the southern region (Shimizu et al. 2024). In Ceará, it was found in seven municipalities in the Steppic Savanna and Dense Ombrophilous Forest. It was only recorded at the Aiuaba Ecological Station.

The species was collected with flowers in January, June, October, and November, and with fruits in the months of February, April, June, September, October, and November.

Uses for this species are not recorded.

There is no vernacular name recorded for this species.

2. Qualea Aubl., Hist. Pl. Guiane 1: 5, t. 1,2 (1775).

Trees, shrubs or subshrubs. Branches cylindrical, tortuous. Stipules deciduous. Petiole glabrous, sericeous or pubescent, with 2 extrafloral nectaries at the base. Leaves opposite; leaf blade narrowly elliptic, elliptic, lanceolate or obovate, base acute, cordate or rounded, apex acute, acuminate, attenuate or rarely emarginate, margin entire, flat or repand, abaxial and adaxial surface glabrous or sericeous, chartaceous or coriaceous, venation brochidodromous. Inflorescence in terminal or terminal and axillary thyrses; bracts deciduous; flower buds ovoid; spur straight or incurved; flowers with one petal; 1 sepal with spur, elliptic or oval; petals yellow or lilac, glabrous; ovary globose, tomentose; style cylindrical, glabrous or with trichomes at the base; stigma terminal, subcapitate or capitate. Capsule ellipsoid or globose, glabrous.

Figure 2
a-i. Species of Vochysiaceae occurring in state of Ceará - a-d. Callisthene fasciculata - a. branch with leaves and inflorescences; b. inflorescence; c. flower; d. fruits; e-f. Callisthene microphylla - e. branch with fruit; f. leaves and fruit details; g-i. Qualea grandiflora - g. leaves; h. inflorescence; i. fruit. (Photos: a-d, g. EB de Souza; e-f. RT de Queiroz; i. MIB Loiola).

2.1. Qualea grandiflora Mart., Nov. Gen. Sp. Pl. (Martius) 1(4): 133, t. 79 (1826). Figs. 1; 2g-i

Tree 4−10 m tall. Petiole 0.5−2 cm long, sericeous; leaf blade 5.3−15 × 3.3−6 cm, lanceolate, obovate or elliptical, base cordate, apex acute or acuminate, margin flat or repand, adaxial surface glabrous, sometimes sericeous on the veins, abaxial surface ferruginous-tomentose on the veins main and secondary, chartaceous. Thyrse 8−16 cm long, terminal, sericeous; peduncles 2−5 cm long, pedicels 0.6−1.3 cm long; flower buds ovoid; spur 1−1.4 cm long, curved; sepal spurred 2−3 × 2−2.2 cm, calyx lobes non-spurred 1−2 × 1−1.7 cm, elliptical or ovate, apex obtuse, base truncated; petal 4−5 × 5−6 cm, yellow; stamen 2−3 cm long, filament 1.6−2 cm long, anther 0.7−0.9 cm long; ovary globose, tomentose; style 1.4−1.8 cm long, cylindrical, glabrous; terminal stigma, subcapitated. Capsules 4.5−6.5 × 1.5−2 cm, ellipsoid.

Examined material: Camocim, 5.I.2013, fr., A.S.F. Castro 2677 (EAC). Guaraciaba do Norte, Chapada da Ibiapaba, 30.IV.1994, fl., A. Fernandes & Matos (EAC 21225). Ibiapina, Açude do Jacaré, 4.VI.2012, fl., M.I.B Loiola et al. 1769 (EAC). Pacujá, Serrinha, 25.VIII.2009, fr., M.F. Mata 2027 (HUEFS). Tianguá, Santa Bárbara, 17.IX.2012, fr., M.I.B. Loiola et al. 1887 (EAC). Ubajara, Parque Nacional de Ubajara, trilha cachoeira do Cafundó, 28.VIII.2012, fr., M.I.B. Loiola et al. 1979 (EAC). Viçosa do Ceará, Serra das Flores, 3.VIII.2005, fr., L.W. Lima-Verde et al. 3403-05 (EAC).

Qualea grandiflora differs from Q. parviflora by its chartaceous leaves (vs. coriaceous), petal yellow (vs. lilac) and capsule ellipsoid (vs. ovoid or globose).

Qualea grandiflora has been confirmed in all Brazilian regions in the phytogeographic domains of Amazon, Caatinga, Cerrado, and Atlantic Forest (Shimizu et al. 2024). In the northeastern region, there are records in the states of Bahia, Ceará, Maranhão, and Piauí (Shimizu et al. 2024). In Ceará, there are records in seven municipalities in Dense Ombrophilous Forest, Neotropical Savanna, Steppic Savanna, and Semideciduous Seasonal Forest vegetation. The specie was registered in the Ubajara National Park.

The species was collected with flowers in March, April, and June and with fruits in March, August, and September.

The species has antiulcer activity (Hiruma-Lima et al. 2006); leaves and bark are used to treat inflammations and wounds (Almeida et al. 1998).

The vernacular name for this species is pau-terra.

2.2. Qualea parviflora Mart., Nov. Gen. Sp. Pl. (Martius) 1(4): 135, t. 81 (1826).Fig. 1

Tree or shrub 2−20 m tall. Petiole 0.5−1.4 cm long, glabrous, sericeous or pubescent; leaf blade 2.3−8.6 × 0.8−4 cm, narrowly elliptic, elliptic or ovate, base acute or rounded, apex acuminate, rounded, attenuate or rarely emarginate, margin flat, adaxial and abaxial surface sericeous to densely sericeous, cinereous trichomes, chartaceous to coriaceous. Thyrse 6−8 cm long, terminal and axillary, pubescent; peduncles absent; pedicels 0.4−1 cm long; floral buds ovoid; spur 4−6 mm long, straight or incurved; sepal spurred 5−6 × 3−6 mm, calyx lobes non-spurred 2−3 × 4−6 mm, ovate, apex obtuse, base truncate; petal 1.4−1.5 × 1.7-2 cm, lilac; stamen 6−8 mm long, filament 6−7 mm long, anther ca. 1 mm long; ovary globose, tomentose; style ca. 7 mm long, cylindrical, trichomes at base; stigma terminal, capitate. Capsules 2−3 × 0.8−1.5 cm, globose or ovoid.

Examined material: Barbalha, FLONA do Araripe-Apodi, proximidades da casa de guarda da Santa Rita, 07°23’02”S, 39°21’16”W, 10.XII.2014, fl., V.M. Mascena 241 (EAC). Cariri, 25.III.1936, fl., P. Luetzelburg 26206 (EAC). Crato, FLONA do Araripe-Apodi, 4.IX.2001, fr., I.R. Costa 386 (EAC). Cruz, 02°52’48”S, 40°19’12”W, 29.X.2013, fr., W. Batista 44 (EAC). Frecheirinha, próximo a BR-222, 24.VIII.2004, fr., L.W. Lima-Verde et al. 3031 (EAC). Granja, entre Martinópole e Granja, cerca de 10 km de Granja, 03°11’15”S, 40°44’38”W, 11.IX.2014, fr., E.B. Souza et al. 3166 (EAC, HUVA). Granjeiro, 7.XII.1933, fl., P. Luetzelburg 26171 (EAC, IPA). Guaraciaba do Norte, Chapada da Ibiapaba, 28.XII.1979, fl., A. Fernandes (EAC 7890). Ipueiras, Buriti, 21.VII.1979, fr., S. Jordy Filho 58 (IPA, ASE). Jardim, proximidades da FLONA do Araripe-Apodi, 22.I.2014, fl. and fr., C.E.B. Proença et al. 4680 (UB). Lavras da Mangabeira, Serra do Boqueirão, 8.IV.1982, fr., M.A. Figueiredo (EAC 11226). Martinópole, estrada para Bom Princípio, 2 km da sede municipal, 03°11’54”S, 40°41’03”W, 21.VII.2017, fl. and fr., E.B. Souza et al. 4717 (EAC, HUEFS). Mauriti, Umburanas, 18.VIII.2016, fr., A.P. Fontana 9624 (HUEFS). Missão Velha, Nova Olinda, 07°6’36”S, 39°38’36”W, 19.VIII.2011, fr., E. Melo 10334 (HUEFS). Mucambo, Pé da Serra Grande, 29.VII.1976, fl. and fr., A. Fernandes (EAC 2904). Pacujá, Planalto Ibiapaba, Serrinha, Distrito Bom Gosto, 23.I.2009, fl., E.B. Souza 1676 (EAC, HUEFS). Reriutaba, estrada para Guaraciaba do Norte, 7.III.1981, fl. and fr., A. Fernandes & E. Nunes (EAC 9886). São Benedito, Faveira, 27.V.1981, fr., A. Fernandes & P. Martins (EAC 10388, SPF 64750). Tianguá, caminho para o Sítio do Bosco, 7.VI.2012, fr., M.I.B. Loiola et al. 1865 (EAC). Viçosa do Ceará, Limoeiro, 15.XII.2006, fl., A.S.F. Castro 1898 (EAC).

Qualea parviflora is easily recognized by its small leaves (2.3−8.6 × 0.8−4 cm), with adaxial surface sericeous to densely sericeous and abaxial surface with cinereous trichomes, petal lilac, and capsule with 2−3 × 0.8−1.5 cm, ovoid or globose. These characteristics differentiate it from Q. grandiflora, which has larger leaves (5.3−15 × 3.3−6 cm), with adaxial surface glabrous, sometimes sericeous on the veins and abaxial surface ferruginous-tomentose on the main and secondary veins, petals yellow, and capsule with 4.5−6.5 × 1.5−2 cm, ellipsoid.

Qualea parviflora has a wide distribution in different regions of Brazil, except for the South, and is found in the Amazon, Caatinga, Cerrado, and Atlantic Forest phytogeographic domains (Shimizu et al. 2024). It has been recorded in 19 municipalities in Ceará in Neotropical Savanna, Steppic Savanna, Dense Ombrophilous Forest, and Semideciduous Seasonal Forest vegetation. It was recorded in the Araripe-Apodi National Forest.

The species was collected with flowers in January, March, July, and December and with fruits in January, March, April, May, June, July, August, September, and October.

Qualea parviflora has antiulcer, anti-diarrheal, antioxidant and anti-hemorrhagic activity (Mazzolin et al. 2010).

The vernacular names for this species are pau-terra-roxo; pau-terra; pau-terra-da-folha-miúda.

Iconography: Azevedo et al. (2015: 36, fig. 5a-c).

3. Salvertia convallariodora A.St.-Hil., Mém. Mus. Hist. Nat. 6: 266 (1820).Fig. 1

Tree 10−14m tall. Branches cylindrical. Stipules deciduous. Petiole 1.5−2.5 cm long, glabrescent to glabrous. Leaves verticillate, 6−7 leaves per whorl; leaf blade 8.2−16 × 5−10.5 cm, obovate, apex emarginate or rounded, base cuneate, margin entire, flat, adaxial surface glabrous and abaxial surface glabrous, coriaceous, venation brochidodromous. Thyrse 25−35 cm long, terminal, pubescent; bracts deciduous; peduncles 2−3.5 cm long; pedicels 1−1.5 cm long; flower buds ellipsoid; spur 0.7−2 cm long, incurved or straight; sepal spurred 3−3.5 cm long, calyx lobes non-spurred 2−2.5 cm long, oblong or obovate, base truncate, apex obtuse; petals 5, with 2.3−3 × 1.2−1.5 cm, subequal, obovate, glabrous, white; stamen ca. 2.5 cm long, filament ca. 1 cm long, anther ca. 1.5 long; ovary pyramidal, tomentose; style ca. 0.4 cm long, clavate, glabrous; stigma lateral, ligulate. Capsule ca. 3.5 × 1 cm, ovoid, glabrescent.

Examined material: Granja, 03°11’15”S, 40°44’40”W, 6.VIII.2014, fl., E.B. Souza et al. 3151 (EAC). Viçosa do Ceará, Serra das Flores, 3.VIII.2005, fl., L.W. Lima-Verde 3403-11 (EAC).

Additional material: BRAZIL. PIAUÍ: Oeiras, 07°01’04”S, 42°08’47”W, 25.I.2014, fr., B.M.T. Walter et al. 6635 (CEN).

Salvertia convallariodora is distinguished from other Vochysiaceae species occurring in Ceará by its verticillate leaves, with 6−7 leaves per whorl and thyrses 25−35 cm long. The species also has peculiar characters, such as ovary pyramidal, style clavate and stigma lateral and ligulate.

Salvertia convallariodora occurs throughout the Central-West region and part of the Northeastern, Northern and Southeastern regions, in the Amazon, Caatinga, and Cerrado phytogeographic domains (Shimizu et al. 2024). It is a species with few records in Ceará, occurring in only two municipalities in Neotropical Savanna vegetation.

The species was collected with buds and flowers in August.

Uses for this species are not recorded.

There is no vernacular name recorded for this species.

Iconography: Gonçalves et al. (2017: 176, fig. 5a-g).

4. Vochysia Aubl., Hist. Pl. Guiane 18 (1775), nom. et orth. cons.

Tree. Branches cylindrical or quadrangular. Stipules persistent or deciduous, not associated with extrafloral nectaries. Leaves opposite or verticillate, narrowly elliptic, elliptic, lanceolate, oblong or obovate, margin entire, flat, revolute or subrevolute, chartaceous or coriaceous, venation brochidodromous. Inflorescence a foliose thyrse of cincinni, terminal or terminal and axillary, cincinni 2−5-florous; bracts deciduous; flower buds ellipsoid or fusiform; spur straight, incurved or recurved; sepals 5, of which 1 with spur; petals 3, one larger and two smaller, yellow; ovary deltoid, glabrous; style straight or curved, glabrous; stigma terminal, subterminal or lateral, capitate, trilobed or triangular. Capsule ovoid or oblong, glabrous.

4.1. Vochysia oppugnata (Vell.) Warm., Fl. bras. (Martius) 13(2): 87 (1875).Fig. 1

Tree. Branches quadrangular or cylindrical. Stipules persistent. Leaves verticillate, 4 leaves per whorl; petiole 1−2 cm long, glabrous; leaf blade 6.6−12.1 × 2.1−4.1 cm, obovate or narrowly elliptic, base attenuate, apex emarginate, margin subrevolute, adaxial and abaxial surfaces glabrous, chartaceous. Thyrse 3.5−8 cm long, terminal, glabrescent; bracts deciduous; peduncles 1−2 cm long; cincinni 2−5-florous; pedicels 0.5−1.7 cm long; flower buds fusiform; spur 0.7−1.1 cm long, incurved or recurved; sepal spurred ca. 1.8 cm, calyx lobes non-spurred 1.2−1.5 cm; central petal 8−10 × 3−4 mm, lateral petals 6−9 × 3−4 mm, yellow; stamen ca. 1.2 cm long, filament ca. 2 mm long, anther ca. 1 cm long; ovary deltoid, glabrous; style ca. 1.3 cm long, curved, glabrous; stigma subterminal, trilobed. Capsule 0.5−1.1 × 0.3−0.5 cm, oblong, glabrous.

Examined material: São Benedito, Serra da Ibiapaba, 6.I.1942, fl., P. Bezerra 353 (EAC); Contendas, 8.II.2018, fr., A.S.F. Castro 3018 (EAC).

Additional material: BRAZIL. BAHIA: Água Quente, base do Pico do Itoibira, 20.VI.1998, fr., FH Nascimento & EA Sousa 4 (HUEFS). Rio de Contas, Serra das Almas, Campo do Queiroz, Mata do Cafundó, 13º30’43”S, 41º57’45”W, I.2006, fl., F.H.F. Nascimento 529 (HUEFS).

Vochysia oppugnata can be confused with V. thyrsoidea because both have leaves verticillate, 4 leaves per whorl, petals yellow, and ovary deltoid. However, the first species has stipules persistent (vs. deciduous), leaves chartaceous (vs. coriaceous), flower buds fusiform (vs. ellipsoid), and capsule oblong (vs. ovoid).

Vochysia oppugnata is endemic to Brazil and occurs only in three regions: Northeastern (Bahia), Central-West (Distrito Federal), and Southeastern (Minas Gerais, Rio de Janeiro and São Paulo), in the Atlantic Forest domain (Shimizu et al. 2024). In Ceará it is represented by only one collection from 1942 and other from 2018, collected in Dense Ombrophilous Forest. This is a new record of occurrence, characterizing a disjunct distribution.

The species was collected with flowers in January and with fruits in February.

Uses for this species are not recorded.

The vernacular name for this species is piroá.

Iconography: Gonçalves et al. (2017: 185, fig. 7f-i).

4.2. Vochysia pyramidalis Mart., Nov. Gen. Sp. Pl. (Martius) 1(4): 148, t. 90 (1826).Fig. 1

Tree 3−5m tall. Branches cylindrical. Stipules persistent. Leaves opposite; petiole 0.7−1.1 cm long, glabrescent; leaf blade 5.5−12.5 × 1.8−4 cm, narrowly elliptic, lanceolate or oblong, base obtuse or rounded, apex acute or acuminate; margin entire, flat, adaxial surface glabrous, abaxial surface ferruginous-tomentose, chartaceous. Thyrse 5.5−23.5 cm long, terminal and axillary, ferruginous-tomentose; bracts deciduous; peduncles 1.3-5 cm long; cincinni 2−4-florous; pedicels 0.5−1 cm long; flower buds ellipsoid; spur 0.3−1 cm long, incurved, cylindrical, pubescent in flower buds, glabrous in open flowers; sepal spurred 0.8−1.5 cm long, calyx lobes non-spurred 0.8−1.3 mm long; central petal ca. 1.3 × 0.4 cm, lateral petals ca. 0.7 × 0.2 cm, yellow; stamen ca. 0.8 cm long, filament ca. 0.3 cm long; anther ca. 0.5 cm long; ovary deltoid, glabrous; style ca. 1.8 cm long, cylindrical, curved, glabrous; stigma lateral, triangular. Capsule 0.8−1 × 0.2 cm, oblong.

Examined material: Ibiapina, próximo do riacho, 22.X.1976, fl., A. Fernandes & F.J.A. Matos (EAC 2951). Ipueiras, Brejo, 4.I.2008, fl., A.S.F. Castro 1998 (EAC). São Benedito, Planalto da Ibiapaba, Sítio Cigarra, 4.XII.1989, fl., M.A. Figueiredo (EAC 19829). Viçosa do Ceará, Uruoca, Chapada da Ibiapaba, 6.I.1987, fl., A. Fernandes & M. Angélica (EAC 14912). Ubajara, Ibiapaba do Norte, 11.X.1992, fl., F.S. Aráujo & M.A. Figueiredo (EAC 23916).

Vochysia pyramidalis differs from V. oppugnata and V. thyrsoidea in having leaves opposite (vs. verticillate), leaf blade abaxial surface, and thyrse ferruginous-tomentose (vs. glabrescent). The species also has, as striking characters, the style curved, stigma lateral, triangular, and capsule oblong.

Vochysia pyramidalis is endemic to Brazil, with records in the Northern, Northeastern, Mid-Western, and Southeastern regions in the Caatinga, Cerrado, and Atlantic Forest phytogeographic domains (Shimizu et al. 2024). It occurs in four municipalities in the state of Ceará, predominantly in Dense Ombrophilous Forest vegetation, but has also been recorded in Steppic Savanna and Forested Savanna.

The species was collected with flowers in January, October, and December.

Uses for this species are not recorded.

The vernacular names for this species are murici-raposa; preguiça; piroá-branco.

Iconography: Sampaio (2017: 97, fig.13a-d).

4.3. Vochysia thyrsoidea Pohl, Pl. Bras. Icon. Descr. 2(1): 24, pl. 115 (1828).Fig. 1

Tree 5-7m tall. Branches quadrangular. Stipules deciduous. Leaves verticillate, 4 leaves per whorl; petiole 4−6 mm long, glabrescent; leaf blade 4−9.2 × 1.9−4.5 cm, elliptic or obovate, apex emarginate, base acute or rounded, margin revolute, adaxial and abaxial surfaces glabrous, coriaceous. Thyrse 14−22 cm long, terminal, glabrescent; bracts deciduous; peduncles 1.5−2.4 cm long; cincinni 2−3-florous; pedicels 0.5−1 cm long; flower buds ellipsoid; spur ca. 1.2 cm long, straight or recurved; sepal spurred 2.5−3 cm long, calyx lobes non-spurred ca. 1.5 cm long; central petal ca. 0.8 × 0.3 cm, lateral petals ca. 0.6 × 0.1 cm, yellow; stamen ca. 1.2 cm long; filament ca. 2 mm long, anther ca. 1 cm long; ovary deltoid, glabrous; style ca. 1.5 cm long, straight, glabrous; stigma terminal, trilobed. Capsule ca. 3 × 1.6 cm, ovoid.

Examined material: Aracati, Gardner (IPA 40587). Crato, Floresta Nacional do Araripe-Apodi, 6.XI.1985, fl., A. Fernandes & Matos (EAC 13964). Em local não determinado, Araripe, fl., F.F. Allemão & M. Cysneiros 592 (R).

Additional material: BRAZIL. BAHIA: Campo Formoso, Morro do Cruzeiro, 15.V.1999, fr., F. França et al. 2934 (MBM).

Vochysia thyrsoidea is a well-defined species and has peculiar characteristics, such as the whorl with 4 leaves coriaceous, flower buds ellipsoid, style straight, and capsule ovoid. This set of characteristics differentiates it from V. oppugnata, which has leaves chartaceous, flower bud fusiform, style curved, and capsule oblong.

Vochysia thyrsoidea is endemic to Brazil with disjunct occurrence recorded in the states of Bahia, Ceará, Distrito Federal, Goiás, Minas Gerais, and São Paulo in the Cerrado phytogeographic domain (Shimizu et al. 2024). It is registered in only two municipalities in Ceará in Forested Savanna and Vegetation Complex of the Coastal Zone (Lowland Semideciduous Forest). The species was recorded in the Araripe-Apodi National Forest.

The species was collected in flower in November.

Uses for this species are not recorded.

The vernacular name for this species is pau-gamela.

Iconography: Azevedo et al. (2015: 38, fig.7a-f).

Acknowledgements

We are grateful for the funding grant awarded by CNPq to the first author. LMS is grateful to CAPES (Coordenação de Aperfeiçoamento de Pessoal de Nível Superior), for the scholarship (Financial Code 001); MIBL to Conselho Nacional de Desenvolvimento Científico e Tecnológico, for the research productivity scholarship granted (Process 306723/2023-9), and Fundação Cearense de Apoio ao Desenvolvimento Científico e Tecnológico (Edital Mulheres na Ciência - Process MLC-0191-00147.01.00/22). The authors thank Elnatan Bezerra de Souza and Rubens Teixeira de Queiroz, for providing photos of the species.

Data availability statement

In accordance with Open Science communication practices, the authors inform that all data are available within the manuscript

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Edited by

• Area Editor:
  Dr. Marcelo Trovó

Publication Dates

• Publication in this collection
  02 Dec 2024
• Date of issue
  2024

History

• Received
  15 Jan 2024
• Accepted
  19 June 2024