ABSTRACT
(Flora of Pernambuco (Brazil): Lythraceae J. St.-Hil.). Lythraceae comprises about 620 species distributed in tropical and subtropical regions of the globe, many of them of ecological and medicinal importance. This work aimed to perform the floristic study of Lythraceae species occurring in Pernambuco (Brazil), in addition to providing an identification key, morphological descriptions, geographical distribution, taxonomic notes, phenophases, photographs and illustrations. Materials obtained from field expeditions and from 10 herbaria were analyzed. Seventeen species and five genera (Ammannia, Cuphea, Lafoensia, Pleurophora and Rotala) were recognized. New records are: C. anagalloidea for Pernambuco, C. mimuloides and P. anomala for the Atlantic Forest and L. glyptocarpa and R. ramosior for the Caatinga. Seven species are endemic from Brazil, two are endemic from the Northeast and one is endemic from Pernambuco. The most frequent species were P. anomala, C. flava and C. racemosa. The species predominate in Caatinga of the State and 15 (88,23%) of them occur in conservation units.
Keywords:
BFG;
Cuphea
; Diversity; Floristic; Myrtales
RESUMO
(Flora de Pernambuco (Brasil): Lythraceae J. St.-Hil.). Lythraceae compreende cerca de 620 espécies distribuídas em regiões tropicais e subtropicais do globo, muitas delas de importância ecológica e medicinal. Este trabalho objetivou realizar o estudo florístico das espécies de Lythraceae ocorrentes em Pernambuco (Brasil), além de fornecer uma chave de identificação, descrições morfológicas, distribuição geográfica, notas taxonômicas, fenofases, fotografias e ilustrações. Materiais obtidos de expedições de campo e 10 herbários foram analisados. Dezessete espécies e cinco gêneros (Ammannia, Cuphea, Lafoensia, Pleurophora e Rotala) foram reconhecidos. São novos registros: C. anagalloidea para Pernambuco, C. mimuloides e P. anomala para a Mata Atlântica e L. glyptocarpa e R. ramosior para a Caatinga. Sete espécies são endêmicas do Brasil, duas são endêmicas do Nordeste e uma é endêmica de Pernambuco. As espécies mais frequentes foram P. anomala, C. flava e C. racemosa. As espécies predominam na Caatinga do estado e 15 (88,23%) delas ocorrem em Unidades de Conservação.
Palavras-chave:
Cuphea
; Diversidade; Flora do Brasil; Florística; Myrtales
Introduction
Lythraceae (Myrtales) comprises about 32 genera and 620 species (Xu & Deng 2017) that are widely distributed in tropical and subtropical regions, with few records in the temperate ones (Cavalcanti 2007). In Brazil, 13 genera and 230 species are recognized, predominantly distributed in Cerrado, Atlantic Forest and Amazon Forest domains, and 168 are reported as endemic to the country (BFG 2022). According to BFG (2022), 17 species in five genera are reported to Pernambuco State, being the second State in the Northeast region with the largest number of species.
Lythraceae species occurs in several habitats, including swamps, dunes, humid tropical forests, but more frequently in savannah, arid and rocky fields. Its representatives are often shrubs with simple leaves, commonly decussed; Inflorescence raceme; Flowers bisexual, floral tube persistent, petals 4-16, dialipetalous; Fruit capsule (Cavalcanti & Graham 2002).
The family includes species with different potentialities. Cuphea brachiata Koehne is considered a dune fixer, as it colonizes open areas in restinga environments allowing the establishment of other species (Pigozzo et al. 2006). Lafoensia glyptocarpa Koehne and Lagerstroemia indica L. are widely used as ornamental (Souza & Lorenzi 2019). Cuphea carthagenensis (Jacq.) J.F. Macbr., is either indicated for prevention and treatment of cardiovascular diseases (Schuldt et al. 2000). Cuphea seeds are antioxidant sources that delay the fat and oil degradation in stored food products, extending the validity of products (Graham 1991). The wood of Lafoensia pacari A.St.-Hil. is used in building (Carvalho 2003).
Several floristic and taxonomic studies were conducted focused on the family were in the Brazilian flora, as the Flora of Grão-Mogol, Minas Gerais (Cavalcanti 2004); the Flora of São Paulo (Cavalcanti & Graham 2002;the taxonomic treatment of the genus Diplusodon Pohl (Cavalcanti 2022); and others. More specifically in the Northeast region, where is located our study area, Lythraceae was monographed in the floras of Ceará (Silva et al. 2023), of Usina São José, Igarassu, Pernambuco (Lima et al. 2020) and genus Cuphea was monographed for the Chapada Diamantina National Park, Bahia (Brauner 2017). Besides that, the family appears in species listings such as the one for the Caatinga of the São Francisco River, Bahia (Cavalcanti 2012).
Despite its economical and ecological importance, Lythraceae collections in Herbaria remain incorrectly or unidentified. Floristic inventories are crucial for the real knowledge of the diversity, especially in those areas under threat (Barbosa et al. 2021), such as the Caatinga (Harold 2024) and Atlantic Forest (Scarano & Ceotto 2015), where our study was conducted. We aim to present the floristic study of Lythraceae species occurring in Pernambuco (Brazil) territory, to provide subsidies for correct identification of species, contributing to the knowledge of the ecological and economic importance of these species, in addition to providing a basis for future studies of evolution and conservation of the family.
Material and methods
Study area - Pernambuco is located in the Center-east of the Northeast region of Brazil, between the coordinates 7º15’45” and 9º28’18” South and 34º48’35” and 41º19’54” West. It has an area of 98.311 km², with greater extension from east to west and is limited by the Atlantic Ocean and by the States of Alagoas, Bahia, Ceará, Paraíba and Piauí. According to Koppen’s (1936) climate classification system, the State is classified as BSh and Aw. It presents heterogeneous soil and phytophysiognomies, with fragments of ombrophilous and seasonal forests, forested highlands, “restingas”, mangrove bodies, dunes, associated ecosystems and Caatinga. Caatinga stands out as predominating about 80% of the total area of the State, with semi-arid climate and is classified into hyperxerophilic and hypoxerophilic areas (Andrade-Lima 2007).
Material collection - Eleven field expeditions were conducted between 2018 and 2020 to collect fertile material, including the following municipalities: Belo Jardim, Buíque, Cabo de Santo Agostinho, Cabrobó, Camaragibe, Ibimirim, Igarassu, Inajá, Ipojuca, Jaboatão dos Guararapes, Pesqueira, Petrolina, Recife, São Lourenço da Mata and Tamandaré. The selected areas for the fieldworks included areas considered priority according to occurrence records to obtain data on the ecological conditions of the species, and preserved areas, in order to encompass the largest possible number of municipalities in Pernambuco. The collected specimens were dehydrated, flowers and fruits were stored in 70% alcohol and leaves stored in silica gel for future studies. The collection and herborization process was based on conventional techniques for taxonomic studies (Bridson & Forman 1998). The specimens were deposited in the PEUFR herbarium, and duplicates will be sent to HVASF, CEN, MO and K (herbaria acronyms follow Thiers, continuously updated).
A total of 680 specimens from ten herbaria were analyzed: CEN, HESBRA, HST, HTSA, HUEFS, HVASF, IPA, JPB, PEUFR and UFP.
Morphological analysis - The species were identified from comparisons to species previously identified by specialist, consults to virtual herbaria platforms [REFLORA (Brazil), Missouri Botanical Garden (MO), the New York Botanical Garden (NY), JSTOR and SpeciesLink] and based on specific bibliography (Graham 1975, Graham 1985, Lourteig 1986, Cavalcanti & Graham 2002, Siqueira-Filho et al. 2015, Graham 2017, Brauner 2017, BFG 2022).
Specific terminologies of morphological descriptions for vegetative and reproductive structures followed Gonçalves & Lorenzi (2011), except for seed shape that followed Harris & Harris (2001). Due to the large amount of examined materials, the best quality fertile specimens from each phytogeographic domain (Caatinga and/or Atlantic Forest) and that represent the phenological period were selected and presented as “selected material examined”. The remaining specimens analyzed are included in an overall list as “examined material”.
The maps of the species richness (figure 1) and distribution patterns (figure 2) were created using DIVA-GIS (Hijmans et al. 2001) and QGIS (QGIS 2021) softwares, respectively. The original or municipal geographic coordinates were obtained through field expeditions, examined collections labels and on SpeciesLink.
Distribution map of Lythraceae species in Pernambuco (Brazil). a. Ammannia auriculata Willd, A. latifolia L., Lafoensia glyptocarpa Koehne. b. Cuphea anagalloidea A. St.-Hil, C. campestris Mart. ex Koehne, C. carthagenensis (Jacq.) J.F. Macbr. c. C. circaeoides Sm. ex Sims., C. ericoides Cham. & Schltdl., C. flava Spreng, C. impatientifolia A. St.-Hil. d. C. micrantha Kunth, C. mimuloides Schltdl. & Cham, C. pulchra Moric., C. racemosa (L. f.) Spreng. e. Pleurophora anomala (A. St.-Hil.) Koehne, P. pulchra J.C. Siqueira, Cotarelli, J.F.B. Pastore & T.B. Cavalc., Rotala ramosior (L.) Koehne.
Results and Discussion
In this study, 17 species in five genera of Lythraceae for Pernambuco were recognized, Ammannia L., Cuphea P. Br., Lafoensia Vand., Pleurophora D. Don., and Rotala L. (table 1). Few cultivated species were found in the study area, but are not included in this floristic treatment, as Lagerstroemia indica L., largely used as ornamental. The species identified as C. strigulosa Kunth, previously mentioned in BFG (2022) and referring to voucher H. C. H. Silva 3 (MO 3412570), represents C. carthagenensis (Jacq.) J.F. Macbr., while C. anagalloidea A. St.-Hil. was not included in the previous survey, being a new record for Pernambuco.
Cuphea is the most representative genus, with 11 species, followed by Ammannia and Pleurophora, with two species each. Seven species are endemic from Brazil. Cuphea campestris Mart. ex Koehne is endemic from the Brazilian Northeast, as is L. glyptocarpa Koehne, but this one is found only in Bahia, Pernambuco and Piauí. Pleurophora pulchra J.C. Siqueira, Cotarelli, J.F.B. Pastore & T.B. Cavalc. is considered rare because it has a restricted distribution to Caatinga of Pernambuco and there are few specimens in the herbaria. The most frequent species in the area were: P. anomala (A. St.-Hil.) Koehne, C. flava Spreng. and C. racemosa (L. f.) Spreng.
Cuphea mimuloides Cham. & Schltdl. and P . anomala (A. St.-Hil.) Koehne are new records for the Atlantic Forest and L. glyptocarpa Koehne and R. ramosior (L.) Koehne are new records for the Caatinga.
In Pernambuco, eight species are found both in Caatinga and Atlantic Forest. Despite presenting greater richness in the Atlantic Forest of Brazil (51 spp. vs. 32 in Caatinga) (BFG 2022), the largest number of species found in the Caatinga of Pernambuco (15 spp. vs. 13 in Atlantic Forest) can be justified by the area percentage (80%) occupied by this biome in the State (Andrade-Lima 2007).
Pernambuco has four areas with greater species richness: Caruaru-Agrestina (figure 1 b) and Maraial-Jaqueira (figure 1 c), that are priorities for conservation because they present high species richness, and Mata da Pimenteira State Park (figure 1 a), besides Areas of Relevant Ecological Interest (ARIE) Dois Irmãos and Mata da Várzea (figure 1 d), that are conservation units. The following species were found in the Mata da Pimenteira State Park and in the ARIE Dois Irmãos and Mata da Várzea: Ammannia auriculata Willd., A. latifolia L., Cuphea campestris Mart. ex Koehne, C. carthagenensis (Jacq.) J.F. Macbr., C. circaeoides Sm. ex Sims, C. flava Spreng., C. impatientifolia A. St.-Hil., C. micrantha Kunth, C. racemosa (L.f.) Spreng., Lafoensia glyptocarpa Koehne, Pleurophora anomala (A. St.-Hil.) Koehne and Rotala ramosior (L.) Koehne.
Taxonomic treatment
Lythraceae J. St.-Hil., Expos. Fam. Nat. 2: 175, 1805.
Terrestrial or aquatic. Herbs, shrubs or trees, branches quadrangulate or cylindrical, glabrous, glabrescent, pubescent, pilose or hirsute. Leaves distichous opposite, decussate or verticillate, sessile or petiolate, blade narrow-triangular, ovate, elliptic, narrow-elliptic, oblong or oblanceolate, margins flat or revolute, membranaceous, papiraceous or chartaceous, venation camptodromous, brochidodromous or hyphodromous. Flowers solitary, dichasium or racemes, axillary or terminal, sessile or pedicellate, bracteoles absent or present. Floral tube campanulate, urceolate or tubular, flat or plicate. Spur absent or present. Petals absent or present. Stamens 4-20, homodynamous, didynamous or heterodynamous, included or exserted, anthers basifixed or dorsifixed, elliptic, oblong, reniform or concavo-convex, vesicles absent or present. Style included or exserted. Capsule globose, fusiform, cylindrical or turbinated. Seeds numerous, flat, globose or concavo-convex, wings absent or present.
Identification key for Lythraceae species from Pernambuco
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1. Branches glabrous.
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2. Branches cylindrical; leaves petiolate; flowers in racemes …… ............................. … 14. Lafoensia glyptocarpa
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2. Branches quadrangulate; leaves sessile; flowers solitary or in dichasium
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3. Leaf base decurrent, apex obtuse; flowers solitary ………...….…… ............................... 17. Rotala ramosior
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3. Leaf base cordate, apex acute or cuneate; flowers in dichasium
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4. Pedicel 1–5 mm long; stamens and style exserted ….……… ........................... …. 1. Ammannia auriculata
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4. Pedicel up to 1 mm long; stamens and style included …… ............................... …..…………. 2. A. latifolia
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1. Branches glabrescent, pubescent, pilose or hirsute.
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5. Leaves verticillate.
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6. Leaf blade narrow-triangular; venation hyphodromous; spur present …..……… ................................ ….. ………………………………………………… .................................. ……………… 7. Cuphea ericoides
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6. Leaf blade oblanceolate; venation brochidodromous; spur absent
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7. Floral tube ca. 5 mm long; dorsal petals ca. 3 × 2 mm … .................................... 15. Pleurophora anomala
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7. Floral tube 14–15 mm long; dorsal petals 5–7 × 5–6 mm ….…… 16. P. pulchra
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5. Leaves opposite distichous or decussate
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8. Leaf blade oblanceolate; venation hyphodromous …… ..................................................... 3. C. anagalloidea
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8. Leaf blade ovate, elliptic, narrow-elliptic or oblong; venation brochidodromous or camptodromous
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9. Venation camptodromous; spur present truncate ........................................................... 13. C. racemosa
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9. Venation brochidodromous; spur absent or present descending or horizontal
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10. Branches pilose; bracteoles and spur absent …… ................................................... 11. C. mimuloides
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10. Branches pubescent or hirsute; bracteoles and spur present
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11. Leaves sessile or short petiolate; base cordate or subcordate
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12. Leaf blade ovate; flowers solitary; petals present ….. 8. C. flava
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12. Leaf blade oblong; flowers in racemes; petals absent … ...... ……………………. 12. C. pulchra
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11. Leaves petiolate; base acute, cuneate or rounded
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13. Leaf blade elliptic or narrow-elliptic; spur descending
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14. Leaf base cuneate; flowers in racemes ………..…………………….. 5. C. carthagenensis
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14. Leaf base acute; flowers solitary.
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15. Capsule cylindrical ……… ................................................ ………. 4. C. campestris
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15. Capsule fusiform …… ..................................................... ………… 10. C. micrantha
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13. Leaf blade ovate; spur horizontal
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16. Leaf base rounded; pubescent on both faces; flowers in racemes …… ..... 6. C. circaeoides
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16. Leaf base cuneate; pilose on both faces; flowers solitary ……..……… 9. C. impatientifolia
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Ammannia L., Sp. Pl. 1: 119, 1753.
Terrestrial or aquatic. Herbs completely glabrous, branches quadrangulate. Leaves decussate, sessile, blade narrow-triangular, margins flat, base cordate, apex acute or cuneate, membranaceous or papiraceous, venation brochidodromous. Dichasium, axillary, pedicellate, bracteoles narrow-triangular or lanceolate. Floral tube urceolate, plicate, appendages of size nearly equal to the sepals, spur absent. Stamens 4, homodynamous, included or exserted, inserted at the middle third of the floral tube, anthers ca. 0.5 × 0.5 mm, dorsifixed, reniform, vesicles absent. Style included or exserted. Capsule globose. Seeds ca. 0.5 × 0.5 mm, concavo-convex, wings absent.
The genus is recognized by the completely glabrous herbs, quadrangular branches, narrow-triangular leaf blade, cordate base, acute or cuneate apex and dichasium inflorescence.
Ammannia is represented by about 25 species and is distributed in Temperate and Tropical zones, occurring primarily in Africa (Graham 1985). According to BFG (2022), in Brazil are recognized four species, of which three are found in the Northeast region. The Atlantic Forest stands out for its diversity of genera in the country. Two species occur in Pernambuco, A. auriculata Willd. and A. latifolia L.
1. Ammannia auriculata Willd., Hort. Berol. 1: 7, pl.7. 1803.
Figures 3 a, b, c
Ammannia auriculata Willd. a. Flowering branches. b. Leaf. c. Flower. A. latifolia L. b. leaf. d. Flowering branches. e. Flower. Cuphea carthagenensis (Jacq.) J.F. Macbr. f. Flowering branches. g. Indumentum. h. Flower. C. circaeoides Sm. ex Sims. i. Flowering branches. j. Indumentum. k. Leaf. l. Flower.
20-50 cm tall. Leaf blade 0.9-6 × 0.1-0.8 cm, membranaceous. Pedicel 1-5 mm long, bracteoles ca. 1 mm long, lanceolate, inserted at the inferior third of the pedicel. Floral tube 2-3 mm long, color not seen, sepals ca. 1 mm long, apex acute. Petals 4, pink or magenta, ca. 2 × 2 mm, wide-elliptic, base subcordate, apex rounded. Stamens exserted. Style ca. 3 mm long, exserted.
Selected material examined: BRAZIL. Pernambuco: Carnaíba, Fazenda Cacimba Velha, 8-V-2008, fl. fr., C. Ferreira 83883 (IPA); Recife, Campus da UFRPE, 16-X-2013, fl. fr., H.C. Gomes 5 (HST, HUEFS, IPA). The species is also recorded in the municipalities of Lagoa Grande and Mirandiba.
Habitat and distribution - It is pantropical (GBIF 2022). In Brazil, the species can be found in the North (TO), Northeast (AL, BA, CE, PE and RN), Central-west (MS and MT) and Southeast (MG) growing in Cerrado, Atlantic Forest and Pantanal environments (BFG 2022). In the study area, it was registered in Atlantic Forest and Caatinga. Ammannia auriculata occurs in wet places, such as margins of weirs, in addition to roadside, anthropized areas and as an invader of sugarcane crops. In Pernambuco, it was collected in the protection unit of Mata da Pimenteira State Park, APA Estuarina do Rio Capibaribe, ARIE Dois Irmãos and ARIE Mata da Várzea.
Notes - It can be identified by the stamens and style exserted. .
Phenology period - Flowers and fruits were collected from January to March, in May, July, August and October.
2. Ammannia latifolia L., Sp. Pl. 1: 119–120, 1753.
Figures 3 b, d-e, 4 a
Ammannia latifolia L. a. Flowering branches. Cuphea campestris Mart. ex Koehne. b. Habit. C. carthagenensis (Jacq.) J.F. Macbr. c. Flowering branches. C. ericoides Cham. & Schltdl. d. Flowering branches. C. flava Spreng. e. Habit. f. Flower. C. micrantha Kunth. g. Flowering branches. C. pulchra Moric. h. Flowering branches. Lafoensia glyptocarpa Koehne. i. Flowering branches. Pleurophora anomala (A. St.-Hil.) Koehne. j. Flowering branches.
20-150 cm tall. Leaf blade 1.4-12.7 × 0.3-1.9 cm, papiraceous. Pedicel up to 1 mm long, bracteoles ca. 2 mm long, narrow-triangular, inserted at the superior third of the pedicel. Floral tube ca. 5 mm long, green, reddish, vinaceous or brown, sepals ca. 1 mm long, apex cuneate. Petals not seen. Stamens included. Style ca. 1 mm long, included.
Selected material examined: BRAZIL. Pernambuco: Ipojuca, RPPN Nossa Senhora do Oiteiro de Maracaípe, Manguezal do Rio Ipojuca, 28-I-2007, fl. fr., M.S. Leite 295 (UFP); Fernando de Noronha, Pedreira, 10-VI-1999, fl. fr., A.M. Miranda 3490 (HST, HUEFS); Petrolina, Margem do Rio São Francisco, 21-II-2019, fl. fr., T. Lacerda 26 (PEUFR). This species is also recorded in the municipalities of Afrânio, Agrestina, Cabo de Santo Agostinho, Cabrobó, Calumbi, Caruaru, Custódia, Floresta, Goiana, Ibimirim, Lagoa Grande, Maraial, Olinda, Ouricuri, Parnamirim, Recife, Salgueiro, Santa Maria da Boa Vista, São Lourenço da Mata, Serra Talhada, Serrita and Timbaúba.
Habitat and distribution - The species is endemic from the Americas, occurring in the United States of America, Brazil, Bahamas, Cuba, Jamaica, Puerto Rico, Guadeloupe, Bolivia, Dominican Republic and Peru (GBIF 2022). In Brazil, it can be found in the North (AP and PA), Northeast (AL, BA, CE, PB, PE and RN) and Southeast (ES) growing in Amazon Forest, Caatinga, Cerrado and Atlantic Forest areas (BFG 2022). In Pernambuco, A. latifolia was registered in Atlantic Forest and Caatinga. The species was collected from rare to frequent, predominantly in wet environments, such as on the margins of rivers, lakes and weirs, but also in dry environments, along roadsides, grazed areas, close to crops and sewer margins. In the study area, it was recorded in the protection unit of RVS Mata do Engenho Uchôa, APA Estuarina do Rio Beberibe, FURB Mata do Passarinho, FURB Mata de Camaçari, Mata do Zumbi State Park, APA de Santa Cruz, RPPN Serro Azul, RPPN Engenho Contestado, RVS Serra do Giz, Mata da Pimenteira State Park, ESEC Serra da Canoa, RVS Serras Caatingueiras, RVS Tatu Bola, ARIE Dois Irmãos and ARIE Mata da Várzea.
Notes - Ammannia latifolia is morphologically related to Rotala ramosior by the glabrous and quadrangulate branches, sessile leaves and includes stamens and style. Ammannia latifolia is distinguished from R. ramosior by the narrow-triangular leaf blade (vs. narrow-elliptic in R. ramosior), cordate base (vs. decurrent), acute or cuneate apex (vs. obtuse) and dichasium (vs. flowers solitary).
Phenology period - Flowers and fruits were collected in January, February and from April to December.
Cuphea P. Browne, Civ. Nat. Hist. Jamaica 216-217, 1756.
Terrestrial or aquatic. Herbs or shrubs, branches cylindrical, glabrescent, pubescent, pilose or hirsute. Leaves distichous opposite, decussate or verticillate, sessile or petiolate, blade narrow-triangular, ovate, elliptic, narrow-elliptic, oblong or oblanceolate, margins flat, membranaceous, papiraceous or chartaceous, venation camptodromous, brochidodromous or hyphodromous. Flowers solitary or racemes, axillary or terminal, pedicellate, bracteoles absent or present. Floral tube tubular, plicate, appendages of size less than half of the sepals, trichome simple, spur absent or present. Petals absent or 6. Stamens 11, heterodynamous, included or exserted, inserted at the superior third of the floral tube, anthers dorsifixed, elliptic or oblong, vesicles absent or present. Style included or exserted. Capsule cylindrical or fusiform. Seeds globose or concavo-convex, wings absent.
The genus is recognized by the presence of trichomes, branches cylindrical, flowers pedicellate, plicate, spur and 6 petals present in most species.
Cuphea is the most representative genus of Lythraceae with about 250 species, and is endemic from the Americas, occurring primarily in the tropics and subtropics (Graham 2017). In Brazil, 110 species are recognized, among which 71 are endemic. They are distributed throughout the country, with a greater predominance in the Southeast (57 spp.), followed by the Central-west (44 spp.) and North and Northeast (37 spp. each one). Cerrado and Atlantic Forest stand out for covering the greatest richness of the genus in the country. However, representatives of this taxon can also be found in the Amazon Forest, Caatinga, Pampa and Pantanal (BFG 2022). In Pernambuco occur 11 species, C. anagalloidea A. St.-Hil., C. campestris Mart. ex Koehne, C. carthagenensis (Jacq.) J.F. Macbr., C. circaeoides Sm. ex Sims, C. ericoides Cham. & Schltdl., C. flava Spreng., C. impatientifolia A. St.-Hil., C. micrantha Kunth, C. mimuloides Cham. & Schltdl., C. pulchra Moric. and C. racemosa (L. f.) Spreng.
3. Cuphea anagalloidea A. St.-Hil., Fl. Bras. Merid., (quarto ed.) 3(23): 122, 1833.
Figures 5 a, b, c, d
Cuphea anagalloidea A. St.-Hil. a. Flowering branches. b. Indumentum. c. Leaf. d. Flower. C. campestris Mart. ex Koehne. e. Flowering branches. f. Indumentum. g. Leaf. h. Flower. C. impatientifolia A. St.-Hil. i. Flowering branches. j. Indumentum. k. Leaf. l. Flower.
Terrestrial. Herbs, ca. 30 cm tall, branches glabrescent, trichomes simple. Leaves decussate, sessile, blade 0.5-1.8 × 0.1-0.2 cm, oblanceolate, base acute, apex obtuse or rounded, membranaceous, glabrescent on both faces, trichomes simple, venation hyphodromous. Flowers solitary, axillary, pedicel 7-12 mm long, glabrescent, trichomes simple, bracteoles absent. Floral tube 4-6 mm long, purple, sepals ca. 0.5 mm long, apex cuneate, spur horizontal, outer surface pubescent, trichomes simple, inner surface pilose above and pubescent below stamen insertion, trichomes simple. Petals 6, light pink, 2 petals 2.5 × 1.5 mm, 4 petals 2-1 mm long, obovate, base acute, apex rounded. Stamens included, filaments pilose, trichomes simple, anthers ca. 0.5 × 0.5 mm, elliptic, vesicles absent. Style ca. 2 mm long, included, pilose, trichomes simple. Capsule cylindrical. Seeds ca. 0.5 × 0.5 mm, concavo-convex.
Selected material examined: BRAZIL. Pernambuco: Tamandaré, Restinga, 6-IX-1954, fl. fr., J.I.A. Falcão 959 (IPA).
Habitat and distribution - Cuphea anagalloidea is endemic to Brazil, distributed in the North (PA and TO), Northeast (AL, BA, PI and SE), Central-west (GO and MT) and Southeast (MG) growing in Amazon Forest, Caatinga, Cerrado and Atlantic Forest areas (BFG 2022). Still according to BFG (2022), it does not occur in Pernambuco, but in this study we record the occurrence in the State. In the region, the species was registered only in Atlantic Forest, collected in restinga regions and found in the protection unit of APA Estuarina do Rio Formoso.
Notes - It can be recognized from the other species by the glabrescent branches, oblanceolate leaf blade, pedicel 7-12 mm long and absent bracteoles.
Phenology period - Flowers and fruits found in September.
4. Cuphea campestris Mart. ex Koehne, Fl. Bras., 13(2): 265, 1877.
Figures 4 b, 5 e, f, g, h
Terrestrial or aquatic. Herbs, 4-50 cm tall, branches pubescent, trichomes simple. Leaves distichous opposite, petioles ca. 1 mm long, blade 1.3-4 × 0.3-0.8 cm, narrow-elliptic, base acute, apex cuneate, membranaceous, pubescent on both faces, trichomes simple, venation brochidodromous. Flowers solitary, axillary, pedicel ca. 2 mm long, pubescent, trichomes simple, bracteoles ca. 1 mm long, lanceolate, pubescent on the margins, trichomes simple, inserted at the superior third of the pedicel. Floral tube ca. 8 mm long, red, vinaceous or purple, sepals ca. 1 mm long, apex acute, spur descending, outer surface pubescent, trichomes simple, inner surface pilose above and pubescent below stamen insertion, trichomes simple. Petals 6, light pink, lilac or purple, 2 petals ca. 7 × 2 mm, 4 petals ca. 5 × 2 mm long, oblanceolate, base acute, apex cuneate. Stamens included or exserted, filaments pilose, trichomes simple, anthers ca. 1 × 0.5 mm, oblong, vesicles 10. Style ca. 4 mm long, included, glabrous. Capsule cylindrical. Seeds ca. 2 × 2 mm, concavo-convex.
Selected material examined: BRAZIL. Pernambuco: Orocó, 9-VI-2005, fl. fr., F. França 5244 (HUEFS). The species was also found in the municipalities of Afrânio, Cabrobó, Caruaru, Ouricuri, Petrolina, Serra Talhada, Serrita and Trindade.
Habitat and distribution - Cuphea campestris is endemic from the Brazilian Northeast, distributed in all States, except in the MA and SE, growing only in Caatinga environments (BFG 2022), as it was registered in Pernambuco. The species was found abundant in open vegetation, regions of wet soil, such as margins of rivers, weirs and lakes, and also in the sun, rocky outcrops and planting areas. In the study area, it occurs in the protection unit of RPPN Serro Azul, Mata da Pimenteira State Park, RVS Tatu Bola and RVS Riacho Pontal.
Notes - Cuphea campestris is morphologically related to C. micrantha by the petiolate leaves, acute base, cuneate apex and descending spur. Cuphea campestris is distinguished from C. micrantha by the leaf blade narrow-elliptic (vs. elliptic in C. micrantha), sepals apex acute (vs. cuneate), stamens exserted (vs. included) and capsule cylindrical (vs. fusiform). The species is viscous and has long trichomes on appendages.
Phenology period - Flowers and fruits were collected from February to June and September.
5. Cuphea carthagenensis (Jacq.) J.F. Macbr., Publ. Field Mus. Nat. Hist., Bot. Ser. 8(2): 124, 1930.
Figures 3 f, g, h, 4 c
Terrestrial. Herbs or shrubs, 10-50 cm tall, branches pubescent, trichomes simple. Leaves decussate, petioles 0.5-2 mm long, blade 0.8-3.7 × 0.3-1.5 cm, elliptic, base cuneate, apex cuneate, papiraceous, pubescent on both faces, trichomes simple, venation brochidodromous. Racemes axillary, pedicel 1-3 mm long, pubescent, trichomes simple, bracteoles ca. 1 mm long, narrow-triangular, pubescent on the margins, trichomes simple, inserted at the superior third of the pedicel. Floral tube 5-7 mm long, purple, sepals ca. 0.5 mm long, apex cuneate, spur descending, outer surface pubescent, trichomes simple and glandular, inner surface pubescent above and glabrescent below stamen insertion, trichomes simple and glandular. Petals 6, rose to purple, 2 petals ca. 3 × 2 mm, 4 petals ca. 3 × 1 mm, elliptic, base cuneate, apex cuneate. Stamens included, filaments pubescent, trichomes simple and glandular, anthers ca. 0.5 × 0.5 mm, oblong, vesicles 5. Style ca. 1.5 mm long, included, glabrous. Capsule cylindrical. Seeds ca. 2 × 2 mm, globose.
Selected material examined: BRAZIL. Pernambuco: São Lourenço da Mata, Estação Ecológica do Tapacurá, Borda de trilha de Mata Atlântica, 2-VII-2018, fl. fr., T. Lacerda 3 (PEUFR); Triunfo, Lagoa Nova, 10-III-1995, fl. fr., A.M. Miranda 2122 (HST). It was also recorded in the municipalities of Arcoverde, Bonito, Cabo de Santo Agostinho, Escada, Igarassu, Lagoa dos Gatos, Maraial, Moreno, Paulista, Recife and Saloá.
Habitat and distribution - According to Graham (2017), this species is the one most widely distributed of the genus, occurring from the United States of America to Argentina. In Brazil, it can be found in all regions, except in MA, PI, RN and RR States, growing in Amazon Forest, Caatinga, Cerrado, Atlantic Forest, Pampa and Pantanal environments (BFG 2022). In the study area, C. carthagenensis was registered in Atlantic Forest and Caatinga. The species occurs often in open or dense vegetation, environments of “brejos de altitude” and “campos rupestres” and areas of forest edge, roadsides and margins of weirs. In Pernambuco, the species was collected in the protection unit of Tapacurá Ecological Station, Mata da Pimenteira State Park, RPPN Engenho Santa Rita, APA Aldeia-Beberibe, ARIE Dois Irmãos, ARIE Mata da Várzea and RVS Matas do Sistema Gurjaú.
Notes - It can be distinguished from the other species by its unique combination of petioles 0.5-2 mm long, leaf blade elliptic, base and apex cuneate, racemes axillary, pedicel 1-3 mm long and bracteoles present. The species has green, vinaceous or brownish branches.
Phenology period - Flowers and fruits were collected in January and from March to November.
6. Cuphea circaeoides Sm. ex Sims, Bot. Mag. 48: , t. 2201, 1820.
Figures 3 i, j, k, l
Terrestrial. Herbs or shrubs, 10-40 cm tall, branches pubescent, trichomes simple. Leaves distichous opposite, petioles 2-27 mm long, blade 2.7-5.6 × 1-2.1 cm, ovate, base rounded, apex acute, membranaceous, pubescent on both faces, trichomes simple, venation brochidodromous. Racemes terminal, pedicel 2-8 mm long, pubescent, trichomes simple, bracteoles ca. 0.5 mm long, linear, pubescent, trichomes simple, inserted at the middle third of the pedicel. Floral tube 5-6 mm long, green, vinaceous or brown, sepals ca. 0.5 mm long, apex acute, spur horizontal, outer surface pubescent, trichomes simple, inner surface pilose above and glabrescent below stamen insertion, trichomes simple. Petals 6, pink or lilac to purple, 2 petals ca. 2 × 1 mm, obovate, 4 petals ca. 2 × 0.5 mm, narrow-elliptic, base acute, apex obtuse. Stamens included, filaments pilose, trichomes simple, anthers ca. 0.5 × 0.5 mm, elliptic, vesicles absent. Style ca. 1 mm long, included, glabrous. Capsule fusiform. Seeds ca. 2 × 2 mm, globose.
Selected material examined: BRAZIL. Pernambuco: Afrânio, entre Afrânio e Caboclo, 21-IV-1971, fl. fr., Academia Brasileira de Ciências 240 (IPA); Goiana, Mata da Usina Santa Teresa, 27-VI-2013, fl. fr., A.M. Miranda 6572 (HST, HUEFS, IPA). It was also recorded in the municipalities of Alagoinha, Arcoverde, Betânia, Bezerros, Custódia, Floresta, Gravatá, Olinda, Ouricuri, Parnamirim, Pesqueira, Petrolina, Recife, São Caetano, Serra Talhada, Sertânia and Triunfo.
Habitat and distribution - The species is endemic to Brazil, occurring in the Northeast (AL, BA, CE, PB, PE and SE) and in the Southeast (MG) growing in Caatinga, Cerrado and Atlantic Forest areas (BFG 2022). In Pernambuco, it was registered in the Atlantic Forest and Caatinga. Cuphea circaeoides occurs occasionally or frequently in open or dense vegetation, in dry places without light, rocky outcrops, forest edges and roadsides. In the study area, it was found in the protection unit of Mata da Pimenteira State Park, FURB Mata do Passarinho, APA Estuarina dos Rios Goiana e Megaó, RPPN Karawa-tá, ARIE Dois Irmãos, ARIE Mata da Várzea and Pedra do Cachorro Natural Monument.
Notes - Cuphea circaeoides is morphologically related to C. impatientifolia by the petiolate leaves, ovate blades and horizontal spur. Cuphea circaeoides is distinguished from C. impatientifolia by the flowers in racemes (vs. solitary flowers in C. impatientifolia), linear bracteoles (vs. elliptic) and acute apex sepals (vs. cuneate). The species has green branches.
Phenology period - Flowers and fruits were collected from March to September and in November.
7. Cuphea ericoides Cham. & Schltdl., Linnaea 2: 366, 1827.
Figures 4 d, 6 a, b, c, d
Cuphea ericoides Cham. & Schltdl. a. Flowering branches. b. Indumentum. c. Leaf. d. Flower. C. flava Spreng. e. Flowering branches. f. Indumentum. g. Leaf. h. Flower. C. pulchra Moric. i. Flowering branches. j. Indumentum. k. Leaf. l. Flower. C. racemosa (L. f.) Spreng. m. Flowering branches. n. Indumentum. o. Leaf. p. Flower.
Terrestrial. Shrubs, 15-150 cm tall, branches pubescent, trichomes simple. Leaves verticillate, sessile, blade 0.2-0.6 × 0.1-0.2 cm, narrow-triangular, base truncate, apex acute, chartaceous, pubescent on both faces, trichomes simple, venation hyphodromous. Flowers solitary, axillary, pedicel 5-6 mm long, pubescent, trichomes simple, bracteoles ca. 1 mm long, ovate, pubescent, trichomes simple, inserted at the superior third of the pedicel. Floral tube 5-8 mm long, purple, sepals ca. 1 mm long, apex acute, spur descending, outer surface pubescent, trichomes simple, inner surface pilose above and pubescent below stamen insertion, trichomes simple. Petals 6, yellow, pink or lilac to purple, 2 petals ca. 4 × 3 mm, 4 petals ca. 3 × 0.2 mm, obovate, base cuneate, apex rounded. Stamens included, filaments pilose, trichomes simple, anthers ca. 0.5 × 0.5 mm, oblong, vesicles absent. Style ca. 3 mm long, included, glabrescent, trichomes simple. Capsule cylindrical. Seeds ca. 2 × 2 mm, globose.
Selected material examined: BRAZIL. Pernambuco: Brejo da Madre de Deus, Fazenda Bituri, 3-III-2005, fl. fr., E.L. Borba 2186 (HUEFS). The species was also recorded in the municipalities of Belo Jardim and Buíque.
Habitat and distribution - The species is endemic to Brazil, occurring in the Northeast (BA and PE) and Southeast (MG) growing in Caatinga and Cerrado environments (BFG 2022). In the study area, it was registered only in Caatinga. Cuphea ericoides can be found occasionally or often in regions of “brejo de altitude”, mountains, rocky outcrops, in sunny areas, forest edges and roadsides. In Pernambuco, the species was collected in the protection unit of Catimbau National Park and RPPN Fazenda Bituri.
Notes - It can be distinguished from the other species by the following set of characters: leaves verticillate, sessile, blade narrow-triangular, base truncate, apex acute, fowers solitary, pedicel 5-6 mm long, bracteoles present. The species has brown branches and fruits.
Phenology period - Flowers and fruits were collected from January to October and in December.
8. Cuphea flava Spreng., Novi Provent. 14, 1818.
Terrestrial. Shrubs, 30-150 cm tall, branches pubescent, trichomes simple. Leaves decussate, sessile or short petiolate, petioles 0-0.5 mm long, blade 0.5-2.3 × 0.2-0.8 cm, ovate, base subcordate, apex cuneate, papiraceous, pilose on the margins, glabrous or glabrescent on the veins and glabrous on both faces, trichomes simple, venation brochidodromous. Flowers solitary, axillary, pedicel 1-4 mm long, pubescent, trichomes simple, bracteoles ca. 1 mm long, ovate, pilose on the margins, trichomes simple, inserted at the middle third of the pedicel. Floral tube ca. 8 mm long, green, sepals ca. 1 mm long, apex acute, spur descending, outer surface pubescent, trichomes simple, inner surface pilose above stamen insertion and pubescent at the base of the ovary, trichomes simple. Petals 6, white or yellow, 2 petals ca. 4 × 2 mm, 4 petals ca. 3 × 1 mm, obovate, base cuneate, apex rounded. Stamens included or exserted, filaments pilose, trichomes simple, anthers ca. 0.5 × 0.5 mm, oblong, vesicles absent. Style ca. 5 mm long, exserted, glabrous. Capsule cylindrical. Seeds ca. 2 × 2 mm, globose.
Selected material examined: BRAZIL. Pernambuco: Igarassu, RPPN Nossa Senhora do Oiteiro de Maracaípe, Borda da trilha para o Cemitério do Outeiro, 7-IV-2019, fl. fr., T. Lacerda 28 (PEUFR). The species was also recorded in the municipalities of Cabo de Santo Agostinho, Camaragibe, Goiana, Ilha de Itamaracá, Ipojuca, Itambé, Jaboatão dos Guararapes, Olinda, Recife, Rio Formoso and Tamandaré.
Habitat and distribution - The species is endemic to Brazil, occurring in the North (PA), Northeast (AL, BA, PB, PE, RN and SE) and Southeast (ES and RJ) growing only in Atlantic Forest areas (BFG 2022), as it was registered in Pernambuco. Cuphea flava can be found occasionally or often in areas of herbaceous and arboreal vegetation, restinga regions, mangroves, edge and interior of forests, rocky environments, cultivation and open fields that are not subject to flooding. In the study area, the species was collected in the protection unit of RPPN Nossa Senhora do Oiteiro de Maracaípe, APA Guadalupe, APA Estuarina dos Rios Sirinhaém e Maracaípe, ARIE Ipojuca Merepe, ARIE Dois Irmãos, ARIE Mata da Várzea, APA Estuarina dos Rios Jaboatão e Pirapama, FURB Mata do Janga and RVS Mata do Jaguaribe.
Notes - Cuphea flava is morphologically related to C. pulchra by the sessile leaves, descending spur and exserted style. Cuphea flava is distinguished from C. pulchra by the blades ovate (vs. oblong in C. pulchra), flowers solitary (vs. flowers in racemes) and petals present (vs. absent). The species is viscous and has green, vinaceous or brownish branches, the trichomes of the branches and the pedicel are present only on one side.
Phenology period - Flowers and fruits were collected during the whole year.
9. Cuphea impatientifolia A. St.-Hil., Flora Brasiliae Meridionalis (quarto ed.) 3(23): 113. 1833.
Figures 5 i, j, k, l
Terrestrial. Herbs, 15-50 cm tall, branches pubescent, trichomes simple. Leaves distichous opposite, petioles ca. 10 mm long, blade 2.3-3 × 1–2 cm, ovate, base cuneate, apex cuneate, membranaceous, pilose on both faces, trichomes simple, venation brochidodromous. Flowers solitary, axillary, pedicel 1-3 mm long, pubescent, trichomes simple, bracteoles ca. 1 mm long, elliptic, pubescent, trichomes simple, inserted at the inferior third of the pedicel. Floral tube ca. 6 mm long, light green with vinaceous veins, white or dark purple, sepals ca. 1 mm long, apex cuneate, spur horizontal, outer surface pilose, trichomes simple, inner surface pilose above and pubescent below stamen insertion, trichomes simple. Petals 6, light pink, lilac or purple, 2 petals ca. 3.5 × 2 mm, 4 petals ca. 3 × 1 mm, obovate, base acute, apex cuneate. Stamens included, filaments pilose, trichomes simple, anthers ca. 0.5 × 0.5 mm, oblong, vesicles absent. Style ca. 3 mm long, included, glabrous. Capsule fusiform. Seeds ca. 3 × 3 mm, concavo-convex.
Selected material examined: BRAZIL. Pernambuco: Petrolina, Próximo à Lagoa de estabilização (CPATSA), 1-VI-2011, fl. fr., M.O. Mangabeira 4412 (HUEFS); Igarassu, Usina São José, Mata de Piedade, 29-VII-2007, fl. fr., D. Araújo 378 (UFP). It was also found in the municipalities of Arcoverde, Caruaru, Custódia, Gravatá, Mirandiba, Salgueiro, Serra Talhada, Triunfo and Verdejante.
Habitat and distribution - Cuphea impatientifolia is endemic to Brazil, distributed in the Northeast (all States) and Southeast (MG) growing in Caatinga and Atlantic Forest areas (BFG 2022), as it was registered in the study area. The species can be found in open or dense vegetation, wet or stony ground, forest edges and abandoned fields. In Pernambuco, it occurs in the protection unit of APA Estuarina do Canal de Santa Cruz, RPPN Karawa-tá and Mata da Pimenteira State Park.
Notes - Cuphea impatientifolia is morphologically related to C. circaeoides, the differences between C. impatientifolia and C. circaeoides were discussed under the comments of that species. It can be recognized by being very viscous, bearing simple trichomes at the apex of the bracteole.
Phenology period - Flowers and fruits were collected from April to July and in November.
10. Cuphea micrantha Kunth, Nova Genera et Species Plantarum (quarto ed.) 6: 196. 1823.
Figures 4 g, 7 a, b, c, d
Cuphea micrantha Kunth. a. Flowering branches. b. Indumentum. c. Leaf. d. Flower. C. mimuloides Schltdl. & Cham. e. Flowering branches. f. Indumentum. g. Leaf. h. Flower.
Terrestrial or aquatic. Herbs or shrub, 10-200 cm tall, branches pubescent or hirsute, trichomes simple or glandular. Leaves distichous opposite or decussate, petioles ca. 1 mm long, blade 2.5-4 × 1-1.3 cm, elliptic, base acute, apex cuneate, membranaceous, pubescent on both faces, trichomes simple, venation brochidodromous. Flowers solitary, axillary, pedicel 2-3 mm long, pubescent, trichomes simple, bracteoles ca. 1 mm long, lanceolate, pubescent, trichomes simple, inserted at the superior third of the pedicel. Floral tube ca. 6 mm long, color not seen, sepals ca. 1 mm long, apex cuneate, spur descending, outer surface pubescent, trichomes simple or glandular, inner surface pilose above and pubescent below stamen insertion, trichomes simple. Petals 6, pink, vinaceous, lilac or purple, 2 petals 4 × 2 mm, 4 petals ca. 3 × 1 mm, elliptic, base acute, apex cuneate. Stamens included, filaments pilose, trichomes simple, anthers ca. 0.5 × 0.5 mm, oblong, vesicles absent. Style ca. 2 mm long, included, glabrous. Capsule fusiform. Seeds ca. 2 × 2 mm, globose.
Selected material examined: BRAZIL. Pernambuco: São Lourenço da Mata, Estação Ecológica do Tapacurá, Borda de trilha de Mata Atlântica, 2-VII-2018, fl. fr., T. Lacerda 2 (PEUFR); Afrânio, Povoado de Caboclo, 2-VI-2007, fl. fr., R.F. Silva 11 (HVASF). The species is also recorded in the municipalities of Bonito, Cabo de Santo Agostinho, Camaragibe, Caruaru, Ferreiros, Igarassu, Maraial, Moreno, Primavera, Recife, Rio Formoso, and Sirinhaém.
Habitat and distribution - Cuphea micrantha is distributed in Brazil, Colombia, Venezuela, Dominican Republic, Cuba, Bolivia, Honduras, Paraguay, El Salvador and Guatemala (GBIF 2022). In Brazil, it can be found in the North (AC, PA, RR and TO), Northeast (all States, except AL) Central-west (DF, GO, MS and MT) and Southeast (MG and SP) growing in Amazon Forest, Caatinga, Cerrado and Atlantic Forest areas (BFG 2022). In the study area, the species was collected in the Atlantic Forest and Caatinga. It occurs abundantly in open or dense vegetation, areas of “brejos de altitudes”, forest edges and interior, roadsides, planted and unplanted crop fields, in addition to wet soils, such as margins of streams, weirs and rivers. In Pernambuco, the species was registered in the protection unit of Tapacurá Ecological Station, RPPN Serro Azul, RPPN Bicho Homem, RPPN Engenho Contestado, APA Guadalupe, APA Aldeia-Beberibe, APA Santa Cruz, ARIE Dois Irmãos, ARIE Mata da Várzea and RVS Mata de Caraúna.
Notes - Cuphea micrantha is morphologically related to C. campestris, and their differences were discussed under the comments of that species. It can be recognized by the yellowish-green or reddish branches and brown anthers.
Phenology period - Flowers and fruits were collected from January to November.
11. Cuphea mimuloides Cham. & Schltdl., Linnaea 5: 570. 1830.
Figures 7 e, f, g, h
Terrestrial or aquatic. Herbs or shrub, branches pilose, trichomes simple. Leaves decussate, petioles ca. 0.5 mm long, blade ca. 1.7 × 0.7 cm, narrow-elliptic, base cuneate, apex cuneate, membranaceous, glabrescent on both faces, trichomes simple, venation brochidodromous. Flowers solitary, axillary, pedicel 8-10 mm long, pilose, trichomes simple, bracteoles absent. Floral tube ca. 5 mm long, color not seen, sepals and appendages not seen, spur absent, outer surface and inner surface glabrescent, trichomes simple. Petals not seen. Stamens and anthers not seen, vesicles absent. Style not seen. Capsule fusiform. Seeds ca. 0.5 × 0.5 mm, concavo-convex.
Selected material examined: BRAZIL. Pernambuco: Pombos, São João, Terreno pantanoso do açude, 30-IV-1932, fl. fr., B. Pickel 2985 (IPA).
Habitat and distribution - Cuphea mimuloides occurs in Cuba, Mexico, Brazil, Costa Rica, Nicaragua, Honduras, Belize, Guatemala, Guyana and United States of America (GBIF 2022). In Brazil, it is registered in the Northeast (AL, BA and PE) growing only in Caatinga environments (BFG 2022), but in this study we confirm the distribution also in Pernambuco’s Atlantic Forest. In the State, the species was collected in wet grounds and was not found in a protection unit.
Notes - The sepals, appendages, petals, stamens, anthers and style were not described because there is only one specimen and it was not possible to analyze the structures in it. Other materials of the same species were researched, but there are few and are deposited in distant herbaria. Cuphea mimuloides is morphologically similar to C. racemosa by sharing the branches pilose, leaves decussate and bracteoles absent, but can be distinguished by the leaf venation brochidodromous (vs. camptodromous in C. racemosa), flowers solitary (vs. flowers in racemes) and spur absent (vs. present).
Phenology period - Flowers and fruits were collected in April.
12. Cuphea pulchra Moric., Pl. Nouv. Amer. 168, pl.98. 1847.
Figures 4 h, 6 i, j, k, l
Terrestrial. Shrubs, ca. 80 cm tall, branches pubescent, trichomes simple and glandular. Leaves decussate, sessile, blade 0.9-2.9 × 0.3-1.2 cm, oblong, base cordate, apex acute or cuneate, chartaceous, pubescent on both faces, trichomes simple and glandular, venation brochidodromous. Racemes terminal, pedicel 7-8 mm long, pubescent, trichomes simple and glandular, bracteoles ca. 1 mm long, ovate, pubescent, trichomes simple, inserted at the middle third of the pedicel. Floral tube 17-22 mm long, red, sepals ca. 1 mm long, apex cuneate, spur descending, outer surface pubescent, trichomes simple and glandular, inner surface hirsute above and pubescent below stamen insertion, trichomes simple. Petals absent. Stamens included or exserted, filaments hirsute, trichomes simple, anthers ca. 1 × 1 mm, oblong, vesicles absent. Style ca. 14 mm long, exserted, pilose, trichomes simple. Capsule fusiform. Seeds ca. 3 × 3 mm, globose.
Selected material examined: BRAZIL. Pernambuco: Ouricuri, 5 km a Oeste de Ouricuri, numa baixada úmida, margem da BR-316, 24-III-1987, fl. fr., J.L.S. Lima 371 (HTSA, HUEFS).
Habitat and distribution - It is endemic to Brazil, occurring in the Northeast (BA, PE and SE) and Southeast (MG) growing in Caatinga and Cerrado environments (BFG 2022). In the study area, it was collected in Caatinga. In Pernambuco, C. pulchra does not occur in a protection unit.
Notes - Cuphea pulchra is morphologically related to C. flava, the differences between them were discussed under the comments of this species.
Phenology period - Flowers and fruits were collected in March.
13. Cuphea racemosa (L. f.) Spreng., Syst. Veg. [Sprengel] 2: 455, 1825.
Figures 6 m, n, o, p
Terrestrial or aquatic. Shrubs, 20-100 cm tall, branches pilose, trichomes simple. Leaves decussate, petioles 3-30 mm long, blade 0.9-7 × 0.5-4.5 cm, elliptic, base cuneate, apex cuneate, papiraceous, pubescent on both faces, trichomes simple, venation camptodromous. Racemes terminal, pedicel 5-10 mm long, pubescent, trichomes simple, bracteoles absent. Floral tube 6-18 mm long, green to purplish-brown, sepals ca. 0.5 mm long, apex cuneate, spur truncate, outer surface pubescent, trichomes simple, inner surface hirsute above and glabrescent below stamen insertion, trichomes simple. Petals 6, pink or lilac to purple, 2 petals ca. 3 × 1 mm, 4 petals ca. 2.5 × 0.5 mm long, obovate, base acute, apex rounded. Stamens included or exserted, filaments hirsute, trichomes simple, anthers ca. 0.5 × 0.5 mm, elliptic, vesicles absent. Style ca. 3 mm long, included, pilose, trichomes simple. Capsule cylindrical. Seeds ca. 1 × 1 mm, globose.
Selected material examined: BRAZIL. Pernambuco: Brejo da Madre de Deus, Fazenda Bituri, 3-III-2005, fl. fr., E.L. Borba 2189 (HUEFS); Recife, Campus da UFPE, VIII-1996, fl. fr., M. Alves s.d. (UFP19810). It was also recorded in the municipalities of Barra de Guabiraba, Bom Conselho, Bonito, Caruaru, Garanhuns, Jaqueira, Maraial, and Pesqueira.
Habitat and distribution - According to Cavalcanti & Graham (2002), this species occurs from Mexico to Argentina. In Brazil, it can be found in the Northeast (AL, BA, CE, PE and SE), Central-west (DF, GO and MS), Southeast (ES, MG, RJ and SP) and South (PR, RS and SC) growing in Amazon Forest, Caatinga, Cerrado, Atlantic Forest, Pampa and Pantanal areas (BFG 2022). In Pernambuco, C. racemosa was registered in Atlantic Forest and Caatinga. The species occurs rarely or often in areas of open or dense vegetation, wet soils, such as margins of rivers, weirs and streams, but also in “brejos de altitude”, stony, sunny and ruderal environments, edge and interior of forests and roadsides. In the study area, it was collected in the protection unit of RPPN Fazenda Bituri, RPPN Pedra D’Antas, RPPN Serro Azul, ARIE Dois Irmãos and ARIE Mata da Várzea.
Notes - Cuphea racemosa is morphologically related to C. mimuloides, the differences between C. racemosa and C. mimuloides were discussed under the comments of that species. It is characterized by being very viscous, the branches are green with vinaceous tones, filaments and style lilac, and the anthers bluish.
Phenology period - Flowers and fruits were collected during the whole year.
Lafoensia Vand., Fl. Lusit. Bras. Spec. 33, 1788.
The genus is recognized by the completely glabrous shrubs or trees, cylindrical branches, elliptic leaf blade, obtuse or rounded base, obtuse or cuspidate apex, campanulate and flat floral tube.
Lafoensia is represented by six endemic species from the Americas, occurring five in South America (Lourteig 1986). In Brazil, seven species are recognized, of which three are endemic. Three species are found in the Northeast region, and the Cerrado stands out for its diversity of genera in the country (BFG 2022). In Pernambuco only L. glyptocarpa Koehne was found.
14. Lafoensia glyptocarpa Koehne, Fl. Bras. 13(2): 353, 1877.
Figures 4 i, 8 a, b, c
Lafoensia glyptocarpa Koehne. a. Flowering branches. b. Leaf. c. Flower. Pleurophora anomala (A. St.-Hil.) Koehne. d. Flowering branches. e. Flower. P. pulchra J.C. Siqueira, Cotarelli, J.F.B. Pastore & T.B. Cavalc. f. Flowering branches. g. Flower. Rotala ramosior (L.) Koehne. h. Flowering branches. i. Leaf. j. Flower.
Terrestrial. Shrubs or trees completely glabrous, 120-3000 cm tall, branches cylindrical. Leaves decussate, petioles ca. 5 mm long, blade 2.7-8.0 × 1.8-5.0 cm, elliptic, margins flat, base obtuse or rounded, apex obtuse or cuspidate, chartaceous, venation brochidodromous. Racemes terminal, pedicel ca. 15-30 mm long, bracteoles absent. Floral tube ca. 25 mm long, vinaceous, campanulate, flat, sepals ca. 8 mm long, apex acute, appendages and spur absent. Petals 10, white or cream, ca. 20 × 5 mm, obovate, base decurrent, apex cuneate. Stamens 20, homodynamous, exserted, inserted at the inferior third of the floral tube, anthers ca. 1 × 5 mm, dorsifixed, concavo-convex, vesicles absent. Style ca. 75-93 mm long, exserted. Capsule 32-75 mm, turbinated. Seeds ca. 20 × 10 mm, flat, wings present.
Selected material examined: BRAZIL. Pernambuco: Garanhuns, 6-XI-2010, fl. fr., M. Oliveira 5265 (IPA); Vicência, Engenho Jundiá, 30-VII-1968, fl., Andrade-Lima 68-5401 (IPA). It was also recorded in the municipalities of Arcoverde, Buíque, Exu, Jataúba, Maraial, Saloá, Tacaratu and Triunfo.
Habitat and distribution - According to BFG (2022), it is endemic from the Brazilian Northeast, so far known only to Bahia, Pernambuco and Piauí, growing only in Cerrado environments (phytogeographic domain not present in Pernambuco), but we recorded it for the Caatinga of the State. The species can be found in places of “brejos de altitude”, “campos rupestres”, rocky outcrops, mountains or roadsides. In the study area, L. glyptocarpa was collected in the protection unit of APA Chapada do Araripe, Mata da Pimenteira State Park, Pedra Furada Park, RPPN Fazenda Bituri and RVS Matas de Água Azul.
Notes - It can be identified by being completely glabrous, leaves petiolate, blade elliptic, racemes terminal, pedicel ca. 15-30 mm long, floral tube ca. 25 mm long and capsule turbinated. The species has branches brownish, grayish or greenish and light yellow filaments and style.
Phenology period - Flowers were collected in January and from July to November and fruits found in April and from August to November.
Pleurophora D. Don, Edinburgh New Philos. J. 12: 112, 1837.
Terrestrial or aquatic. Shrubs, branches cylindrical, pubescent, trichomes simple. Leaves verticillate, sessile or short petiolate, petioles 0-1 mm long, blade oblanceolate, margins revolute, base acute, apex cuneate, papiraceous, pubescent on both faces, trichomes simples, venation brochidodromous. Flowers solitary, axillary, pedicel pubescent, trichomes simple, bracteoles narrow-elliptic or linear, pubescent, trichomes simple. Floral tube tubular, plicate, sepals apex acute, appendages of size less than half of the sepals, trichome simple, spur absent, outer surface pubescent, trichomes simple, inner surface glabrous. Petals 6. Stamens 6, heterodynamous, exserted, inserted at the base of the floral tube, filaments glabrescent, trichomes simple, anthers dorsifixed, reniform, vesicles absent. Style exserted, glabrescent or pubescent. Capsule cylindrical. Seeds concavo-convex, wings absent.
The genus is recognized by the presence of trichomes, branches cylindrical, leaves verticillate, blade oblanceolate, margins revolute, base acute, apex cuneate and spur absent.
Pleurophora is represented by six species currently accepted, although more than twice as many have been described. It is endemic from the Americas, occurring in Argentina, Bolivia, Brazil, Chile and Paraguay (Siqueira-Filho et al. 2015). According to BFG (2022), in Brazil are recognized three species, of which two are endemic. Two species are registered for the Northeast region and the Caatinga stands out for its diversity of genera in the country. In Pernambuco were found the species P. anomala (A. St.-Hil.) Koehne and P. pulchra J.A. Siqueira, V.M. Cotarelli, J.F.B. Pastore & T.B. Cavalcanti.
15. Pleurophora anomala (A. St.-Hil.) Koehne, Fl. Bras. 13(2): 307, pl.57. 1877.
Terrestrial or aquatic. Shrubs, 20-100 cm tall. Leaf blade 0.5-3.3 × 0.1-0.7 cm. Pedicel ca. 1 mm long, bracteoles ca. 1 mm long, inserted at the superior third of the pedicel. Floral tube ca. 5 mm long, vinaceous-green, sepals ca. 1 mm long. Petals white, light pink or lilac to purple, 2 petals ca. 3 × 2 mm, obovate, base obtuse, apex rounded, 4 petals ca. 2 mm long, narrow-elliptic, base cuneate, apex cuneate. Anthers ca. 0.5 × 1 mm. Style ca. 4 mm long, pubescent, trichomes simple. Seeds ca. 0.5 × 0.5 mm.
Selected material examined: BRAZIL. Pernambuco: Pesqueira, Assentamento Rosário, Margem da BR-232, Próximo ao distrito de Mimoso, 9-IV-2019, fl. fr., T. Lacerda 31 (PEUFR); Recife, Curado, 15-VIII-1958, fl. fr., Andrade-Lima 585962 (IPA). The species is also recorded in the municipalities of Afrânio, Alagoinha, Altinho, Arcoverde, Belo Jardim, Bodocó, Bonito, Brejo da Madre de Deus, Buíque, Cabrobó, Carpina, Caruaru, Cruzeiro do Nordeste, Custódia, Floresta, Gravatá, Maraial, Mirandiba, Orocó, Ouricuri, Petrolândia, Petrolina, Salgueiro, São Bento do Una, São Caetano, Serra Talhada, Sertânia, Surubim and Vitória de Santo Antão.
Habitat and distribution - It is endemic to Brazil, occurring in the Northeast (AL, BA, CE, PB, PE, PI, RN and SE) and Southeast (MG) growing only in Caatinga areas (BFG 2022), but in this study we confirm the occurrence also in the State’s Atlantic Forest. The species can be found occasionally or very abundant in sunny and rupicolous environments, places of open or dense vegetation, wet spots, such as on the margins and close to streams, rivers, weirs, lakes and canals and areas of natural pasture, agricultural vegetation and roadsides. In Pernambuco, P. anomala was collected in the protection unit of Mata da Pimenteira State Park, RVS Riacho Pontal, RVS Serras Caatingueiras, RVS Serra do Giz, ARIE Dois Irmãos, ARIE Mata da Várzea, Serra da Canoa Ecological Station and RPPN Bituri.
Notes - Pleurophora anomala is related to P. pulchra by the leaves verticillate, blade oblanceolate, flowers solitary and bracteoles present. However, it can be distinguished by the floral tube green (vs. green to red in P. pulchra), petals white, light pink to lilac (vs. red), floral tube ca. 5 mm long (vs. ca. 15 mm long) and dorsal petals ca. 3 × 2 mm (vs. ca. 7 × 6 mm). The species is viscous and has green to violaceous branches, indument on the entire adaxial leaf face and more concentrated on the veins of the abaxial leaf face and green anthers. It is invasive in sugarcane, forage and nectar-polliniferous. Pleurophora anomala is popularly known as Vassourinha, Vassourinha-de-alagado, Marcela-branca or Vassourinha-do-brejo.
Phenology period - Flowers and fruits were collected from January to November.
16. Pleurophora pulchra J.C. Siqueira, Cotarelli, J.F.B. Pastore & T.B. Cavalc., Syst. Bot. 40(1): 185, 2015.
Terrestrial. Shrubs, 30-100 cm tall. Leaf blade 0.6-2.5 × 0.2-0.5 cm. Pedicel 1-3 mm long, bracteoles 2-3 mm long, inserted at the middle third of the pedicel. Floral tube 14-15 mm long, green at the base and becoming red towards the apex, sepals 2-4 mm long. Petals red, 2 petals 5-7 × 5-6 mm, wide-elliptic, base decurrent, apex emarginate, 4 petals 1-2 mm long, narrow-triangular, base truncate, apex acute. Anthers ca. 1 × 1 mm. Style ca. 20 mm long, glabrescent, trichomes simple. Seeds 0.4-1 × 0.5-0.8 mm.
Selected material examined: BRAZIL. Pernambuco: Cabrobó, Serra do Bendó, Eixo Norte, 22-IX-2009, fl. fr., A.P. Fontana 6195 (HVASF, PEUFR). It is also recorded in the municipality of Salgueiro.
Habitat and distribution - The species is endemic to Pernambuco, growing in Caatinga environments (BFG 2022), as it was found in the study area. Pleurophora pulchra occurs with few individuals, in rocky environments and along highways. In Pernambuco, the species was collected in the protection unit of RVS Serras Caatingueiras.
Notes - Pleurophora pulchra is similar to P. anomala and the differences were discussed under the comments of that species.
Phenology period - Flowers and fruits were collected in April, June and September.
Rotala L., Mant. Pl. 2: 143-144, 175, 1771.
Rotala is represented by about 45 species predominantly from Africa and Asia, besides few species from Americas (Cook 1979). The genus is considered naturalized in Brazil and three species are recognized, growing in Atlantic Forest, Amazon Forest and Cerrado areas and of which one is found in the Northeast (BFG 2022). In Pernambuco occurs the species R. ramosior (L.) Koehne.
17. Rotala ramosior (L.) Koehne, Fl. Bras. 13(2): 194, 1877.
Figures 8 h, i, j
Terrestrial or aquatic. Herbs completely glabrous, 7-40 cm tall, branches quadrangulate. Leaves decussate, sessile, blade 0.4-2.7 × 0.1-0.6 cm, narrow-elliptic, margins flat, base decurrent, apex obtuse, membranaceous, venation brochidodromous. Flowers solitary, axillary, sessile or short pedicel 0-0.5 mm long, bracteoles 2-3 mm long, linear, inserted at the base of the floral tube. Floral tube 2-4 mm long, green, reddish or vinaceous, urceolate, plicate, sepals ca. 0.5 mm long, apex cuneate, appendages of size nearly equal to the sepals, spur absent. Petals white, light pink or lilac to purple not seen. Stamens 4, didynamous, included, inserted at the middle third of the floral tube, anthers ca. 0.5 × 0.5 mm, basifixed, reniform, vesicles absent. Style ca. 0.5 mm long, included. Capsule globose. Seeds ca. 0.5 × 0.5 mm, concavo-convex, wings absent.
Selected material examined: BRAZIL. Pernambuco: Cabo de Santo Agostinho, Pedra do Cayango, Entre os Engenhos Megahype e Cayango, 14-IX-2008, fl. fr., M. Sobral-Leite 873 (UFP); Serra Talhada, Parque Estadual Mata da Pimenteira, Lagoa Pimenteira II, Margem do açude, 22-III-2012, fl. fr., W. Cordeiro 223 (HESBRA). It is also recorded in the municipalities of Afrânio, Arcoverde, Cabrobó, Caruaru, Custódia, Igarassu, Maraial, Mirandiba, Recife, Sertânia, Sirinhaém and Vitória de Santo Antão.
Habitat and distribution - It is distributed in the United States of America, Mexico, Brazil, Costa Rica, Taiwan, Canada, Japan, Nicaragua, Colombia and Panama (GBIF 2022). According to BFG (2022), the species can be found in the North (PA), Northeast (AL, BA, PE, PI and RN), Central-west (GO, MS and MT) and Southeast (MG). It was recorded for the Atlantic Forest of Pernambuco (Lima et al. 2020) and in this study we confirm the occurrence also in the State’s Caatinga. Rotala ramosior occurs little or very often in places of open or dense vegetation, wet soils, such as on the margins of streams, rivers, lakes or weirs and areas of planting, pasture, rocky outcrop or road. In Pernambuco, the species was collected in the protection unit of RVS Serras Caatingueiras, Mata da Pimenteira State Park, ARIE Dois Irmãos, ARIE Mata da Várzea, RPPN Serro Azul and RPPN Engenho Contestado.
Notes - Rotala ramosior is morphologically similar to Ammannia latifolia, and the differences between them were discussed under the comments of that species. It is characterized by having pink to vinaceous branches.
Phenology period - Flowers and fruits were collected from January to March, from May to October and in December.
Acknowledgements
We thank Universidade Federal Rural de Pernambuco (UFRPE), for providing transportation for the field trips and Laboratório de Sistemática Integrativa de Angiospermas (LASI) and its team, for all assistance, especially to Swami Costa, for preparing the maps. We also thank Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - Brasil (CAPES) - Finance Code 132217/2018-0, for the master fellowship. We are also grateful to herbaria curators of CEN, HESBRA, HST, HTSA, HUEFS, HVASF, IPA, JPB, PEUFR and UFP, for the assistance and making available the materials to be analyzed and Felipe Martins for the illustrations.
List of material examined
Academia Brasileira de Ciências 151 (6), 247 (4), 279 (15), 280 (4), 426 (6), 713 (15); Albuquerque, N. A. 477 (10); Alcântara, G. D. 51 (7); Almeida, N. s.d. (10); Alves, J. L. H. s.d. (8); Alves-Araújo, A. 1027 (17), 1034 (15), 1037 (1), 543 (17), 860 (8); Andrade, I. M. 57 (7); Andrade-Lima 48-144 (8), 49-251 (4), 49-357 (14), 552066 (8), 552215 (2), 552245 (2), 56-2589 (6), 57-2714 (13), 585943 (10), 585965 (1), 61-3998 (7), 65-4287 (7), 65-4288 (7), 66-4884 (7), 68-5439 (14), 716405 (9), 716487 (13), 716561 (7), 718103 (7), 737489 (7), 75-8124 (7), 7719 (8), 78-8530 (15), 78-8534 (15), 79-9190 (15), 79-9475 (4), 79-9550 (13), 80-8927 (2); Araújo, D. 1687 (8), 378 (10), 417 (10), 848 (2), 878 (15), 884 (2); Araújo, D. A. 1687 (8); Araújo, E. L. 014 (9); Araújo, F. 54 (6); Araújo, M. J. 13 (15); Ataíde, M. 27 (15), 639 (9); Baracho, G. S. 267 (7), 278 (14); Barbosa, R. 2 (5); Barreto, R. 12 (6); Barros, I. C. L. 151 (5), 132 (8), s.d. (8); Belo, D. s.d. (13); Bezerra, G. J. 37 (5), 59 (5); Bocage, A. -73382 (14), -73424 (14), 1155 (14); Brito, Z. 58 (15); Campelo, L. 15 (7); Cano, O. 13 (5); Cantarelli, J. 449 (15), 595 (8); Carvalho-Sobrinho, J. G. 1905 (15), 2163 (6), 2245 (15), 2302 (6); Carvalho, H. 3868 (15); Cavalcanti, D. 146 (8), 278 (15), 279 (6), 601 (2); Cavalcanti, G. s.d. (4); Coelho, A. s.d. (8); Coelho, M. M. 103 (2), 194 (6), 51 (2), 67 (15); Coelho, P. s.d. (8); Cordeiro, W. 145 (2), 174 (2), 261 (2), 303 (17), 439 (2), 58 (15), 62 (17), 726 (17), 99 (15); Correia, M. 269 (13), 273 (13); Costa e Silva, M.B. 2147 (15), REC-152 (5); Costa Soares, C. M. 115 (13); Costa-Queiroz, S. 1 (15); Costa, A. C. G. 09 (7); Costa, J. T. 80 (5); Costa, K. C. 344 (6), 377 (6); Cotarelli, V. M. 1065 (16), 1343 (1), 1343 (17), 1552 (16), 1830 (16), 1831 (16), 1832 (16), 1833 (16), 1866 (15), 1967 (15), 1987 (15), 2017 (15), 2019 (2), 2112 (2), 2172 (16), 2312 (2), 2536 (2), 860 (15); Cruz, J. 11 (6); Diniz, R. E. M. 36 (15); Duarte, R. F. 41 (2); Falcão, J. I. A. 1005 (15), 825 (10), 958 (8), s.d. (3); Falcão, M. 114 (8); Félix, L. P. 6640 (8), 7966 (2), s.d. (14); Ferrari, I. S. 3 (2); Ferrucci, M. S. 1134 (8); Filardi, F. L. R. 914 (13), 916 (7); Filho, J. B. s.d. (8); Fonseca, A. S. S. 37 (2); Fontana, A. P. 6080 (15), 6125 (15), 6259 (14), 6730 (15), 7213 (17); Forstrentes, W. 1614 (10); Fotius, G. 3343 (15), 3364 (4), 3407 (9), 3462 (2), 3473 (15), Francisco, L. V. 64 (1); França, F. 5243 (15); Freire, E. 121 (13), 127 (13); Gallindo, F. 201 (8); Gardner 1000 (2), 1001 (1), 1002 (15), 1003 (8), s.d. (15), s.d. (6); Giulietti, A. s.d. (15); Giulietti, A. M. 2127 (14), s.d. (15); Gomes de Lima, J. E. 141 (6), 142 (15), 149 (15), 217 (13), 58 (8); Gomes-Silva, F. 589 (8); Gomes, A. P. S. 451 (13); Gomes, M. L. 200 (15); GrillA, M. 124 (10); Guedes, M. L. 1685 (13); Harley, R. M. 54127 (9); Heringer, E. P. s.d. (6), s.d. (4); Irapuan, J. 25 (10), 25 (5); Jesus, M. 3 (13); Josinaldo Alves 20 (13), 546 (13), 637 (10), 692 (8); Krause, L. 170 (6); Lacerda, T. 1 (10), 25 (15), 28 (8), 30 (15), 32 (15), 34 (5), 38 (2); Laurênio, A. 1140 (15), 583 (17), 959 (8); Leite, M. S. 13 (13), 130 (17), 130 (2), 163 (2), 289 (10), 449 (10), 583 (2), 588 (13); Lima, A. 56-2589 (6), 57-2714 (13), s.d. (17), s.d. (6); Lima, J. C. -11788 (1); Lima, J. L. S. 4628 (15); Lima, O. 2007 (8); Lima, R. s.d. (13), 732 (13); Lima, R. S. 8 (10); Lima, V. C. 183 (6), 1986 (15), 453 (14), 56 (13); Lins e Silva, A. C. B. 225 (6); Lira, O. C. 105 (14), 228 (15), s.d. (15), s.d. (6), s.d. (8); Lira, S. S. 378 (8), 530 (8), 572 (8); Locatelli, E. s.d. (13); Lourteig 563 (8); Lucena, M. F. A. 152 (8), 728 (8), 75 (13), 336 (5), 1402 (15), 1450 (9), 336 (10), 504 (5); Luiz, J. s.d. (8); Luz, A. S. 83182 (5); Machado, I. C. s.d. (7); Maciel-Júnior, L. 248 (4), 272 (6); Maciel, J. R. 59 (13), 1317 (15), 625 (2), 661 (15); Mangabeira, M. O. 4412 (6); Marcon, A. B. 193 (13); Mariz, G. 03085 (8), 503 (8); Marques, J. S. 186 (8); Matos, S. S. 211 (9); Melo, A. V. 567 (13); Melo, J. I. M. 100 (13); Melo, M. R. C. S. 109 (13), 201 (13), 47 (13), 9 (15); Melquíades, A. 180 (9), 180 (10); Menezes, T. G. C. 137 (15), 75 (15); Miranda, A. M. 1546 (5), 1786 (7), 1932 (15), 2037 (8), 2222 (6), 2259 (15), 2586 (4), 2670 (17), 2673 (15), 2768 (15), 2800 (14), 3558 (2), 576 (8), 6081 (1), s.d. (14); Moura, A. C. A. 301 (14); Moura, F. 301 (14); Nádia, T. C. L. s.d. (8); Nascimento, I. S. 476 (4); Nascimento, L. M. 313 (7), 359 (7), 435 (13); Nascimento, R. M. 2 (6); Oliveira, A. C. P. 2470 (9), 2473 (15), 2909 (6), 2911 (5), 3137 (15); Oliveira, C. R. S. 176 (7), 207 (7), 331 (7); Oliveira, L. B. 21 (6); Oliveira, M. 268 (8), 288 (13), 536 (8), 3580 (14), 3691 (5), 3692 (14), 3789 (15), 3874 (15), 3939 (6), 4274 (6), 4414 (15), 4827 (15), 5143 (13), 5355 (17), 5646 (15), 5656 (15), 5689 (6); Paulo Germano s.d. (10); Pereira, R. 1125 (14), 209 (10), 2110 (7), 2806 (15), 301 (4), 327 (8), 358 (8), 681 (4), FRC-135 (10), FRC-21 (10), s.d. (13); Pessoa, E. 66 (10); Pessoa, L. M. 232 (15), 400 (5); Pessoa, M. C. 163 (15); Pick, R. A. 1221 (7), 220 (7); Pickel, B. 1293 (4), 138 (10), 1949 (8), 20 (8), 2177 (13), 2395 (10), 2733 (10), 2788 (2), 3181 (10), 3327 (8), 3574 (9), 3575 (6), 542 (15), 557 (10), 558 (10), 568 (2), 792 (8), 965 (6); Pinheiro, K. 495 (9), 541 (9), 638 (9), 679 (9); Pontual, I. 1669 (7), 338 (7), 68-834 (8); Pontual, I. B. 1332 (13); Primo, L. M. s.d. (8); R. L. 18 (2); Ramalho Campelo, C. s.d. (15); Ramos, T. 27 (15); Rocha, E. A. 1496 (7); Rodal, M. J. N. 210 (13), 620 (13), 695 (13); Rodrigues-Lima, L. 42 (9), 46 (15); Rodrigues, E. 20 (13), 47 (13); S/C s.d. (9); Sacramento, A. 136 (8), 193 (8), 31 (8), 370 (8), 402 (8), 73 (8), 81 (8); Sales, M. 1080 (15), 2011 (7), 232 (13); Sales, M. R. C. 169 (13); Santos, A. P. B 20 (15), 32 (15); Santos, E. A. 4 (13); Santos, L. L 355 (15); Santos, V. 36 (8); Sevilha, A. C. 2478 (8) Silva, A. M. 16 (13), 42 (10); Silva, E. I. 31 (13) Silva, H. C. H. 3 (5); Silva, L. B. 23 (2), 108 (17) Silva, L. F. 236 (10); Silva, L. R. 445 (8); Silva, M J. 493 (7); Silva, R. A. 2894 (6), 2894 (9); Silva, S. I. 255 (13); Silva, V. C. 18 (7); Silva, V. D. 11 (17), 46 (15); Siqueira-Filho, J. A. 2253 (17), 2529 (15), 2554 (2), 267 (7), 2696 (7), 3414 (7), 492 (13); Sobral-Leite, M. 1011 (17), 1181 (17), 1190 (13), 1191 (10), 130 (17), 446 (17), 484 (15), 510 (17), 588 (13), 797 (17), 1149 (2), 1153 (2), 1200 (2), 435 (10), 483 (2), 496 (10), 543 (10), 583 (2), 880 (2); Sobrinho, M. S. 324 (5); Souza, A. C. 3 (5); Souza, E. 41 (15), 53 (2); Souza, E. B. 60 (5); Sousa, L. G. 59 (15); Souza, M. s.d. (5), s.d. (8); Souza, S. A. O. 2394 (6); Tadeu, J. s.d. (8); Tavares, S. 20 (8), 37 (8), 606 (5), 751 (5), 751 (10); Tenório, E. 52 (13); Tonizza, M. C. s.d. (1), s.d. (2); Vasconcellos Sobrinho s.d. (10); Viana, J. L. 200 (5); Vicente, A. 11 (8); Villarouco, F. M. O. 188 (13), 65 (13); Xavier, L. P. JPB2055 (15), JPB295 (8), JPB3056 (5), JPB381 (15).
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Edited by
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Associate Editor:
Renata Sebastiani
Publication Dates
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Publication in this collection
26 May 2025 -
Date of issue
2025
History
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Received
11 Jan 2023 -
Accepted
11 Sept 2024
















