Diversity and distribution of Leguminosae on an island in the Atlantic Forest, Rio de Janeiro, Brazil

the journal of biodiversity data ANNOTATED LIST OF SPECIES 1238 Academic editor: Ana Carolina Mezzonato-Pires Received: 25 February 2025 Accepted: 27 October 2025 Published: 17 December 2025 Copyright © The authors. This is an open-access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International – CC BY 4.0) Abstract. Taxonomic knowledge is essential for assessing biodiversity. When combined with species’ spatial distribution, it serves as a tool for understanding ecological and evolutionary processes. In this study in the Rio de Janeiro State, Brazil, we associate the taxonomic diversity of shrub–tree Leguminosae with species locations across an island continuum with different Atlantic Forest phytophysiognomies. A total of 428 specimens, representing 61 species, were recorded and mapped. Our survey identified areas requiring increased sampling effort and highlighted priority sites for conservation, based on diversity, taxonomic representation, and the presence of threatened species. The highest species richness and specimen abundance were concentrated below 500 m a.s.l., within Lowland and Submontane Dense Ombrophilous Forests. Notably, only four species were represented by more than 20 specimens, while nearly 50% of species had just two specimens in the island’s vegetation sampling. Key words. Biodiversity conservation, Fabaceae, Ilha Grande, spatial distribution of Leguminosae, taxonomic backbone Castilhori MF, Manão CYG, Castelar JVS, Callado CH (2025) Diversity and distribution of Leguminosae on an island in the Atlantic Forest, Rio de Janeiro, Brazil. Check List 21 (6): 1238–1256. https:// doi.org/10.15560/21.6.1238 INTRODUCTION The continuing decline in biodiversity represents one of the most pressing environmental challenges of the twenty-first century. Quantifying this loss depends on robust taxonomic knowledge, which, when combined with information on species’ spatial distributions, is fundamental for elucidating ecological and evolutionary processes and for effectively addressing global environmental change (Jetz et al. 2012; Sandall et al. 2023). The Atlantic Forest, a coastal biome of Brazil, is recognised as one of the world’s five major biodiversity hotspots (Myers et al. 2000; Giulietti et al. 2005; Mittermeier et al. 2005; Scarano and Ceotto 2015; Almeida 2016; Rezende et al. 2018; Marques et al. 2021) and among the three most climate-vulnerable biomes globally (Bellard et al. 2014). In this context, advancing taxonomic and geographical knowledge is imperative for developing effective strategies to safeguard species and ecosystems from extinction. In this study, we highlight the remnant Atlantic Forest of Ilha Grande, Rio de Janeiro, Brazil. Distributed across 193 km2, the vegetation is a continuum of areas of Dense Ombrophilous Forest, from Lowland to Montane formations, and associated ecosystems such as Restinga (Open Restinga and Restinga Forest combined), Mangrove and Swamp Forests, and Rocky Outcrops (Callado et al. 2009). Ilha Grande has a long history of human occupation, with the first recorded inhabitants being hunter-gatherers, identified from sambaquis (shell mounds) dating back to around 3000 years BP (Oliveira 2002). The indigenous people of the Tamoios formation occupied this territory until the Brazilian colonial period (1530–1822), when coffee, sugar cane and slave-“fattening” farms were installed in its coves. Later, institutions for sanitary control (Lazaretos) and prison institutions were built (1884), the latter remaining in operation until 1994 (Santiago et al. 2009). Currently, almost all Ilha Grande is legally protected in Protected Areas: Reserva Biológica Estadual da Praia do Sul, Parque Estadual da Ilha Grande, Área de Proteção Ambiental dos Tamoios and Reserva de Desenvolvimento Sustentável do Aventureiro. However, intense tourism and disorderly human occupation continue to have impacts on the remaining forest areas. Despite the human activities on Ilha Grande, its importance was once again recognized by UNESCO in 2019 with the title of the only Brazilian site that is simultaneously a cultural and biodiversity heritage site. 21 (6) · https://doi.org/10.15560/21.6.1238 21 (6): 1238–1256. https://doi.org/10.15560/21.6.1238 Diversity and distribution of Leguminosae on an island in the Atlantic Forest, Rio de Janeiro, Brazil Marcelo Fraga Castilhori1, 2, Carla Y Gubáu Manão2, 3, João Victor de Souza Castelar1, Cátia Henriques Callado1, 2 1 Programa de Pós-graduação em Biologia Vegetal, Laboratório de Anatomia Vegetal, Departamento de Biologia Vegetal, Instituto de Biologia Roberto Alcantara Gomes, Universidade do Estado do Rio de Janeiro, Rio de Janeiro, Brazil 2 Parque Botânico da Ilha Grande, Ecomuseu Ilha Grande, Universidade do Estado do Rio de Janeiro, Rio de Janeiro, Brazil 3 Herbário RFA, Departamento de Botânica, Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil Corresponding author: Cátia Henriques Callado ([email protected]) Check List 21 (6) · https://doi.org/10.15560/21.6.1238 Castilhori et al. · Leguminosae diversity across phytophysiognomies 1239 The flora of Ilha Grande had already aroused economic interest in the beginning of the 17th century when its wood was mentioned as important to the portuguese court (Callado et al. 2009; Santiago et al. 2009). However, the oldest botanical samples from Ilha Grande were recorded in the P herbarium (Muséum National d’Histoire Naturelle, Paris, France) in 1862 (Naudeau s/no.). Only at the end of the 1980s were the first systematized inventories carried out on Ilha Grande (Araújo and Oliveira 1988), which intensified from 2002 onwards, especially in areas of Dense Ombrophilous Forest (Oliveira 2002; Manão 2011; Rosa 2013; Barros et al. 2022). Nonetheless, few studies have been developed to evaluate the distribution of species in the Ilha Grande (Antonini and Nunes-Freitas 2004; Santo 2016). The distribution of species is the result of interactions among biotic and abiotic factors and helps to understand the different existing phytophysiognomies, in addition to highlighting human actions in a given region (Costa and Santos 2011). In Brazil, studies of distribution of species aim to contribute to vegetation restoration actions (Giehl et al. 2007; Castilhori et al. 2021), enabling better management and replication in similar areas. Such studies can encompass broad geographic areas (Giulietti and Hensold 1990; Kemenes 2003; Marchioretto et al. 2008), only Protected Areas (Giehl et al. 2007; Ribeiro and Lima 2009; Marchioretto et al. 2011; Rando and Pirani 2011; Castilhori et al. 2021) or even address the distribution of specific taxonomic groups (Antonini and Nunes-Freitas 2004; Santo 2016; Silva et al. 2017; Fernandes and Milward-de-Azevedo 2021). In this context, we highlight the Leguminosae (or Fabaceae), the third largest family of angiosperms in the world (BFG 2015, 2021, 2024; LPWG 2017), with 3033 species indicated for Brazil, among which, 1012 occur in the Atlantic Forest (Flora e Funga do Brasil 2024), particularly in the Southeast Region of Brazil (Pereira and Assis 2000; Forzza et al. 2014; Castilhori et al. 2021; Machado et al. 2021; Sabino et al. 2021). Leguminosae is the second largest botanical family, in terms of number of species, on Ilha Grande (Barros et al. 2022). Thus, this study aimed to progress in the taxonomic study of shrub-tree Leguminosae and to map the distribution of native species in the continuum of different phytophysiognomies of the Atlantic Forest in the Ilha Grande to infer preferential aspects of taxon distribution and evaluate the current state of sampling in this island environment. STUDY AREA Ilha Grande is located in the municipality of Angra dos Reis in southern Rio de Janeiro State, with central coordinates 23°10’33”S, 044°12’27”W (Figure 1). It is the sixth largest coastal island of Brazil (Oliveira 2002; Manão 2011), separated from the mainland by a channel approximately 2 km wide (Esbérard et al. 2006). Geologically, Ilha Grande was formed from the uplift of the Serra do Mar at the end of the Cretaceous Period (Gama et al. 2009), giving shape to an extremely rugged relief with the highest points being Pico da Pedra D’água (1011 m above sea level) and Pico do Papagaio (989 meters above sea level.). The island is covered by a complex vegetation mosaic (Figure 1), with Dense Ombrophylous Forest being the predominant phytophysiognomy (Callado et al. 2009). The climate is humid tropical with the annual average temperature of 20–26 °C, an average annual rainfall of 2000 mm, and abundant rainfall in summer (Coe et al. 2018). Figure 1. Study Area. A. Map of Brazil showing the domain of the Atlantic Forest, with the state of Rio de Janeiro highlighted. B. Municipality of Angra dos Reis. C. Phytophysiognomic continuum of Ilha Grande. Check List 21 (6) · https://doi.org/10.15560/21.6.1238 Castilhori et al. · Leguminosae diversity across phytophysiognomies 1240 METHODS In this study, the name Leguminosae (or Fabaceae) was used for the family, according to Lewis and Schite 2003. Sampling of native shrub-tree species of Leguminosae was carried out using research authorization No. 008/2021 granted by the Instituto Estadual do Ambiente (INEA), across all existing phytophysiognomies on Ilha Grande and deposited in the Herbarium of Universidade do Estado do Rio de Janeiro (HRJ). In addition to the specimens collected during this study, materials located in the following herbaria were also included: Herbário do Instituto de Pesquisas Jardim Botânico (RB), Herbário do Museu Nacional (R), Herbário da Universidade Federal Rural do Rio de Janeiro (RBR), Herbário da Faculdade de Formação de Professores da Universidade do Estado do Rio de Janeiro (RFFP) and Herbarium Bradeanum (HB) (Thiers et al. 2016). The computerized collections Splink (Specieslink 2024) and Jabot (JBRJ 2024) were also analysed. All specimens listed in this study were examined and had their botanical determinations updated (Supplementary Material, Examined material). Maps were prepared using geographic coordinates taken during the in situ surveys or from exsiccates labels recorded in the consulted herbaria. In the absence of these data, points were plotted by approximation according to location references available in the exsiccates labels, such as roads, trails, beaches, and villages. The names, spellings, and currently valid authorship of the taxa were based on the Flora and Funga of Brazil databases (2024). Fruit types were defined according to data from Flora and Funga of Brasil (2024), except for those of the genus Inga Mill., which followed Barroso et al. (1999). To indicate the conservation status of the species, the Official List of Threatened Species of Brazilian Flora (Brasil 2022, Portaria no. 148). All recorded specimens (Supplementary Material, Examined material) were plotted on species distribution maps built with QGis software (QGis Develompent Team 2022). The zoning of the phytophysiognomies of Ilha Grande followed Callado et al. (2009). For each taxon, a single sample was maintained as a reference in the main text. The remaining samples used in the construction of the distribution maps were made available in supplementary material. The habits of the recorded species were determined based on in situ observations, indications obtained from the exsiccates labels and data available in the Flora e Funga do Brasil database (2024). RESULTS A total of 61 native species shrub-tree of Leguminosae, subordinate to 34 genera, were recorded for Ilha Grande (Table 1). Species richness was highest below 500 meters above sea level, wirh only 14 specimens, representing 10 species, recorded in the Dense Ombrophilous Forest between 500 and 1032 meters above sea level. In Submontane Dense Ombrophilous, 222 specimens of native were recorded and 73 were found in the Lowland Dense Ombrophilous Forest. Altogether, 428 specimens were georeferenced and plotted on georeferenced and maps (Figure 2). Sampling was concentrated in the vicinity of Vila do Abraão (Figure 2a), Vila Dois Rios (Figure 2b), and Praia do Aventureiro (Figure 2c), mainly along trails and the Abraão-Dois Rios road (Figure 2). Collections rarely extend more than 50 m into the forest from these access routes. Approximately 30% of the samples occurred within a 1 km radius of these three localities, excluding those close to the road. The mapping clearly reveals irregular sampling density and the absence of records in the north-western portion of the island (Figure 2). Table 1. Distribution of shrub-tree Leguminosae according to the phytophysiognomies of Ilha Grande: Montane Dense Ombrophilous Forest (MDO); Submontane Dense Ombrophilous Forest (SDO); Lowland Dense Ombrophilous Forest (LDO); Open Restinga (ORe); Restinga Forest (ReF); Swamp Forest (SF); Mangrove Forest (MF) and Rocky Outcrop (RO); (*) Threatened species. Species MDO SDO LDO ReF ORe SF MF RO Total specimens Distribution map 1Anadenanthera colubrina (Vell.) Brenan 1 15 13 1 ———— 30 Figure S1 2Andira anthelmia (Vell.) Benth. — 2 4 — — 1 — — 7 Figure S1 3Andira fraxinifolia Benth. — — 2 6 7 — — — 15 Figure S1 4Andira ormosioides Benth. — — 1 — 1 — — — 2 Figure S1 5Bauhinia forficata Link — 2 —————— 2 Figure S1 6Bauhinia longifolia (Bong.) Steud. —10 1 ————— 11 Figure S1 7Cassia ferruginea (Schrad.) Schrad. ex DC. —12 2 ————— 14 Figure S2 8Chamaecrista ensiformis (Vell.) H.S. Irwin & Barneby — 1 — — — — — — 1 Figure S2 9Copaifera langsdorffii Desf. 1 1 —————— 2 Figure S2 10 Copaifera lucens Dwyer 1 ——————— 1 Figure S2 11 Copaifera trapezifolia Hayne 1 1 —————— 2 Figure S2 Check List 21 (6) · https://doi.org/10.15560/21.6.1238 Castilhori et al. · Leguminosae diversity across phytophysiognomies 1241 Species MDO SDO LDO ReF ORe SF MF RO Total specimens Distribution map 12 Dahlstedtia pinnata (Benth.) Malme — 7 1 — — — — — 8 Figure S2 13 Dalbergia ecastaphyllum (L.) Taub — — — — 9 — 4 2 15 Figure S3 14 Erythrina speciosa Andrews — 5 6 — — — — — 11 Figure S3 15 Erythrina verna Vell. — 1 —————— 1 Figure S3 16 Exostyles venusta Schott — 1 —————— 1 Figure S3 17 Guilandina bonduc L. ———— 4 ——— 4 Figure S3 18 Hydrochorea pedicellaris (DC.) M.V.B. Soares, Iganci & M.P. Morim 1 5 2 — — — — — 8 Figure S4 19 Hymenaea courbaril L. 1 ——————— 1 Figure S4 20 Indigofera suffruticosa Mill. — 1 1 — 2 — — — 4 Figure S4 21 Inga bullata Benth. — 6 1 — — — — — 7 Figure S4 22 Inga capitata Desv. — 2 —————— 2 Figure S4 23 Inga edulis Mart. —11 2 6 — — — — 19 Figure S4 24 Inga flagelliformis (Vell.) Mart. — 2 — — — — — — 2 Figure S5 25 Inga lanceifolia Benth. 2 1 — — — 1 — — 4 Figure S5 26 Inga laurina (Sw.) Willd. — 1 — — 1 — — — 2 Figure S5 27 Inga marginata Willd. — 5 — — — 1 — — 6 Figure S5 28 Inga maritima Benth.* — — — 1 1 — — — 2 Figure S5 29 Inga sellowiana Benth. ——— 2 ———— 2 Figure S5 30 Inga sessilis (Vell.) Mart. — 4 — — — — — — 4 Figure S6 31 Inga striata Benth. — 1 —————— 1 Figure S6 32 Inga subnuda subsp. luschnathiana (Benth.) T.D.Penn. —17 3—1——1 22 Figure S6 33 Inga tenuis (Vell.) Mart. — 1 — — — — — — 1 Figure S6 34 Inga vera subsp. affinis (DC.) T.D.Penn. — 1 —————— 1 Figure S6 35 Jupunba brachystachya (DC.) M.V.B. Soares, M.P. Morim & Iganci — 9 5 10 12 — — 2 38 Figure S7 36 Jupunba langsdorffii (Benth.) M.V.B. Soares, M.P. Morim & Iganci ——————— 2 2 Figure S7 37 Lonchocarpus cultratus (Vell.) A.M.G.Azevedo & H.C.Lima — 12 —————— 12 Figure S7 38 Marlimorimia contorta (DC.) L.P. Queiroz & P.G. Ribeiro — 2 1 — — — — — 3 Figure S7 39 Mimosa bimucronata (DC.) Kuntze — 2 6 — — 1 2 — 11 Figure S7 40 Myrocarpus frondosus Allemão — 2 — — — — — — 2 Figure S7 41 Ormosia arborea (Vell.) Harms. — 1 1 4 2 — — — 8 Figure S8 42 Ormosia fastigiata Tul. 1 ——————— 1 Figure S8 43 Piptadenia gonoacantha (Mart.) J.F.Macbr. — 8 2 1 — — — — 11 Figure S8 44 Pityrocarpa schumanniana (Taub.) L.P. Queiroz & L.M. Borges — 1 —————— 1 Figure S8 45 Platymiscium floribundum Vogel — 2 —————— 2 Figure S8 46 Pterocarpus violaceus Vogel — 5 —————— 5 Figure S8 47 Schizolobium parahyba (Vell.) Blake — 15 3 ————— 18 Figure S9 48 Senna macranthera (DC. Ex Collad.) H.S.Irwin & Barneby — 4 4 — — — — — 8 Figure S9 49 Senna multijuga (Rich.) H.S.Irwin & Barneby — 17 6 ————— 23 Figure S9 50 Senna pendula (Humb. & Bonpl.ex Willd.) H.S.Irwin & Barneby — 1 1 — 3 — — — 5 Figure S9 51 Senna silvestris (Vell.) H.S.Irwin & Barneby — 4 — — — — — — 4 Figure S9 52 Sophora tomentosa L. ———— 9 ——— 9 Figure S10 53 Stryphnodendron polyphyllum Mart. 313 2 ————— 18 Figure S10 54 Swartzia acutifolia Vogel — 1 —————— 1 Figure S10 55 Swartzia flaemingii Raddi — 1 1 — — — — — 2 Figure S10 56 Swartzia myrtifolia Sm. — 1 —————— 1 Figure S10 57 Tachigali paratyensis (Vell.) H.C. Lima — 11 3 ————— 14 Figure S11 58 Vatairea heteroptera (Allemão) Ducke 2 — — — — — — — 2 Figure S11 59 Zollernia glabra (Spreng.) Yakovlev — 4 — 1 — — — — 5 Figure S11 60 Zollernia ilicifolia (Brongn.) Vogel ——— 2 ———— 2 Figure S11 61 Zygia latifolia (L.) Fawc. & Rendle — 5 — — — — — — 5 Figure S11 Total 14 222 73 34 51 467 428 Check List 21 (6) · https://doi.org/10.15560/21.6.1238 Castilhori et al. · Leguminosae diversity across phytophysiognomies 1242 Jupunba brachystachya (DC.) M.V.B. Soares, M.P. Morim & Iganci, a small shrubby species found sparsely, occurs in the largest number of phytophysiognomies (Submontane Dense Ombrophilous Forest, Lowland Dense Ombrophilous Forest, Restinga Forest, Open Restinga and Rocky Outcrop) and accounts for the largest number of specimens (38). Anadenanthera colubrina (Vell.) Brenan (30 specimens), Senna multijuga (Rich.) H.S.Irwin & Barneby (23 specimens), Inga subnuda subsp. luschnathiana (Benth.) T.D.Penn. (22 samples), Schizolobium parahyba (Vell.) Blake (18 samples) and Stryphnodendron polyphyllum Mart. (17 specimens) complete the group of the six most frequently sampled species (Table 1). Specifically, S. parahyba has a concentration of approximately 10 individuals on Aventureiro beach due to ongoing anatomical studies. It is worth noting that some areas under anthropogenic impact or at the beginning of natural succession have greater species richness, as observed on the banks of the Abraão–Dois Rios road, where species of the genera Bauhinia L. and Cassia L. were found in areas of Submontane Dense Ombrophilous Forest and the Dense Lowland Ombrophilous Forest. Mangrove Forest, in turn, was the phytophysiognomy with the lowest number of Leguminosae. Dalbergia ecastaphyllum (L.) Taub and Mimosa bimucronata (DC.) Kuntze were recorded in this phytophysiognomy, but always in areas of transition with Restinga Forest. Family Leguminosae Juss. Subfamily Caesalpinioideae DC. Anadenanthera colubrina (Vell.) Brenan Figure 3A Material examined. BRAZIL — Rio de JaneiRo • Angra dos Reis, Ilha Grande, road to Vila Dois Rios; 23°09’48”S, 044°10’44”W; 295m alt.; 28.IX.2013; M.F. Castilhori 1092 & L.P.G. Rosa, leg.; Montane Dense Ombrophilous Forest; HB95645. Identification. Tree 5–20 m high, lenticellate branch; bipinnate leaf, 10-20 pairs of pinnae, petiole with extrafloral nectary, oblong leaflet, 0.3–0.6 cm long; globose spike inflorescence; white flower; follicle fruit, 10–30 cm long, compressed plane, crenate margin. Remarks. Phenology: collected from flowers inJanuary, February, March, November, and December, and from fruit in February to November. Cassia ferruginea (Schrad.) Schrad. ex DC. Material examined. BRAZIL — Rio de JaneiRo • Angra dos Reis, Ilha Grande, road to Vila Dois Rios; 23°10’15”S, 044°11’01”W; 130 m alt.; 18.XI.2014; M.F. Castilhori 1275 & I.E. Santo, leg.; Submontane Dense Ombrophilous Forest; HB96197. Figure 2. General distribution of arboreal Leguminosae sampled on Ilha Grande. Circles delimit radii of 1 km: (a) Vila do Abraão, (b) Vila Dois Rios and (c) Praia do Aventureiro. Straight line: Estrada Abraão - Dois Rios. Check List 21 (6) · https://doi.org/10.15560/21.6.1238 Castilhori et al. · Leguminosae diversity across phytophysiognomies 1243 Identification. Tree 4–10 m high, glabrous branches; pinnate leaf, 36–70 leaflets, nectary absent; leaflet 0.5–1.0 cm long; inflorescence in raceme; flower yellow, anther reddish; fruit lomentum, 30–45 cm long, straight, cylindrical. Remarks. Phenology: collected from flowers in November, and from fruit in January, April, July, and December. Chamaecrista ensiformis (Vell.) H.S.Irwin & Barneby Material examined. BRAZIL — Rio de JaneiRo • Angra dos Reis, Ilha Grande, Praia dos Castellanos; 23°09’56”S, 044°05’59”W; 90 m alt.; 21.III.2010; A.A.M. Barros 3973 & G.M. Vabo, leg.; Submontane Dense Ombrophilous Forest; RFFP13400. Identification. Tree 4 m high, lenticellate branch; pinnate leaf, leaflets 6–16, leaflet 2–7 cm long; inflorescence in raceme; flower yellow; fruit legume 11–13 cm long, flat compressed, oblong, elastic dehiscence. Remarks. Phenology: collected from flowers in March. Guilandina bonduc L. Figure 3G Material examined. BRAZIL — Rio de JaneiRo • Angra dos Reis, Ilha Grande, REBIO Praia do Sul; 23°10’41”S, 044°18’35”W; 15 m alt.; 26.X.2023; M.F. Castilhori 2181, D. Barbosa & S. Nascimento, leg.; Open Restinga; HRJ13988. Identification. Climbing shrub 1–2 m high, branch with erect trichomes associated with abundant spines; bipinnate leaf, 6–16 leaflets, oval to elliptical, 2–6 cm long; inflorescence in raceme; pale yellow flower; legume fruit, 4–8 cm long, dry, densely covered with spines, globose seed, 15–20 cm long, grey. Remarks. Phenology: collected flowers in February, and from fruit in September and October. Hydrochorea pedicellaris (DC.) M.V.B.Soares, Iganci & M.P.Morim Figure 3H Material examined. BRAZIL — Rio de JaneiRo • Angra dos Reis, Ilha Grande, Abraão; 23°07’54”S, 044°10’26”W; 38 m alt.; 20.XI.2014; M.F. Castilhori 1331 & I.E. Santo, leg.; Lowland Dense Ombrophilous Forest; HB96210 Identification. Tree 8–20 m high, puberulent branch; bipinnate leaf; pinnae 5–7 pairs, leaflet 0.3–1 cm long, oblong lanceolate; inflorescence globose spike; pink flower; lomentiform fruit 8–10 cm long, compressed flat. Remarks. Phenology: collected flowers and fruit in November. Inga bullata Benth. Figure 4B Material examined. BRAZIL — Rio de JaneiRo • Angra dos Reis, Ilha Grande, trail to Parnaioca; 23°11’38”S, 044°14’13”W; 220 m alt.; 06.XI.2014; M.F. Castilhori et al., 1225, leg.; Submontane Dense Ombrophilous; HB96147 Identification. Shrub to tree 2–5 m high, hirsute to glabrous branch, lenticellate; pinnate leaf, falciform stipule, 4–6 leaflets, obovate, 4–10 cm long, hairy rachis, canaliculate, extrafloral nectary between pairs of leaflets; white flower, androecium with more than 10 stamens, exsert, striking, sweet odor; capituliform inflorescence; nucoid legume 10–20 cm long, smooth and glabrous, yellow when ripe, compressed. Remarks. Phenology: collected flowers inJanuary, February, September, and November, and from fruit in March, September, and November. Figure 3. Leguminosae in Ilha Grande. A. Anadenanthera columbrina. B. Andira fraxinifolia. C. Bauhinia longifolia. D. Dahlstedtia pinnata. E. Dalbergia ecastaphyllum. F. Erythrina speciosa. G. Guilandina bounduc. H. Hydrochorea pedicellaris. Check List 21 (6) · https://doi.org/10.15560/21.6.1238 Castilhori et al. · Leguminosae diversity across phytophysiognomies 1244 Inga capitata Desv. Material examined. BRAZIL — Rio de JaneiRo • Angra dos Reis, Ilha Grande, Angra dos Reis, Ilha Grande, Vila Dois Rios, Morro do Cavalinho; 23°11’30”S, 044°11’33”W; 136 m alt.; 18.VIII.2012; A.A.M. Barros 4766 et al., leg.; Submontane Dense Ombrophilous Forest; RFFP15011. Identification. Tree 10 m high, lenticellate branch; pinnate leaf, sickle–shaped stipule, 4–6 leaflets, elliptical, 5–15 cm long, marginated rachis; extrafloral nectary between pairs of leaflets; capituliform inflorescence; white flower, androecium with more than 10 stamens, exserted, striking, sweet odor; nucoid legume 10–20 cm long, oblong. Remarks. Phenology: collected fruit in August. Inga edulis Mart. Figure 4C Material examined. BRAZIL — Rio de JaneiRo • Angra dos Reis, Ilha Grande, trail to Parnaioca; 23°10’54”S, 044°11’45”W; 60 m alt.; 05.II.2014; M.F. Castilhori et al. 1076, leg.; Submontane Dense Ombrophilous Forest; HB95531. Identification. Tree 5–20 m high, lenticellate branch; pinnate leaf, 6–10 leaflets, elliptical to obovate, 5–18 cm long, winged rachis, dark extrafloral nectary, elliptical between pairs of leaflets; inflorescence spiciform inflorescence; white flower, androecium with more than 10 exposed, striking stamens, sweet odor; nucoid legume 10–20 cm long, extended margin, longitudinal grooves, woody. Remarks. Phenology: collected in flower inFebruary, March, April, July, August, October, and December, and from fruit in February, March, April, July, August, October, November, and December. Inga flagelliformis (Vell.) Mart. Material examined. BRAZIL — Rio de JaneiRo • Angra dos Reis, Ilha Grande, RAPELD project; 23°09’48”S, 044°11’03”W 23.II.2019; A.A.M. Barros et al., 5691, leg.; Submontane Dense Ombrophilous Forest; RFFP20328. Identification. Tree; glabrous branch; pinnate leaf, 4–6 leaflets, elliptical, 10–20 cm long, cylindrical rachis, extrafloral nectary between pairs of leaflets; umbel inflorescence; white flower, androecium with more than 10 exposed, striking stamens, sweet odor; nucoid legume 10–20 cm, oblong, flat and compressed, without striations, leathery. Remarks. Phenology: collected from fruits in November. Inga lanceifolia Benth. Material examined. BRAZIL — Rio de JaneiRo • Angra dos Reis, Ilha Grande, Pico do Papagaio; 23°09’11”S, 044°11’21”W; 707 m alt.; 15.IX.2012; L.P.G. Rosa s.n. leg.; Montane Dense Ombrophilous Forest; HRJ12320. Identification. Tree 3 m high, lenticellate, striated branch; pinnate leaf, 4–6 leaflets, elliptical, 6–8 cm long, cylindrical rachis, extrafloral nectary between pairs of leaflets; congested, glomeruliform inflorescence; white flower, androecium with more than 10 prominent, striking stamens, sweet odor; nucoid legume 10–20 cm, oblong, flat and compressed, leathery valves. Remarks. Phenology: collected from flowers in September, and fruit in August and September. Inga laurina (Sw.) Willd. Figure 4D Material examined. BRAZIL — Rio de JaneiRo • Angra dos Reis, Ilha Grande, Vila Dois Rios; 23°10’53”S, 044°11’31”W; 3 m alt.; 25.IV.2020; M.F. Castilhori 2020, leg.; Open Restinga; HRJ13920. Identification. Tree 3 m high, lenticellate branch; pinnate leaf, 4 leaflets, obovate, 4–15 cm long, winged Figure 4. Leguminosae in Ilha Grande. A. Indigofera suffruticosa. B. Inga bullata. C. Inga edulis. D. Inga laurina. E. Inga maritima. F. Inga subnuda. G. Inga vera. Check List 21 (6) · https://doi.org/10.15560/21.6.1238 Castilhori et al. · Leguminosae diversity across phytophysiognomies 1245 rachis, extrafloral nectary between pairs of leaflets; spiciform inflorescence; white flower, androecium with more than 10 exposed, striking stamens, sweet odor; nucoid legume 3–10 cm long, smooth, yellow when ripe. Remarks. Phenology: collected from flowers in November and from fruit in April. Inga marginata Willd. Material examined. BRAZIL — Rio de JaneiRo • Angra dos Reis, Ilha Grande, Praia do Sul, Rio Capivari; 23°10’20”S, 044°17’17”W; 2 m alt.; 06.IX.1991; R. Ribeiro 2017, leg.; Mata Alagadiça; RB855316. Identification. Tree 4 m high, lenticellate branch; pinnate leaf, leaflet 4–6; elliptical to lanceolate, 8–16 cm, winged rachis, extrafloral nectary between pairs of leaflets; spike-like inflorescence; white flower, androecium with more than 10 prominent, striking stamens, sweet odor; nucoid legume 6–12 cm long, smooth, leathery. Remarks. Phenology: collected from flowers in January, September, and November, and from fruit in November. Inga maritima Benth. Figure 4E Material examined. BRAZIL — Rio de JaneiRo • Angra dos Reis, Ilha Grande, Lopes Mendes; 23°10’12”S, 044°07’34”W; 2 m alt.; 9.X.2013; M.F. Castilhori et al. 1056, leg.; Restinga Forest; HB95532. Identification. Shrub 2 m high, lenticellate branch, villous; pinnate leaf, 6 leaflets, elliptical; winged rachis; spiciform inflorescence; white flower, androecium with more than 10 exposed, striking stamens, sweet odor; nucoid legume 5–10 cm long, smooth, yellow when ripe. Remarks Phenology: collected from flowers in July and October. Conservation status: Endangered. Inga sellowiana Benth. Material examined. BRAZIL — Rio de JaneiRo • Angra dos Reis, Ilha Grande, • REBIO Praia do Sul; 23°10’30”S, 044°17’29”W; 10 m alt.; 11.IV.1984; D.S.D. Araújo et al., 6217, leg.; Restinga Forest; RB371309. Identification. Shrub up to 3 m tall, glabrous, lenticellate branch; pinnate leaf, 4–6 leaflets; obovate to elliptical, 5–10 cm, cylindrical rachis, extrafloral nectary between pairs of leaflets; umbel inflorescence; white flower, androecium with more than 10 exposed, striking stamens, sweet odor; nucoid legume 3–15 cm long, smooth, leathery, yellow when ripe. Remarks. Phenology: collected from flowers and fruit in April. Inga sessilis (Vell.) Mart. Material examined. BRAZIL — Rio de JaneiRo • Angra dos Reis, Ilha Grande, road to Vila Dois Rios; 23°10’16”S, 044°11’02”W; 115 m alt.; 28.X.2013; M.F. Castilhori 1093 & L.P.G. Rosa, leg.; Submontane Dense Ombrophilous Forest; HB95644. Identification. Tree 6–8 m high, tomentose, angular branch; pinnate leaf, 8–10 leaflets, elliptical, winged rachis, extrafloral nectary present between the pairs of leaflets; white flower, androecium with more than 10 exsert, striking stamens, sweet odor; spiciform inflorescence; nucoid legume 10–20 cm long, curved, velvety, blackish trichomes on the edges that are thick. Remarks. Phenology: collected from flowers in September, and from fruit in August, September, October, and November. Inga striata Benth. Material examined. BRAZIL — Rio de JaneiRo • Angra dos Reis, Ilha Grande, Sítio Forte; 23°08’55”S, 044°16’57”W; 500 m alt.; 24.V.1979; D.S.D. Araújo 3069, N.C. Maciel, leg.; Montane Dense Ombrophilous Forest; RB 861858. Identification. Tree up to 3 m tall, hispid to villous, striate branch; pinnate leaf, 8–10 leaflets; elliptical, 5–10 cm, rachis winged, extrafloral nectary between pairs of leaflets; spiciform inflorescence; white flower, androecium with more than 10 exposed, striking stamens, sweet odor; nucoid legume 21 cm long, linear, dense hairy, woody yellow when ripe. Remarks. Phenology: collected from flowers in May. Inga subnuda subsp. luschnathiana (Benth.) T.D.Penn. Figure 4F Material examined. BRAZIL — Rio de JaneiRo • Angra dos Reis, Ilha Grande, road to Vila Dois Rios; 23°10’27”S, 044°11’07”W; 65 m alt.; 28.IX.2013; M.F. Castilhori 1096 & L.P.G. Rosa, leg.; Submontane Dense Ombrophilous Forest. Check List 21 (6) · https://doi.org/10.15560/21.6.1238 Castilhori et al. · Leguminosae diversity across phytophysiognomies 1246 Identification. Tree to shrub 2–8 m high, villous branch; pinnate leaf, 6–8 leaflets, obovate to elliptical; winged rachis, extrafloral nectary between pairs of leaflets; spiciform inflorescence; white flower, androecium with more than 10 prominent, striking stamens, sweet odor; nucoid legume, 14–20 cm long, tomentose, striated, thickened margin. Remarks. Phenology: collected from flowers from May to December, and from fruits in January, February, March, May, September, November, and December. Inga tenuis (Vell.) Mart. Material examined. BRAZIL — Rio de JaneiRo • Angra dos Reis, Ilha Grande, Pico do Papagaio; 23°09’00”S, 044°11’11”W; 570 m alt.; 30.IV.1996; R.R. Oliveira et al. s.n., D. Flores; C.A.L. Oliveira, leg.; Montane Dense Ombrophilous Forest; FCAB5443. Identification. Small tree 6 m high, hairy, lenticelated branch; leaf 6–10 leaflets, elliptical to obovate; winged rachis, extrafloral nectary between pairs of leaflets; capituliform inflorescence; white flower, androecium with more than 10 exposed, striking stamens, sweet odor; nucoid legume 18–20 cm long, flat, linear, lenticelated, yellow when ripe. Remarks. Phenology: collected flowers in April. Inga vera subsp. affinis (DC.) T.D.Penn. Figure 4G Material examined. BRAZIL — Rio de JaneiRo • Angra dos Reis, Ilha Grande, trail between Palmas and Abraão; 23°08’29”S, 044°08’46”W; 160 m alt.; 03.XI.2000; D. Fernandes 575.; Submontane Dense Ombrophilous Forest; RB351749. Identification. Tree 4 m high, tomentose, brown, lenticellate branch; pinnate leaf, 8–10 leaflets, elliptical to oval; winged rachis, extrafloral nectary present between pairs of leaflets; spike-shaped inflorescence; white flower, androecium with more than 10 prominent, striking stamens, sweet odor; nucoid legume 10–20 cm long, linear, tomentose, leathery, thickened margin. Remarks. Phenology: collected from flowers in November. Jupunba brachystachya (DC.) M.V.B.Soares et al. Figure 5A, D Material examined. BRAZIL — Rio de JaneiRo • Angra dos Reis, Ilha Grande, trail to Caxadaço; 23°10’35”S, 044°10’57”W; 155 m alt.; 18.XI.2014; M.F. Castilhori 1272 & I.E. Santo, leg.; Submontane Dense Ombrophilous Forest; HB96194. Identification. Small tree 2–5 m high, pubescent branch; bipinnate leaf, pinnae 3–6, leaflets 6–10, quadrangular to obovate, 2.5–10 cm long, extrafloral nectary between the pinnae; axillary spiciform inflorescence; greenish white flower, exsert stamens; spiral legume fruit, 5–15 cm long, bicolor white and black seed. Remarks. Phenology: collected from flowers from February to November, and from fruit in from February to December. Figure 5. Leguminosae in Ilha Grande. A, D. Jupunba brachystachya. B, E. Jupunba langsdorfii. C. Lonchocarpus cultratus. Check List 21 (6) · https://doi.org/10.15560/21.6.1238 Castilhori et al. · Leguminosae diversity across phytophysiognomies 1253 Remarks. Phenology: collected from flowers and fruits in June to November. Swartzia acutifolia Vogel Material examined. BRAZIL — Rio de JaneiRo • Angra dos Reis, Ilha Grande, REBIO Praia dos Sul; 23°10’37”S, 044°18’09”W; 7 m alt.; 11.XII.1992; R. Oliveira s.n., D. Flores, D. Toffoli & P. Delamonica, leg.; Lowland Dense Ombrophilous Forest; FCAB7124. Identification. Tree, glabrescent branch; pinnate leaf, 15–20 leaflets, elliptical to lanceolate; racemose panicle inflorescence; white flower; nucoid legume fruit. Remarks. Phenology: collected from fruit in December. Swartzia flaemingii Raddi Material examined. BRAZIL — Rio de JaneiRo • Angra dos Reis, Ilha Grande, Aventureiro, 23°11’29”S, 044°19’11”W; 97 m alt.; 17.I.2003 D. Velludo 36, leg.; Submontane Dense Ombrophilous Forest; RB 553376. Identification. Tree to shrub 2–3 m high; strigose branch; pinnate leaf, leaflets 11–20, oblong, 5–8.5 cm long; inflorescence raceme or panicle; flower white; fruit ellipsoide nucoid legume, seed 1, aril yellow. Remarks. Phenology: collected from flowers in January and August. Swartzia myrtifolia Sm. Material examined. BRAZIL — Rio de JaneiRo • Angra dos Reis, Ilha Grande, REBIO Praia do Sul, Rio Capivari; 23°11’03”S, 044°15’06”W; 70 m alt.; 13.VIII.1985; R. Ribeiro 671, leg.; Submontane Dense Ombrophilous Forest; RB 867289. Identification. Small tree, hairy branch; petiole marginated; leaflets 5–7, ovate, lanceolate or elliptical; inflorescence axillary, hairy peduncle; flower with globose floral bud, yellow corolla, glabrous legume fruit, 3–6 cm long, oblong-elliptical. Remarks. Phenology: collected from fruit in August. Vatairea heteroptera (Allemão) Ducke Material examined. BRAZIL — Rio de JaneiRo • Angra dos Reis, Ilha Grande, Pedra D’ Água; 23°09’09”S, 044°13’34”W; 880 m alt.; 01.VIII.2013; M.F. Castilhori 999 & L.P.G Rosa, leg.; HB95477. Identification. Tree 20 m high, buttresses at the base, glabrescent branch; pinnate leaf, leaflet 9–19 pairs, elliptical; panicle inflorescence; lilac flower; samara fruit, 10–12 cm long. Remarks. Phenology: collected from fruit in August. Zollernia glabra (Spreng.) Yakovlev Figure 8C, G Material examined. BRAZIL — Rio de JaneiRo • Angra dos Reis, Ilha Grande, Pouso; 23°09’35”S, 044°08’17”W; 2 m alt.; 25.XI.202; M.F. Castilhori 2195 leg.; Restinga Forest; HRJ14300. Identification. Shrub to tree 2 m high, glabrous, lenticulate branch; simple leaf 3.5–18.0 cm long, ovate to elliptical, stipule 0.6–1.7 cm long, persistent, falcate; inflorescence raceme, flower pink, drupe fruit, 1.5–3.5 cm long, globose. Remarks. Phenology: collected from flowers in September, and from fruit in November and December. Zollernia ilicifolia (Brongn.) Vogel Material examined. BRAZIL — Rio de JaneiRo • Angra dos Reis, Ilha Grande, REBIO Praia do Sul; 23°10’46”S, 044°16’07”W; 5 m alt.; 01.IX.2017; M.F. Castilhori 1931 & J.V. Castelar, leg.; Restinga Forest; HRJ13207. Identification. Shrub 5–20 m high, glabrous, lenticulate branch; unifoliate leaf, 4–9 cm long, obovate to elliptic, stipule 0.4–0.9 cm long, persistent, falcate; inflorescence raceme, flower pink, drupe fruit, 2.8–3.0 cm long, globose. Remarks. Phenology: collected from fruit in September and October. DISCUSSION Dense Ombrophilous Forest is the predominant phytophysiognomy on Ilha Grande (Callado et al. 2009). However, the rugged relief, proximity to rivers or the sea, and history of use confer distinct environmental gradients and varied microclimates, in addition to different successional stages (Oliveira and Coelho-Netto 2001; Oliveira 2002; Salgado and Vasquez 2009). These environmental gradients favor the occurrence of different species and directly contribute to greater plant diversity. Callado et al. (2009), for example, referred Check List 21 (6) · https://doi.org/10.15560/21.6.1238 Castilhori et al. · Leguminosae diversity across phytophysiognomies 1254 to the existence of 972 species for Ilha Grande, while Barros et al. (2022), indicated that 509 of these species are trees. In these studies, Leguminosae appear as the fourth (Callado et al. 2009) or the second (Barros et al. 2022) most representative botanical family in terms of the number of species on Ilha Grande. Comparing the exclusively arboreal species identified in this work to studies on the diversity of arboreal Leguminosae in coastal forests of Rio de Janeiro (Machado 2021), Ilha Grande appears to be slightly more similar to the species indicated for the localities of Niterói and Maricá and Ilha da Marambaia. Floristic and phytosociological studies in areas of Submontane Dense Ombrophilous Forest and Montane Dense Ombrophilous Forest on Ilha Grande (Manão 2011; Rosa 2013, respectively) corroborate the greater species richness for Leguminosae below 500 m a.s.l. (Table 1). Lowland Dense Ombrophilous Forest is the most anthropogenically impacted phytophysiognomy on Ilha Grande, with profoundly altered areas (Callado et al. 2009), which may have influenced its lower richness of native species in relation to the other two areas. The occurrence of Inga maritima, a species Endangered (Brasil 2022) in both Open Restinga and Restinga Forest of Praia de Lopes Mendes, constitutes the only species of the family endangered on Ilha Grande and reinforces the need for conservation measures for this region that it was previously impacted by a landing strip for small aircraft. Furthermore, the uneven distribution of samples, with lower representation in the northwestern portion of the island, likely reflects the access restrictions and logistical challenges in that area. This pattern contrasts with regions located closer to the support base of the Rio de Janeiro State University, where the available infrastructure favors a greater number of collections. The results of the present study validate the existence of 61 native species of shrub-tree Leguminosae in an island environment of 193 km2 and highlight their spatial organization along a continuum of eight different phytophysiognomies of the Atlantic Forest. Although the floristic list is increased by four species in relation to the last work carried out on Ilha Grande (Barros et al. 2022), the plot of recorded specimens identifies under-sampled areas that correspond to the most difficult to access areas. In this sense, the results demonstrate the importance of studying species distribution to better understand the ecology of Leguminosae. Furthermore, they help identify areas requiring greater sampling effort and priority areas for conservation, either based on the criterion of greater taxon representation or the presence of endangered species. Lastly, verification and taxonomic correction in the herbaria revealed that the species Abarema cochliacarpos (Gomes) Barneby & J.W. Grimes and Tachigali denudata (Vogel) Oliveira-Filho are not recorded for Ilha Grande. ACKNOWLEDGEMENTS We thank Ricardo Reis for his support during the collections for his support with the manuscript writing standards, the Instituto Estadual do Ambiente (INEA) for authorization of the research (No. 070002/001050/2021), CEADS and ECOMUSEU Ilha Grande for logistical support. ADDITIONAL INFORMATION This study was part of M.F. Castilhori’s doctoral thesis in the Postgraduate Program in Plant Biology at the Universidade do Estado do Rio de Janeiro. Conflict of interest The authors declare no competing interests exist. Ethical statement No ethical statement is reported. Funding This study was financed in part by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES; Financial Code 001) and the Fundação Carlos Chagas Filho de Amparo à Pesquisa do Estado do Rio de Janeiro (FAPERJ; E-26/210.045/2021, E-26/200.961/2021 and E-26/204.346/2024). Author contributions Conceptualization: MFC, CYGM, CHC. Data curation: MFC, CHC. Formal analysis: MFC, CYGM, CHC. Funding acquisition: CHC. Investigation: MFC. Methodology: MFC, CYGM. Resources: CHC. Supervision: CHC. Project administration: MFC, CHC. Validation: MFC, CYGM, CHC. Visualization: MFC, CYGM, JVSC, CHC. Writing – original draft: MFC, CHC. Writing – review and editing: MFC, CYGM, JVSC, CHC. Author ORCID iDs Marcelo Fraga Castilhori https://orcid.org/0000-0003-2784-3254 Carla Y Gubáu Manão https://orcid.org/0000-0002-7705-9751 João Victor de Souza Castelar https://orcid.org/0000-0003-0110-6643 Cátia Henriques Callado https://orcid.org/0000-0003-2190-3375 Check List 21 (6) · https://doi.org/10.15560/21.6.1238 Castilhori et al. · Leguminosae diversity across phytophysiognomies 1255 Data availability All data that support the findings of this study are available in the main text and Supplementary material. 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