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Malpighiaceae in southern Brazil: Identification keys for genera and updates on species richness

Abstract

Recent studies have contributed to a better understanding of the circumscription of some genera of Malpighiaceae and the overall diversity of the family in Brazilian tropical ecosystems. In southern Brazil, however, this important group of lianas and shrubs still has not been fully categorized. The present study provides identification keys based on floral and fruit morphology for the 23 genera reported from southern Brazil. The importance of the number of styles and the position and shape of the stigmas for the recognition of genera was confirmed. Illustrations of morphological characteristics used in the keys and notes about taxonomy, species richness and distribution are provided. It should be noted that the occurrence of some Malpighiaceae genera that are represented by only one species in the southern region, such as Bronwenia, Camarea and Heladena, are not yet cited in the Flora do Brasil 2020 database. A total of 95 species were confirmed and documented by voucher specimens. Banisteriopsis pseudojanusia, Heteropterys dusenii and Peixotoa catarinensis, occur exclusively in southern Brazil and are classified as “Critically Endangered”, “Vulnerable” and “Endangered” in the Brazilian Red List, respectively.

Key words
Atlantic Forest; Cerrado; lianas; Malpighiales; Pampa

Resumo

Embora recentes estudos tenham contribuído para uma melhor compreensão da circunscrição de gêneros e da diversidade da família Malpighiaceae em ecossistemas tropicais brasileiros, na Região Sul, esse importante grupo de lianas e arbustos ainda apresenta dificuldades para a sua identificação. O presente estudo traz chaves de identificação baseadas na morfologia de flores e frutos para 23 gêneros de Malpighiaceae registrados no sul do Brasil. A importância do número de estiletes e da posição e forma dos estigmas para o reconhecimento de gêneros foi confirmada. Adicionalmente, são apresentadas ilustrações relativas às características morfológicas incluídas nas chaves, além de observações sobre a taxonomia, riqueza de espécies e distribuição dos gêneros abordados. Cabe destacar que a ocorrência de alguns gêneros representados na Região Sul por apenas uma espécie, como é o caso de Bronwenia, Camarea e Heladena, não é mencionada na Flora do Brasil 2020, em construção. Ao total, 95 espécies de Malpighiaceae foram confirmadas para a região e documentadas por materiais testemunho. Banisteriopsis pseudojanusia, Heteropterys dusenii e Peixotoa catarinensis ocorrem exclusivamente no sul do Brasil e são classificadas como “Criticamente em Perigo”, “Vulnerável” e “Em Perigo”, respectivamente, no Livro Vermelho da Flora do Brasil.

Palavras-chave
Mata Altântica; Cerrado; lianas; Malpighiales; Pampa

Introduction

Malpighiaceae is composed of approximately 75 genera and 1300 species, including a great diversity of neotropical lianas, shrubs and trees (Anderson et al. 2006Anderson WR, Anderson C & Davis CC (2006) Malpighiaceae. Available at <http://herbarium.lsa.umich.edu/malpigh.>. Accessed on 14 May 2019.
http://herbarium.lsa.umich.edu/malpigh....
). Most of this species richness is found in Brazil and is frequently associated with more-open vegetation types, such as Cerrado and Campos Rupestres (Anderson 2004Anderson WR (2004) Malpighiaceae. In: Smith N, Mori SA, Henderson A, Stevenson DWM & Heald SV. Flowering Plants of the Neotropics. Princeton Press, New Jersey. Pp. 229-232.; Mamede et al. 2015Mamede MCH, Sebastiani R, Almeida RF de, Francener A& Amorim AMA (2015) Malpighiaceae. In: Lista de Espécies da Flora do Brasil. Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br/jabot/floradobrasil/FB155>. Accessed on 31 March 2020.
http://floradobrasil.jbrj.gov.br/jabot/f...
), or rain forests (Almeida et al. 2016Almeida RF de, Francener A & Amorim AMA (2016) A generic synopsis of Malpighiaceae in the Atlantic Forest. Nordic Journal of Botany 34:285-301. ). Despite being more diverse in tropical ecosystems, some species of Malpighiaceae reach extratropical latitudes. In southern Brazil, Malpighiaceae occur in both forested and non-forested ecosystems (Amorim et al. 2013Amorim, AM, Kutschenko, DC, Judice DM & Barros FSM (2013) Malpighiaceae. In: Martinelli G & Moraes MA (orgs.) Livro Vermelho da Flora do Brasil. Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, Rio de Janeiro. Pp. 635-639.). Nonetheless, as mentioned, the family includes a great diversity of lianas which have not been fully studied in extratropical regions, despite their great floristic and ecological importance (Durigon et al. 2019Durigon J, Sperotto PS, Ferreira PPA, Dettke GA, Záchia RA, Farinaccio MA, Seger GDS & Miotto STS (2019) Updates on extratropical region climbing plant flora: news regarding a still-neglected diversity. Acta Botanica Brasilica 33: 1-10.).

The south of Brazil (i.e., states of Paraná, Santa Catarina, and Rio Grande do Sul) has been recognized for its flora, which includes species of tropical and temperate origins (Durigon & Waechter 2011Durigon J & Waechter JL (2011) Floristic composition and biogeographic relations of a subtropical assemblage of climbing plants. Biodiversity and Conservation 20: 1027-1044.). It is a convergence area of different floristic elements and, notably, the southern distribution limit of several tropical species (Rambo 1961Rambo B (1961) Migration routes of the South Brazilian rainforest. Pesquisas, Botânica 12: 1-54.). Three biomes can be identified, with different extents over the area (IBGE 2004Instituto Brasileiro de Geografia e Estatística (IBGE) (2004) Mapa da vegetação do Brasil e Mapa de Biomas do Brasil. Available at <http://www.ibge.gov.br>. Accessed on 09 October 2018.
http://www.ibge.gov.br...
): the Atlantic Rainforest, distributed across the three states; the Cerrado (Brazilian savannah), represented by only a small area in northeastern Paraná; and the Pampa, which in Brazil is restricted to Rio Grande do Sul, covering the state’s southern half. The Atlantic Forest and Cerrado are widely recognized as biodiversity conservation hotspots (Mittermeier et al. 2005Mittermeier RA, Gil PR, Hoffman M, Pilgrim J, Brooks T, Mittermeier CG, Lamoreux J, Fonseca GAB (2005) Hotspots revisited: earth’s biologically richest and most endangered terrestrial ecoregions. Cemex-Conservation International and Agrupacion Sierra Madre, Mexico City. 392 p.), and several efforts have been employed to understand the diversity of Malpighiaceae in these biomes (Francener et al. 2015Francener A, Damasceno-Júnior GA & Gomes-Klein VL (2015) Tetrapterys Cav. (Malpighiaceae) from Brazilian Midwest. Acta Botanica Brasilica 29: 143-156., 2017Francener A, Almeida RF de & Mamede MCH (2017) Taxonomic novelties in Byrsonima (Malpighiaceae) from the state of Minas Gerais, Brazil. Phytotaxa 291: 133-140.; Almeida et al. 2016). On the other hand, little is known about the Malpighiaceae species richness in the Pampa, which is still considered a neglected biome (Overbeck et al. 2007Overbeck GE, Müller SC, Fidelis AT, Pfadenhauer J, Pillar VP, Blanco CC, Boldrini II, Both R & Forneck ED (2007) Brazil’s neglected biome: The South Brazilian Campos. Perspectives in Plant Ecology, Evolution and Systematics 9: 101-116.), despite having several endemic species and, in some places, a highly species-rich flora (Boldrini 2009Boldrini II (2009) A flora dos campos do Rio Grande do Sul. In: Pillar VP, Müller SC, Castilhos ZMS & Jacques AVA (eds.) Campos Sulinos, conservação e uso sustentável da biodiversidade. Ministério do Meio Ambiente, Brasília. Pp. 63-77.).

Malpighiaceae is one of the ten most threatened botanical families of the Brazilian flora (Martinelli et al. 2013Martinelli G, Valente ASM, Maurenza D, Kutschenko DC, Judice, DM, Silva DS, Fernandez EP, Martins EM, Barros FSM, Sfair JC, Santos Filho LFA, Abreu MB, Moraes MA, Monteiro NP, Pietro PV, Fernandes RA, Hering RLO, Messina T & Penedo TSA (2013) Avaliações de risco de extinção de espécies da flora brasileira. In: Martinelli G & Moraes MA (orgs.) Livro Vermelho da Flora do Brasil. Andrea Jakobsson, Instituto de Pesquisas Jardim Botânico do Rio de janeiro, Rio de Janeiro. Pp. 60-84.; Martins et al. 2018Martins E, Martinelli G & Loyola R (2018) Brazilian efforts towards achieving a comprehensive extinction risk assessment for its known flora. Rodriguésia 69: 1529-1537.) and, in the Rio Grande do Sul Endangered Plant Species List (Rio Grande do Sul 2014Rio Grande do Sul (2014) Lista de Espécies da Flora Ameaçada de Extinção no Rio Grande do Sul. Available at <https://secweb.procergs.com.br/livlof/?id_modulo=2&id_uf=23&ano=2013>. Accessed on 09 April 2018.
https://secweb.procergs.com.br/livlof/?i...
), eight species of the family are cited as threatened. The rapid advance of monocultures is accelerating the loss of habitats and species in southern Brazil (Overbeck et al. 2007Overbeck GE, Müller SC, Fidelis AT, Pfadenhauer J, Pillar VP, Blanco CC, Boldrini II, Both R & Forneck ED (2007) Brazil’s neglected biome: The South Brazilian Campos. Perspectives in Plant Ecology, Evolution and Systematics 9: 101-116.). Rio Grande do Sul has been considered a new frontier of mineral exploration, and most of the projects are focused on the most conserved areas of the Pampa Biome, with great potential impact on biodiversity (Durigon & Seifert Jr. 2018Durigon J & Seifert Jr. CA (2018) Em defesa das vocações territoriais: mineração versus arranjos produtivos locais e biodiversidade. In: Machado RCF & Silva A (orgs.) Mulheres, Organização e Produção Agroecológica: Floresta Nacional de Tefé. CRV, Curitiba. Pp. 171-182.).

Despite recent taxonomic and phylogenetic studies concerning the family Malpighiaceae (Almeida et al. 2016Almeida RF de, Francener A & Amorim AMA (2016) A generic synopsis of Malpighiaceae in the Atlantic Forest. Nordic Journal of Botany 34:285-301. , 2017Almeida RF de, Amorim AMA, Correa AMS & Van den Berg C (2017) A new infrageneric classification for Amorimia (Malpighiaceae) based on morphological, phytochemical and molecular evidence. Phytotaxa 313:231-248.; Almeida 2018Almeida RF de (2018) Taxonomic revision of Amorimia W.R. Anderson (Malpighiaceae). Hoehnea 45: 238-306. ), in the South Region of Brazil, floristic studies of the family are still lacking. Monographs of Malpighiaceae genera and extensive taxonomic revisions generally do not account for specimens that are in border areas between biogeographical regions and do not cover particular morphological data in addition to not providing taxonomic tools applicable at a regional scale. These gaps in the taxonomy of Malpighiaceae weaken the circumscription of taxa, making it difficult to recognize the group in southern Brazil and, ultimately, prevent the measurement of regional species diversity. The objective of the present study is to provide identification keys for Malpighiaceae genera in southern Brazil, along with notes on its current taxonomy, as well as updates on the species richness and their distribution in these three southern states.

Materials and Methods

The study area included the states of Paraná, Santa Catarina, and Rio Grande do Sul, encompassing the three different biomes, Atlantic Forest, Cerrado, and Pampa (Fig. 1).

Figure 1
Location of study area and the three biomes sampled in southern Brazil – a. map of South America, with emphasis on Brazil; b. map of the southern region of Brazil; c. geopolitical division of southern Brazil and the distribution of the three biomes. RS: Rio Grande do Sul; SC: Santa Catarina and PR: Paraná.

The list of genera of Malpighiaceae occurring in the study area and their morphological characteristics were obtained from the analysis of specimens collected from 13 field expeditions conducted between December 2011 and January 2014 as well as specimens found in the largest herbarium collections in southern Brazil, as well as, other important collections of Brazilian flora (CRI, FLOR, FUEL, HAS, HBR, HUCS, ICN, MBM, PACA, PEL, R, RB, SMDB, UPCB). The terminology used in the keys follow Anderson (1981)Anderson WR (1981) Malpighiaceae. In: The botany of Guayana Highland. Part XI. Memoirs of the New York Botanical Garden 32: 21-305., and the character states were determined only from the species found in southern Brazil. The notes about genera taxonomy are based on specialized literature on the family (see Supplementary Material), the distribution of confirmed species was determined based on the collected material, the occurrence data obtained from the databases (CRIA 2020CRIA (Centro de Referência em Informação Ambiental) SpeciesLink (2020) Available at <http://splink.cria.org.br/>. Accessed on 31 March 2020.
http://splink.cria.org.br/...
; REFLORA, 2020), and from the consulted herbarium collections.

Results and Discussion

In our extensive survey on southern Brazilian Malpighiaceae, 23 of the 45 genera recognized for Brazil were recorded (Flora do Brasil 2020Flora do Brasil 2020 under construction. Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br>. Accessed on 31 March 2020.
http://floradobrasil.jbrj.gov.br...
, under construction), which represents 52.3% of the family diversity at the generic level. The placement of the genera into clades according to Anderson et al. (2006)Anderson WR, Anderson C & Davis CC (2006) Malpighiaceae. Available at <http://herbarium.lsa.umich.edu/malpigh.>. Accessed on 14 May 2019.
http://herbarium.lsa.umich.edu/malpigh....
and the relevant references about the genera can be found in the Supplementary Material (Tab. S1, available on <https://doi.org/10.6084/m9.figshare.16876165.v1>).

The importance of the number of styles and the position and shape of the stigmas for the recognition of genera was confirmed, in addition to several characteristics of dried or fleshy fruits. Two identification keys for Malpighiaceae genera, based on floral and fruit morphology, respectively, are presented as follows. These also include some vegetative characters, especially related to the indument, leaves and stipules.

Key to the genera of Malpighiaceae from southern Brazil based on floral characters

  • 1. Styles with terminal stigmas (Figs. 2e, 2g-i, 2l, 3d, 3j, 4i, 5f, 5k) ................... 2

  • 1̓. Styles with stigmas on ventral surface (Figs. 3g, 4e-f, 5h) ................... 14

    • 2. Styles slender and subulate, tapering to minute stigmas (Figs. 2l, 3j); inflorescence terminal ................... 3

    • 2̓. Styles slender to stout, of uniform diameter along their length or enlarged at apex; stigmas capitate, truncate or subpeltate; inflorescence terminal or axillary ................... 4

      • 3. Leaves eglandular; pedicels sessile (Fig. 2k); petals posterior (Fig. 5a) with erect claw (Fig. 2k); calyx bearing a pair of oblong large glands on each sepal ................... Byrsonima

      • 3̓. Leaves bearing marginal glands on base of lamina; pedicels pedunculate; all petals with patent claw; calyx eglandular or with rounded glands on base of the sinus between adjacent sepals (Fig. 3i) ................... Galphimia

        • 4. Stigmas capitate with any sort of dorsal extension (Fig. 5k) or stigmas subpeltate (Fig. 2i) ................... 5

        • 4̓. Stigmas capitate or truncate without dorsal extension ................... 7

          • 5. Calyx with 4 pairs of stalked peltate glands (Fig. 3k) ................... Heladena

          • 5̓. Calyx eglandular or with 4 or 5 pairs of sessile glands ................... 6

            • 6. Plants with stellate hairs; calyx eglandular; inflorescence a thyrse; bracts and bracteoles eglandular; sepals completely concealing the petals at pre-anthesis stages (Fig. 5j); scandent shrubs to lianas ................... Thryallis

            • 6̓. Plants with T-shaped hairs; calyx bearing 4 or 5 pairs of glands; inflorescence an axillary pseudoraceme, unbranched or sometimes ternate at base; bracts and bracteoles bearing glands; sepals leaving petals exposed at pre-anthesis stages; shrubs to small tree ................... Bunchosia

              • 7. Gynoecium with 2 or 3 styles (Figs. 2g, 5f) ................... 8

              • 7̓. Gynoecium with only 1 style (Figs. 2e, 3d) ................... 12

                • 8. Pedicels pedunculate (peduncle ≥ 4 mm long) ................... Tetrapterys

                • 8̓. Pedicels sessile or rarely short pedunculate (peduncle 0.5–3.0 mm long) ................... 9

                  • 9. Androecium with 5 stamens and 5 staminodia (Fig. 5e); pair of stipules from opposite leaves connate to form a large cordate structure (Fig. 5d) ................... Peixotoa

                  • 9̓. Androecium with 9 or 10 stamens; stipules absent or if present, triangular ................... 10

                    • 10. Styles densely hairy for up to half their length; all stamens with glandular connective; calyx eglandular or calyx glands attached below the free part of the sepals on receptacle ................... Diplopterys

                    • 10̓. Styles glabrate or with strigose indument in the base; some stamens with glandular connective or all stamens with connectives not glandular; calyx eglandular or calyx glands attached on or below the free part of sepals on receptacle ................... 11

                      • 11. Calyx eglandular or bearing pairs of glands on the free part of the sepals; some stamens with glandular connective; solitary umbel or ultimate unit of inflorescence an umbel; petals white, yellow or pink ................... Banisteriopsis

                      • 11̓. Calyx glands attached below the free part of the sepals on receptacle; all stamens with connectives not glandular; panicles (ultimate unit of inflorescence a pseudoraceme); petals yellow ................... Bronwenia

                        • 12. Androecium with 5 or 6 stamens, without staminodia (Fig. 4h); climbers, rarely shrubs ................... Janusia

                        • 12̓. Androecium with 3 or 4 stamens, staminodia present; subshrubs or shrubs ................... 13

                          • 13. Three stamens and 2 staminodia (Fig. 2d); lamina with sericeous indument ................... Aspicarpa

                          • 13̓. Four stamens and 2 staminodia (Fig. 3c); lamina with hirsute indument.................. Camarea

                            • 14. Styles bearing lateral appendages in their apices (Fig. 5h); petiole biglandular at apex (Fig. 5g) or eglandular with glands on abaxial base of lamina; stamens strongly unequal; stipules epipetiolar, triangular ................... Stigmaphyllon

                            • 14̓. Styles without lateral appendages in their apices; petiole eglandular or bearing glands between base and middle of its length; stipules triangular, filiform or subulate or stipules absent ................... 15

                              • 15. Stigmatic surface transversely expanded (Fig. 3a); three posterior anthers smaller than the other seven ................... Callaeum

                              • 15̓. Stigmatic cavity; subequal anthers ................... 16

                                • 16. Styles 2 (Fig. 3g), rarely 3, the third (anterior position) style rudimentary or absent; styles with two ventral protuberances; bracts concave (Fig. 3f); petals abaxially densely sericeous ................... Dicella

                                • 16̓. Styles 3; styles without ventral protuberance; bracts flat; petals abaxially tomentose or sericeous ................... 17

                                  • 17. Stipules epipetiolar, subulate between middle and apex of petiole (Fig. 4d); epipetiolar stipules; flowers sessile; inflorescence a solitary umbel ................... Hiraea

                                  • 17̓. Stipules epipetiolar or not, triangular, very small, at base, middle or apex of petiole; stipules interpetiolar or absent; flowers sessile or pedunculated; pseudoraceme, corymb, panicle or another type of compound inflorescence, never solitary umbel ................... 18

                                    • 18. Sepals strongly revolute in anthesis (Fig. 5c) ................... ...................................... Niedenzuella

                                    • 18̓. Sepals appressed, recurved or slightly revolute at apex in anthesis ................... 19

                                      • 19. Lamina abaxially densely and persistently golden, silvery or metallic-sericeous ............................................................................ Heteropterys [in part]

                                      • 19̓. Lamina glabrate or abaxially glabrate, pubescent, tomentose or very thinly sericeous ................... 20

                                        • 20. Petals abaxially carinate (Fig. 4B); flowers pink or white, rarely yellow ......................................................... Heteropterys [in part]

                                        • 20̓. Petals abaxially not carinate; flowers yellow, rarely lilacs or pink ................... 21

                                          • 21. Bracteoles subopposite inserted just below the apex of peduncle ................... Heteropterys [in part]

                                          • 21̓. Bracteoles opposite or subopposite inserted at apex of peduncle ................... 22

                                            • 22. Petiole bearing a pair of glands at base; bracteoles eglandular; leafy-bracteous, paniculiform sinflorescence ................... Heteropterys [in part]

                                            • 22̓. Petiole eglandular or bearing glands slightly above of its base; bracteoles eglandular or bearing glands; ultimate units of inflorescence a pseudoraceme (Fig. 4l) or a corymb ................... ...................................... 23

                                              • 23. Leaves abaxially eglandular or bearing glands on margin; petals abaxially densely sericeous ................... Carolus

                                              • 23̓. Leaves abaxially bearing glands between midrib and margin; petals abaxially glabrous, tomentose or sericeous ................... 24

                                                • 24. Stipules epipetiolar or absent; bracteoles slightly concave ................... Alicia

                                                • 24̓. Stipules interpetiolar (Fig. 4k); bracteoles flat ................... 25

                                                  • 25. Bracts sessile, linear or triangular, eglandular; petals yellow, pink or lilac ................... Mascagnia

                                                  • 25̓. Bracts short-petiolate, lanceolate, bearing marginal glands on each side near base (Fig. 2c); petals yellow ................... Amorimia

Key to the genera of Malpighiaceae from southern Brazil based on fruit caracters

  • 1. Fruit fleshy at maturity (Figs. 2j, 2m) ................... 2

  • 1̓. Fruit dry at maturity (Figs. 2f, 3b, 3h, 4c, 4g, 4j, 5b, 5i, 5l) ................... 3

    • 2. Leaves and bracteoles eglandular; calyx bearing 5 pairs of glands; inflorescence a terminal pseudoraceme ................... Byrsonima

    • 2̓. Leaves bearing glands impressed in abaxial surface of blade; some bracteoles bearing abaxial glands; calyx bearing 4 pairs of glands; inflorescence an axillary pseudoraceme, unbranched or sometimes ternate at base ................... Bunchosia

      • 3. Fruit syncarpic, densely sericeous when immature (Fig. 3h) ................... Dicella

      • 3̓. Fruit schizocarpic, sericeous when immature ................... 4

        • 4. Mericarps with winged pericarp ................... 5

        • 4̓. Mericarps with unwinged pericarp ................... 20

          • 5. Mericarps with dorsal wing dominant (Figs. 2f, 3e, 4c, 4j); nut smooth or bearing winglets or crests on each side ................... 6

          • 5̓. Mericarps with lateral wings dominant (Figs. 3b, 4g, 5b, 5i) ................... 13

            • 6. Wing of mericarp with the abaxial edge thickened (Fig. 4c) ................... Heteropterys

            • 6̓. Wing of mericarp with the adaxial edge thickened (Figs. 2f, 3e, 4j) ................... 7

              • 7. Pedicels pedunculate (peduncle ≥ 4 mm long) ................... 8

              • 7̓. Pedicels sessile or rarely short pedunculate (peduncle 0.5–3.0 mm long) ...................................... 9

                • 8. Mericarp with dorsal wing well developed (Fig. 4j) ................... Janusia

                • 8̓. Mericarp with dorsal ridge or rudimentary dorsal wing (Fig. 5f) ......................................................... Aspicarpa

                  • 9. Stem with a pair of stipules from opposite leaves connate to form a large cordate structure (Fig. 5d) ................... Peixotoa

                  • 9̓. Stem with triangular and minute stipules or stipules absent ................... 10

                    • 10. Nut with winglets and with or without an appendage at base of adaxial edge of dorsal wing ................... 11

                    • 10̓. Nut smooth or with discontinuous winglets and without appendages at base of adaxial edge of dorsal wing .......................................................................................................................... 12

                      • 11. Nut bearing a winglet on each side; a pair of basal glands on each side of the midrib; inflorescence paniculate; dorsal wing without appendage ................... Bronwenia

                      • 11̓. Nut bearing 2–3 winglets on each side (Fig. 3e); lamina bearing minute and numerous glands on margin; petiole bearing glands in its apex; umbel or pseudoraceme; dorsal wing with appendage ...................................... Diplopterys

                        • 12. Petiole eglandular or rarely with a pair of glands near the apex, whose diameter is less than 0.5 mm ................... Banisteriopsis

                        • 12̓. Petiole with a pair of prominent glands at its apex (Fig. 5g), whose diameter is equal to or greater than 0.8 mm ................... Stigmaphyllon

                          • 13. Mericarps with 4 lateral wings or with 2 lateral wings and one dorsal wing, both well developed ................... 14

                          • 13̓. Mericarps with two lateral wings, free or continuous at base and dorsal wing much reduced (winglet or dorsal crest) or absent ................... 15

                            • 14. Sepals strongly revolute in anthesis (Fig. 5c); stigma ventral ................... Niedenzuella

                            • 14̓. Sepals erect or recurved in anthesis; stigma terminal ................... Tetrapterys

                              • 15. Pedicels sessile; stipules subulate, epipetiolar (Fig. 4d) ................... Hiraea

                              • 15̓. Pedicels pedunculate; stipules interpetiolar, triangular or filiform ................... 16

                                • 16. Mericarps bearing two lateral wings with free margins at the base and apex (Figs. 3b, 4g) ................... 17

                                • 16̓. Mericarps bearing two lateral wings with confluent margins at the base (Fig. 5b) ................... 19

                                  • 17. Bracts with 1 or 2 basal glands (Fig. 2c); bracteoles eglandular, rarely with 1 pair of glands ................... Amorimia

                                  • 17̓. Bracts and bracteoles eglandular ................... 18

                                    • 18. Bracteoles flat; glands on or embedded in margin; elongated pseudoracemes; stipules interpetiolar ................... Carolus

                                    • 18̓. Bracteoles concave; glands at or on margin of lamina; umbels, corymbs or short pseudoracemes; stipules epipetiolar ......................................................... Callaeum

                                      • 19. Stipules interpetiolar; membranous wings (Fig. 5b); bracts and bracteoles flat ................... Mascagnia

                                      • 19̓. Stipules epipetiolar; chartaceous or papyraceous wings; bracts flat, lanceolate and bracteoles concave (Fig. 2a) ...................................... Alicia

                                        • 20. Mericarps equinate; lamina bearing a pair of pedunculate glands at base, near the margin ...................................... Camarea

                                        • 20̓. Mericarps smooth or slightly rugose; lamina eglandular or with sessile glands ................... 21

                                          • 21. Fruit with enlarged winglike sepals (Fig. 5l); sepals eglandular; plants with stellate hairs ...................................... Thryallis

                                          • 21̓. Fruit without winglike sepals; sepals bearing pairs of glands; plants with a T-shaped hairs ...................................... 22

                                            • 22. Calyx with 4 pairs of stalked peltate glands (Fig. 3k) ................... Heladena

                                            • 22̓. Calyx eglandular or with rounded glands in base of each sinus between adjacent sepals (Fig. 3i) ................... Galphimia

Species richness and occurrence

The 23 genera of Malpighiaceae found in southern Brazil comprise 95 species, among them, trees, shrubs, subshrubs (upright, rhizomatous or xylopodiferous), and climbers. Almost all species (92) were recorded in the Atlantic Forest, in the state of Paraná. More than a third of the 95 species were found in the states of Santa Catarina (35) and Rio Grande do Sul (31). The complete list of species, occurrence data and voucher specimens can be found in the Table 1. Notes about genera taxonomy, species richness by genera, and distribution data are presented in the following.

Table 1
List of Malpighiaceae species in southern Brazil, respective states and biomes of occurrence and vouchers. States: PR: Paraná; SC: Santa Catarina; RS: Rio Grande do Sul. Biomes: ATL: Atlantic Forest; CER: Cerrado; PAM: Pampa.

1. Alicia

This is one of the generic segregates of Mascagnia s.l. (Anderson 2006aAnderson WR (2006a) Eight segregates from the neotropical genus Mascagnia (Malpighiaceae). Novon 16: 168-204.) and comprises two species, both South American lianas. Alicia anisopetala (A.Juss.) W.R.Anderson is the only species of the genus in southern Brazil, occurring in the Atlantic Forest, in the states of Santa Catarina and Paraná.

2. Amorimia

This is one of the generic segregates of Mascagnia s.l. (Anderson 2006aAnderson WR (2006a) Eight segregates from the neotropical genus Mascagnia (Malpighiaceae). Novon 16: 168-204.) and comprises fifteen species of South American lianas. Amorimia exotropica (Griseb.) W.R.Anderson is the only species of the genus in southern Brazil, occurring in the Atlantic Forest in Paraná, Santa Catarina and Rio Grande do Sul. Recently, Almeida et al. (2017) proposed two new subgenera for Amorimia; A. exotropica was placed in the subgenus Amorimia. As the epithet makes reference to, this species is distributed outside the tropics and, according to Almeida (2018)Almeida RF de (2018) Taxonomic revision of Amorimia W.R. Anderson (Malpighiaceae). Hoehnea 45: 238-306. , it should be regarded as Endangered in Brazil. It is easily recognized by the hairy petals on the abaxial surface, bracts with 1–2 basal glands (Fig. 2c) and fruits that split into three lateral-winged mericarps.

Figure 2
Morphological details of selected genera of Malpighiaceae found in southern Brazil – a–b. Alicia – a. bracts and bracteoles, b. gynoecium; c. Amorimia –bract; d-f. Aspicarpa – d. androecium, e. gynoecium, f. fruit with three mericarps; g-j. Banisteriopsis – g. gynoecium, g. detail of apex of the style; i-j. Bunchosia – i. gynoecium, j. fruit; k-m. Byrsonima – k. sessile flower (lateral view), showing posterior petal with erect claw (PC: pedicel; PP: posterior petal; AL: antero-lateral petal; bt: bract; bl: bracteole; cl: claw), l. gynoecium, m. fruit. (a-b. E.L.C. Soares 307; c. E.L.C. Soares 319; d-f E.L.C. Soares 380; g-h E.L.C. Soares 453; i-j E.L.C. Soares 355; k-m E.L.C. Soares 481).
Figure 3
Morphological details of selected genera of Malpighiaceae found in southern Brazil – a–b. Callaeum – a. gynoecium; b. mericarp (ventral view); c–d. Camarea – c. Androecium, d. gynoecium; e. Diplopterys – fruit with two mericarps; f–h. Dicella – f. bracts, g. gynoecium, h. fruit; i–j. Galphimia – i. calyx (two sepals were removed), j. fruit; k. Heladena – fruit with three mericarps. (a-b. E.L.C. Soares 336; c-d. E.L.C.Soares 448; e. E.L.C.Soares 505; f-h. E.L.C. Soares 491; i-j. E.L.C.Soares 502; k. E.L.C.Soares 504).
Figure 4
Morphological details of selected genera of Malpighiaceae found in southern Brazil – a–c. Heteropterys – a. flower (lateral view, petals were removed, PN: peduncle, PC: pedicel, bt: bract, bl: bracteole), b. carenate petals, c. fruit with three mericarps; d–g. Hiraea – d. epipetiolar stipule, e. gynoecium, f. detail of apex of the style, g. mericarp (ventral view); h–j. Janusia – h. gynoecium, i. gynoecium, j. mericarp (lateral view); k–l. Mascagnia – k. interpetiolar stipule, l. pseudoraceme. (a. E.L.C. Soares 500; b. E.L.C. Soares 316; c. E.L.C. Soares 321; d-g. E.L.C. Soares 390; h-j. E.L.C. Soares 395; k. E.L.C. Soares 423; l. E.L.C. Soares 406).
Figure 5
Morphological details of selected genera of Malpighiaceae found in southern Brazil – a–b. Mascagnia – a. flor (frontal view; PP: posterior petal, PL: postero-lateral petal, AL: antero-lateral petal), b. mericarp (ventral view); c. Niedenzuella – flower (lateral view, petals were removed); d–f. Peixotoa – d. stipule, e. androecium, f. gynoecium; g–h. Stigmaphyllon – g. petiole abaxially biglandular near apex, h. gynoecium; i. Tetrapterys – mericarp (ventral view); j–l. Thryallis – j. bud, k. gynoecium, l. expanded calyx. (a-b. E.L.C. Soares 406; c. E.L.C. Soares 476; d-f. E.L.C.Soares 451; g-h. E.L.C. Soares 479; i. E.L.C. Soares 490; j-l. E.L.C. Soares 506).

3. Aspicarpa

The genus currently comprises ten species but does not seem to be monophyletic (Cameron et al. 2001Cameron KM, Chase MW, Anderson WR & Hills HG (2001) Molecular systematics of Malpighiaceae: Evidence from plastid rbcL and matK sequences. American Journal of Botany 88: 1847-1862.; Davis & Anderson 2010Davis CC & Anderson WR (2010) A complete generic phylogeny of Malpighiaceae inferred from nucleotide sequence data and morphology. American Journal of Botany 97(12): 2031-2048.). Aspicarpa pulchella (Griseb. ex Mart.) O’Donell & Lourteig is the only species of the genus in southern Brazil, occurring in grasslands of three different biomes (Atlantic Forest, Cerrado and Pampa) and registered in Paraná, Santa Catarina and Rio Grande do Sul. A. pulchella is characterized by a herbaceous stem emerging from a perennial base, and the presence of a single style, three stamens, and two staminodia (Figs. 2d, 2e)

4. Banisteriopsis

As traditionally defined by Gates (1982)Gates B (1982) Banisteriopsis, Diplopterys (Malpighiaceae). Flora Neotropica Monograph 30: 1-238., this genus seems to be polyphyletic (Cameron et al. 2001Cameron KM, Chase MW, Anderson WR & Hills HG (2001) Molecular systematics of Malpighiaceae: Evidence from plastid rbcL and matK sequences. American Journal of Botany 88: 1847-1862.; Davis et al. 2001Davis CC, Anderson WR & Donoghue MJ (2001) Phylogeny of Malpighiaceae: evidence from chloroplast ndhF and trnl-F nucleotidesequences. American Journal of Botany 88(10): 1830-1846.). Banisteriopsis comprises 57 species, with 47 species recorded in Brazil. In southern Brazil, nine species were registered, including trees, climbers, and subshrubs, which are distributed almost exclusively in the Cerrado, in the state of Paraná. One exception is B. pseudojanusia (Nied.) B.Gates, which occurs in the Atlantic Forest in Paraná and Santa Catarina.

5. Bronwenia

This genus comprises ten species: eight of them were previously segregated from Banisteriopsis subgenus Banisteriopsis, and the other two species have been described as new species (Anderson & Davis 2007Anderson WR & Davis CC (2007) Generic adjustments in neotropical Malpighiaceae. Contributions from the University of Michigan Herbarium 25: 137-166.). Bronwenia ferruginea (Cav.) W.R.Anderson & C.Davis, a liana, is the only species of the genus in southern Brazil, occurring in the Atlantic Forest, in the state of Paraná. The species, and consequently the occurrence of this genus is not yet cited in the Flora do Brasil 2020Flora do Brasil 2020 under construction. Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br>. Accessed on 31 March 2020.
http://floradobrasil.jbrj.gov.br...
database (Flora do Brasil 2020Flora do Brasil 2020 under construction. Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br>. Accessed on 31 March 2020.
http://floradobrasil.jbrj.gov.br...
, under construction).

6. Bunchosia

This pan-American genus comprises about 75 species of trees or shrubs, with few species reaching the subtropics of Argentina and Brazil (Anderson et al. 2006Anderson WR, Anderson C & Davis CC (2006) Malpighiaceae. Available at <http://herbarium.lsa.umich.edu/malpigh.>. Accessed on 14 May 2019.
http://herbarium.lsa.umich.edu/malpigh....
). In southern Brazil, two species were registered from the Atlantic Forest in Paraná, Santa Catarina, and Rio Grande do Sul.

7. Byrsonima

This is one of the largest and most taxonomically complex genera of Malpighiaceae, including about 135 pan-American species of tree, shrubs or subshrubs (Anderson et al. 2006Anderson WR, Anderson C & Davis CC (2006) Malpighiaceae. Available at <http://herbarium.lsa.umich.edu/malpigh.>. Accessed on 14 May 2019.
http://herbarium.lsa.umich.edu/malpigh....
). In southern Brazil, sixteen species were found: thirteen in the Cerrado and the other three in the Atlantic Forest in Paraná, Santa Catarina, and Rio Grande do Sul.

8. Callaeum

A revision of this genus was published by Johnson (1986)Johnson DM (1986) Revision of the neotropical genus Callaeum (Malpighiaceae). Systematic Botany 11: 335-353.; Anderson (2006b)Anderson WR (2006b) A new species of Callaeum (Malpighiaceae) from Puebla, Mexico. Acta Botanica Mexicana 74: 179-183. proposed one new species, giving a total of 11 pan-American species. Callaeum psilophyllum (A.Juss.) D.M.Johnson, a liana, is the only species registered in southern Brazil, occurring in the Atlantic Forest in Paraná, Santa Catarina, and Rio Grande do Sul.

9. Camarea

This genus comprises eight species and has its center of diversity in the Central Brazilian Plateau (Mamede 1990Mamede MCH (1990) Revisão do gênero Camarea Saint-Hilaire (Malpighiaceae). Hoehnea 17: 1–34.).Camarea affinis A.St.-Hil, a xylopodiferous subshrub, is the only species registered in southern Brazil, occurring in the Cerrado, in the state of Paraná. The species, and consequently the occurrence of the genus in southern Brazil, are not included in the Flora do Brasil 2020Flora do Brasil 2020 under construction. Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br>. Accessed on 31 March 2020.
http://floradobrasil.jbrj.gov.br...
database (Flora do Brasil 2020Flora do Brasil 2020 under construction. Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br>. Accessed on 31 March 2020.
http://floradobrasil.jbrj.gov.br...
, under construction).

10. Carolus

This is one of the generic segregates of Mascagnia s.l. (Anderson 2006aAnderson WR (2006a) Eight segregates from the neotropical genus Mascagnia (Malpighiaceae). Novon 16: 168-204.) and includes six species. Carolus chlorocarpus (A.Juss.) W.R.Anderson, a liana, is the only species in southern Brazil, occurring in the Atlantic Forest in Paraná.

11. Dicella

This genus comprises seven South American species (Anderson et al. 2006Anderson WR, Anderson C & Davis CC (2006) Malpighiaceae. Available at <http://herbarium.lsa.umich.edu/malpigh.>. Accessed on 14 May 2019.
http://herbarium.lsa.umich.edu/malpigh....
). Two species of lianas of the section Dicella occur in southern Brazil, in the Atlantic Forest in Paraná, Santa Catarina and Rio Grande do Sul. Dicella nucifera Chodat has a subtropical distribution, with populations in the central-western portion of southern Brazil, as well as southern Paraguay and northeastern Argentina (Chase 1981Chase MW (1981) A revision of Dicella (Malpighiaceae). Systematic Botany 6: 159-171.). Most of the Brazilian collections are from the interior of the continent, and the few coastal populations occur in sympatry with Dicella bracteosa Griseb.

12. Diplopterys

This genus was redefined by Anderson & Davis (2006)Anderson WR & Davis CC (2006) Expansion of Diplopterys at the expense of Banisteriopsis (Malpighiaceae). Harvard Papers in Botany 11: 1-16. to include all species previously positioned in Banisteriopsis subgenus Pleiopterys (Gates 1982Gates B (1982) Banisteriopsis, Diplopterys (Malpighiaceae). Flora Neotropica Monograph 30: 1-238.), totaling 31 South American species. In southern Brazil, two species of lianas were found in the Atlantic Forest in Paraná, where the genus reaches the southern limit of its distribution.

13. Galphimia

This pan-American genus comprises 26 species. In southern Brazil, Galphimia australis Chodat, a shrub or upright subshrub, was the only species registered, occurring manly in more-open areas of the Atlantic Forest in Paraná, Santa Catarina and Rio Grande do Sul, and in the Pampa in Rio Grande do Sul.

14. Heladena

Heladena multiflora Nied., a liana or upright subshrub, is the single species described for this genus. In southern Brazil, the species was found in forested ecosystems in the Atlantic Forest, in Paraná and Rio Grande do Sul. Dettke et al. (2018)Dettke GA, Crespão LMP, Siquerolo LV, Siqueira EL & Caxambú MG (2018) Floristic composition of the Seasonal Semideciduous Forest in Southern Brazil: Reserva Biológica das Perobas, State of Paraná. Acta Scientiarum. Biological Sciences 40: 1-14. cited the species as a unique record and a new occurrence for Paraná, and Durigon et al. (2019)Durigon J, Sperotto PS, Ferreira PPA, Dettke GA, Záchia RA, Farinaccio MA, Seger GDS & Miotto STS (2019) Updates on extratropical region climbing plant flora: news regarding a still-neglected diversity. Acta Botanica Brasilica 33: 1-10. confirmed its occurrence in Rio Grande do Sul, based on material collected in the present study. These recent records increase the number of subpopulations known in Brazil, which were represented by only four locations (Amorim et al. 2013Amorim, AM, Kutschenko, DC, Judice DM & Barros FSM (2013) Malpighiaceae. In: Martinelli G & Moraes MA (orgs.) Livro Vermelho da Flora do Brasil. Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, Rio de Janeiro. Pp. 635-639.) and contribute to a better understanding of the distribution area of this species, classified as “Endangered” in the Brazilian Red List (Martinelli et al 2013). It is important to highlight that this species, and consequently the occurrence of the genus in southern Brazil, is not included in the Flora do Brasil 2020Flora do Brasil 2020 under construction. Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br>. Accessed on 31 March 2020.
http://floradobrasil.jbrj.gov.br...
database (Flora do Brasil 2020Flora do Brasil 2020 under construction. Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br>. Accessed on 31 March 2020.
http://floradobrasil.jbrj.gov.br...
, under construction).

15. Heteropterys

This is the largest and most taxonomically complex genus of Malpighiaceae. The last taxonomic revision was carried out by Niedenzu (1928)Niedenzu F (1928) Malpighiaceae. In: Engler A. (ed.) Das Pflanzenreich IV 141. Wilhelm Engelmann, Leipizig. Pp. 1-870.. In Brazil, about 97 species are recorded, at least half from the Atlantic Forest. In southern Brazil, 20 species were registered, including climbers, shrubs and subshrubs; it is the genus with the largest number of species in the region, most of them in the Atlantic Forest in Paraná. It should be noted that Heteropterys dusenii Nied. occurs exclusively in southern Brazil and is classified as “Vulnerable” in the Brazilian Red List (Martinelli et al. 2013Martinelli G, Valente ASM, Maurenza D, Kutschenko DC, Judice, DM, Silva DS, Fernandez EP, Martins EM, Barros FSM, Sfair JC, Santos Filho LFA, Abreu MB, Moraes MA, Monteiro NP, Pietro PV, Fernandes RA, Hering RLO, Messina T & Penedo TSA (2013) Avaliações de risco de extinção de espécies da flora brasileira. In: Martinelli G & Moraes MA (orgs.) Livro Vermelho da Flora do Brasil. Andrea Jakobsson, Instituto de Pesquisas Jardim Botânico do Rio de janeiro, Rio de Janeiro. Pp. 60-84.). In addition, H. hypericifolia A.Juss., which presents a disjunct distribution in the states of Mato Grosso do Sul and Rio Grande do Sul, is mentioned in the Rio Grande do Sul Endangered Plant Species List (Rio Grande do Sul 2014Rio Grande do Sul (2014) Lista de Espécies da Flora Ameaçada de Extinção no Rio Grande do Sul. Available at <https://secweb.procergs.com.br/livlof/?id_modulo=2&id_uf=23&ano=2013>. Accessed on 09 April 2018.
https://secweb.procergs.com.br/livlof/?i...
) as having insufficient data, and the few records found in the present study is a strong indication that the species may be under serious threat locally.

16. Hiraea

About 60 to more than 70 species are reported to this genus, of which about 20 are listed for Brazil (Anderson et al. 2006a, Almeida et al. 2016, Flora do Brasil 2020Flora do Brasil 2020 under construction. Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br>. Accessed on 31 March 2020.
http://floradobrasil.jbrj.gov.br...
, under construction). In southern Brazil, two species of lianas were recorded in the Atlantic Forest in all the states sampled, and one species (Hiraea hatschbachii C.E.Anderson) was only found in Paraná. This species was recognized and described by Anderson (2014)Anderson C (2014) Hiraea cuneata, H. macrophylla, and four new species confused with them: H. hatschbachii, H. occhionii, H. reitzii and H. restingae (Malpighiaceae). Edinburgh Journal of Botany 73: 361-378. when examining the collections of H. cuneata Griseb. H. hatschbachii differs from H. cuneata by showing adaxially glabrous sepals, eglandulous petioles, and solitary 4-flowering umbels. Although the Flora do Brasil database lists H. cuneata for the state of Paraná, this was not confirmed by the present study. Therefore, additional collection efforts are needed to better understand the distribution of this recently revised species. On the other hand, the same database does not cite H. fagifolia A.Juss. for the states of Santa Catarina and Rio Grande do Sul, in which the species was found frequently in the Atlantic Forest.

17. Janusia

Despite recent taxonomic advances, more studies are necessary to determine the natural delimitation of this genus, which does not seem monophyletic (Anderson & Davis 2007Anderson WR & Davis CC (2007) Generic adjustments in neotropical Malpighiaceae. Contributions from the University of Michigan Herbarium 25: 137-166.; Davis & Anderson 2010Davis CC & Anderson WR (2010) A complete generic phylogeny of Malpighiaceae inferred from nucleotide sequence data and morphology. American Journal of Botany 97(12): 2031-2048.). Janusia comprises about 15 South American species (Sebastiani & Mamede 2015). In southern Brazil, four species were registered, including climbers, shrubs and one xylopodiferous subshrub. Most of species occurs in the Cerrado and Atlantic Forest in Paraná. J. guaranitica A.Juss. is also found in the Atlantic Forest, in the state of Santa Catarina, and in the Pampa biome, in the Rio Grande do Sul state.

18. Mascagnia

This genus had its circumscription drastically reduced (Anderson 2006aAnderson WR (2006a) Eight segregates from the neotropical genus Mascagnia (Malpighiaceae). Novon 16: 168-204.; Anderson & Davis 2007Anderson WR & Davis CC (2007) Generic adjustments in neotropical Malpighiaceae. Contributions from the University of Michigan Herbarium 25: 137-166., 2013Anderson WR & Davis CC (2013) Combination of Mascagnia and Triopterys (Malpighiaceae). Memoirs of the New York Botanical Garden 108: 191-203.; Anderson & Corso 2007Anderson WR & Corso S (2007) Psychopterys, a new genus of Malpighiaceae from Mexico and Central America. Contributions from the University of Michigan Herbarium 25: 113-135.) and currently includes about 45 species (Anderson et al. 2006a). In southern Brazil, five species of lianas were registered in the Atlantic Forest, all of which occur in the state of Paraná, and two (Mascagnia australis C.E.Anderson and M. divaricata (Kunth) Nied.) also present in the states of Santa Catarina and Rio Grande do Sul.

19. Niedenzuella

This genus consists of 16 species previously placed in Mascagnia and Tetrapterys (Anderson 2006aAnderson WR (2006a) Eight segregates from the neotropical genus Mascagnia (Malpighiaceae). Novon 16: 168-204.). In southern Brazil, five species of lianas were registered in the Atlantic Forest, most of them in the Paraná state. The occurrence of Niedenzuella poeppigiana (A.Juss.) W.R.Anderson in the state of Paraná is not currently cited in the Flora do Brasil 2020Flora do Brasil 2020 under construction. Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br>. Accessed on 31 March 2020.
http://floradobrasil.jbrj.gov.br...
database (Flora do Brasil 2020Flora do Brasil 2020 under construction. Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br>. Accessed on 31 March 2020.
http://floradobrasil.jbrj.gov.br...
, under construction), despite having been cited in previous versions.

20. Peixotoa

This pan-American genus comprises 29 species, 28 of which were registered in Brazil (Anderson 1982Anderson C (1982) A monograph of the genus Peixotoa (Malpighiaceae). Contributions from the University of Michigan Herbarium 15: 1-92.). In southern Brazil, five species were found, including lianas and shrubs. They occur in the Atlantic Forest and Cerrado, especially in the Paraná state, with the exception of Peixotoa catarinensis C.E.Anderson, which is registered only in the coastal zone of Santa Catarina, and is classified as Endangered in the Brazilian Red List (Martinelli et al. 2013Martinelli G, Valente ASM, Maurenza D, Kutschenko DC, Judice, DM, Silva DS, Fernandez EP, Martins EM, Barros FSM, Sfair JC, Santos Filho LFA, Abreu MB, Moraes MA, Monteiro NP, Pietro PV, Fernandes RA, Hering RLO, Messina T & Penedo TSA (2013) Avaliações de risco de extinção de espécies da flora brasileira. In: Martinelli G & Moraes MA (orgs.) Livro Vermelho da Flora do Brasil. Andrea Jakobsson, Instituto de Pesquisas Jardim Botânico do Rio de janeiro, Rio de Janeiro. Pp. 60-84.).

21. Stigmaphyllon

The taxonomic revision of this genus was carried out by Anderson (1997)Anderson C (1997) Monograph of Stigmaphyllon (Malpighiaceae). Systematic Botany Monographs 51: 1-313. and was then revisited to include species previously placed in Ryssopterys (Anderson 2011Anderson C (2011) Revision of Ryssopterys and transfer to Stigmaphyllon (Malpighiaceae). Blumea 56: 73-104.). Stigmaphyllon comprises about 120 species distributed in the tropics worldwide (Anderson 2011Anderson C (2011) Revision of Ryssopterys and transfer to Stigmaphyllon (Malpighiaceae). Blumea 56: 73-104.); about 49 species are recorded in Brazil (Almeida et al. 2016). In southern Brazil, five species of climbers were found in the Atlantic Forest; S. bonariense (Hook. & Arn.) C.E.Anderson, S. ciliatum (Lam.) A.Juss. and S. jatrophifolium A.Juss. also occur in the Pampa biome. It is noteworthy that Pampa biome is not included among the “phytogeographic domains” in which the species occurs in the Flora do Brasil database (Flora do Brasil 2020Flora do Brasil 2020 under construction. Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br>. Accessed on 31 March 2020.
http://floradobrasil.jbrj.gov.br...
, under construction), and, consequently, no species of Stigmaphyllon are cited for the Pampa biome in this important database.

22. Tetrapterys

This genus is not monophyletic, and its delimitation is still unclear (Davis & Anderson 2010Davis CC & Anderson WR (2010) A complete generic phylogeny of Malpighiaceae inferred from nucleotide sequence data and morphology. American Journal of Botany 97(12): 2031-2048.). A possible segregation of Tetrapterys s.l. to a new genus was mentioned by Almeida et al. (2016), however, there is no recent taxonomic review of the genus. Tetrapterys comprises approximately 70 species (Anderson 2013Anderson WR (2013) Origins of Mexican Malpighiaceae. Acta Botanica Mexicana 104: 107-156.). In southern Brazil, seven species were registered, including lianas and upright subshrubs. Most of them occur in the Atlantic Forest, while T. xylosteifolia A.Juss. was also present in the Pampa biome, and T. jussieuana Nied. and T. salicifolia (A.Juss.) Nied. occur exclusively in the Cerrado, in the Paraná state. It should be noted that the occurrence of T. xylosteifolia in Paraná is not included in the Flora do Brasil 2020Flora do Brasil 2020 under construction. Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br>. Accessed on 31 March 2020.
http://floradobrasil.jbrj.gov.br...
database (Flora do Brasil 2020Flora do Brasil 2020 under construction. Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br>. Accessed on 31 March 2020.
http://floradobrasil.jbrj.gov.br...
, under construction). Nevertheless, the species is cited in the Flora Vascular do Paraná (Kaehler et al. 2014Kaehler M, Goldenberg R, Evangelista PHL, Ribas OS, Vieira AOS & Hatschbach GG (2014) Plantas vasculares do Paraná. UFPR, Curitiba. Pp. 131-132. ), and there are several records identified by specialists in the species link database (CRIA, 2020).

23. Thryallis

This pan-American genus comprises five species, all of them are found in Brazil (Anderson 1995Anderson C (1995) Revision of Thryallis (Malpighiaceae). Contributions from the University of Michigan Herbarium 20: 3-14.). Thryallis brachystachys Lindl., a liana or shrub, is the only species registered in southern Brazil, occurring in forest edges of the Atlantic Forest in Paraná and Santa Catarina.

Acknowledgements

We thank CNPq for the PROTAX (Programa de Capacitação em Taxonomia, Edital MCT/CNPq/MEC/CAPES Nº 52/2010) grant, as well as PhD fellowships of ELC Soares. We also thank Guilherme Peres Coelho for assistance with the figures.

References

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Edited by

Area Editor: Dra. Tatiana Carrijo

Publication Dates

  • Publication in this collection
    03 Dec 2021
  • Date of issue
    2021

History

  • Received
    04 May 2020
  • Accepted
    28 Dec 2020
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