Rodriguésia 63(1): 227-242. 2012 http://rodriguesia.jbrj.gov.br
An overview of recent taxonomic studies on Euphorbiaceae s.l. in Brazil Panorama dos recentes estudos taxonômicos em Euphorbiaceae s.l. no Brasil
Ricardo de S. Secco1, Inês Cordeiro2, Luci de Senna-Vale3, Margareth F. de Sales4, Letícia Ribes de Lima5, Débora Medeiros3, Bárbara de Sá Haiad3, Arline Souza de Oliveira3, Maria Beatriz Rossi Caruzo2, Daniela Carneiro-Torres6 & Narcisio C. Bigio7
Abstract The Euphorbiaceae sensu lato are distributed mainly in the tropics, in various types of vegetation and habitats, being one of the largest, most complex and diverse families of angiosperms. It has recently been divided into four families, according to classification systems based on molecular phylogeny: Euphorbiaceae sensu stricto, Phyllan- thaceae, Putranjivaceae, and Picrodendraceae. There is a proposition to establish Peraceae still under discussion. There were also changes in the taxonomic position of genera widely distributed in the Brazilian territory, such as Amanoa, Drypetes, Pera, Phyllanthus, Podocalyx, Pogonophora, and Richeria, among others. In addition, new species have been proposed and the limits of taxa distribution are expanding in Brazil. Thus, the authors provide an overview of recent studies and advances in the taxonomy of Euphorbiaceae s.l. in the Northern, Northeastern, Southeastern and Southern regions of Brazil, concentrating on review works and regional floras, as well as the changes that resulted in setting a new taxonomic family. Key words: Croton, Phyllanthus, taxonomy of Phanerogams, plant distribution. Resumo As Euphorbiaceae sensu lato distribuem-se especialmente nos trópicos, nos mais variados tipos de vegetação e habitats, sendo uma das maiores, mais complexas e diversificadas famílias das Angiospermas. Foi recentemente dividida em quatro famílias e segundo os atuais sistemas de classificação, baseados em estudos filogenéticos envolvendo dados moleculares, está assim classificada: Euphorbiaceae sensu stricto, Phyllanthaceae, Putranjivaceae e Picrodendraceae, havendo uma proposição de estabelecer Peraceae, táxon ainda em discussão. Houve também mudanças no posicionamento taxonômico de gêneros de ampla distribuição no território brasileiro, tais como Amanoa, Drypetes, Pera, Phyllanthus, Podocalyx, Pogonophora e Richeria, entre outros. Além disso, novas espécies vêm sendo propostas e a amplitude dos limites de distribuição de diversos táxons vem aumentando no Brasil. Diante do exposto, os autores fornecem um panorama geral dos estudos mais recentes e avanços na taxonomia de Euphorbiaceae s.l. nas regiões Norte, Nordeste, Sudeste e Sul do país, concentrando-se nos trabalhos de revisão e floras regionais, bem como nas mudanças que resultaram em nova configuração taxonômica da família. Palavras-chave: Croton, Phyllanthus, taxonomia de fanerógamos, distribuição de plantas.
Introduction This family is considered as one of the most The Euphorbiaceae is composed of 334 genera important Angiosperm groups, especially for (Webster 1994) and over 8,000 species (Radcliffe- comprising genera such as Hevea Aublet and Manihot Smith 2001), which are distributed mainly in the Miller. According to Schultes (1987), the rubber-tree tropics in the several types of vegetations and (Hevea brasiliensis (Willd. ex A. Juss.) Muell.Arg.) habitats. It is one of the most complex, large and has drastically changed human life within a single diverse families of Angiosperms. Wurdack et al. century, because it is a natural source of rubber, (2004) consider Euphorbiaceae sensu lato as a whereas the manioc, also known in Brazil as “aipim”, pantropical family, composed of 340 genera and “macaxeira” or “cassava”, (Manihot esculenta approximately 8,000‒9,000 species. Crantz) is among the 13 most important human foods.
1 Museu Paraense Emilio Goeldi, Av. Magalhães Barata 376, 66040-170, Belém, PA, Brazil. 2 Instituto de Botânica, C.P. 3005, São Paulo, SP, Brazil. 3 Museu Nacional, UFRJ, Quinta da Boa Vista, São Cristóvão, 20940-040, Rio de Janeiro, RJ, Brazil. 4 Universidade Federal Rural de Pernambuco, 52171-900, Recife, PE, Brazil. 5 Universidade Federal de São Carlos, Araras, SP, Brazil. 6 Universidade Estadual de Feira de Santana, 44031-460, Feira de Santana, BA, Brazil. 7 Universidade Federal de Rondônia, Depto. Biologia, Campus BR 364, km 9,5 Sentido Acre, 78900-000, Porto Velho, RO, Brazil. 228 Secco, R.S. et al.
Many authors studied the taxonomy of this Webster (1975) proposed an infrafamiliar family, standing out Jussieu (1824), Baillon (1858), classification for the Euphorbiaceae, subdividing it Bentham (1878, 1880) Mueller (1873), Hutchinson into five subfamilies, which were separated into two (1969), Jablonski (1967), and Pax & Hoffmann groups, according to the number of ovules per ovary (1914, 1931). Recently, a significant contribution locule: Phyllanthoideae and Oldfieldioideae, both was made by Webster (1975, 1987, 1994), which was with two ovules and Acalyphoideae, Crotonoideae mainly related to the family’s classification and to a and Euphorbioideae, with one ovule. revision of members of the Euphorbiaceae in the New In a revised and extended classification, World (Webster 1993; Webster & Armbruster 1991). Webster (1994) proposed that Euphorbiaceae s.l. Recently, the family has been through several encompasses five subfamilies: Phyllanthoideae, taxonomic changes, mainly based on phylogenetic Oldfieldioideae, Crotonoideae, Acalyphoideae, and studies with molecular data (APG II 2003; Wurdack Euphorbioideae. He also proposed new tribes and et al. 2004, 2005; APG III 2009), with marked subtribes in the same study. differences in relation to the traditional classification However, according to the classification revised and proposed by Webster (1994), in which system proposed by the Angiosperm Phylogeny Euphorbiaceae was divided into five subfamilies. Group (APG II 2003), there are no molecular However, although the Euphorbiaceae s.l. have evidences that allow maintaining Euphorbiaceae been studied by a group that comprises botanists s.l. as a single group. The family was then from several countries, including Brazil, and carries divided into four: Euphorbiaceae sensu stricto, out research in taxonomy, anatomy, phytochemistry, comprising the subfamilies with uniovulate economic botany, and molecular systematic, which ovary locules (Euphorbioideae, Crotonoideae, already resulted in the division of the family into four and Acalyphoideae); Phyllanthaceae, including (APG II 2003), the knowledge of this family still the Phyllanthoideae (biovulate ovary locules), has considerable gaps, even regarding morphology. Picrodendraceae, including the Oldfieldioideae Webster (1987) emphasizes that to propose a (biovulate ovary locules) and Putranjivaceae safer classification for this family, at least detailed (biovulate ovary locules, comprising Drypetes Vahl morphological and anatomical studies involving and Putranjiva Wall.). many of its genera are required. Wurdack et al. (2005), using molecular data of This situation is also observed in Brazil, where the plastidial regions rbc-L and trnL-F, assessed the many genera of Euphorbiaceae s.l. exhibit high circumscription of Euphorbiaceae s.s. (Acalyphoideae, number of species, such as Phyllanthus L., Mabea Crotonoideae and Euphorbioideae), as well as the Aublet, Dalechampia Plumier ex L., and Croton monophyletism of the suprageneric taxa recognized L. These genera still have confusing and outdated in recent classifications. Hence, they proposed two taxonomy in some regions of the country, though new subfamilies in Euphorbiaceae s.s.: Peroideae some advancements have been made in the past years, and Cheilosoideae. These studies corroborated the mainly with the publication of the Species List of the findings of the Angiosperm Phylogeny Group (APG Brazilian Flora (Cordeiro et al 2010; Secco et al 2010). II 2003), i.e., only the subfamilies with uniovulate Therefore, the present study aims at presenting an ovary locules (Euphorbioideae, Acalyphoideae and overview of the most recent studies on Euphorbiaceae Crotonoideae) are considered as Euphorbiaceae s.s.; s.l. in four Brazilian regions as well as changes in the they re-established Peroideae Baill. ex Hassk. as classification of the group, and at pointing out research a subfamily and admitted that the classification of opportunities that could lead to a more complete Euphorbiaceae s.s. needs revision. knowledge, in particular of the less studied genera. Euphorbiaceae s.s. and the other families segregated from Euphorbiaceae s.l. are included Euphorbiaceae Systematics in Malpighiales, according to the Angiosperm in the Light of Modern Phylogeny Group (APG II 2003; APG III 2009), Classification Systems Wurdack et al. (2005) and Wurdack & Davis (2009). In Engler’s system (1964) the Euphorbiaceae Hence, it was necessary to move some genera to were classified in the class Dicotyledoneae, independent families, as for example, Amanoa subclass Archychlamydeae, order Geraniales and Aublet (to Phyllanthaceae), Podocalyx Klotzsch (to suborder Euphorbiineae. Picrodendraceae), and Drypetes (to Putranjivaceae). Cronquist (1981) classified them in the class Those changes have been debated and have not Magnoliopsida, subclass Rosideae, order Euphorbiales. always received the approval of specialists in
Rodriguésia 63(1): 227-242. 2012 Euphorbiaceae studies in Brazil 229 Euphorbiaceae, as in the case of Pera Mutis, in Phyllanthus, Dalechampia, Manihot, and Mabea Peraceae, which still needs further studies according still need updated taxonomic revision, aiming at a to APG III (2009). better knowledge of their species boundaries. This division of Euphorbiaceae s.l. into Next, we provide a general overview of what four families was maintained by the Angiosperm has been studied on the matter by different research Phylogeny Group (APG III 2009), which stated groups in Euphorbiaceae s.l. in the northern, also that the recognition of Peraceae Klotzsch northeastern, southeastern, and southern regions would be necessary to maintain Euphorbiaceae as of Brazil. a monophyletic group, since Rafflesiaceae would be placed between the subfamilies Peroideae I. Euphorbiaceae in the Northern and Acalyphoideae. Yet it would depend on Region future studies, what would hinder the tentative In the Brazilian Amazon, pioneer studies establishment of the family (Peraceae). were led by Jacques Huber, Adolpho Ducke, and Judd et al. (2009) recognized four families: Walter Egler, with contributions by Murça Pires Euphorbiaceae s.s., Phyllanthaceae, Picrodendraceae, and William Rodrigues. From the 1980s on, there and Putranjivaceae, all in the clade Malpighiales; has been great increase in Euphorbiaceae research Phylanthaceae exhibits two ovules per locule and in the region, mainly involving the subfamily seeds without aril. Picrodendraceae also has two Crotonoideae (sensu Webster), starting with studies ovules per locule, but it does not have divided styles by Secco (1985), discussing varieties of Sagotia and nectariferous disk. racemosa Baill. Secco (1987) assessed systematic Wurdack & Davis (2009) proposed the and evolutionary aspects of Sandwithia Lanj., and elevation of Peroideae to the family status compared them with those of Sagotia Baill. and (Peraceae). Although approved by the APWeb, Anomalocalyx Ducke. Secco (1988) described the this proposal is still under debate (Stevens 2001). second species of Sandwithia, S. heterocalyx Secco, Reveal & Chase (2011) did not consider Peraceae an uncommon taxon that exhibits a pistillate flower as a family, considering it as a synonym of with free calyx, contrary to the genus’ pattern. Euphorbiaceae. In the 1990s stands out the study by Secco (1990a), who revised Anomalocalyx, Dodecastigma Geographic Distribution of Ducke, Pausandra Radlk., Pogonophora Miers Euphorbiaceae ex Bentham, and Sagotia, providing a consistent update of Pausandra, a genus with very confusing Euphorbiaceae s.s. has pantropical distribution, taxonomy until then, in particular because of its with higher occurrence in tropical regions. According sparse distribution in the Amazon and its little to Judd et al. (2009), this family is composed conspicuous, usually deciduous flowers. The of 222 genera and 6,100 species. These authors revision of Richeria Vahl (Secco & Webster 1990) state that Phyllanthaceae has predominantly invalidated the varieties proposed previously for tropical distribution, with 55 genera and 1,745 R. grandis Vahl, as well as updated the synonymy. species. According to Souza & Lorenzi (2008), Patterns of geographic distribution of some Picrodendraceae is a pantropical taxon and has Crotonoideae in the Amazon were presented approximately 30 genera and 100 species, and by Secco (1990b; 1993a), who assessed mainly Putranjivaceae is predominantly paleotropical, with Pausandra species and started a study on Croton only three genera and approximately 200 species. species that occur in the Brazilian Amazon. In his Wurdack & Davis (2009) stated that study, he described some liana species, comparing Euphorbiaceae s.s. is the largest Malpighiales family, them with C. pullei Lanj., whose palynological with more than 246 genera and approximately aspects continued to be discussed by Carreira et 6,300 species. al. (1996). A taxonomic treatment for Nealchornea Huber was made by Secco (1997). Euphorbiaceae Studies in Brazil Starting the studies on Alchorneae (Hurusawa) Croton L. is the most studied genus of Hutchinson, Secco (1993b) proposed the species Euphorbiaceae s.l. in Brazil and abroad, not only Alchornea fluviatilis Secco, collected in the Carajás from the perspective of classic taxonomy, but Mountain Range (state of Pará) and in the Pindaré also of phylogenetic systematics made mainly River (state of Maranhão); the species is uncommon with molecular data. Complex genera such as in the genus because it is monoecious. Later,
Rodriguésia 63(1): 227-242. 2012 230 Secco, R.S. et al. analyzing collections from Bolívia, he described that occur in the Brazilian Amazon by including Alchornea anamariae Secco and proposed a new Croton urucurana Baill., C. draconoides Muell.Arg., combination for Alchornea Swartz, based on C. trombetensis Secco; Berry & Rosa, C. sampatik Conceveiba pubescens Britton, later renamed as Muell.Arg., C. palanostigma Kl., C. pullei Lanj., and Alchornea brittonii Secco. This research resulted C. matourensis Aublet. in a revision of the Neotropical Alchorneae, Currently, Euphorbiaceae research in the published in Flora Neotropica (Secco 2004a), Amazon is focused on Pera (see Bigio & Secco 2012, updating Aparisthmium Endlicher, Alchornea, and in this volume), a genus which had never before been Conceveiba Aublet, resulting in a total of 37 species. subjected to revision, and which still has a confusing Next, Secco & Giulietti (2007) discussed aspects and outdated taxonomy, in addition to Micrandra of the biology of Conceveiba guianensis Aublet Bentham, Dalechampia and Amanoa. collected in the Mocambo Reserve (Belém, Pará), in Pera is a Neotropical taxon of the subfamily which only individuals with pistillate flower, fruits, Acalyphoideae, with ca. 30 species distributed from and seedlings were found. Cuba and Central America to southern Brazil, and An update of the taxonomy and highest diversity in the Amazon (Webster 1994). phytogeography of Hevea was proposed by Pires It is the only member of the tribe Pereae (Klotz. & et al. (2002). Taxonomic treatments were also made Gcke.) Pax & K. Hoffmann, a position supported for Alchorneopsis Muell. Arg., Cleidion Blume and unanimously by many who have made treatment Polyandra Leal (Secco 2001), Adenophaedra (Muell. to the genus (Baillon 1858; Pax & Hoffmann 1919; Arg.) Muell. Arg., and Tetrorchidium Poeppig (Secco Jablonski 1967; Webster 1994). It exhibits a unique 2003), as well as for 15 Euphorbiaceae genera of morphological character among the Euphorbiaceae: the Ducke Reserve (Secco 2005). In addition, other the unisexual pseudanthia (cymose inflorescence), studies on Euphorbiaceae present in Neotropical with reduced flowers involved by a showy involucral floras are Flora of Venezuelan Guiana (Secco in bract (yellow, cream, white or sometimes red), which Webster et al 1999), Flora of the Ducke Reserve is fragrant and more or less globose. The most recent (Secco & Cordeiro 1999), Checklist of Flora of studies were published by Gillespie & Ambruster Acre (Secco et al. 2008b), Checklist of the Plants (1997), comprising only 5 species from the Guianas, of the Guiana Shield, Checklist of Flora of Bolivia and by Webster et al. (1999), comprising 6 species (Secco et al. in press), Euphorbiaceae of the Flora included in the Flora of Venezuelan Guiana. In Brazil of Caxiuanã Reserve (Trindade & Secco 2009), stands out the study by Leal (1951), who proposed and Rare Plants of Brazil (Secco & Rosário some new species. Bigio (2009) made a taxonomic 2009). A taxonomic update of the rubber-tree treatment of 13 species from the Brazilian Amazon. Hevea brasiliensis was made by Secco (2008a), Recently, two new species were proposed: P. by requested of EPAMIG (Agricultural Research manausensis Bigio & Secco (Bigio & Secco 2010) and Company of Minas Gerais). P. eiteniorum Bigio & Secco (Bigio & Secco 2011). Secco (2008b) concluded the revision of Croton Molecular systematics led to great species that occur in the Brazilian Amazon, including advancement in studies on Euphorbiaceae. most of those addressed by Mueller (1873) and However, morphological characteristics hinder the bringing about taxonomic novelties such as Croton insertion of some genera in this family, as it is the faroensis R. Secco, C. dissectistipulatus R. Secco case of Pera, which exhibits inflorescences inside (Secco 2004b), an uncommon species because it an involucral bract, and its flowers and floral parts has only three stamens, C. subasperrimus R. Secco, in the pseudanthia. By having this inflorescence P. Berry & C. Rosário, C. diasii Pires ex Secco & type, Pera is similar to Dalechampia, though the P.E.Berry, and C. trombetensis Secco, P.E.Berry & latter belongs to Euphorbiaceae s.s. N. Rosa. He also recorded new occurrences in the Micrandra is an example of an Amazonian region, such as Croton gossipiifolius Vahl, Croton genus that needs revision. According to Webster nervosus Kl., Croton sampatik Muell.Arg., C. (1994), it is composed of seven species, and guianensis Aublet, C. spiraeifolius Jabl., C. urucurana belongs to Euphorbiaceae, subfamily Crotonoideae, Baill., C. draconoides Muell.Arg., C. mollis Benth., tribe Micrandreae, subtribe Micrandrinae, C. tessmannii Mansf., and C. schiedianus Schl. together with Cunuria Baillon and Micrandropsis Guimarães & Secco (2009) updated the species Rodrigues. According to APG system (APG of Croton section Cyclostigma and Croton section III 2009), Micrandra would be subordinated to Luntia subsection Matourense (Euphorbiaceae s.s.) Euphorbiaceae s.s. The most recent study on this
Rodriguésia 63(1): 227-242. 2012 Euphorbiaceae studies in Brazil 231 genus remains Schultes (1952). Therefore, the The taxonomic treatment of Dalechampia genus needs revision, though Berry in Webster et by Webster & Armbruster (1991) is still the most al. (1999) provided a synopsis of its species in the up-to-date. These authors provided a detailed study Flora of the Venezuelan Guiana. on the morphology, described briefly the genus, Schultes (1979) published additional notes sections and subsections, and two new species that on Micrandra, focusing the economic potential of they proposed. However they did not provide full the genus, especially as a source of latex for rubber descriptions and illustrations for other species of the production. It is important to revision this genus, genus and did not cite most collections examined, especially because of the close relationship that it providing only the citation of the types. Therefore, has with Cunuria, which, according to Baldwin Secco & Bigio (unpublished data) are revising and & Schultes (1947), has 4 species and a variety updating the species of Brazil, providing detailed distributed in the Amazon. However, Schultes keys, descriptions, and illustrations, since they were (1952) considered them as synonyms of Micrandra, not yet subjected to complete taxonomic treatment. proposing thus new combinations. Webster (1994) Webster (1994) states that Amanoa belongs drew attention to this fact, but kept Cunuria as a to Euphorbiaceae, subfamily Phyllanthoideae, tribe valid genus. The relationships between Micrandra, Amanoeae (Pax & Hoffamann) Webster, together Cunuria, and Micrandropsis, the latter recently with Pentabrachion Müll.Arg., a monotypic genus collected also in the state of Pará (it had been of western Africa. However, according to the APG previously found only in the state of Amazonas), system (APG III 2009), the taxon would be part of need to be better established. the family Phyllanthaceae. According to Webster (1994), Dalechampia Most of the 16 Amanoa species occur in the is a genus of Euphorbiaceae, subfamily Neotropics (13); only three are endemic to Africa. Acalyphoideae, tribe Plukenetieae (Bentham) The genus was never subjected to a complete Hutchinson, positioned isolatedly in the subtribe revision; the most recent study is still the one Dalechampineae, and it is composed of 100 species, carried out by Hayden (1990), which provided out of which 90 are distributed in tropical America. an identification key for the Neotropical species, In the APG system (APG II 2009), this genus is described four new species, proposed a new included in Euphorbiaceae s.s. It has an uncommon name (A. sinuosa Hayden), and commented on A. morphological characteristic to Euphorbiaceae, guianensis Aublet, a type species and its varieties. which is the specialized inflorescence of the type The taxonomic treatment by Hayden (1990) needs bilabiate, bisexual pseudanthia, with two involucral update, especially because many samples available bracts, usually colorful, with separate staminate in the herbaria of the IAN, INPA, and MG were not and pistillate flowers. This type of pseudanthia analyzed and new collections were accomplished is different from that found in Pera. Webster & lately, making it necessary to make a new approach Armbruster (1991) recognized six sections for the to the genus. Webster et al. (1999) updated only 90 Neotropical species of the genus: Dalechampia the species present in the Flora of the Venezuelan section Rhopalostylis Pax & Hoffmann, D. section Guiana. Secco (2005) has made a treatment for two Dioscoreifoliae Pax & Hoffmann, D. section species for the Flora of the Ducke Reserve, Manaus, Cremophyllum (Scheidweiler) Baillon, D. section state of Amazonas, and Secco et al. (2010) provided Coriaceae Pax & Hoffmann, D. section Tiliifolae an updated checklist of the species present in Brazil. Webster & Armbruster, and D. section Dalechampia. In the herbaria of MG and IAN, the most According to the survey carried out in the herbaria representative of eastern Amazon, there are many MG and IAN, and based on Webster & Armbruster collections lacking updated identification of (1991), 17 species occur in the Brazilian Amazon; Amanoa; some contain only fruits or other not seen only Dalechampia section Coriaceae has probably no with entire reproductive parts, so new collections representatives in the region. Gillespie & Armbruster are needed. Besides, the complete material (with (1997) revised 18 species of Dalechampia for the flowers and fruits) needs to be analyzed by a Flora of the Guianas, providing only 4 illustrations, specialist, since many samples were identified whereas Armbruster in Webster et al. (1999) revised by comparison, and not always by specialists. briefly the 12 species of the Flora of the Venezuelan This hinders the recognition of some species with Guiana. In Brazil, the study by Rodrigues (2007) is more restricted distribution in the Amazon, such the most up-to-date, and included four species of as Amanoa glaucophylla Muell. Arg., A. gracilima Distrito Federal. W.J. Hayden, and Amanoa neglecta W.J. Hayden.
Rodriguésia 63(1): 227-242. 2012 232 Secco, R.S. et al.
Gymnanthes Swartz does also need further II. Euphorbiaceae in the studies. This genus has approximately 40 species, Northeastern Region according to Webster (1994), and belongs to the Studies on Euphorbiaceae s.l. in northeastern subfamily Euphorbioideae, tribe Hippomaneae, Brazil focused mainly the floras of the states of subtribe Hippomaninae. According to the APG II Bahia and Pernambuco. The diversity of this family system, the genus belongs to Euphorbiaceae s.s. and is still poorly known, though some relevant studies includes also Actinostemon Mart. ex Kl. Although the have been carried out, aiming at solving mainly greatest specialists in Euphorbiaceae, such as Pax problems in the tribes Euphorbieae, Hippomaneae, & K. Hoffman (1912), Baillon (1858) and Mueller Hureae, and Crotoneae, particularly in the genus (1866), have kept Gymnanthes and Actinostemon Croton. In this aspect, a pioneer study by Bezerra as separate genera, Webster & Huft (1988) and & Fernandes (1982) stands out, which comprises Webster (1994) synonymized Actinostemon with six Croton species, locally known as ‘marmeleiros’, Gymnanthes, since the characters to separate it emphasizing the presence of Croton sonderianus from Gymnanthes are not convincing from the Muell.Arg. as the most abundant in the region. taxonomic perspective. Despite that, Webster et al. Studies comprising the entire family were (1999) considered Actinostemon as an independent carried out by Cordeiro (1995) for the Flora of Pico genus, separated from Gymnanthes. Esser (1999), in das Almas, Chapada Diamantina, and Carneiro et a partial taxonomic treatment of the Hippomaneae of al. (2002) for the Euphorbiaceae of “Inselbergs of Malaysia, mentioned the inclusion of Actinostemon Milagres” (state of Bahia). For the whole northeastern in Gymnanthes proposed by Webster (1994), but did region, Cordeiro & Carneiro-Torres (2006) listed not keep it as a synonym of Gymnanthes. Previously, 211 species and 45 genera in the plant checklist of authors such as Jablonski (1967, 1969) and Gillespie northeastern Brazil. A comprehensive study on the (1993) kept Actinostemon and Gymnanthes separated, family Euphorbiaceae for reserves in the Caatinga though Jablonski (1967) stated that the distinction of of Pernambuco was carried out by Lucena (2009), the two genera based on the number of the stamens is who studied 54 species. A synopsis of this family, very ambiguous. The taxonomic treatment proposed comprising 25 species and 11 genera of Serra de by Jablonski (1969) is still insufficient, as the author Itabaiana National Park (state of Sergipe), was did not describe or illustrate all species, detailing published by Lucena et al. (2009). Lucena & Alves more the two new species described (Actinostemon (2010) added 29 Euphorbiaceae taxa, with comments, unciformis Jabl., of Bahia, and A. appendiculatus to the list of this family in the northeastern region. Jabl., of northeastern, including Pernambuco, and Taxonomic treatments for genera and tribes southeastern Brazil, including Espírito Santo and were also carried out for Chapada Diamantina, probably Rio de Janeiro). Webster (pers. comm.) located within Bahia and Pernambuco. Carneiro- stated that he was inclined to reconsider Actinostemon Torres et al. (2003b) studied the genus Phyllanthus as an autonomous genus, though he had included it (currently Phyllanthaceae) reporting 11 species, in Gymnanthes (Webster 1994). He admits that and Carneiro-Torres et al. (2002) studied the tribe Actinostemon is very close to Gymnanthes, and most Euphorbieae and recognized 16 species. Still representatives of Hippomaneae (ex. Sebastiania regarding Phyllanthus, Silva & Sales (2007), based Sprengel, Actinostemon) have not been well on Silva (2004), found 17 species in the state of delimited morphologically. Pernambuco and 11 species in the Caatinga biome Studies by Melo (2006), revising Sebastiania, (Silva & Sales 2004). A synopsis of Phyllanthus included some species of this genus in Gymnanthes/ with 36 species for northeastern Brazil was also Actinostemon. According to this author, only the published by Silva & Sales (2008). Silva (2009) species of Sebastiania section Sebastiania should described two new species of Phyllanthus. The study be included in Sebastiania. of Cnidoscolus Pohl for the state of Pernambuco The problems exposed above led to a study was carried out by Melo & Sales (2008). Santos & on species of Gymnanthes (and Actinostemon), Sales (2009) studied also the tribe Hippomaneae in emphasizing those found in the Amazon, as a the state of Pernambuco. contribution to the taxonomic delimitation of the Revision and morphological phylogeny studies aforementioned taxa, since both Webster (1994) were carried out and other are in developmental phase and Esser (1999) admit that the generic delimitation focusing mainly the genera of the tribe Hipomaneae. of the Neotropical representatives of Hippomaneae The revision of Sebastiania, currently circumscribed is still under dispute. to 19 species, was developed by Melo (2006). Studies
Rodriguésia 63(1): 227-242. 2012 Euphorbiaceae studies in Brazil 233 on morphological phylogeny of Sebastiania, including III. Euphorbiaceae in the 17 species and 13 species of other Hippomaneae Southeastern Region genera and the use of carpological characters in the IIIa. Research in São Paulo delimitation of genera of the tribe Hippomaneae In the case of the research group of the state of were developed by Souza (2011) and submitted São Paulo, the most recent studies focused mainly for publication. Revisions and morphological and Euphorbiaceae s.s. and Phyllanthaceae, comprising molecular phylogeny studies of Gymnanthes and revisions, floras, and descriptions of new species. Stillingia Garden ex L. are being carried out. For the Floras: The survey carried out in Cadeia tribe Hureae studies on morphological phylogeny and do Espinhaço by Lima (2001) and Lima & Pirani a synopsis of the species of this tribe have been carried (2003) reported 22 species: Croton antisyphiliticus out by Oliveira (2010), who analyzed 17 species Mart., C. argyroglossum Baill., C. atrorufus Muell. of the genera Algernonia Baill. (11 species), Hura Arg., C. buxifolius Muell.Arg., C. campestris A. St.- L. (1 species), and Ophthalmoblapton Allemão (4 Hil., C. celtidifolius Baill., C. floribundus Spreng., species); Tetraplandra was considered as a subgenus C. glutinosus Muell.Arg., C. grewioides Baill., of Algernonia. C. julopsidium Baill., C. lobatus L., C. lundianus The best studied Euphorbiaceae genus in Muell.Arg., C. migrans Casar., C. muscicarpa Muell. northeastern Brazil is doubtless Croton, in particular Arg., C. mysinites Baill., C. nummularius Baill., C. in the semi-arid region. Local and regional floras siderophyllus Baill., C. subferrugineus Muell.Arg., and revisions of sections have been published. The C. timandroides Muell.Arg., C. urucurana Baill., revision of Croton section Argyroglossum in Brazil C. aff. desertorum Muell.Arg., and C. aff. petraeus was carried out by Gomes (2006) and Gomes et Muell.Arg. al. (2010). A morphological phylogeny study on The study on the tribe Crotoneae in the state of Croton section Ocalia is being concluded. Silva et São Paulo (Caruzo 2005; Caruzo & Cordeiro 2007) al. (2009) studied 15 species of Croton of the micro- reported 37 Croton species and two Astraea Klotzsch region of Vale do Ipanema, in the Caatinga biome, species. Among the Croton species, several occur also state of Pernambuco. In this state, Silva et al. (2010) in other countries, and some are broadly distributed published a synopsis of Croton, reporting 35 species all over the Neotropics, such as C. glandulosus L, C. of the genus. Attention should be given to Croton in hirtus L’Herit, C. pedicellatus Kunth, and C. triqueter the Caatinga biome, including the phytogeography Lam.; others are moderately distributed in South of this genus, of which 68 species were recognized America: they occur in Bolivia, Paraguay, Argentina, by Carneiro-Torres (2009). Lucena (2000) analyzed Uruguay, and center-western, southeastern and 14 species that occur in Zona do Litoral and Zona da southern Brazil, such as C. urucurana Baill., C. Mata of Pernambuco. Still regarding Croton, a study didrichsenii Webster, and C. rottlerifolius Baill.; on trichomes of species that occur in Pernambuco in Argentina, Paraguay, and Brazil, such as C. was carried out by Lucena & Sales (2006). Other antisyphiliticus Mart.; in Bolivia, Paraguay, center- studies are in the phase of data collection, standing western, southeastern and southern Brazil, such as out the studies of Croton in Chapada do Araripe, C. gracilipes Baill.; in Paraguay and most part of state of Ceará, and Vale do Catimbau National Park, Brazil, such as C. floribundus Spreng.; in Argentina, Buique, state of Pernambuco, and the flora of the Uruguay, southeastern and southern Brazil, such as C. state of Rio Grande do Norte. lanatus Lam.; in Bolivia, center-western, southeastern Recently, some new species have been described and southern Brazil, such as C. grandivelus Baill.; for northeastern Brazil. A new species of Gymnanthes in Paraguay, center-western, southeastern and was proposed by Esser et al. (2010). Gomes et al. southern Brazil, such as C. serratifolius Baill.; in (2010) described a new species of the genus Croton, Argentina, southeastern and southern Brazil, such and Carneiro-Torres et al. (2011) described three new as C. glechomifolius Muell. Arg.; and in Bolivia, species for this genus in the Caatinga. northeastern and southeastern Brazil, such as C. Although Euphorbiaceae studies have advanced tricolor Klotzsch ex Baill. Among the species recently, there are still many groups that need to be exclusively Brazilian are: C. lundianus (Didr.) Muell. studied. Among Hippomaneae, the genus Sapium Arg, which occurs all over Brazil; C. piptocalyx P. Browne deserves attention for the difficulty of Muell. Arg., found in northeastern, center-western delimiting its species. The species of Croton that and southeastern Brazil; C. macrobothrys Baill., occur in forests and restingas still need to be further found in northeastern, southeastern and southern investigated. Brazil; C. fuscescens Spreng., C. hemiargyreus
Rodriguésia 63(1): 227-242. 2012 234 Secco, R.S. et al.
Muell.Arg. and C. sphaerogynus Baill., present in Reviews: Among revisions of the genus is the northeastern and southeastern regions; C. fuscus study by Lima (2006) on Croton section Lamprocroton (Didr.) Muell.Arg., C. leptobotryus Müll.Arg., C. (Muell. Arg.) Pax. This section is entirely Neotropical, sancti-crucis S. Moore and C. sclerocalyx (Didr.) with an interesting disjoint distribution between South Muell.Arg., in center-western and southeastern; C. America and Mexico, and is characterized by the alchorneicarpus Croizat, C. ceanothifolius Baill., shrubby habit of its species, indumentum of lepidote C. celtidifolius Baill., C. dichrous Muell.Arg., C. trichomes, absence of cymules on the inflorescence, heterodoxus Baill, C. organensis Baill., C. salutaris and bifid style branches (Lima & Pirani 2008). Baill., C. serpyllifoius Baill. and C. vulnerarius Twenty-six species were recognized in the section, Baill., in southeastern and southern; C. compressus among which three were new: C. imbricatus L.R. Lam. and C. erythroxyloides Baill., with distribution Lima & Pirani, C. muellerianus L.R. Lima & Pirani, restricted to southeastern Brazil; and C. priscus and C. pygmaeus L.R. Lima & Pirani; the first Croizat endemic to the state of São Paulo. endemic to the state of Bahia, the second to Paraná and Besides, a Rapid Color Guide, containing the third endemic to Rio Grande do Sul. In addition to photos of the tree Croton species of the Atlantic the taxonomic treatment and phylogenetic analysis of Forest, was produced and distributed by the Field the section, morphological studies of leaf trichomes Museum of Natural History (Caruzo 2005). and pollen grains of some of its species have been The Flora of Grão Mogol, state of Minas Gerais also carried out. (Cordeiro 2004), reported 15 genera and 38 species of Caruzo (2010) made another revision on Euphorbiaceae s.l.: Acalypha L. (1), Alchornea (1), Croton section Cleodora (Klotzsch) Baill. Caruzo Bernardia Houstoun ex Miller (1), Chamaesyce S.F. et al. (2011) assessed the phylogenetic relationships Gray (2), Croton (11), Dalechampia (1), Euphorbia of this section, using the nuclear region ITS and the L. (2), Mabea (1), Manihot (1), Maprounea plastidial regions trnL-F and trnH-psbA, separately Aublet (1), Pera (1), Phyllanthus (5), Richeria (1), and combined. In the individual phylogenies and Sebastiania (7), and Stillingia (1). in that with combined data, species of the section Among ongoing flora studies are the one of the Cleodora sensu Webster (1993) emerged, almost state of São Paulo, in which the family Euphorbiaceae in their totality, together with other species that had s.s. is represented by 36 genera and ca. 139 species, as been originally not included in the section, forming follows: Acalypha (14), Actinostemon (3), Adelia L. a strongly supported clade characterized by two (1), Alchornea (3), Algernonia (1), Aparisthmium (1), morphological synapomorphies: female flowers with Astraea (2), Bernardia (3), Caperonia A.St.-Hil. (3), imbricate sepals and style united at the base or above. Caryodendron Karsten (1), Chaetocarpus Thwaites Croton section Cleodora is an exclusively Neotropical (1), Chiropetalum A. Jussieu (2), Cnisdoscolus Pohl section that has an interesting disjoint distribution (2), Croton (37), Dalechampia (12), Euphorbia (17), among Central America, Amazon, and Atlantic Forest. Jatropha L. (2), Joannesia Vellozo (1), Mabea (2), Eighteen species were recognized for this section; Manihot (9), Microstachys A. Juss. (5), Maprounea three of which were new (Caruzo et al. 2008; Caruzo (1), Margaritaria (1), Micrandra (1), Pachystroma et al. 2010a,b). The section is composed of arboreous Muell. Arg. (1), Pausandra Radlk. (1), Pera Mutis or shrubby species, usually latescent, covered by (2), Philyra Klotzsch (1), Plukenetia L. (1), Ricinus L. trichomes that are adpressed/star-shaped, star-shaped/ (1), Romanoa Trevisan (1), Sapium (2), Sebastiania lepidote, lepidote or more rarely star-shaped and (3), Tetraplandra Baillon (1), Tetrorchidium (1), and multiradiate; the leaves exhibit a pair of acropetiolar Tragia Plumier ex L. (6); and the Phyllanthaceae by or basilaminar glands; inflorescences are terminal, 5 genera and 5 species: Gonatogyne Muell. Arg. (1), rarely axillary, basal cymules, usually bisexual; male Hieronyma Fr. All. (1), Margaritaria (1), Richeria flowers are campanulate, rarely subcampanulate, (1) and Savia McPherson (1). Among these genera, valvate or slightly imbricate, with stamens between monographs on Croton and Astraea have already 15‒25; female flowers are campanulate to urceolate, been prepared by Caruzo & Cordeiro (2007), whereas in general conspicuously imbricate, styles 4-fidus those on Dalechampia, Plukenetia, Romanoa, and or multifidus, united at the base or above, usually Tragia are still in preparation (R. Secco, personal forming a crown. In this new circumscription of communication). the section Cleodora, two new subsections were The monograph on the genus Phyllanthus recognized: Sphaerogyni, characterized by sepals (Phyllanthaceae), represented by 16 species in the with quincuncial estivation in its pistillate flowers, state of São Paulo, has already been concluded and Spruceani, characterized by sepals with imbricate (Martins & Lima 2011, in press). or reduplicate-valvate estivation.
Rodriguésia 63(1): 227-242. 2012 Euphorbiaceae studies in Brazil 235 The revison of Microstachys, associated with de Janeiro. Oliveira & Senna (1983) also studied a phylogeny of Hippomaneae, is in progress (I. leaf anatomy for the characterization of Anabaenella Cordeiro, personal communication). tamnoides (Juss.) Pax & Hoffm. In Sebastiania Brasiliocroton P.E. Berry & Cordeiro, a Spreng., Oliveira & Penna (1985) studied the taxa S. new genus, and the reestablishment of Astraea: brasiliensis var. erythroxyloides (Muell.Arg.) Muell. according to Berry et al. (2005a), the erect stamens Arg. (currently Sebastiania brasiliensis Spreng.), on the flower bud of Brasiliocroton mamoninha and S. glandulosa var. fallax (Müll.Arg.) Pax P.E. Berry & Cordeiro are the main morphological (currently Microstachys corniculata (Vahl) Griseb.), characteristic that differentiates it from Croton. approaching their taxonomy and leaf anatomy. However, the best justification for the establishment Palynological studies were associated with taxonomic of this new genus is its position as sister-group of studies on Sebatiania Spreng. (Gonçalves-Esteves most Croton species, with the exception of Croton et al. 1986). Oliveira & Sá-Haiad (1988) pointed lobatus L. and C. praetervisus Muell.Arg., which out structural similarities and differences between belong to Croton section Astraea (Klotzsch) Baill. Euphorbia heterophylla L. and E. cyathophora In order to make Croton a monophyletic group, Murray. Emmerich (1987) devoted herself to the Berry et al. (2005b) incorporated into the taxon taxonomic study of Mabea fistulifera Mart. Oliveira other small genera formally placed in the tribe et al. (1989) studied the Euphorbiaceae that occur Crotoneae and excluded from it species of the in the restingas of the state of Rio de Janeiro from a section Astraea, which was reestablished by the floristic and taxonomic perspective. authors as the genus Astraea Klotzsch. In addition to those, stand out revisions of Chaetocarpus Sw. (Alves 1982) and taxonomic/ IIIb. Research in Rio de Janeiro floristic studies, such as those on Alchornea Sw. that In the state of Rio de Janeiro, research has occur in Rio de Janeiro (Silva 1993), Dalechampia been focused on Euphorbiaceae s.l. Studies started Plum. (Maia et al. 2002), Pera glabrata (Penna in the 1970s with the taxonomic revision of the 1981), and Croton L. (Sá-Haiad 1987). The latter was genera Algernonia Baill. and Tetraplandra Baill. pioneering in Brazil regarding the use of scanning Following these studies, Emmerich (1981) proposed microscopy for studying Euphorbiaceae species. new species (A. glaziouii and A. paulae) and a It is important to emphasize the multidisciplinary new combination: A. gibbosa (Pax & K. Hoffm.) approach of these studies, in which leaf anatomy Emmerich. Oliveira (1985) studied Sebastiania and pollen grains were used as subsidies for a better corniculata (Vahl) Müll.Arg. (currently a synonym taxonomic revision of the species. of Microstachys corniculata (Vahl) Griseb.). In the 1990s, studies focused the taxonomy In the 1980s, studies focused taxonomic of genera and species found in different Brazilian problems at different infrageneric levels. The validity ecosystems, studying the morphology of taxa such of infraspecific taxa, accepted at that time, was tested as Croton urticifolius Lam., Sebastiania bidentata by several researchers. Senna-Valle (1983) showed (Mart. & Zucc.) Pax (currently Microstachys that the infraspecific taxa of Pachystroma ilicifolium bidentata (Mart. & Zucc.) Esser, following Oliveira Müll. Arg. (currently Pachystroma longifolium & Ferraz (1991)), Caryodendron grandifolium (Nees) I.M.Johnst.) were no more than phenotypic (Muell.Arg.) Pax (currently Caryodendron expressions of characters, such as shape and leaf janeirense Muell. Arg., following Senna-Valle dimensions; these variations occur frequently in a & Campos (1992)), and nomenclatural types in single individual. These taxonomic studies were the family (Oliveira & Senna 1991). Silva (1993) almost invariantly associated with anatomical and studied varieties of Alchornea triplinervia (Spreng.) palynological studies, which observed the presence Muell.Arg., and only two out of the eight previously of structures that could be used in the taxonomic established varieties were accepted. In regional characterization of the Euphorbiaceae, such as the studies, Euphorbiaceae was recorded in florulas, such presence of Rosanoff’s crystals in Actinostemon as the Flora of Mato Grosso, mainly through studies concolor (Spreng.) Muell. Arg. (Emmerich & Senna- on species of Sebastiania Spreng. and Sapium P. Br Valle 1983). In the same genus, Senna-Valle (1983, (Oliveira 1991); the Flora of the Cairuçu Reserve - 1984, 1985) studied A. communis var. spathulatus Parati, Rio de Janeiro; the Flora of the Restinga de (Muell. Arg.) Pax (currently A. klotzschii (Didr.) Pax), Jurubatiba National Park – Rio de Janeiro (Oliveira A. verticillatus (Klotzsch) Baill., and A. concolor et al. 1997); and the Flora of the Atlantic Forest (Spreng) Muell. Arg. – all present in the state of Rio of southeastern Brazil. With a multidisciplinary
Rodriguésia 63(1): 227-242. 2012 236 Secco, R.S. et al. approach, stand out the studies on the sexual system and Brasiliocroton. The type of inflorescence, and pollination types of some Euphorbiaceae of as well as some morphological characters of its sandy coastal plains (restinga) (Oliveira et al. reproductive structures, distinguishes B. mamoninha 1989) and studies on Caperonia, Chamaesyce from the other two genera. The average size of the Gray, Croton L., Euphorbia, Sebastiania Spreng. pollen grains of the studied species of Astraea and and Stillingia Gard. Senna-Valle (1999) studied B. mamoninha differs from that of representatives of the complex Sapium glandulatum (Vell.) Pax Croton section Cyclostigma, which have large pollen (currently Sapium glandulosum (L.) Morong), grains. The presence of dorsiventrality, paracytic assessing phytochemical, anatomic, palynological, stomata, collateral vascular bundles, branched non- pharmacological, and taxonomic aspects. articulated laticifers and crystal idioblasts as well From the 2000s on, the number of monographs, as spheroidal, unaperturate and with a croton-type dissertations, and theses that focused the family ornamentation pattern pollen grains in the species increased. Medeiros et al. (2002, 2007, 2009, studied indicates morphological similarities among 2011) devoted themselves to the taxonomic and the three genera. The interpretation of the stipite of morphological study of Croton species present in the complex trichomes as emergences is a structural rocky fields of Minas Gerais. Initially, the study novelty for the group. The presence of innumerable was restricted to the São José Mountain Range, characteristics associated with defense mechanisms, Tiradentes, as part of the Flora of São José Mountain such as trichomes, idioblasts with crystal and/or with Range (Alves & Kolbek 2009). The species richness lipophilic content, laticifers and extrafloral nectaries, of Croton section Medea identified in this study especially in the genus Croton, together with the led to the morphological and taxonomical study possibility of both anemophilous and entomophilous of the section, using specimens from southeastern pollination, may represent one of the explanations for Brazil. In these studies, the focus was the taxonomic its great success, which is manifested by a pantropical validation of rare species, which were lacking in distribution and by the large number of species herborized collections. Some of these species were (Sá-Haiad et al. 2009). This group of researchers rediscovered (such as Croton gnidiaceus Baill., C. contributed with the study on Peraceae for the list josephinus Muell.Arg. and C. staechadis Baill.), of Brazil (Bigio et al. 2010). Currently, the group is as they have been no longer collected since the devoted to the study of species of Euphorbiaceae s.l. type material was described. As a result from this present in the Flora of the State of Rio de Janeiro. approach and due to the high endemism in rocky fields, three new species were described for the IV. Euphorbiaceae in the Southern section. The section could be recognized through a Region matrix of relevant morphological characters for its In the southern region, studies with a more circumscription. Pinto (2004) studied the biological general approach stand out: Smith & Downs (1959) diversity of Euphorbiaceae in Serra da Tiririca State made a preliminary assessment of the Euphorbiaceae Park, Niterói, state of Rio de Janeiro, and recorded 21 of Santa Catarina; Rambo (1960) presented a list genera and 41 species. Sá-Haiad et al. (2009) carried of 15 genera and 64 species of Euphorbiaceae in out studies aiming at contributing to the knowledge the state of Rio Grande do Sul, based on data of a of relationships among Croton, Brasiliocroton, private herbarium owned by the author; Klein (1977) and Astraea. Morphological, anatomical, and provided morphological and ecological data of 27 palynological analyses were used to describe and Croton species in the state of Santa Catarina; Allem compare species. The main objective of this study (1978) made a preliminary taxonomic assessment of was to search for structural elements to be used in ca. 40 Croton species in Rio Grande do Sul. analyses of the lineage of crotonoids. Vegetative Allem (1979) proposed four new species and a characteristics – both morphological and anatomical section (Calycireduplicatae Allem) of Croton for Rio – as well as the presence of reduced petals on pistillate Grande do Sul. According to this author, these species flowers – are similar among the studied specimens would be represented by dioecious plants, what of Croton section Cyclostigma and Brasiliocroton would be something extraordinary, since the Croton mamoninha, differing from Astraea. The presence species known so far are all monoecious, usually with and protodermal origin of the idioblasts with pistillate flowers on the base and staminate flowers lipophilic content and the interruption of the palisade on the rest of the rachis. However, in his descriptions, by collenchyma layers on the adaxial side of the mid Allem (1979) states that the staminate and pistillate vein are common to the Croton L. species analysed flowers are in separate inflorescences, but on the
Rodriguésia 63(1): 227-242. 2012 Euphorbiaceae studies in Brazil 237 same individuals, hence, those species would be, in References fact, monoecious. Allem, A.C. 1978. Preliminares para uma abordagem Probably, the greatest contribution to the taxonômica do gênero Croton L. (Euphorbiaceae) taxonomic study of the Euphorbiaceae of southern no Rio Grande do Sul. Boletim do Museu Botânico Brazil was made by Smith et al. (1988), in a taxonomic Municipal 34: 1-32. treatment entitled Flora Ilustrada Catarinense. These Allem, A.C. 1979. Quatro novas espécies de Croton L. authors used the old subdivision of the family, (Euphorbiaceae) do Rio Grande do Sul. Boletin de approaching the subfamilies Phyllanthoideae (with la Sociedad Argentina de Botânica 18: 63-81. two ovules in each locule of the ovary), in which Alves, M. V. S. 1982. Chaetocarpus Tw. (Euphorbiaceae). stood out genera such as Phyllanthus and Richeria, Espécies do continente americano. Dissertação de among others, and Crotonoideae (with one ovule in Mestrado. Universidade Federal do Rio de Janeiro, each locule of the ovary), in which most genera were Rio de Janeiro. 84p. placed, such as Acalypha, Croton, Euphorbia, etc. Alves, R.J.V. & Kolbek, J. 2009. Summit vascular flora of Serra de São José, Minas Gerais, Brazil. Checklist 5: 35-73. It was proven that studies on Euphorbiaceae APG II. 2003. An update of the Angiosperm Phylogeny s.l are growing, particularly in the northern, Group classification for the orders and families of northeastern, and southeastern regions of Brazil, flowering plants: APG II. Botanical Journal of the with still some gaps in the central and southern Linnean Society 141: 399-436. regions. Research in these two last regions will lead APG III. 2009. An update of the Angiosperm Phylogeny to a broader knowledge of this important family. Group classification for the orders and families of There are many studies on Brazilian species flowering plants: APG III. Botanical Journal of the of Croton L. However, species of the states of Mato Linnean Society 161: 105-121. Grosso, Goiás, part of the Brazilian Amazon (Pará, Baillon, H. 1858. Étude générale du groupe des Amazonas, Amapá, and Roraima), Rio Grande do Euphorbiacées. Victor Masson, Paris. Sul, and Paraná, especially herbs, need revision. Baldwin, J. & Schultes, R.E. 1947. A conspectus of There is a need for taxonomic revision/update the genus Cunuria. Botanical Museum Leaflets of also for Phyllanthus, Dalechampia, Drypetes, Harvard University 12: 325-351. Mabea, and Sapium Jacq.; the latter is still a great Bentham, G. 1878. Notes on Euphorbiaceae. Journal challenge. In addition, we point out the need of Linnean Society of London, Bot. 17: 185-267. for more comprehensive taxonomic studies on Bentham, G. 1880. Euphorbiaceae. In: G. Bentham & J. the Brazilian Hippomaneae, in particular due to D. Hooker, J (eds.). Genera Plantarum 3: 239-340. the changes in this taxon in the past few years, Berry, P.E.; Cordeiro, I.; Wiedenhoeft, A.C.; Vitorino- what lead to the establishment of novelties such Cruz, M.A. & Lima, L.R. 2005a. Brasiliocroton, a as Pseudosenefeldera Esser, Dendrothrix Esser, new crotonoid genus of Euphorbiaceae from eastern Brazil. Systematic Botany 30: 356-364. Rhodothyrsus Esser, and Pleradenophora Esser. Berry, P.E.; Hipp, A.L.; Wurdack, K.J.; Ee, B.V & Riina, We noticed a lack of published studies in Brazil R. 2005b. Molecular phylogenetics of the giant on molecular systematics of the Euphorbiaceae genus Croton and tribe Crotoneae (Euphorbiaceae s.s., Phyllanthaceae, Picrodendraceae, and s.s.) using ITS and trnL-trnF DNA sequence data. Putranjivaceae. We believe that graduate programs American Journal of Botany 92: 1520-1534. can provide the means to form new taxonomists Bezerra, P. & Fernandes, F. 1982. Marmeleiros do and to choose modern methods for the taxonomic Nordeste. Anais do XXXIII Congresso Nacional treatment of the genera that need additional studies, de Botânica. Pp. 145-156. in order to define the species better. Special attention Bigio, N.C. 2009. As espécies de Pera Mutis (Euphorbiaceae) should be given to Peraceae, since this group is still na Amazônia brasileira. Dissertação de Mestrado. being debated. Universidade Federal Rural da Amazônia, Museu Paraense Emílio Goeldi, Belém. 117p. Acknowledgements Bigio, N.C.; Secco, R.; Oliveira, A.S.; Senna-Valle, L.; Medeiros, D. & Pinto, L.J.S. Peraceae. 2010. In: Inês Cordeiro, Margareth Sales, and Ricardo Forzza, R.C. et al. (org.). Catálogo de plantas e Secco thank the Brazilian Research Council fungos do Brasil. Vol. 2. JBRJ, Rio de Janeiro. Pp. (CNPq) for the productivity fellowships granted 1437-1438. for the study of Euphorbiaceae in the southeastern, Bigio, N.C. & Secco, R.S. 2010. Uma nova espécie de northeastern, and northern regions of Brazil. The Pera (Peraceae), endêmica de Manaus, Amazonas, PhD Alessandro Rosário formatted the text. Brasil. Rodriguésia 61: 77-81.
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Artigo recebido em 20/06/2011. Aceito para publicação em 24/11/2011.
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