Rodriguésia 63(1): 089-100. 2012 http://rodriguesia.jbrj.gov.br
Poaceae: an overview with reference to Brazil Poaceae: uma visão geral com referência ao Brasil
Hilda Maria Longhi-Wagner1,2
Abstract A summary of the classification of Poaceae into subfamilies according to different proposals is provided, as well as data on morphology and chorology of grasses, especially those that occur in Brazil. Key words: Gramineae, classification, morphology, geographical distribution, Brazil. Resumo É fornecido um sumário da classificação de Poaceae em subfamílias, de acordo com diferentes propostas, bem como dados sobre a morfologia e corologia de gramíneas, especialmente daquelas ocorrentes no Brasil. Palavras-chave: Gramineae, classificação, morfologia, distribuição geográfica, Brasil.
Introduction Classification in subfamilies and The family Poaceae (Gramineae) encompasses its relationship with morphology approximately 800 genera and 10,000 species and chorology (Watson & Dallwitz 1992), which predominate in the Several classification systems of Poaceae have flora and physiognomy of open vegetations all over been proposed, based mainly on the morphology the world. The economic importance of this family of the spikelets. Two large groups were accepted: is unquestionable for it has many different types of (1) spikelets with two anthecia (see McClure & use, and it includes the major cereals and forages. Soderstrom 1972), the basal anthecium staminate Because of the great diversity of Poaceae or neuter (very rarely bisexual) and the apical and its wide variability (Watson & Dallwitz bisexual, with the articulation between rachilla 1992), researchers have traditionally divided it and pedicel below the glumes; and (2) spikelets into subfamilies. This division was initially based exhibiting from one to many anthecia, all bisexual on morphological, anatomical, cytological, and or at least one basal bisexual, and the articulation chromosomic characters, with molecular data being above the glumes, corresponding, respectively, to added more recently. Hence, the number of accepted the series Panicaceae and Poaceae by Bentham & subfamilies and the characters used to determine Hooker (1880). These groups were later accepted them have varied through time. In the seminal as the subfamilies Panicoideae and Festucoideae, publication of GPWG (2001), which proposed a respectively (Hitchcock 1950). Prat (1936) added new subfamilial classification of Poaceae based on the subfamily Bambusoideae, also based on morphological and molecular data, the authors made anatomical and cytological characters, especially a comparison between their classification and the the presence of fusiform cells in the mesophyll. main previous proposals of classification. Later, Pilger (1956) accepted a larger number of The objective of this study is to review subfamilies: Andropogonoideae, Anomochlooideae, different aspects of the taxonomy of Poaceae, Eragrostoideae, Festucoideae, Micrairoideae, including morphology and subfamily classification, Olyroideae, Oryzoideae, and Panicoideae. and to consider the relationship between chorology Parodi (1961) proposed a classification and habitat for different groups. An overview of of the Argentinean grasses in the subfamilies the taxonomic studies of Poaceae in Brazil and of Bambusoideae, Oryzoideae, Phragmitoideae, the possibilities within this field of research is also Festucoideae, Eragrostoideae, and Panicoideae. provided. This classification was used for a long time by
1 Universidade Federal do Rio Grande do Sul, Depto. Botânica, Av. Bento Gonçalves 9500, Prédio 43433, bl. 4, 91501-970, Porto Alegre, RS, Brazil. 2Author for correspondance: [email protected] 90 Longhi-Wagner, H.M. agrostologists of Argentina, Uruguay, and southern independent of Bambusoideae, following Parodi Brazil, where, due to their role in cattle raising (a (1961), Stebbins & Crampton (1961) and Valls very important economic activity in this region), (1980); (ii) the division of part of Bambusoideae the study of grasses has been well developed. into the subfamilies Anomochlooideae, Puelioideae, At the same time, Stebbins & Crampton (1961) and Pharoideae, constituting the group with the proposed to divide the North American grasses into most basal forest grasses; (iii) the acceptance of the six families, similar to those outlined by Parodi subfamily Aristidoideae encompassing the genera (1961), except for Phragmitoideae, which was Aristida L., Sartidia De Winter and Stipagrostis named Arundinoideae. Nees, which had been included in Arundinoideae Valls (1980), in his list of the grass genera of by Clayton & Renvoize (1986). Brazil, accepted the same six families mentioned Later, based on molecular phylogeny, by Stebbins & Crampton (1961), but named Sánchez-Ken et al. (2007) acknowledged the Festucoideae as Pooideae, and Eragrostoideae as inclusion of the tribe Micraireae in the subfamily Chloridoideae, according to the nomenclatural Micrairoideae, which had been included in rules prevailing at that time. Arundinoideae by Clayton & Renvoize (1986) and Later, Clayton & Renvoize (1986) published incertae sedis (at least the genus Micraira F. Muell.) Genera Graminum, also recognizing these in GPWG (2001), increasing to 13 the number of six subfamilies. Pooideae, Chloridoideae, and subfamilies accepted. The genus Isachne Benth. Panicoideae consisted of the study by Valls (ca. three species in Brazil), traditionally included (1980), but with a different circumscription for in the tribe Paniceae, subfamily Panicoideae, Bambusoideae and Arundinoideae. Bambusoideae appeared unexpectedly in the clade Micrairoideae included the tribes Oryzeae and Erharteae (Sánchez-Ken et al. 2007). (Oryzoideae, in Parodi 1961; Stebbins & Crampton Regarding the number of accepted subfamilies 1961; Valls 1980), and Arundinoideae included the of Poaceae, more recent studies, using new DNA tribes Micraireae and Aristideae. Aristideae was sequences and larger samples, identify only 12 accepted in Chloridoideae by Valls (1980). The subfamilies again, maintaining Micrairoideae, sixth subfamily accepted by Clayton & Renvoize but placing the subfamily Centothecoideae (1986) was Centhotecoideae, which Valls (1980) (represented in Brazil only by the forest genus considered as part of the tribe Centhoteceae, Orthoclada P. Beauv.) back in the traditional subfamily Arundinoideae. Panicoideae (Sánchez-Ken & Clark 2007, 2010). In addition to their seminal work Genera Among the data used by GPWG (2001), Graminum (Clayton & Renvoize 1986), Clayton et characters are related to fruit and embryo (Reeder al. (2002) were responsible for the development of 1957), leaf anatomy (Ellis 1986), biochemistry of the project GrassBase – The Online World Grass photosynthesis (Brown 1977; Hattersley 1986), Flora (
Rodriguésia 63(1): 089-100. 2012 Poaceae: an overview 91 which is highly specialized. In addition, there Pooideae), which is exclusively C3, and the other is the presence of fusiform and arm-cells in the clade is PACCAD (Panicoideae, Arundinoideae, mesophyll, characters which are present in most Chloridoideae, Centothecoideae, Aristidoideae, members of other forest Poaceae subfamilies. and Danthonioideae). This second clade also has These two genera are exclusively Neotropical and Kranz syndrome in most of its members, and it is occur mainly in the Atlantic Forest. Anomochloa assumed that this syndrome evolved more than marantoidea Brong., known from a few disturbed once in Poaceae (GPWG 2001). More recently, areas in southern Bahia, is included in the list of Sánchez-Ken et al. (2007) denominated the clade endangered species of Brazil published by the PACCAD as PACCMAD, because the subfamily Ministry of the Environment (MMA 2008). The Micrairoideae was included. Clark (2009) named genus Streptochaeta has two species in tropical this clade PACMAD, as recent studies suggested Brazil (Filgueiras & Gonçalves 2006); among the inclusion of Centothecoideae in Panicoideae them, only one, S. spicata Nees, occurs up to the (Sánchez-Ken & Clark 2007). northern coast of Rio Grande do Sul, following the Bambusoideae: this subfamily has also southern boundary of the Brazilian Atlantic Forest. C3 photosynthesis and occurs predominantly Pharoideae: this is the next divergent lineage in forests. However, contrarily to the above- of Poaceae, in which occur the typical reproductive mentioned subfamilies, Bambusoideae is highly structures of the family: the spikelets. Its spikelets diversified, including approximately 1,400 are formed by two glumes and one anthecium species (Clark 2009). For Brazil, Filgueiras & (lemma and palea), both in staminate and pistillate Santos-Gonçalves (2004) estimated 232 native spikelets, and the plants are monoecious. The two species; this number was updated by Filgueiras et types of spikelets are assembled in pairs in each al. (2012) as 266 species, among which 159 are node of the inflorescence; staminate spikelets are endemic. In its current state, it includes bamboos considerably smaller than pistillate ones. A strong that are herbaceous, monoecious, polycarpic, with morphological synapomorphy is the presence of a staminate and pistillate spikelets in the same plant twist on the pseudopetiole of the leaf, which makes in the tribe Olyreae, and bamboos that are lignified, the abaxial surface of the leaf face up, unlike other with bisexual spikelets, and which are usually Poaceae. This subfamily occurs in the tropics of the monocarpic multiannual, in the tribes Bambuseae Old and New World and is represented in Brazil (tropical) and Arundinarieae (temperate), according by the genus Pharus P. Browne, with four species to Clark (2009) and Soreng et al. (2011 onwards). (Filgueiras & Santos-Gonçalves 2004; Filgueiras Members of the tribe Arundinarieae occur et al. 2012). mainly in montane regions of Asia and in lower Puelioideae: this is the next divergent lineage degree, North America, without native members in (GPWG 2001), which also includes forest grasses, Brazil. Olyreae is an almost exclusively Neotropical represented by the genera Guaduella Franch. and tribe and includes several species that are either native Puellia Franch., the former with six species and or endemic to Brazil, with the exception of Olyra the latter with five. These are endemic to Africa, latifolia L., which occurs also in Africa (whether and are distributed from Serra Leone to Angola it is native or introduced is still under discussion), (Clayton & Renvoize 1986). and Burgersiochloa Pilg., a monotypic genus from The above-mentioned subfamilies are the New Guinea (Clayton & Renvoize 1986). The most basal in the phylogeny of Poaceae; they occur diversity center of Olyreae is located in the Atlantic in forests with a predominantly tropical distribution. Forest of southern Bahia and northern Espírito Their members usually exhibit fusiform and Santo (Soderstrom et al. 1988; Oliveira 2006), arm-cells in the mesophyll and absence of Kranz and new species are constantly found, as studies anatomy, and have clearly (or presumably, in are intensified in these areas (Oliveira et al. 2008 the case of Anomochlooideae and Puelioideae, a,b). The tribe Bambuseae exhibits two diversity according to GPWG 2001) C3 photosynthesis. centers, one in Asia and one in the Neotropics, with According to GPWG (2001), all the remaining several members in the Atlantic Forest, many of subfamilies form a large clade that diversified early which are endemic. In this tribe many new species and includes two smaller clades. One of these is have also been described (Sendulsky 1995, 1997; the clade BEP (including Bambusoideae, also Clark 1996, 2004; Filgueiras & Londoño 2006; predominantly forest grasses, Ehrartoideae, and Santos-Gonçalves et al. 2011), and some revisions
Rodriguésia 63(1): 089-100. 2012 92 Longhi-Wagner, H.M. have been made (Guala 2003; Santos-Gonçalves as a subtribe (Aveninae). Pooideae includes ca. 3,300 2005; Viana 2010). Probably, the tribes Olyreae and species (GPWG 2001), and it is the main subfamily Bambuseae are also well represented in the Amazon of microthermic grasses. In southern Brazil and Forest, though there are few records due to the lower neighboring countries, they contribute significantly sampling intensity in the region. to the production of forage during winter, which The subfamily Bambusoideae needs more is a crucial period for natural pastures. Its species studies in Brazil, including intensive collecting exhibit usually membranous ligule, spikelets with in the Atlantic Forest and Amazon, which would the basal bisexual anthecia and articulation above probably reveal new and endemic species, both the glumes, which persist in the inflorescence after among lignified and herbaceous bamboos. An the fall of mature anthecia, lemmas with five or analysis of the distribution and conservation of more nerves, and C3 photosynthesis. Burman (1985) herbaceous bamboos was published by Oliveira et reported 128 species for Brazil, whereas 135 are al. (2006), and an analysis of the native bamboos listed in Filgueiras et al. (2012), 25 of which are of Brazil was published by Filgueiras & Santos- endemic. The highest species richness occurs in Gonçalves (2006). Furthermore, there are different the natural grasslands of Rio Grande do Sul state, a research groups studying the commercial use of few species extending to the “campos de altitude” lignified bamboos (Almeida & Teixeira 2006). or, in smaller numbers, to the rocky fields (“campos Many governmental measures have been rupestres”) of southeastern Brazil. The main winter implemented in order to advance the knowledge crops of southern Brazil belong to this subfamily, e.g. and use of bamboos. One of the most important wheat, barley, rye and oat. Although this subfamily measures was the creation of the Center for the is already well-documented in Brazil (Kämpf 1975; Study of Bamboos and Natural Fibers (Centro Longhi-Wagner 1987; Zanin et al. 1992), one aspect de Estudos de Bambus e Fibras Naturais) at the that still requires further studies is the northern University of Brasília and the National Bamboo limit of its distribution within the country, which Network (Rede Nacional do Bambu). was published only for the genus Stipa L. (Longhi- Erhartoideae: this subfamily was included Wagner & Zanin 1998). in Bambusoideae by Clayton & Renvoize (1986), Danthonioideae: this subfamily was in part due to the presence of fusiform and arm- described by Barker & Linder in GPWG (2001), cells in some of its members. These types of grouping part of the genera previously accepted cells do not occur in other Poaceae, except for in Arundinoideae sensu lato. It includes ca. 250 the genera Homolepis Chase and Streptostachys species, all C3 (GPWG 2001). It is represented in Desv. (Panicoideae), according to GPWG (2001). Brazil by the genera Cortaderia Stapf, with three This subfamily exhibits bisexual spikelets with native species and Danthonia DC., with five native characteristically very reduced or even absent species (Santos & Boechat 1989), as well as the glumes. It comprises ca. 120 species (GPWG 2001) genus Hakonechloa Makino ex Honda, with one that inhabit moist sites, such as marshes and river subspontaneous species (Filgueiras et al. 2012). banks. Burman (1985) reported the occurrence of Some native species occur in southern Brazil. A few ca. 19 species for Brazil, a number very close to native species have been found in low abundance the 20 species reported by Filgueiras et al. (2012), in high-altitude fields of southeastern Brazil; one including two cultivated species. This subfamily of them, Danthonia montana Pilg., is found all the needs to be studied in detail, particularly the genera way to the high altitude fields of Bahia (Morro do Luziola Juss. and Leersia Sol. ex Sw. Chapéu). This is the northernmost limit known for Pooideae: the circumscription of this family microthermic grasses in Brazil (Longhi-Wagner & underwent few modifications in the system of Oliveira 2002). GPWG (2001), compared with Clayton & Renvoize Arundinoideae: this subfamily had its (1986). However, the circumscription of tribes in the circumscription drastically reduced in the study two aforementioned classifications was profoundly by GPWG (2001); it was not considered to be changed in comparison with previous classifications, well-defined, needing further studies, since no such as that of Parodi (1961), in particular with the morphological synapomorphy was found to support division of Poeae (sensu lato, in Parodi 1961) its monophyly. In the systems of Parodi (1961, as into the tribes Poeae sensu stricto, Bromeae, and Phragmitoideae) and Stebbins & Crampton (1961, Meliceae, and with the inclusion of Aveneae in Poeae as Arundinoideae), as well as of Valls (1980), this
Rodriguésia 63(1): 089-100. 2012 Poaceae: an overview 93 subfamily includes many genera with Brazilian forests (Longhi-Wagner 1999). It is possible that native species, such as Cortaderia, Gynerium other Aristida C3 species could be found, as the Willd. ex P. Beauv., Danthonia, and Phragmites number of sampled species increases. This is an Adans. In the classification proposed by Watson interesting topic for studies on this genus in Brazil. & Dalwitz (1992), Arundinoideae included even Chloridoideae: the circumscription of this the tribe Stipeae, which is currently within the subfamily did not change much in the system of subfamily Pooideae. In Clayton & Renvoize (1986), GPWG (2001), compared to the system of Clayton it includes the tribes Aristideae and Micraireae, & Renvoize (1986). In relation to previous systems, both later accepted as independent subfamilies such as that of Parodi (1961) and Valls (1980), (GPWG 2001; Sanchez-Ken et al. 2007). the greatest modification was the separation of Arundinoideae had always been a subfamily Aristida in a different subfamily, as mentioned with a difficult circumscription, and its current form above. It is interesting to emphasize that, in Clayton
(GPWG 2001) includes 33 to 38 species, all C3. It & Renvoize (1986), this genus was included in has only one representative in Brazil, Phragmites Arundinoideae. Members of Chloridoideae exhibit australis (Nees) Döll, a species considered spikelets with one or more basal bisexual anthecia, cosmopolitan, of swamps and riverbanks. with persistent glumes, as in Pooideae, but the Micrairoideae: includes ca. 170 species ligule is usually pilose or membranous-ciliate, distributed mainly in Australia and Asia, less and the lemma is 3-nerved. Besides, it is a very commonly in the Neotropics, in open savannas or homogeneous subfamily in terms of presence and edges of forest. Most of them have C3 photosynthesis, type of Kranz syndrome (Brown 1977), with only except for species of the genera Eriachne R. Br. and one African species of Eragrostis Wolf and one of
Pheidochloa S.T. Blake, which do not occur in Brazil Merxmuellera Conert known as C3 (GPWG 2001). (GPWG 2001). This subfamily is represented in Members of this subfamily are usually found in tropical Brazil by four species of the genus Isachne grasslands, often in arid environments, and very R. Br., a genus which includes approximately 100 occasionally occurr on forest edges, but not inside species that are mainly distributed in Asia (Clayton & forests. It includes ca. 1,400 species (GPWG 2001). Renvoize 1986). Isachne was included by Filgueiras For Brazil, Burman (1985) estimated 157 species, et al. (2012) in the subfamily Panicoideae, to which it including species of Aristida. Filgueiras et al. previously belonged. These plants have stoloniferous (2012) listed 161 species, but excluded Aristida, habits, occur in forest edges, and exhibit morphology which is currently accepted in Aristidoideae. similar to the tribe Paniceae, namely spikelets with Recently, Columbus et al. (2007) published deciduous glumes and two anthecia, but with both a phylogenetic analysis of the subfamily bisexual and coriaceous anthecia. Isachne is one of Chloridoideae, modifying the circumscription of the grass genera that need urgent revision in Brazil. some tribes. Aristidoideae: this subfamily was originally Panicoideae: the subfamily Panicoideae described by Caro (1961), but it was not adopted belongs to a Poaceae lineage that developed by later authors. GPWG (2001) accepted this spikelets with the articulation between the rachilla subfamily with three genera: Aristida L., with and the pedicel located below the glumes (both ca. 250 pantropical species, of which 36 occur in glumes and mature anthecia are deciduous), and Brazil (Filgueiras et al. 2012); Sartidia Winter, with only two anthecia, the upper one is bisexual and the four African species and C3 photosynthesis; and lower one is generally staminate or neuter. These Stipagrostis Nees, with ca. 50 African and Asian characteristics distinguish it from most Poaceae, species, C4 photosynthesis, and only one Kranz- in which the spikelets usually have basal bisexual type sheath around the vascular bundles. Aristida anthecia and persistent glumes. was reported as a genus with C4 photosynthesis by According to GPWG (2001), Panicoideae
GPWG (2001) and other authors, with a double includes 3,270 species, most with C4 photosynthesis Kranz sheath (Brown 1977). However, recently of three different biochemical subtypes (Brown
Cerros-Tlatilpa & Columbus (2009) discovered 1977; Hattersley 1986), but it also includes C3 an Aristida species C3 without Kranz anatomy, species. Among these, there are stoloniferous A. longifolia Trin., which occurs from Central species of the tribe Paniceae that occur inside America to central and southeastern Brazil, in forests, covering the forest ground (species of cerrado, caatinga, and shaded sites next to riparian Oplismenus P. Beauv., Pseudechinolaena Stapf
Rodriguésia 63(1): 089-100. 2012 94 Longhi-Wagner, H.M. and Ichnanthus P. Beauv., among others), or modifications (Denham et al. 2002; Denham 2005; with scandent habits (Lasiacis (Griseb.) Hitchc.). Zuloaga et al. 2007; Morrone et al. 2008a; Sede et However, apparently there is no strong relationship al. 2008; Zuloaga & Morrone 2009). between habitat and photosynthesis type. There According to Filgueiras et al. (2012), the are C3 species that occur typically in open and remarkable number of 238 species of Panicoideae arid environments, for example those of the genera are endemic to Brazil. Apochloa Zuloaga & Morrone and Renvoizea Zuloaga & Morrone, which previously belonged to Poaceae in Brazil Panicum L. group Lorea (Sede et al. 2008). Those As previously mentioned among Poaceae species inhabit outcrops in the “campos rupestres” subfamilies, only Puelioideae is not represented in of Cadeia do Espinhaço, southeastern Brazil. Brazil, though Micrairoideae is still not included Likewise, Echinolaena inflexa (Poir.) Chase, a in the list of Brazilian grasses (Filgueiras et al. grass considered as typical of the Cerrado, is C . 3 2012). In subfamilies that include megathermic It is important to highlight that most species of the species of predominantly tropical distribution tribe Paniceae occur in grasslands, cerrados, rocky (i.e., all that were previously mentioned, except fields, and aquatic environments (Allem & Valls Danthonioideae and Pooideae) there is usually a 1987) all over Brazil. pronounced decrease in species richness towards Burman (1985) estimated the existence of southern Brazil, as well as a decrease in population 838 Panicoideae species in Brazil, and Filgueiras density and size. An example is the tribe Olyreae et al. (2012) listed 778 species. It is by far the most of herbaceous bamboos, of which, according to diversified subfamily of Poaceae in Brazil and its the data currently available only the genera Olyra species predominate in grasslands, even in southern L. and Parodiolyra Soderstr. & Zuloaga reach the Brazil, where the microthermic subfamily Pooideae state of Rio Grande do Sul, the first genus with is highly diversified. Due to its importance as a key two species and the second with one, following element of Brazilian grasslands and cerrados, and the southern limit of the Atlantic Forest. As for the due to its high richness of genera and species, this mainly tropical tribe Bambuseae, which includes subfamily needs further taxonomic revisions in lignified bamboos, the decrease in diversity addition to those already made (e.g. Canto-Dorow towards southern Brazil is less pronounced: 2001; Boechat 2005; Oliveira 2004; Zuloaga & Schmidt & Longhi-Wagner (2009) recorded five Morrone 2005; Zanin & Longhi-Wagner 2006). genera with 27 native species in Rio Grande do Sul. The published surveys of Brazilian floras are also Some genera of grasses that are more important, such as Valls et al. (2001) and Oliveira diversified in the tropics present two different & Valls (2001), who studied the important genera patterns of distribution in Brazil, one including Axonopus P. Beauv. and Paspalum L., respectively. species predominantly or exclusively tropical Moreover, it must be taken into account that and another including species predominantly Panicoideae includes genera of great importance for or exclusively extratropical. Examples of these natural pastures, most notably Paspalum, which has patterns are Aristida (Aristidoideae), and Eragrostis been the subject of multidisciplinary studies aimed (Chloridoideae). The distribution patterns of the at its commercial use (Quarín et al. 1997; Casa et al. species of Eragrostis native to Brazil were analyzed 2002; Souza-Chies et al. 2005; Machado et al. 2005; by Boechat & Longhi-Wagner (2000). Miz & Souza-Chies 2006; Pozzobon et al. 2008). On the other hand, members of Pooideae and The changes that have been proposed Danthonioideae have a distribution predominantly for the internal classification of the subfamily extratropical in Brazil, with a pronounced decrease Panicoideae are focused mainly on the inclusion of in species richness, density, and population size the subfamily Centothecoideae, on changes in the from south to north. circumscription of genera and, to a lesser degree, One of the first studies of Brazilian grasses was tribes (Sánchez-Ken & Clark 2010). For example, published by Raddi (1823), a member of the group the genus Panicum has been subjected to several of scientists that arrived in Brazil with the Princess phylogenetic studies that led to the transference Maria Leopoldina of Austria. Raddi remained in of many of its species to other genera of the tribe Brazil from 1817 to 1818, collecting plants in the Paniceae or even to new genera, whereas the state of Rio de Janeiro and its surroundings. His circumscription of Paspalum has undergone minor collections served as a basis for the publication of
Rodriguésia 63(1): 089-100. 2012 Poaceae: an overview 95 Agrostografia brasiliensis (Raddi 1823), which was It is important to emphasize that only a few recently analyzed by Baldini & Longhi-Wagner Brazilian states include Poaceae in their published (2006) and Longhi-Wagner & Baldini (2007). floras (Smith et al. 1981-1982; Renvoize 1984, Nees (1829) published the study Agrostologia 1988; Filgueiras 1995; Longhi-Wagner et al. 2001). brasiliensis, which included 83 genera and 403 However, a great number of floristic surveys of species, emphasizing the high number of species smaller areas have been published and provide waiting to be described. In Flora brasiliensis, the the basis for studies in other fields of research, grasses were studied by Döll (1871-1877) and by such as phytosociology, floristic, and ecology (e. Hackel (1883), including several new species. g. Burman et al. 1987; Rocha et al. 2001; Rocha & The historical collections, on which the Secco 2004; Longhi-Wagner & Todeschini 2004; publications cited above were based, are deposited in Welker & Longhi-Wagner 2007; Ferreira et al. different European herbaria and are relatively well- 2009), contributing substantially to the knowledge known. However, there are other historical collections, of the Brazilian agrostological flora. also in European herbaria, that remain to be studied Several genera revisions for Brazil have and that can reveal taxonomic and nomenclatural already been published (Sendulsky & Soderstrom novelties, as pointed out by Baldini (2009). 1984; Filgueiras 1984, 1989; Boechat & Longhi- Valls (1980) estimated the occurrence of more Wagner 1995, 2001; Longhi-Wagner 1999; Oliveira than 180 Poaceae genera in Brazil and Burman & Valls 2002, Guglieri et al. 2006b; Zanin & (1985) published an estimate of 197 genera and Longhi-Wagner 2006). Due to the continental 1,368 species. In the 1990s, many new species, dimensions of Brazil, revisions comprising the and even genera, were published (Longhi-Wagner whole country are frequently hindered by the 1994; Filgueiras 1996; Sendulsky 1997; Boechat & difficulty of collecting with the necessary intensity. Longhi-Wagner 1998; Filgueiras et al. 2001; Zanin It is important, however, that these revisions are & Longhi-Wagner 2003; Guglieri et al. 2006a; carried out, even if intensive sampling is conducted Oliveira & Longhi-Wagner 2007; Oliveira et al. only in a part of a taxon’s range (more accessible to 2008 a,c; Oliveira & Valls 2009, among others). In researchers), and the work is complemented by the the List of Species of the Brazilian Flora, Filgueiras analysis of herbarium material for the remaining et al. (2012) mentioned 210 genera of Poaceae, of areas. Such studies will have a far-reaching impact which 21 are endemic, and 1,414 species, of which and will bring advancements in the knowledge of 468 are endemic. Poaceae in Brazil. Checklists of species of Poaceae were published for a number of regions of Brazil. Final Considerations Longhi-Wagner & Renvoize (2006) listed 99 As the subfamily classification of Poaceae genera and 421 species for northeastern Brazil, tends to stabilize, current phylogenetic studies and Rezende et al. (2008) listed 91 genera and focus mainly on the analysis of the circumscription 510 for the Cerrado (Central Brazil). In addition, of tribes (e. g. Columbus et al. 2007) and genera Morrone et al. (2008b) listed 203 genera and 1,528 (e. g. Zuloaga et al. 2007; Morrone et al. 2008a; species (540 endemic) for the Uruguay, Argentina, Sede et al. 2008; Rua et al. 2010). Unfortunately, Paraguay and the three states of southern Brazil. although these analyses are very important, few Checklists like those mentioned above or even studies with this focus are being conducted in less comprehensive lists provide important data for Brazil (e. g. Oliveira 2006; Essi et al. 2008). phytogeographic analysis and lay the ground for Likewise, ontogenetic analyses (Sajo et al. 2007, surveys and conservation studies. However, as they 2008; Nakamura & Scatena 2009; Nakamura et do not provide analytic keys or illustrations, they al. 2009) and population analyses (Oliveira et al. need to be complemented by taxonomic revisions 2008b) are scarce in Brazil, even though they are that allow species identification. Publications extremely valuable. of regional floras and taxonomic revisions are Data obtained with the above-mentioned very useful and urgently needed. Once these data approaches are essential for biodiversity analysis, become available, they would be the basis for the survey of hotspots, and for conservation programs. implementation of public policies aimed at the The Official List of Endangered Species of Brazil sustained use and conservation of native resources (MMA 2008) contains 14 Poaceae, of which seven in different regions of the country. are from the Atlantic Forest, one is shared between
Rodriguésia 63(1): 089-100. 2012 96 Longhi-Wagner, H.M. the Atlantic Forest and the Pampa, five are from Boechat, S.C. & Longhi-Wagner, H.M. 1995. O gênero the Cerrado, and only one is from the Amazon. Sporobolus (Poaceae: Chloridoideae) no Brasil. However, a much higher number of species could Acta Botanica Brasilica 9: 17-73. be under threat, but were not assessed due to data Boechat, S.C. & Longhi-Wagner, H.M. 1998. Duas novas deficiency. Among these are many Amazonian espécies de Eragrostis (Poaceae, Chloridoideae) do Brasil. Iheringia 51: 177-187. bamboo species, known only through their type Boechat, S.C. & Longhi-Wagner, H.M. 2000. Padrões material, which have never been collected again. de distribuição dos táxons brasileiros de Eragrostis Hence, the study of Poaceae in Brazil must (Poaceae - Chloridoideae). Revista Brasileira de focus mainly on intensive field collection in several Botânica 23: 177-194. areas, such as the Amazon and other areas that Boechat, S.C. & Longhi-Wagner, H.M. 2001. O gênero have been relatively poorly sampled. Taxonomic Eragrostis (Poaceae) no Brasil. Iheringia 55: 23-169. revisions and floristic studies should also be Brown, W.V. 1977. The Kranz syndrome and its subtypes continued and have their publication stimulated. in grass systematic. Memoirs of the Torrey Botanical The training of human resources in different Club 23: 331-422. regions of the country is also important, especially Burman, A.G. 1985. Nature and composition of grass in the northern and northeastern regions, where flora of Brazil. Willdenowia 15: 211-233. specialists in Poaceae are still lacking. There is Burman, A.G.; Sendulsky, T.; Longhi-Wagner, H.M. also a need for more agrostologists to be enlisted & Renvoize, S.A. 1987. Gramineae. In: Giulietti, as supervisors in graduate programs in Botany. A.M.; Menezes, N.L.; Pirani, J.R.; Meguro, M.; Wanderley, M.G.L. (eds.). Flora da Serra do Cipó, Finally, it is necessary to promote cooperation Minas Gerais: caracterização e lista das espécies. between researchers using different approaches, Boletim de Botânica 9: 117-123. so that Brazilian agrostologists can contribute not Canto-Dorow, T.S. 2001. O gênero Digitaria no Brasil. only to a more complete survey on this important Tese de Doutorado. Universidade Federal do Rio family in the country, but also to the advancement Grande do Sul, Porto Alegre. 386p. of its knowledge at a global level. Caro, J.A. 1961. Las especies de Aristida (Gramineae) del Centro de la Republica Argentina. Kurtziana 1: Acknowledgements 123-206. Casa, A.M.; Mitchell, S.E.; Lopes, C.R. & Valls, J.F.M. The Brazilian National Council for Scientific 2002. RAPD analysis reveals genetic variability and Technological Development (CNPq) granted us among sexual and apomictic Paspalum dilatatum a research fellowship. Anonymous reviewers made Poiret biotypes. Journal of Heredity 93: 300-302. invaluable suggestions to the manuscript. Cerros-Tlatilpa, R. & Columbus, T. 2009. C3 photosynthesis in Aristida longifolia: implications for photosynthetic References diversification in Aristidoideae (Poaceae). American Allem, A.C. & Valls, J.F.M., 1987. Recursos forrageiros Journal of Botany 96: 1379-1387. nativos do pantanal mato-grossense. EMBRAPA, Clark, L.G. 1992. Chusquea sect. Swallenochloa Depto. Difusão e Tecnologia, Brasília. 339p. (Poaceae: Bambusoideae) and allies in Brazil. Almeida, J.G. & Teixeira, A.A. 2006. Estruturação da Brittonia 44: 387-422. rede de pesquisa e desenvolvimento em bambu. Clark, L.G. 1996. Four new species of Chusquea Seminário Nacional de Bambu: Anais, Brasília. (Poaceae: Bambusoideae) from Brazil and Ecuador. Baldini, R.M. & Longhi-Wagner, H.M. 2006. Poaceae Brittonia 48: 250-262. Raddianae: an updated nomenclatural and Clark, L.G. 2004. Two new species of Aulonemia and taxonomical evaluation of G. Raddis´s Brazilian Chusquea (Poaceae: Bambusoideae) from Brazil. Poaceae. Taxon 55: 469-482. Revista Brasileira de Botânica 27: 31-36. Baldini, R.M. 2009. An outlook at the historical collections Clark, L.G. 2009. Subfamilial classification of the of Neotropical Poaceae in the European herbaria. In: Poaceae, with emphasis on the Bambusoideae. In: Moura, C.W.N.; Silva, T.R.S.; Giulietti-Harley, A.M. Moura, C.W.N.; Silva, T.R.S.; Giulietti-Harley, & Santos, F.A.R. (eds.) Botânica brasileira: futuro e A.M. & Santos, F.A.R. (eds.) Botânica brasileira: compromissos. Resumos. Pp. 229-233. futuro e compromissos. Resumos. Pp. 218-222. Bentham, G. & Hooker, J.D. 1880. Genera Plantarum. Clayton, W.D. & Renvoize, S.A. 1986. Genera Vol. 3, pt. 1. L Reeve, Londini. Graminum. Grasses of the world. Her Maj. Stat. Boechat, S.C. 2005. O gênero Ichnanthus (Poaceae- Office, London. 389p. Panicoideae-Paniceae) no Brasil. Iheringia 60: Clayton, W.D.; Vorontsova, M.S.; Harman, K.T. & 189-248. Williamson, H. 2002 onwards. World grass species:
Rodriguésia 63(1): 089-100. 2012 Poaceae: an overview 97 descriptions, identification, and information Altoparadisium and a reevaluation of Arthropogon retrieval. Available in
Rodriguésia 63(1): 089-100. 2012 98 Longhi-Wagner, H.M.
Longhi-Wagner, H.M. 1999. O gênero Aristida (Poaceae) Nakamura, A.T.; Longhi-Wagner, H.M. & Scatena, V.L. no Brasil. Boletim do Instituto de Botânica 12: 2009. Desenvolvimento de óvulo, fruto e semente 113-179. de espécies de Poaceae (Poales). Revista Brasileira Longhi-Wagner, H.M. & Baldini, R. 2007. Synopsis de Botânica 32: 165-176. Poacearum in Josephii Raddii Agrostografia Nakamura, A.T. & Scatena, V.L. 2009. Desenvolvimento brasiliensis. Kew Bulletin 62: 381-405. pós-seminal de espécies de Poaceae (Poales). Acta Longhi-Wagner, H.M.; Bittrich, V.; Wanderley, M.G.L. Botanica Brasilica 23: 212-222. & Shepherd, J.G. 2001. Poaceae. In: Wanderley, Nees, C.C. 1829. Agrostologia brasiliensis. In: Martius, M.G.L.; Shepherd, G.J. & Giulietti, A.M. (eds.). C.F.P. Flora Brasiliensis Enumeratio Plantarum 2. Flora fanerogâmica do estado de São Paulo. Vol. 1. J.G. Cottae, Stuttgartiae. Pp. 1-292. Hucitec, São Paulo. Oliveira, R.C. 2004. Estudo taxonômico das espécies de Longhi-Wagner, H.M. & Oliveira, R.P. 2002. New grass Paspalum grupo Plicatula (Poaceae- Panicoideae- records for Bahia State, Brazil. Kew Bulletin 57: Paniceae) no Brasil. Tese de Doutorado. Universidade 971- 977. Estadual de Campinas, Campinas. 366p. Longhi-Wagner, H.M. & Renvoize, S.A. 2006. Poaceae. Oliveira, R.C. & Valls, J.F.M. 2001. Paspalum. In: In: Barbosa, M.R.V.; Sothers, C.; Mayo, S.; Gamarra- Wanderley, M.G.L.; Shepherd, G.J. & Giulietti, Rojas, C.F.L. & Mesquita, A.C. (eds.). Checklist A.M. (eds.). Flora fanerogâmica do estado de São das plantas do nordeste brasileiro. Vol. 1. Brasília. Paulo. Vol. 1. Hucitec, São Paulo. Pp. 191-228. Pp. 91-97. Oliveira, R.C. & Valls, J.F.M. 2002. Taxonomia de Longhi-Wagner, H.M. & Todeschini, B.H. 2004. Flora de Paspalum L. grupo Linearia (Gramineae - Paniceae) Grão Mogol, Minas Gerais: Gramineae (Poaceae). do Brasil. Revista Brasileira de Botânica 25: 371-389. Boletim de Botânica da Universidade de São Paulo Oliveira, R.C. & Valls, J.F.M. 2009. Duas novas espécies 22: 143- 165. de Paspalum L. (Poaceae: Paniceae) do Brasil. Longhi-Wagner, H.M. & Zanin, A. 1998. Padrões de Revista Brasileira de Botânica 32: 89-94. distribuição geográfica das espécies de Stipa L. Oliveira, R.P. 2006. Estudos taxonômicos, filogenéticos (Poaceae - Stipeae) ocorrentes no Brasil. Revista e biossistemáticos em Raddia Bertol. (Poaceae: Brasileira de Botânica 21: 167-175. Bambusoideae: Olyreae). Tese de Doutorado. McClure, F.A. & Soderstrom, T.R. 1972. The Universidade Estadual de Feira de Santana, Feira agrostological term anthecium. Taxon 21: 153-154. de Santana. 311p. Machado, A.C.C.; Valls, J.F.M.; Peñaloza, A.P.S. & Oliveira, R.P. & Longhi-Wagner, H.M. 2007. New Santos, S. 2005. Novos biótipos pentaplóides do species of Streptostachys (Poaceae: Paniceae) from grupo Dilatata de Paspalum L. (Gramineae) no Sul Brazil. Kew Bulletin 62: 493-497. do Brasil. Ciência Rural 35: 56-61. Oliveira, R.P.; Longhi-Wagner, H.M. & Jardim, J.G. Miz, R.B. & Souza-Chies, T.T. 2006. Genetic relationships and variation among biotypes of 2006. Diversidade e conservação de bambus dallisgrass (Paspalum dilatatum Poir.) and related herbáceos (Poaceae: Bambusoideae: Olyreae) da species using random amplified polymorphic DNA Mata Atlântica, Brasil. Seminário Nacional de markers. Genetic Resources and Crop Evolution Bambu: Anais. Brasília. Pp. 62-66. 53: 541-552. Oliveira, R.P.; Borba, E.L. & Longhi-Wagner, H.M. MMA ‒ Ministério do Meio Ambiente. 2008. Lista 2008a. Morphometrics of herbaceous bamboos oficial das espécies da flora brasileira ameaçadas of the Raddia brasiliensis Complex (Poaceae- de extinção. Instrução Normativa de setembro Bambusoideae): implications for the taxonomy 2008, anexo I. of the genus and new species from Brazil. Plant Morrone, O.; Denham, S.S.; Giussani, L.M. & Zuloaga, Systematics and Evolution 270: 159-182. F.O. 2008a. Parodiophyllochloa, a new genus Oliveira, R.P.; Borba, E.L.; Longhi-Wagner, H.M.; segregated from Panicum (Paniceae, Poaceae) based Pereira, A.C.S. & Lambert, S.M. 2008b. Genetic on morphological and molecular data. Systematic and morphological variability in the Raddia Botany 33(1): 66-76. brasiliensis Complex (Poaceae: Bambusoideae). Morrone, O.; Zuloaga, F.O.; Longhi-Wagner, H.M.; Plant Systematics and Evolution 274: 25-35. Izaguirre, P.; Beyhaut, R.; Cialdella, A.M.; Giussiani, Oliveira, R.P.; Longhi-Wagner, H.M.; Leite, K.R.B. & L.; Denham, S.S.; Guglieri, A.; Boldrini, I.I.; Zanin, Hollowell, V. 2008c. Pariana multiflora (Poaceae, A.; Salariato, D. & De Gennaro, D. 2008b. Poaceae. Bambusoideae, Olyreae), a new species from In: Zuloaga, F.O.; Morrone, O. & Belgrano, M.J. Eastern Brazil, with notes on the leaf anatomy of Catálogo de las plantas vasculares del Cono the genus. Systematic Botany 33: 262-266. Sur (Argentina, sur de Brasil, Chile, Paraguay y Parodi, L. 1961. La taxonomia de las gramíneas Uruguay). Monographs in Systematic Botany from argentinas a la luz de las investigaciones más the Missouri Botanical Garden 1: 705-1063. recentes. Recent Advances in Botany 1: 125-130.
Rodriguésia 63(1): 089-100. 2012 Poaceae: an overview 99 Pilger, R. 1956. Gramineae II. In: Engler, A. & Prantl, Panicoideae clade (Poaceae), based on ndhF and K. (eds.). Die Natürlichen Pfanzenfamilien. Vol. 14. rpl16 intron sequences and morphological data. Duncker & Hum blot, Berlin. 225p. Aliso 23: 487-502. Pozzobon, M.T.; Machado, A.C.C.; Vaio, M.; Valls, Sánchez-Ken, J.G. & Clark, L.G. 2010. Phylogeny and J.F.M.; Peñaloza, A.P.S.; Santos, S.; Côrtes, a new tribal classification of the Panicoideae s.l. A.L.A. & Rua, G.H. 2008. Cytogenetic analyses (Poaceae) based on plastid and nuclear sequences in Paspalum L. reveal new diploid species and and structural data. American Journal of Botany accessions. Ciência Rural 38: 1292-1299. 97: 1732-1748. Prat, H. 1936. La systématique des Graminées. Annales Santos, A.M.P.V. & Boechat, S.C. 1989. Flora Ilustrada des Sciences Naturelles Botanique 10: 165-258. do Rio Grande do Sul XX. Gramineae ‒ tribo Quarín, C.L.; Valls, J.F.M. & Urbani, M.H. 1997. Danthonieae. Boletim do Instituto de Biociências Cytological and reproductive behavior of Paspalum 344: 1-57. atratum, a promising forage grass for the Tropics. Santos-Gonçalves, A.P. 2005. Estudos taxonômicos Vol. 31. Tropical Grasslands, St. Lucia. Pp. 114-116. e morfoanatômicos em Colanthelia (Poaceae: Raddi, G. 1823. Agrostografia brasiliensis. Attir. Accad. Bambusoideae: Bambuseae). Tese de Doutorado. Luchese Sci, Lett. Ed Artii 2: 329-383 (Reimpresso Universidade Estadual de Campinas, Campinas. 161p. como Agrostografia brasiliensis. Pp. 1-58, pl. 1. Tip. Santos-Gonçalves, A.P.; Filgueiras, T.S. & Clark, L.G. G. Nerici, Lucca, 1823). 2011. Atractantha shepherdiana, a new species of Reeder, J.R. 1957. The embryo in grass systematics. woody bamboo from Brazil. Systematic Botany American Journal of Botany 44: 756-769. 36: 310-313. Renvoize, S.A. 1984. The grasses of Bahia. Royal Schmidt, R. & Longhi-Wagner, H.M. 2009. A tribo Botanic Gardens, Kew. Pp 1-300. Bambuseae (Poaceae, Bambusoideae) no Rio Renvoize, S.A. 1988. Hatschbach’s Paraná grasses. Grande do Sul, Brasil. Revista Brasileira de Royal Botanic Gardens, Kew. Pp 1-76. Biociências 7: 71-128. Rezende, A.V.; Walter, B.M.T.; Fagg, C.W.; Felfili, J.M.; Sede, S.M.; Morrone, O; Giussani, L.M. & Zuloaga, Silva Júnior, M.C.; Nogueira, P.E.; Mendonça, R.C. F.O. 2008. Phylogenetic studies in the Paniceae & Filgueiras, T.S. 2008. Flora vascular do bioma (Poaceae): a realignment of section Lorea of cerrado. In: Sano, S.M.; Almeida, S.P. & Ribeiro, Panicum. Systematic Botany 33: 284-300. J.F. (eds.). Cerrado: ecologia e flora. Embrapa, Sendulsky, T. 1995. Merostachys multiramea (Poaceae: Brasília. Pp. 1028-1059. Bambusoideae: Bambuseae) and similar species Rocha, A.E.S.; Bastos, M.N.C. & Secco, R.S. 2001. from Brazil. Novon 5: 76-96. O gênero Panicum L. (Poaceae) na Restinga da Sendulsky, T. 1997. Twelve new species of Merostachys Praia da Princesa, APA de Algodoal/Maiandeua, (Poaceae: Bambusoideae: Bambuseae) from Brazil. Maracanã, Pará, Brasil. Boletim do Museu Paraense Emílio Goeldi 17: 297-314. Novon 7: 285-307. Rocha, A.E.S. & Secco, R.S. 2004. Contribuição à Sendulsky, T. & Soderstrom, T.R. 1984. Revision of taxonomia de Axonopus P. Beauv. (Poaceae) no the South American genus Otachyrium (Poaceae: Estado do Pará, Brasil. Acta Botanica Brasilica Panicoideae). Smithsonian Contributions to Botany 18: 295-304. 57: 1-24. Rua, G.H.; Speranza, P.R; Vaio, M. & Arakaki, M. 2010. Smith, L.B.: Wasshausen, D.C. & Klein, R.M. 1981- A phylogenetic analysis of the genus Paspalum 1982. Gramíneas. In: Reitz, R. (ed.). Flora ilustrada (Poaceae) based on cpDNA and morphology. Plant catarinense. Herbário Barbosa Rodrigues, Itajaí. Systematics and Evolution 288: 227-243. 1407p. Sajo, M.G.; Longhi-Wagner, H.M. & Rudall, P.J. 2007. Soderstrom, T.R.; Judziewicz, E.J. & Clark, L.G. 1988. Floral development and embryology in the early- Distribution patterns of neotropical bamboos. divergent grass Pharus. International Journal of Proceedings of a workshop on neotropical Plant Sciences 1688: 181-191. distribution patterns. Academia Brasileira de Sajo, G.M.; Longhi-Wagner, H.M. & Rudall, P. Ciências, Rio de Janeiro. Pp. 121-157. 2008. Reproductive morphology of the early- Soreng, R.J.; Davidse, G.; Peterson, P.M..; Zuloaga, F.O.; divergent grass Streptochaeta. Plant Systematics Judziewicz, E.J.; Filgueiras, T.S. & Morrone, O. 2011 and Evolution 275: 245-255. onwards. A world-wide phylogenetic classification Sánchez-Ken, J.G.; Clark, L.G.; Kellogg, E.A. & Kay, of Poaceae (Gramineae). Available in
Rodriguésia 63(1): 089-100. 2012 100 Longhi-Wagner, H.M.
base em ITS (rDNAnu). In: Congresso Brasileiro Welker, C.A.D. & Longhi-Wagner, H.M. 2007. A família de Melhoramento de Plantas, 3. Anais. Embrapa Poaceae no Morro Santana, Rio Grande do Sul, Trigo, Sociedade Brasileira de Melhoramento de Brasil. Revista Brasileira de Biociências 5: 53-92 Plantas, Passo Fundo. Pp. 1-7. Zanin, A. & Longhi-Wagner, H.M. 2003. Taxonomic Stebbins, G.L. & Crampton, B. 1961. A suggested novelties in Andropogon (Poaceae- Andropogoneae) revision of the grass genera of temperate North for Brazil. Novon 13: 368-375. America. Recent Advances in Botany 1: 133-144. Zanin, A. & Longhi-Wagner, H.M. 2006. Sinopse do Valls, J.F.M. 1980. Gramíneas nativas e sua importância gênero Andropogon L. (Poaceae – Andropogoneae) forrageira: situação do estudo no Brasil. EMBRAPA, no Brasil. Revista Brasileira de Botânica 29: 289-299. Plantas forrageiras, Brasília. Pp. 7-23. Zanin, A.; Mujica-Salles, J. & Longhi-Wagner, H.M. Valls, J.F.M.; Longhi-Wagner, H.M. & Boldrini, I.I. 1992. A tribo Stipeae no Rio Grande do Sul e Brasil. 2001. Axonopus. In: Wanderley, M.G.L.; Shepherd, Boletim do Instituto de Biociências 51: 1-174. G.J. & Giulietti, A.M. (eds.). Flora fanerogâmica Zuloaga, F.O.; Giussiani, L.M. & Morrone, O. 2007. do estado de São Paulo. Vol. 1. Hucitec, São Paulo. Hopia, a new monotypic genus segregated from Pp. 129-141. Panicum (Poaceae). Taxon 56: 145-156. Viana, P.L. 2010. O gênero Aulonemia Goudot (Poaceae: Zuloaga, F.O. & Morrone, O. 2005. Revisión de las Bambusoideae) no Brasil. Tese de Doutorado. especies de Paspalum para America del Sur Austral. Universidade Federal de Minas Gerais, Belo Missouri Botanical Garden, Saint Louis. 297p. Horizonte. 272p. Zuloaga, F.O. & Morrone, O. 2009. Phylogenetic Watson, L. & Dallwitz, M.F. 1992. The grass genera of studies in the Paniceae (Poaceae – Panicoideae): the world. Cambridge University Press, Cambridge. Ocellochloa, a new genus from the new world. 1081p. Systematic Botany 34: 684-692.
Artigo recebido em 06/05/2011. Aceito para publicação em 07/10/2011.
Rodriguésia 63(1): 089-100. 2012