Diversity of Cyperaceae in Brazil

2009·5.5 MB·English

by  Marccus Alves

Diversityof CyperaceaeinBrazil
Marcais Alves1, Ana Claudia Araújo2, Ana Paula Praia3, Fabio Vitta4, Sonia Hefler\
Rafael Trevisan6, André dos Santos Bragança Gil7, Shirley Martins' & Wayt Thomas®
Abstract
(Diversity ofCyperaceae in Brazil) The purpose ofthis catalogue was tocombine the available datafrom
publications,theses,databases,andherbariumspecimensfromaround 120Herbaria,andcolectionssampled
inBrazilianvegetationduringthelast 15yearstoproducethemostcompletelist,aspossibleas,ofCyperaceae
We
species for Brazil. cataloguedca. 1,700 names for678 species in42 genera occurring in Brazil. These
valuesrepresentca. 15percentofthespeciesand40percentofthegenerafoundintheworld. Bothsubfamilies
ofCyperaceaearefoundinBrazilwithCyperoideaebeingthemostdiverseatbothgenericandspecificleveis.
Although low'er species were recorded forthe tribes Cryptangieae, Sclerieae, and Trilepideae, these tribes
representamuchhigherpercentageoftheworld'stotaisforgeneraandspecies.Themostdiversegeneraare
Rhynchospora(157spp.),Cyperus(101 spp.),Scleria(82spp.)andEleocharis(69spp.).Fifteengenerahave
onespeciesinBrazil,althoughfiveofthemaremonospecific.Themostspecies-richregionsinBrazilarethe
NorthandSoutheast.TherearenogeneraendemictoBrazil.Thereare,however,around200endemicspecies,
ofw-hich40areinthegenusRhynchospora.Taxonomicandnomenclaturalproblemsfoundarepointedunder
the species. For each catalogued species, the principal synonyms, bibliographic references, distribution
withinBraziFsfiveregions,vegetationtype,andcitationofselectedmaterialexaminedareprovided.
Keywords:monocotyledons,conservation,Cyperales,neotropies,taxonomy.
Resumo
(DiversidadedeCyperaceaenoBrasil) Estaobrafoielaboradaapartirdareuniãodeinformaçõesoriundasde
publicações,teses,bancosdedadosevisitasrealizadasacercade 120herbários,alémdeamostrascoletadas
nos últimos anos em diversos biomas no Brasil. Estão aqui catalogados cerca de 1.700 nomes para as 678
espéciesregistradasdeCyperaceae no Brasiledistribuídasem42gêneros. Estes valoresrepresentamcerca
de 15%dototaldeespéciese40%dototaldegênerosdafamíliaocorrentesnomundo. Asduassubfamílias
de Cyperaceaeestãorepresentadas no Brasil, sendoqueCyperoideaeéamaisdiversaemtermosgenéricos
eespecíficos. Noentanto, quandoconsiderado em termoscomparativoscom a floraciperológicamundial,
Cryptangieae, Sclerieae eTrilepideae são as tribos com maiorriqueza de gêneros e espécies no Brasil. Os
gêneroscommaiorriquezadeespéciesno BrasilsãoRhynchosporaVahl(157 spp.), CyperusL. (101 spp.),
ScleriaBcrg.(82espécies)eEleocharisR. Br.(69spp.).Quinzegênerosestãorepresentadosporapenasuma
espécie no Brasil, sendo cinco deles monoespecíficos. As regiões Norte e Sudeste despontam como as de
maiorriquezagenéricaeespecífica.NãoforamconstatadosgênerosendêmicosdoBrasil.Foramcatalogadas
cercade200espéciesendêmicasdopaís,dasquais40pertencemaRhynchospora.Osproblemastaxonômicos
e nomenclaturais encontrados estão indicados para cada espécie. Para cada espécie catalogada são
apresentados os principais sinônimos, referências bibliográficas, distribuição geográfica nas cincoregiões
político-geográficasdopaís,materialexaminadodereferênciaecomentáriosgeraisquandopertinentes.
Palavras-chave:monocotiledôneas,conservação,Cyperales,neotrópicos,taxonomia.
Artigorecebidoem 11/2008.Aceitoparapublicaçãoem04/2009.
'UniversidadeFederaldePernambuco,Lab.Morfo-taxonomiaVegetal,Av.MoraesRegos.n.,CDU,Recife,PE,50670-
901,[email protected],www.ufpe.br/taxonomia
•Royal BotanicGardens,Kew,The Herbarium, Richmond, Surrey,TVV93AE. U.K.
UniversidadeFederaldeSergipe,Depto.Biologia,Lab.SistemáticaVegetal,SãoCristóvão,SE,49000-000,Brazil.
"UniversidadeFederaldosValesdoJequitinhonhaeMucuri.Depto.CiênciasBiológicas,Diamantina,MG,Brazil.
5UniversidadeFederaldeRioGrande,Av.Itáliasn,km8,CampusCarreiros,RS,96201-900,Brazil.
'PhDstudent, Pós-Graduaçãoem Botânica, Universidade Federal do RioGrandedoSul, Av. BentoGonçalves9.500,
CampusdoVale.PortoAlegre-RS.90501-970.Brazil.
'PhDstudent,Pós-GraduaçãoemBiologiaVegetal,UniversidadeEstadualdeCampinas/UNICAMP,InstitutodeBiologia.
BarãoGeraldo,Campinas,SP, 13083-970,Brazil.
PhDstudent.Pós-GraduaçãoemBiologiaVegetal,UniversidadeEstadualJúliodeMesquita,Unesp CampusRioClaro
RioClaro.SP, 13083-970,Brazil.
"TheNew YorkBotanicalGarden,NewYork, 10458-5126,USA.
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Introduction (Alves et al. 2007, 2008a, b). The main goal
Cyperaceae has around 5,000 species, of this Catalogue is to combine the available
well distributed and with highest diversity in complementarydatafrom publications,theses,
ÁfricaandtheNeotropics(Goetghebeur 1998; databases,andherbariumspecimenstoproduce
Govaerts et al. 2007). Many species are the most complete specimen-backed list of
known from savanna marshes and other species ofCyperaceae as possible for Brazil.
wetlandvegetation,butmanyothersare found
in forests and drierhabitats (Souza & Lorenzi MaterialsandMethods
2006; Govaertsetal. 2007;Guaglianoneetal. This catalogue is a compilation of
2008).Bruhl(1995)recognizedtwosubfamilies, information collected by the authors over the
and Goetghebeur (1998) recognised four last 15 years. In most cases, the taxa reported
subfamilies. The major clades on these are herearebasedonspecimenscollectedinBrazil
partially supported in most recent molecular with vouchersdepositedin Brazilian herbaria.
phylogenies(e.g.,Simpsonetal. 2007).Simpson In a few cases, taxa are included which were
etal. (2008),however,supporttherecognition reported in theliteratureasoccurring in Brazil
of only two subfamilies: Mapanioideae and butforwhichspecimenshavenotbeen located.
Cyperoideae(=Cyperoideae+Sclerioideae+ In addition, selected taxa reported in the
Caricoideae). literatureasoccurringoutsideofBrazil, but in
In Brazil, Cyperaceae are very well the border regions around the country were
represented. Nees (1842) published the first alsoincluded.
complete study of Cyperaceae of Brazil in Brazil is divided in Five geographical
Martius’s masterpiece - Flora Brasiliensis, regions: North (N), Northeast (NE), Central
andreportedmorethan 300names. Sincethen, West(CO), Southeast(S) and South (S). Each
however, much has changed. During the last oftheseregioncomprisesdifferentvegetation
few years, there has been considerable types. The North region (including the States
research on the taxonomy of Cyperaceae in ofAcre, Amapá, Amazonas, Pará, Rondônia,
Brazil (Araújo 2001; Alves 2003; Gil 2004; Roraima,andTocantins)isthelargestareaand
Prata 2004; Trevisan 2005, 2009; Vitta 2005; essentially covers the Amazon Basin - it is
Hefler2007). This activity has also produced mostly covered by Amazonian forest but has
useful checklists and floristicsstudies (Muniz also areas of savanna vegetation. The
&
1987,2001;Muniz Shepher 1987;Lucenocí Northeast (including the States of Alagoas,
al. 1997;Martinsetal. 1998;Prata2002;Thomas Bahia,Ceará,Maranhão,Paraíba,Pernambuco,
et al. 2006; Alves 2006; Alves et al. 2007; Piauí, Rio Grande do Norte, and Sergipe) is
Araújoetal. 2007;Giletal. 2007;Trevisan et the driest area in the country and includes the
al. 2007; Ferreira& Egger2008;Guaglianone “caatinga” and parts of the “cerrado” and
et al. 2008; Silveira & Longhi-Wagner 2008; Atlantic Forest. The Central West region
Thomas&Alves,2008;Alves& Martins2009; (including the States of Goiás, Mato Grosso,
Silvaetal.2(X)9),highlightingnewspeciesand andMatoGrossodoSul)hastheworId’slargest
new recordsforBrazil (Koyama 1972a; Kral & fresh-waterwetland,the Pantanal, alargearea
Thomas 1988; Simpson 1989a, 1995; Luceno of “cerrado” vegetation. and a large area to
& Alves 1997;Alves etal. 2002; Moore etal. the north covered by Amazonian forest. The
2002; Araújo et al. 2003, 2004, 2008; Rocha Southeast(includingtheStatesofEspíritoSanto,
& Luceno 2002). Minas Gerais. Rio de Janeiro, and São Paulo)
As an important contribution to the is the most urbanized area in the country and
knowledgeoftheBrazilianfloraandthediversity includes a variety of ecosystems including
ofCyperaceaeoftheworld,theCatalogueproposed AtlanticForestand“cerrado”.Finally,theSouth
here was First presented as a series of drafts (includingtheStatesofParaná, SantaCatarina,
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DiVersityofCyperaceaeinBrazil 773
andRioGrandedoSul)comprisesthesmallest thesesanddissertations).Theabbreviationsof
areacomposed ofsubtropical vegetation. classic works follows those proposed inTL-2
Up to five vouchers are cited for the by Stafleu & Cowan (1976-1988), while
geographic regions of Brazil, and they are in authors of species follow www.ipni.org. For
alphabetical order of collectors. Herbarium each species, a list of selected bibliographic
acronyms follow Thiers (continuously referencesispresented in whichthe taxon has
updated).Thefollowingherbariawerevisited been described, illustrated, orat leastcited.
by the authors and most of them are Foreach referenced species, notes on its
represented by at least one cited specimen: geographic distribution, habitats, taxonomic
AAU,ALCB,ASE, B, BHCB, BLA, BM, BP, delimitation,infraspecifictaxa,andnomenclatural
BOTU,C,CAY,CEN,CESJ,CEPEC,CEUL, questions were addedwheneverpertinent.An
CGMS, CH, CPAP, CTES, CVRD. DAV, E, indication of the endemic status in Brazil is
EAC, EAN, ESA, ESAL,F, FLOR, FUEL, G providedunderthecommentsoneachspecies.
GENT,GH,GUA, HAMAB,HAS, HB, HBR, For some species, however, the data are
HC, HCF, HEPH, HMS, HRB, HRCB, HRJ,
insufficienttosupportsuchconclusions.
HST, HTO, HUEFS, HUEM, IAC, IAN,
IBGE, ICN. INPA, IPA, JPB, K, L, LE. LIL, Results and Discussion
LPB, M. MA, MAC, MBM, MBML, MEXU,
MGMHN,MICH,MO,MPUC,MVFA,NLH, After analyzing approximately 40,000
specimens, we have catalogued 678 species
NY, P. PACA, PEL, PEUFR, PMSP, R, RB,
RFA, RFFP, RUSU, S, SI, SP, SPF, TAES, ofCyperaceaeoccuiringin Brazil(Appendix).
TCD, TEPB, TEX, U, UB, UC, UEC, UFG These are grouped into 42 genera and are
UFP, UFRN, UFRR. UPCB. UPS, US, VEN, distilledfromca. 1,700namespreviouslycited
VIC, VIES, WAG W, WU, Z, and ZT. Other for Brazil. .Using Goetghebeur (1998) and
herbaria (ECOSUR, HPZ, HRR, HUEPG, Govaerts etal. (2007), thesefiguresrepresent
SJPR. and UESC) listed at www.ufrgs.br/ almost 20% of the species and 40% of the
taxonomia were also visitedby the authors. genera in the family. Both two subfamilies
The most used synonyms are included (Simpson etal. 2007, 2008 or all four. ifit is
under the currently accepted name. They are based in Goetghebeur 1998) are represented
also listed alphabetically with a reference to in Brazil with Cyperoideae the most diverse
the accepted name in this article. bothatthespecificandgenericleveis. However,
Specieswhichoccurinborderingregions whenconsideredrelativetothediversityofthe
ofadjacentcountries butforwhich records of world’s Cyperaceae, Brazil is richest in
collections from Brazil have not been found, Cryptangieae,Sclerieae,andTrilepideaeatboth
are included here marked with “(?)” in the the generic and specific leveis.
geographic region of possible occurrence. Data presented in this work corroborate
Thus, the authors accept the taxa and think it in part the estimation of Luceno & Alves
likely those species occur in Brazil. For each (1997) for Brazil. They estimated that there
ofthese taxa predicted to occur in Brazil, the were 44 genera and 500-600 species of
literature citation noting its presence in an Cyperaceae in Brazil. Severalgeneraaccepted
adjacent area is given. by Goetghebeur (1998) and Luceno & Alves
The protologue ofeach taxon, when not (1997)aresynonymizedhereorrecircumscribed.
accesseddirectlybytheauthors,wasobtained Micropapyrus and Syntrinema are included
from the information available on the in Rhynclwspora. The monospecific genus
W3Tropicos (www.tropicos.mobot.org) and Websteria has been submerged within
IPNI (www.ipni.org) websites and other Eleocharis (Roalson & Hinchliff 2007). The
taxonomic works (including Govaerts et al. circumscriptionofLagenocarpusfollowsthat
2007;Guaglianoneetal. 2008,andunpublished proposed byVitta(2005)withthe recognition
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andseparationofCryptangiumwhich,in tum, the exception ofthree species, Pleurostachys
includes the genus Didymiandrum. is entirely restricted to the Atlantic forest.
The most species-rich genera in Brazil Several genera are disjunct between the
are Rhynchospora (157 spp.), Cyperus (101 Amazonian andAtlanticCoastal forests.These
spp.), Scleria (82 spp.), Eleocharis (69 spp.), including Becquerelia, Calyptrocarya,
Bulbostylis (50 spp.), Carex (30 spp.), Diplacrum, Hypolytrum and Pleurostachys.
Hypolytrum (28 spp.), Pleurostachys (21 A remarkable number (around 50% of
spp.), Lagenocarpus and Pycreus (16 spp. species) of Cyperaceae are found in moist
each), Fimbristylis (15 spp. each), and forests of both the Amazon Basin and the
Mapania (12 spp. each). Fifteen genera are Coastal forests.Generadiverseinthesehabitats
represented by a single species in Brazil. Of include PleurostachysandHypolytrum. Some
these, Diplasia, Egleria, Exochogyne, generasuchasPleurostachysandHypolytrum,
Koyamaea and Remirea are monospecific. havedisjunctionbetween theAmazonandthe
There are no genera endemic to Brazil. Atlantic forests. A disjunction between the
On theotherhand, 203 areendemic speciesto NeblinaMassifandtheCoastal areaoftheState
Brazil, of which 40 are belonging to ofBahia, is reported here forthe first time for
Rhynchospora, 28 to Cyperus, 25 to Scleria, Oreobolus goeppingeri Suess.
22 to Bulbostylis, 19 to Pleurostachys, 16 to The diversity ofCyperaceae on BraziFs
Eleocharis and 1 1 to Hypolytrum. oceanic islands is relatively low. Bulbostylis
Severalgeneraaredistinguishedfortheir nesiotis is endemic to the Ilhas de Trindade
high percentage of endemic species, with at (state of Espírito Santo) and Cyperus
Ieast 50 percent of their species endemic atlanticus is endemic to oceanic islands
to Brazil. These include the following: includingtheFernandodeNoronhaarchipelago
Abildgaardia (25%), Cryptangium (87%), (state of Pernambuco).
Machaerina (100%), Pleurostachys (90%) Several genera and many species are
and Trilepis (75%). The endemic species of exclusively associated with aquatic and
these genera, with the exception of wetland habitats. Among the genera are
Abildgaardia, are specific to the Atlantic Bolboschoenus, Egleria and Oxycaryum.
Coastal forest or the rupestrian fields. Eleocharis and Fuirena are aquatic or
Ofthefivegeographical regionsofBrazil wetland genera with the largest number of
used here, the richest in number ofgenera is speciesin Brazil.
theNorth with34genera. Forspeciesrichness, Several species (around 110 spp.) are
the mostdiverseregionsaretheSoutheastand widely distributed in all five geographical
Northwitharound350speciespresentineach. regions of Brazil, and they are in generally
This similarity in numbers probably does not weedy and invasive plants. Cyperus, with 25
reflect the actual diversity of the family in spp., Rhynchospora, with 23 spp., Eleocharis
Brazil, because the North is much larger and with14 spp, and Scleria with 1 1 spp. have the
much lesswell-collected. higest number ofwidespread species.
Some generaare apparently restricted to Although Cyperaceae are often thought
a single vegetation type. Mapania, for of as weedy and invasive plants, there are a
example, is restricted to the Amazon region. large number ofspecies that are specific to a
Bisboecklera, Cephalocarpus, Diplasia, and narrowrangeofhabitatsand,therefore,appear
Everardia are endemic to theAmazon forest. to be excellent bioindicators of healthy
Androtrichum and Remirea are restricted to ecosystems. Also, many sedges are used by
Coastaldunes.Machaerinaisfoundexclusively local people in handicrafts (Simpson & Inglis
in clearings at high elevations in the Coastal 2001), asthebasis forperfumes(i.e., Cyperus
mountains of the South and Southeast. With articulatus, Zoghbi et al. 2006).
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DiversityofCyperaceaeinBrazil 775
Manyspeciesareverypoorlyrepresented represent almost 20% ofthe species and40%
in Brazilian herbaria. Greatereffort is needed ofthegeneraaroundtheworld.Whencompared
incollecting thesespeciestobeterunderstand to the diversity of the world’s Cyperaceae,
their distribution and ecology. These poorly Brazil is richest in the genera Cryptangieae,
knownspeciesincludethoseknovvnfromasingle Sclerieae, andTrilepideae at both the generic
collection (e.g., Rhynchospora paranaensis) andspecificleveis.Thereare203endemicspecies
or those collected in areas difficult to reach ofCyperaceaeinBrazil,occurringspecifically
(e.g., Koyamaea neblinensis). at the Atlantic Coastal forest and rupestrian
Some species (e.g., Bulbostylisbrevifolia) fields. Some genera are apparently restricted to
have not been collected for many years. asinglevegetationtypeinBrazilasBisboecklera
Although this may be the result of lack of totheAmazon forest;AndrotrichumtoCoastal
collecting effort, many species are rare, have dunes and Machaerina to Coastal mountains
a limited distribution, and have not been ofthe South and Southeast regions.
encountered for a long time. These data indicate a high diversity of
Even though these are all criteria for Cyperaceae in Brazil, with many species
includingaspeciesontheIUCNRedList,few occurringatimpactedhabitats.Thisisaserious
Cyperaceae were included in the most recent problem considering that a large number of
version of the list of endangered species of Cyperaceae species are useful as potential
the Brazilian flora (Biodiversitas 2008). Tvvo bioindicators of healthy ecosystems and as
species Çryptangiwn hurnite andHypolytrum handicraft material, andalsopresentpotential
(
lucennoi) are listed as Endangered and 17 as pharmacologicalactivity.
Vulnerable: Bulbostylis distichoides Lye, B. This becomes more serious when we
latifolia Kral&M.Strong,B. nesiotis(Hemsl.) perceive that many Cyperaceae species are
C.B. Clarke, B. smithii Barros, Çryptangiwn very poorly represented in Brazilian herbaria
clausenii C.B.Clarke, C. coma/um Boeck., while someothershave notbeencollected for
Cyperus atlanticus Hemsl., Hypolytrum many years. Although this may be the result
amorimiiM.Alves&WAV.Thomas,H.bahiense of lack of collecting effort, as many species
M. Alves & W.W. Thomas, H. lucennoi M. are rareorhavealimiteddistribution,thiscan
Alves& W.W.Thomas,H. paraenseM.Alves be an indicative of a potential extinction. In
& W.W. Thomas, Lagenocarpus bracteosus this sense, although there are clear enough
C.B. Clarke, Pleurostachys angustifolia criteriaforincludingmoreCyperaceaespecies
Boeck., RhyncosporaparanaensisA.C.Araújo on the IUCN Red List, until now few were
& W.W. Thomas, R. wanningii Boeck., and included in the most recent version ofthe list
ofendangered species ofthe Brazilian flora.
Trilepis tenuis Vitta.
Recently, Alves et al. (2009) increased This shortage oflisted species is due not
the numberofrare species ofCyperaceae for only to the lack of collections, but also to
taxonomic uncertainty as to what species
Brazilto47. Nevertheless,thenumberremains
shouldberecognized. Inthissenseweconclude
low. This shortage oflisted species is due not
that: i) more extensive good collections of
onlytothelackofcollections,buttotaxonomic
Cyperaceae are necessary to betterunderstand
uncertainty as to what species should be
recognized. It is our hope that this catalogue their distribution and ecology in Brazil; ii)
scientific investigations in different botanical
will help stimulate better recognition of
areas should be produced in order to clear
threatened species of Cyperaceae.
taxonomicalproblems.Wehopethatthecatalogue
presented here could stimulate these
CONCLUSIONS
investigations, improvingthebrazilianstrategies
Wecatalogued42generaand678species toprotectendangeredandunknownCyperaceae
of Cyperaceae occurring in Brazil. These species and respective lineages.
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Acknowledgements In:Barbosa,L.&Santos,N.(orgs.).Abotânica
The authors are thankfull to all herbaria noBrasil: pesquisa,ensinoepolíticaspúblicas
curators, whose open their collection to be ambientais.SBB,SãoPaulo.Pp.286-290.
Alves,M.;Araújo,A.C.;Prata,A.P.;Vitta,F.;Hefler,
studiedduringtheselongyears,ourInstitutions &
S.;Trevisan,R.;Gil,A.B.;Martins.S. Thomas,
and supervisors, for their strong support and
WAV.2008aAvançosedesafiosnoconhecimento
knowledge during graduation and post-
da diversidade de Cyperaceae no Brasil. In:
graduationprograms,manybotanists fortheir Loiola,M.etal. (orgs.).Atualidades,desafios
collections, which helped to provide material e perspectivas da Botânica no Brasil. SBB,
forthiswork, R.GuaglianoneandN. Brummit Natal.Pp. 121-122.
for the criticai review. Also the authors Alves,M.;Araújo.A.C.;Prata,A.P.;Vitta,F.;Hefler,
recognizethe importantfinancial supportfroni S.;Trevisan,R.;Gil,A.B.;Martins,S.&Thomas,
NationalCounsilofTechnologicalandScientific WAV. 2008b. Documenting Cyperaceae in
Development (CNPq), Coordenação de diversetropicalcountries:theexempleofBrazil.
Monocts IV, Abstracts. Natural History
AperfeiçoamentodePessoaldeNível Superior
(CAPES), Mellon Foundation, IAPT, Red MuseumofDenmark,Copenhagen. Pp.5.
Alves,M.&Martins,S.2009.Cyperaceae.In:Alves,
Latinoamericana de Botânica and Margaret
&
Mee Trust Foundation, fellowship programs. M.;Araújo,M.F.;Maciel,J. Martins,S.(eds.).
FloradeMirandiba.APNE,Recife.Pp. 137-145.
Special acknowledgments to The New York Alves,M.&Thomas,WAV.2002.Fournewspeciesof
Botanical GardenandRoyal BotanicGardens,
HypolytrumRich.(Cyperaceae)frornBraziland
Kew (Herbaria and Library) for providing CostaRica.FeddesRepertorium 1 13:261-270.
opportunities to visit them, improving our Alves,M.;Thomas,WAV.&Wanderley,M.G.2002.
knowledge during the contact with their NewspeciesofHypolytrumRich.(Cyperaceae)
researchers and all staff in many ways. The fromtheNeotropics.Brittonia54: 124-135.
First and second authors also are thankfull to Alves,R.&Martins,L.2004.Restabelecimentode
Taxonomy Program/CNPq to pemit a special espécies endêmicas da ilha de Trindade.
taxonomical study on Cyperaceae at Rio de Albertoa3:45-52.
JaneiroBotanicGardenandFederalUniversity Angely,A. 1965.FloraanalíticadoParaná. Phyton,
ofRioGrandedoSul University,respectively. SãoPauIo.728p.
Araújo, A.C. 2001. Revisão taxonômica de
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