How Rich is the Flora of Brazilian Cerrados? Author(s): A. A. J. F. Castro, F. R. Martins, J. Y. Tamashiro and G. J. Shepherd Source: Annals of the Missouri Botanical Garden, Vol. 86, No. 1 (Winter, 1999), pp. 192-224 Published by: Missouri Botanical Garden Press Stable URL: http://www.jstor.org/stable/2666220 Accessed: 18-06-2015 18:22 UTC Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at http://www.jstor.org/page/ info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Missouri Botanical Garden Press is collaborating with JSTOR to digitize, preserve and extend access to Annals of the Missouri Botanical Garden. http://www.jstor.org This content downloaded from 143.106.108.174 on Thu, 18 Jun 2015 18:22:09 UTC All use subject to JSTOR Terms and Conditions HOW RICH IS THE FLORA A. A. J. F. Castro,2 F. R. Martins,*3J. E OF BRAZILIAN CERRADOS?1 Tamashiro,3and G. J. Shepherd3 ABSTRACT An attempt is made to summarize what is known about the richness of the total terrestrialangiosperm flora of the "cerrados" (as a complex of formations)in Brazil, based on published surveys and species lists. A "refined" list of arboreal and shrubby species was compiled froma total of 145 individual lists from78 localities, taking into account synonymyand recent taxonomic changes. The refinedlist had 1709 references to taxonomic entities at the species level (973 identified with confidence and 31 with aff or cf.), 572 references to generic entities (363 genera identifiedwith confidence), and 210 references to the familylevel (88 families identifiedwith confidence). There are many unidentified arboreal and shrubby taxa at the specific, generic, and familylevels, indicating that a considerable amount of taxonomic research remains to be done on the cerrado flora, and that this flora may be much richer than is generally assumed. Depending on the assumptions made, these data suggest a total of around 1000 to 2000 arboreal and shrubby species and 2000 to 5000 herbaceous ones, yielding estimates for the total cerrado flora (terrestrialangiosperms) rangingfrom 3000 to 7000 species. These limits, especially the upper one, are dubious, but give an idea of the magnitude of the angiosperm flora in the Brazilian cerrados. Surveys of cerrados are very unevenly distributed,and studies of relatively unknown sites may reveal much more diversitythan that presently known. On readingaccounts of floristicstudies on cer- component have been rare in Brazil (Mantovani & rados in Brazil, one rapidlycomes to realize that Martins, 1993). Comparing the non-woody compo- the majorityof authors,either implicitly or explic- nent in differentlocalities in Brazil, Mantovani itly,consider the cerrado flora to be well knownand (1983) found a local richness that varied between to have low richness.For example,Rizzini (1963, 165 species in the Serra Dourada (state of Goias) 1971) estimatedaround 600 species and a little and 640 in the municipality of Lagoa Santa (state over200 generafor the whole cerrado arboreal and of Minas Gerais). In an area of 343.42 ha of a cer- shrubbyflora, but Heringeret al. (1977) cited 193 rado in the Reserva Biologica de Moji GuaGu (state arborealand shrubbyspecies and confirmedless of Sao Paulo), Mantovani and Martins (1993) found than 150 genera.Even recentstudies (e.g., Leitao 403 species of non-woody angiosperms. The her- Filho, 1992; Ratteret al., 1997) have estimatedthe baceous-subshrubby angiosperm flora of the cer- numberof arboreal-shrubbyspecies forthe cerra- rados thereforeappears to be richer than the ar- dos as being around800. Eiten (1990) has been boreal-shrubby flora, but its richness varies with one of the few authorsto suggestthat the thick- physiognomy(Mantovani, 1987). stemmedarboreal-shrubby flora contains more than It can also be argued that the maximum physi- 1000 species and that the denser physiognomies ognomic and floristic expression, together with may reach more than 150 arborealand shrubby maximum spatial continuity,should occur in the species per hectare.Castro (1994; see Ratteret al., "nuclear" (Labouriau, 1966), "central" (Rizzini, 1997) made an extensivesurvey of the literature in 1963), or "core" (Eiten, 1972; Ferri, 1977a) areas. orderto gathersupport for the idea thatthe arboreal An implication of this reasoning is that marginal and shrubbyflora of the Brazilian cerrados is much and disjunct areas (Ratter et al., 1988a) should richerthan previously assumed. have a relatively impoverished flora in comparison It could be arguedthat as the cerradosconsist to the nuclear area, although they may be supple- of physiognomiesthat are predominantlygrass- mented by floristicelements fromthe surrounding lands, the greatestfloristic richness should be en- vegetation formations (Eiten, 1972; Fernandes & counteredin thenon-woody (herbaceous-subshrub- Bezerra, 1990; Rizzini, 1963). These elements, by) componentof the vegetation.Surveys of this which occur preferentiallyin other formationsand I Researchwas developedin the Cursode Pos-Graduaca-oem BiologiaVegetal, Instituto de Biologia,Universidade Estadual de Campinas,Brazil. We thankEsmeralda Zancheta Borges for preparation of the map in Figure1. 2 Departamentode Biologia,Centro de Ciencias da Natureza,Universidade Federal do Piauf,Campus da Ininga, Teresina64049-550, PI, Brazil. 3Departamento de Botanica,Instituto de Biologia,Universidade Estadual de Campinas,Caixa Postal6109, Campinas 13083-970, SP, Brazil. * To whomcorrespondence should be sent. ANN. MISSOURI BOT. GARD. 86: 192-224. 1999. This content downloaded from 143.106.108.174 on Thu, 18 Jun 2015 18:22:09 UTC All use subject to JSTOR Terms and Conditions Volume 86, Number1 Castro et al. 193 1999 Flora of BrazilianCerrados sporadicallyin cerrados,were called "accessory" lectionsin the field.Which areas have been well species or elementsby Rizzini(1963). Species that collected? To what extenthave collectionsbeen occur exclusivelyin cerradoformations, or showa widelyspread or have theyconcentrated in certain markedpreference for cerrados, were termed "char- areas? In whichregions are collectionssparse or acteristic"species ("especies peculiares ou pro- nonexistent?These representpriority areas forfu- prias") by Rizzini (1963). He consideredthat only turework and shouldbe clearlyidentified. thewoody species occurringin the "cerraddo"(for- Answersto these questionsare essentialfor any est physiognomy)could be designatedcharacteris- attemptto establishplans forconservation and fur- tic species,that is, essentiallythe arboreal-shrubby ther investigation of the cerradovegetation, tasks species. However,because ofa largenumber of ac- whichare, sadly,increasingly urgent given the rap- cessoryspecies (sensuRizzini, 1963), marginaland id destructionand exploitationof this vegetation disjunctcerrado areas generallyshow considerable typein Brazil today. floristicrichness. In the presentstudy we triedto summarizewhat Althoughthis mightsuggest that cerrados in is knownfrom floristic studies in the cerradosof marginalor disjunctareas should graduallygrade Brazil,at least forthe arboreal-shrubbycomponent intoother formations, they are usuallyfairly easily of the vegetation,and to providesome indications distinguishedfrom other formations in the neigh- of wherefurther work might most usefully be in- borhoodby theirphysiognomy and floristiccom- vested to improveour knowledgefor conservation position,often with a numberof commonwell- and rationalsustainable exploitation. known species. The fact that a species occurs, grows,and reproducessuccessfully in an area MATERIAL AND METHODS showsthat it is adapted to the local conditions.It This studywas based on materialpublished and is possible to construct a continuum of species theses defendedup to 1992, supplementedby 12 rangingfrom those restrictedto verylocal areas unpublishedfield surveys from the statesof Piaui withina singlevegetation type to thosethat are very and Sao Paulo (Castroet al., in press).A literature widespreadand occur in several differentforma- surveyuncovered 135 publicationsand thesesthat tions.The distinctionbetween "characteristic" and includedfloristic surveys of cerrados.Of these,92 "accessory" species thereforeseems ratherartificial wereselected for the presentstudy. It is likelythat and debatable, lacking in ecological significance. otherstudies exist, but theywere not localized or The spatial and temporal abundance of the different could notbe obtained. species also cannot be ignored. While it might be An initial surveywas based on Garcia et al. argued that a species typical of other formations (1981), Huber (1974), Lemos (1976), Pinto(1979), and occurring only sporadically in cerrados should and Silva (1982), togetherwith publications by Ei- be considered an accessory species (e.g., Ficus ten (1972), Ferri(1963, 1971, 1973, 1977b, 1979), spp.), it is possible to find examples of species that Goodland (1979), Labouriau (1966), Marchetti occur with very low abundance but only in cerrado (1988), and Marchettiand Machado (1982). A sec- areas (such as Eugenia aurata), along with numer- ond surveywas based on thecitations in thesepub- ous intermediate situations (such as Copaifera licationsand on thesesdefended, as