V CONFERENCIAINTERNACIONALde Leguminosas (VILC)

Editado porRenée H. Fortunato

V CONFERENCIAINTERNACIONALde Leguminosas (VILC)

Editado porRenée H. Fortunato

Primera edición: noviembre 2015

La ilustración de tapa es reproducción de A. Burkart “LAS LEGUMINOSAS ARGENTINAS SILVESTRES Y CULTIVADAS”, Segunda Edición. ACME AGENCY, Soc. de Resp. Ltda., Buenos Aires, 1952.

Diseño de tapa: Andrea Hamid/Andy Sfeir Producción, Coordinación y Diseño: Andrea Hamid/Andy Sfeir

© 2015 - Ediciones CICCUS Medrano 288 (C1179AAD) (54-11) 4981.6318 / 4958.0991 ciccus@ciccus.org.ar www.ciccus.org.ar

Hecho el depósito que marca la ley 11.723Prohibida la reproducción total o parcial del contenido de este libro en cualquier tipo de soporte o formato sin la autorización previa del editor.

Impreso en ArgentinaPrinted in Argentina

Ediciones CICCUS ha sido merecedora del reconoci-miento Embajada de Paz, en el marco del Proyec-to-Campaña “Despertando

Conciencia de Paz”, auspiciado por la Or-ganización de las Naciones Unidas para la Ciencia y la Cultura (UNESCO).

Fortunato, Renée V Conferencia Internacional de Leguminosas, VILC / Renée Fortunato et al. ; editado por Renée Fortunato. - 1a ed . - Ciudad Autónoma de Buenos Aires : Fundación CICCUS, 2015. 224 p. ; 26 x 18 cm.

ISBN 978-987-693-131-1

1. Biología. 2. Botánica. I. Fortunato, Renée, ed. II. Título. CDD 570

Index

Arturo Erhardo Burkart (1906-1975) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

V Conferencia Internacional De Leguminosas (Vilc)Renée H . Fortunato . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

Agradecimientos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

Comité Organizador . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

Diversity, endemisms and the history of southern brazilian legumesJoão R . V . Iganci, Silvia T . S . Miotto, Toby R . Pennington & Roseli Lopes Da Costa Bortoluzzi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

Progress with leguminosae in WageningenL .J .g . Van Der Maesen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

Estado actual de los tratamientos de la familia leguminosae para cuatro proyectos de floras con alta diversidad en México IIMaría de Lourdes Rico Arce . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

A survey of the Mimosoideae of Serra do Cipó, Minas Gerais, BrazilLeonardo Maurici Borges & José Rubens Pirani . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41

Diversity and geographical distribution patterns of the Genus Mimosa (Imosoideae) in the United States, Mexico, and Central AmericaRosaura grether, Sara L . Camargo-Ricalde, Angélica Martínez-Bernal, Susana Montaño-Arias & Ma . Eugenia Fraile . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57

A floristic analysis of papilionoid legumes of Southern AfricaA . N . Moteetee & B-E . van Wyk . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69

A review of the taxonomy and biogeography of the Genus Crotalaria (leguminosae)Marianne M . le Roux, Ben-Erik van Wyk & andAnnah N . Moteetee . . . . . . . . . . . . . . . . . . . 89

Variations on a theme: novel rhizobial strains of burkholderia, methylobacterium and microvirga demonstrate the diversity of root nodule bacteria/legume symbiosesJulie K . Ardley, giovanni garau, Ron J . Yates, Matt Parker, graham W . O’Hara, Wayne G. Reeve, Sofie De Meyer, Rob Walker, Michael J. Dilworth, Anne Willems, Elizabeth Watkin, Sunil Ratnayake & John G. Howieson . . . . . . . . . . . . . . . . . . . . . . . . . . . 121

Legume nodulation and rhizobial diversity in ChinaTao Wang, Xin Hua Sui, Wen Feng Chen, Chang Fu Tian & Wen Xin Chen . . . . . . . . . . . . 137

Roots of leguminosae: structure and functionRosana Malpassi, Teresa Kraus, Luciana Bianco, Mónica grosso, Sara Basconsuelo & Júlio Ramos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169

Characterization of the varieties of Acacia Caven (Mimosoideae) by means of molecular and morphological markersCarolina L . Pometti, Juan C . Vilardi, Ana M . Cialdella & Beatriz O . Saidman . . . . . . . . . . 187

Análisis molecular del género Acacia (Leguminosae, Mimosoideae), con énfasis en el grupo mirmecófiloSandra Luz gómez-Acevedo, Lourdes Rico-Arce, Alfonso Delgado-Salinas, Susana Magallón y Luis Eguiarte-Fruns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 201

7

Arturo Erhardo Burkart (1906-1975)

Antonio KrApovicKAs †*

Visité el Instituto Darwinion por pri-mera vez en 1940 para identificar mis pri-meras colecciones, cuando era estudiante del segundo año de la Facultad de Agro-nomía y Veterinaria de Buenos Aires . Había aprobado el curso de Botánica que dictaba el Ing . Agr . Lorenzo Parodi, exce-lente profesor que nos entusiasmó y nos transmitió su gusto por las plantas .

En el Darwinion nos recibió el Ing. Burkart y a partir de ese momento su Ins-tituto pasó a ser nuestro segundo hogar .

Teníamos una mesa de trabajo, una lupa, libre acceso a la biblioteca y al her-bario y su disposición a atender nuestras dudas .

Arturo Erhardo Burkart nació en la ciudad de Buenos Aires el 25 de setiembre de 1906 . Sus padres el industrial Norberto Burkart y Olga Rebling . Como buen hijo de ale-manes cursó la escuela primaria en la Belgrano Schule, de la ciudad de Buenos Aires . En 1925 ingresó en la Fa-cultad de Agronomía y Veterinaria de la UBA, egresando, tras un brillante desempeño, en 1928 .

Acontecimientos 1930 Golpe militar 1932-35 Guerra, Chaco Paraguayo 1936 39 Guerra civil, España

En 1926, alumno del 2º año, publica su primer trabajo en la Revista del Centro de Estudiantes de su facultad, “Una forma nueva de Xanthium para la flora argentina”.

En 1928, el año de su graduación, aparece “Notas so-bre Leguminosas platenses” en Physis, con el que inicia su producción sobre la familia de su predilección: las Le-guminosas .

Recibió el Premio Mitre al mejor trabajo de un estu-diante en 1928, por su estudio “Las Leguminosas cultiva-das y adventicias en la República Argentina”, publicada al año siguiente .

Ese mismo año conoce a Angel L . Cabrera y junto con Román Pérez Moreau realizan excursiones botánicas, especialmente a remo por el Delta del río Paraná . El co-nocimiento adquirido lo indujo posteriormente a realizar la Flora Ilustrada de Entre Ríos .

En 1929 y 1930, gracias a una beca de la Universidad, viajó a Alemania donde estudió genética con Erwin Baur y con Curt Stern . Aprovechó su estadía en Berlín para estudiar en su Herbario material de Leguminosas y del

género de Compuestas Chaptalia.Su trabajo de tesis elaborado en Ale-

mania “Investigaciones genéticas sobre una nueva mutación en Drosophyla me-lanogaster determinante de excepcioness hereditarias” es el primer trabajo sobre el tema publicado en Argentina, en 1931 .

A su regreso de Alemania es desig-nado Ayudante de genética entre 1930 y 1937, bajo la dirección de S . Horovitz .

En el período 1931-1936 fué Jefe T .P . de Botánica en la cátedra de L .R . Parodi .

El 28 de diciembre de 1936, a los 30 años de edad el Ing . Burkart fué designado director del Darwinion, instituto de origen

privado que creó Cristóbal Hicken en 1911 . Hicken perte-necía a una familia pudiente y pudo atesorar una espléndi-da biblioteca y un herbario de más de 150 .000 ejemplares .

Los Hicken tuvieron una activa participación en la vida cultural de la Argentina . Su hermano Ricardo Hicken fué un experto en lunfardo y conocido autor de sainetes . Su primo Pablo C . Ducros Hicken un estudioso del cine argentino, su colección privada de películas es la base de la Cinemateca Argentina .

A los 33 años Burkart fué Profesor Titular de Forra-jicultura en la Facultad de Agronomía de La Plata, donde ejerció entre los años 1939 y 1961 . Allí realizo estudios sobre biología floral, genética y mejoramiento de la alfal-fa . Su descubrimiento de alfalfas resistenies sl “nematode del tallo”, sus métodos de selección y sus cálculos de fe-cundación cruzada, marcaron inportantísimos jalones en la fitotecnia de Medicago sativa .

Se ocupó también de otras forrajeras y del estudio de praderas .

Es en el Darwinion donde Burkart pudo desarrollar su vocación por la taxonomía vegetal . Allí produjo su gran contribución sobre este tema . Sus trabajos tratan sobre diversas familias: las Compuestas, con su monografía so-bre Chaptalia, las Gramíneas, con su magnífico trabajo en la Flora Ilustrada de Entre Ríos y varias más . Pero son las Leguminosas la familia a la que dedicó su mayor em-peño desde el inicio de su carrera .

La mayor parte de sus trabajos se refieren a las Legu-minosas argentinas y sudamericanas . Publicó revisiones totales o parciales de Lathyrus, Caesalpinia, Centrose-ma, Hedisareas, Prosopis, Indigofera, Clitoria, Mimosa, Adesmia, Galactia, Vicia, Dioclea, Piptadenieas etc .

† Lamentamos su reciente fallecimiento durante la edición de este libro.

* instituto de Botánica del nordeste (iBonE), Av. sargento cabral 2131, 3400 corrientes, Argentina.

8

En 1943 expuso sus conocimientos sobre esta fami-lia en un libro: “Las Leguminosas Argentinas silvestres y cultivadas”, del cual apareció una segunda edición au-mentada en 1952 .

Publicó más de 170 trabajos, principalente de botá-nica y en especial sobre Leguminosas llegando a ser un especialista reconocido mundialmente .

En 1957 fué designado Profesor de Plantas Vas-culares, en la Facultad de Ciencias,de la Universidad de Buenos Aires . Fue un excelante profesor que jamás escatimó trabajo o tiempo en su labor docente . Dirigió tesis en la Facultad de Agronomía de La Plata y en la Facultad de Ciencias de Buenos Aires . Muchos jóve-nes se iniciaron con él en la investigación botánica, entre los cuales pueden recordarse al Ing .Agr . Osvaldo Boelcke . la Dra . Ada I . Pastore, la Lic . Nélida S . Tron-coso, la Prof . Nélida Bacigalupo, el Dr . Román Pé-rez Moreau (h .), la Dra . Ana María Ragonese, la Lic . Lidia D . Beavo, el Ing .Agr . Ramón Palacios, la Sra .

Juana S . de Lichtenstein, el Sr . Eduardo grondona y muchos más .

El 25 de abril de 1975 la muerte lo sorprendió a los 68 años de edad, en un momento cumbre de su carrera cientí-fica, siendo Director del Instituto de Botánica Darwinion, Profesor Emérito de la Universidad de Buenos Aires (1972) y cuando acababa de recibir el Premio Houssay dela Organización de los Estados Americanos en 1974, como reconocimiento a sus estudios agronómicos sobre la alfalfa .

Escribió casi toda su obra en castellano, que era el idioma de su país y el de sus alumnnos y discípulos . Su larga bibiografía apareció principalmente en Darwiniana y en algunas otras revistas argentinas, por lo cual resulta fácil de localizar y consultar

Fué un hombre responsable y disciplinado . Ejerció la dirección del Instituto Darwinion, hasta su muerte, con cor-dura, sensatez y respeto . Su instituto fué un oasis de tran-quilidad en una época de vertiginosos y drásticos cambios .

Acontecimientos durante la formación de A. Burkart

1925 regresa a Bélgica Lucien Hauman1926 fallece Carlos Spegazzini1929 fallece Moisés S .Bertoni1931 fallece Miguel Lillo1932 fallece Teodoro Stuckert1933 fallecen Cristóbal M .Hicken y Augusto Scala

1934 fallecen Angel gallardo y Carlos Thays1936 fallece Cornelius Osten1937 fallecen Ladislao Holmberg, Emilio Hassler y Pedro Jörgensen Hansen se suicida Horacio Quiroga1938 se suicidan Leopoldo Lugones y Alfonsina Storni

Arturo Erhardo Burkart (1906-1975)

1926 su primer trabajo, sobre Xanthium1928 Ingeniero Agrónomo “Notas sobre Leguminosas platenses”1929-1930 Beca en Alemania1930-1937 Ayudante de genética 1931-1936 Jefe T .P . de Botánica1931 Tesis sobre Drosophila1936 Director del Instituto Darwinion

1939-1961 Profesor de Forrajicultura, Fc .Agronomía, La Plata1943 “Las Leguminosas Argentinas silvestre y cultivadas”1952 “ “ 2º edic .1957 Profesor de Botánica, Fc . de Ciencias, B .Aires1972 Profesor Emérito

Generaciones de botánicos argentinos

Parodi 1895 Castellanos 1896 Ruiz Leal 1898

Pérez Moreau 1905 Burkart 1906173 Cabrera 1908

Ragonese 1909 Meyer 1910 Descole 1910 O’Donell 1912 Lourteig 1913 Covas 1915

9

V CONFERENCIA INTERNACIONAL DE LEGUMINOSAS (VILC)

rEnéE H. FortunAto

La V Conferencia Internacional de Leguminosas (VILC) se llevó a cabo en el Centro de Convenciones de la Universidad Católica Argentina (UCA), Ciudad de Buenos Aires, Argentina, entre los días 8 y 14 de Agosto de 2010; 11 años después de la IV ILC en Canberra, Australia, 2000 . El propósito principal fue reunir a los especialistas a nivel global en la Familia Leguminosae y generar el ámbito de difusión de los avances relevantes al inicio del Siglo XXI .

Ante la trayectoria y reconocimiento nacional e in-ternacional en el estudio y comprensión de las Legumi-nosas Argentinas y Sudamericanas, la V Conferencia fue en Honor a la Memoria del Prof . Ing . Agr . Arturo Burkart (1906-1975), quien fuera Director del Instituto de Botáni-ca Darwinion (IBODA): Academia Nacional de Ciencias Exactas y Naturales-CONICET (1936-1975) .

Sobre su semblanza y trayectoria se realizó la Confe-rencia Magistral de apertura dictada por el senior inves-tigador Prof . Antonio Krapovickas, Presidente Honorario del Comité Organizador (VCIL) .

La V Conferencia Internacional tuvo un total de 392 participantes de 23 países (Argentina, Bolivia, Brasil, Ca-nadá, Chile, China, Colombia, Corea, Costa Rica, Egipto, España, Estados Unidos de América, Francia, India, In-glaterra, Italia, Japón, México, Polonia, Sudáfrica, Tai-wán, Uruguay y Venezuela), con la participación sobre-saliente de representantes latinoamericanos . El programa incluyó 12 simposios y cuatro reuniones satélites .

En este contexto la Conferencia Magistral de clausura la efectuó la Dra . Lidia Poggio: Parámetros Citológicos: Aportes a Modelos de Evolución Cromosómica en el Contexto Filogenético de Leguminosae, en coautoría con Alexandra M . gottlieb & Renée H . Fortunato .

La reunión permitió el intercambio y vínculos entre especialistas de diferentes continentes y la difusión de la información actual desarrollada en los distintos campos científicos y de importancia económica, incluso para gru-pos sociales emergentes, especialmente ante los urgentes requerimientos de desarrollo de generar estrategias de Conservación para un real uso sustentable .

Sobre esta base se diagramaron los Simposios:

• Filogenia y nueva clasificación de Leguminosas (Caesalpinioideae, Mimosoideae y Papilionoideae)

• Evolución del Bioma/Biogeografía • Acacia (género conflictivo en su reconocimiento ta-

xonómico) • Flora • Morfo-anatomía y Ecofisiología • Interacción Planta-Suelo• Recursos Genéticos • Recursos Genéticos no Tradicionales • Soja en el Siglo XXI • VII Encuentro Internacional de Especialistas de Ara-

chis (maní)

Además se incluyeron las reuniones satélites sobre Etno- y Paleobotánica, Especialistas de la Región Cha-queña y de Madera, Sistema Nacional de Datos Biológi-cos, Ministerio de Ciencia, Tecnología e Innovación Pro-ductiva (MINCyT) y Productos Forestales no Madereros .

Asimismo se han expuesto 12 sesiones de paneles de diferentes tópicos de interés: Filogenia y Nueva Cla-sificación de Leguminosas, Fitogeografía, Desarrollo y Estructura, Biología Reproductiva, Biología de Semilla y Plántula, Anatomía, Anatomía de Madera, Citología, Fitoquímica, Palinología, Polinización, Paleobotánica, genética, Sistemas de Cruzamiento, Mejoramiento, Es-pecies-Poblaciones - Híbridos - Poliploides, Botánica Económica, Conservación, Recursos genéticos, Floras, Sistemática y Taxonomía, Interrelación Planta - Animal - Microorganismos, Evolución Funcional en Nódulos Ra-diculares, Interrelación Planta - Suelo, Ecología, Ecolo-gía Molecular .

Como antecedente se señala que es la primera en Hispanoamérica, la cual para contar con una mayor par-ticipación de especialistas de la región, algo no ocurrido en las cuatro anteriores, se realizó por primera vez tra-ducción simultánea: Inglés-Español, Español-Inglés, In-glés-Portugués, Portugués-Inglés .

Se quiere resaltar que la Conferencia ha sido decla-rada de “INTERÉS DEL SENADO DE LA NACIÓN ARGENTINA” (exp. 5 nº3414/09).

La actual edición contiene los aportes de las contri-buciones enviadas por los Coordinadores de Simposios y que fueron aceptadas por el Comité Organizador del VCIL .

10

AGRADECIMIENTOS

Se destaca en los agradecimientos que la traducción simultánea: Inglés-Español, Español-Inglés, Inglés-Por-tugués, Portugués-Inglés, ha sido posible gracias al apor-te brindado por The Andrew W. Mellon Foundation, con la colaboración de Argenbio, asociación que además ha facilitado la ejecución del Simposio Soja en el Siglo XXI . Es de señalar que también se reconoce a The Andrew W. Mellon Foundation por haber financiado en parte, la par-ticipación de expositores de Simposios de representantes de países subdesarrollados .

gracias a la predisposición de los organizadores del VII Encuentro Internacional de Especialistas de Arachis (maní) ha sido posible aunar su reunión en conjunto con la V Conferencia Internacional de Leguminosas, ante esto se agradece a los organizadores por posibilitar la ejecu-ción como parte de este evento .

Asimismo, en nombre del Comité Organizador se destaca el apoyo brindado por el Director y al personal del Instituto Darwinion, especialmente en el permiso y la asistencia efectivizada durante la visita de los participan-tes del VCIL al herbario del Instituto . Asimismo, se dis-tingue los auspicios recibidos a nivel Nacional de INTA, Agencia Nacional de Promoción Científica y Tecnológi-ca, CONICET, Argenbio, y de la Sociedad Argentina de Botánica, Sociedad que sin su sostén y trabajo hubiese sido imposible cristalizar la inscripción y depósitos de los subsidios recibidos. A nivel internacional de The Andrew W . Mellon Foundation, Missouri Botanical garden y Ro-yal Botanic Gardens, Kew. Sin el aporte de todos ellos no habría sido posible plasmar esta Conferencia Interna-cional en Argentina . Asimismo se hace mención de los avales recibidos de: Ministerio de Ciencia, Tecnología e

Innovación Productiva, Academia Nacional de Agrono-mía y Veterinaria, Academia Nacional Ciencias Exactas, Físicas y Naturales, Instituto Nacional de Semillas, de Universidades Nacionales (UBA, UNC, UNNE) y pri-vadas (Universidad de Morón y Católica de Argentina) . Asimismo se agradece a todos los participantes que sin su presencia este evento no hubiese sido posible ejecutar .

Las Instituciones editoriales son: Ministerio de Cien-cia, Tecnología e Innovación Productiva (MINCyT), Mi-nisterio de Agricultura, ganadería y Pesca, (MAgyP), Instituto Nacional de Tecnología Agropecuaria (INTA), Consejo Nacional de Ciencia y Tecnología (CONICET), Sociedad Argentina de Botánica (SAB) y Missouri Bo-tanical garden, USA . Estas memorias además están dis-ponibles vía electrónica en la página web de INTA con enlace en MAgyP, CONICET, MINCyT, SAB y MBg .

Como reseña final se cuenta que “La idea” de nueva-mente congregar a los especialistas se generó posterior a la reunión de Leguminosas realiza en Botany 2008, (Con-greso Anual organizado por American Society of Plant Taxonomy y Botanical Society of America), Vancouver, Canadá . Al regreso del evento fue propuesto a los investi-gadores de Argentina el lanzamiento de la V Conferencia; idea que tuvo aceptación a pesar del compromiso, si bien no pertenecíamos a una misma Institución, nos uni-dos como si así lo fuéramos, y este fue el resultado del trabajo en conjunto en distintos puntos del país . Espera-mos que lo diagramado haya sido de interés y brinde los nuevos avances alcanzados en el inicio del Siglo XXI .

Comité Organizador

11

COMITÉ ORGANIZADOR:

Presidente-Honorario: Antonio Krapovickas † (ibone@agr .unne .edu .ar)

Presidente: Renée H . Fortunato (fortunato .renee@inta .gob .ar)

Secretarios: María Pía Mom (mpmom@bg .fcen .uba .ar) y María Magdalena Brizuela (brizuela@bg .fcen .uba .ar)

Relaciones Institucionales: Ramón A . Palacios (palacios@bg .fcen .uba .ar)

Tesorero: Beatriz O . Saidman (saidman@bg .fcen .uba .ar)

Protesorero: Cecilia Bessega (cecib@ege .fcen .uba .ar)

Antonio Krapovickas † 1921-2015 .

41

A SURVEY OF THE MIMOSOIDEAE OF SERRA DO CIPÓ, MINAS GERAIS, BRAZIL

LEonArdo MAurici BorgEs1 & José ruBEns pirAni1

AbstrAct

A survey of Leguminosae Mimosoideae as part of the project “Flora da Serra do Cipó” is presented. The area is located at the Espinhaço Range of mountains in Minas Gerais State, Southeastern Brazil. 61 species belonging to 14 genera were found, four species are endemic to Serra do Cipó, and 12 to the Espinhaço Range, mostly restricted to its Southern portion. The completion of the Mimosoideae treatment almost tripled the number of species previously accounted for the area. Our data highlight the need to develop more field collections and floras, either in well explored regions as the Serra do Cipó, and, especially, still poor studied areas, in order to improve the knowledge about the Brazilian flora, one of the richest of the world.

Keywords: Leguminosae, Mimosoideae, Brazil, Serra do Cipó, floristics.

AcKnowledgments. The authors would like to acknowledge the curators of the herbaria BHCB, MBM, ESA, RB, SP, SPF and UEC (acronyms following Thiers 2009) for loan of specimens, Klei R. Souza for the illustrations, C. Siniscalchi, F.U. Yamamoto, D. Zappi and D.W. Milliken for photographs of specimens in the field, C. Siniscalchi for help with the English version, C. Siniscalchi & P. Fiaschi for comments on the manuscript. L.M. Borges was supported by FAPESP (proc. 2005/54316-2, proc. 2006/58598-5). J.R. Pirani is supported by CNPq.

1 Departamento de Botânica, Instituto de Biociências, Universidade de São Paulo, Rua do Matão, 277. São Paulo. SP. Brazil. 05508-900. aquitemcaqui@gmail.com

Introduction

leguminoSae in brazilIn Brazil, Leguminosae is represented by 221 gen-

era and 2802 species (Lima et al . 2014), being the rich-est family in species and the second more diverse in endemic species (Forzza et al . 2010a) . Its occurrence is widespread among all Brazilian phytogeographic domains - Amazonian, Atlantic Forest, Caatinga, Cer-rado, Pampas and Pantanal (IBgE 2010) . The highest diversity is found in the Amazonian Domain, where 166 genera and 1128 species have been recorded so far (Lima et al . 2014) . Besides, the family is the rich-est one either in the Amazonian and Caatinga domains, the second richest in the Atlantic Forest, Cerrado and Pantanal domains, and the third in the Pampas Domain (Forzza et al . 2010a) .

Mimosoideae, with 78 genera and 3270 species (Lewis et. al 2005), presents a cosmopolitan distribution, with centres of diversity in tropical areas (Schrire et al. 2005) . It is distinguished from the other subfamilies by the general presence of bipinnate leaves, valvate aestiva-tion of the corolla, attractive stamens and seeds with a U-shapped pleurogram .

Thirty-two genera and 802 species of Mimosoideae are found in Brazil, and more than 45% of its species (361) belong to Mimosa L . (Lima et al . 2014, Dutra & Morim 2014), a genus highly diversified at the Cerrado Domain (Barneby 1991, Simon & Proença 2000) . The second largest genus of Mimosoideae of Brazil is Inga, with 131 species distributed mainly through the Atlan-tic Forest and Amazonian domains (garcia & Fernandes 2014; Pennington 1997) .

In between two Leguminosae-rich domains, the Cer-rado and Atlantic Forest, the Espinhaço Range extends as a mountainous region whose flora presents high levels of endemism (giulietti & Pirani 1988) .

eSpinhaço range anD the Serra Do cipó

The Espinhaço Range mountain chain occupies an approximately 1200 km long and 50 to 100 km wide zone in the Brazilian Central Plateau, with elevations up to 800 m above sea level (Almeida-Abreu & Renger 2002; Moreira 1965) . It spreads in S-N direction, from Serra do Ouro Branco, near Belo Horizonte in Minas gerais state, to the northern limits of Bahia, Pernambuco and Piauí states (Almeida-Abreu & Renger 2002 (Fig . 1 .) . It is formed mainly by Quartzitic rocks from the Espin-haço Supergroup, dating back to the Proterozoic (Almei-

42

Fig. 1. Map of elevations of the southern Portion of the Espinhaço Range with localities indicated. Modified from rapini 2000.

43

habit vegetational formations

taxon ar ab sb CAM CER SDTF SF DIST

Abarema langsdorfii x x

A. obovata x x

Albizia polycephala x x

Anadenanthera colubrina x x x x

Calliandra asplenioides x x

C. brevipes x x

C. dysantha x x x

C. fasciculata x x

C. linearis x x

C. parvifolia x x

Enterolobium gummiferum x x

Inga cylindrica x x

I. edulis x x

I. sessilis x x

I. striata x x

I. subnuda x x

I. vera x x

I. vulpina x x x

Leucochloron incuriale x x

Mimosa adenotricha x x x

M. arenosa x x

M. barretoi x

M. bimucronata x x

M. bombycina x x

M. calocephala x x

M. diplotricha x x

M. dolens x x x

M. filipes x x x

M. foliolosa x x

M. gemmulata x x

M. gracilis x

M. hirsutissima x x x

M. macedoana x x

M. maguirei x x

M. nuda x x

M. paucifolia x x

M. pigra x x

M. pithecolobioides x x

M. radula x x x

M. setosa x x

M. somnians x x

M. sordida x x

M. stylosa x x

44

above 900 m ASL on rocky areas, the “campos rupestres” dominate as an herbaceous-arbustive vegetation with ele-vated endemism degree (giulietti & Pirani 1988, giulietti et al. 1987). At lower altitudes and related to substrate nature and conditions, other vegetation types occur, such as cerrado, seasonally dry tropical forest, semideciduous forest, riverine forest, and swamps (locally called “bre-jos”). Cerrado vegetation occurs in most of the low el-evation areas as well as in the dryer western slopes and sometimes as scattered patches in the highlands . Season-ally dry tropical forests occur exclusively in areas where limestone outcrops are frequent . Semideciduous forests are mainly located at the wet eastern slopes of the range, but are also present as “capões de mata”, small forest patches found in areas with suitable pedological condi-tions within the “campos rupestres” (Meguro et al. 1996).

In order to preserve the Serra do Cipó and its peculiar flora and fauna, both of which were important subject of many scientific studies throughout the years, the National Park of Serra do Cipó and its associated encompassing area, the Environment Protection Area of Morro da Pe-dreira, were created in 1984, the first comprising an area of 31632 ha, and the latter 100107 ha .

da-Abreu 1995), with NNW-SSE trending and west-verg-ing thrusts (gontijo 1993) .

The Range can be subdivided into two regions sepa-rated by a gap in the continuity of elevations: one located in Minas gerais and the other in Bahia . This gap is re-puted to have acted as a barrier to migration of mountain plants across the divide, resulting in distinct patterns of plant species distribution found in several groups (Harley 1988) .

At the southern portion of the Espinhaço Range, a set of elevations known as Serra do Cipó (Fig. 1.), is delim-ited to the east by the Cipó River and its eastern tributar-ies, and to the north by the Paraúna River . The climate is characterized by annual temperature means ranging between 18 and 20ºC, with a maximum of 27.2º C and a minimum of 15 .4º C, and rainfall average of about 1400 mm per year .

The Serra do Cipó vegetation is very complex, clearly related to the large variations in relief, soil and micro-climate conditions, with a rich flora bearing an endemic contingent intermingled with several taxa from the two surrounding phytogeographic domains, the Cerrado to the west, and the Atlantic Forest to the east. At elevations

Table 1. species list for the Mimosoideae of serra do cipó and the respective habit and vegetational for-mation. ar - arboreous; ab - arbustive; sb - subarbustive; CAM - “campo rupestre”; CER - Cerrado; SDTF - Seasonally Dry Tropical Forest; SF - Semideciduous Forest; DIST - Disturbed area.

M. velloziana x x

M. xanthocentra x x

Piptadenia adiantoides x x

P. gonoacantha x x

P. macradenia x x

P. paniculata x x

Plathymenia reticulata x x

Pseudopiptadenia contorta x x

P. leptostachya x x

P. warmingii x x

Senegalia martiusiana x x

S. polyphylla x x

S. riparia x x

S. tenuifolia x x

Stryphnodendron adstringens x x

S. gracile x x

S. polyphyllum x x

Zygia latifolia x x

total 61 33 19 9 18 27 3 17 4

45

The Flora of Serra do Cipó project lead to the pub-lication of a preliminary check-list of 1520 species of vascular plants and bryophytes (giulietti et al . 1987) . At that time, nine genera with 23 species of Mimosoideae were recorded (a number updated to account for nomen-clatural changes). The publication of a series of floristic treatments of taxonomic groups at Serra do Cipó started in 1987 and to date 92 articles are available .

Based on the data on the Mimosoideae of Serra do Cipó (Borges & Pirani 2013), we provide here com-ments on genera, their respective list of species and also a comparison with the preliminary results of Giulietti et al. (1987), as well as relating to the results for the native Mimosoideae of Brazil extracted from Lima et al . (2014) .

Results

DiverSity of mimoSoiDeae at Serra Do cipóMimosoideae is represented at Serra do Cipó by 62

species distributed in 14 genera: Abarema (2 spp .), Al-bizia (1 sp .), Anadenanthera (1sp .), Calliandra (6 spp .), Enterolobium (1 sp .), Inga (7 spp .), Leucochloron (1 sp .), Mimosa (27 spp .), Piptadenia (4 spp .), Plathymenia (1 sp .), Pseudopiptadenia (3 spp .), Senegalia (4 spp .), Stryph- nodendron (3 spp .) and Zygia (1 sp.). Table 1 shows the spe-cies list and indicates habit and vegetation formations for each .

Abarema Pittier - The genus contains 49 species re-stricted to the Neotropical region (Barneby & grimes 1996, Iganci & Morim 2009, Lewis & Rico-Arce 2005). The reddish endocarp and the bicoloured seed are dis-tintive characteristics of the genus (Barneby & grimes 1996) . There are 24 species in Brazil, and the genus is distributed through the Amazon, Atlantic Forest and Cer-rado domains (Iganci & Morim 2014) . It is represented at Serra do Cipó by two species.

Abarema langsdorffii (Benth .) Barneby & J .W .grimes (Fig . 2 .A) occurs in tropical rain forest, cer-rado, “campo rupestres” and “restingas” of Bahia and of the southern and south regions of Brazil (Barneby & grimes 1996; Iganci & Morim 2012) . At Serra do Cipó it is found only in the semideciduous forest of the eastern slopes

Abarema villosa Iganci & M .P . Lima (Fig . 2 .B), as well as the former, was only collected from the western slopes of Serra do Cipó in semideciduous forest . It has a small distribution area comprising part of Espirito Santo, Minas gerais and Rio de Janeiro states (Iganci & Morim 2009) .

Albizia Durazz. is a pantropical genus with 120-140 species (Lewis & Rico-Arce 2005). Amongst other in-goide legumes, the diagnostic characters of Albizia in-clude sympoidal growth, lack or armament, and palmate or palmate-pinnate leaflets (Barneby & Grimes 1996). The genus is represented in the majority of the Brazilian states by ten species occurring in the Amazonian, Catin-ga, Cerrado and Atlantic Forest domains .

The occurrence of the genus in the cerrado of Serra do Cipó is limited to Albizia polycephala (Benth .) Killip ex Record (Fig 2.C), which occurs throughout Caatinga, Cerrado, semideciduous forest and seasonally dry tropi-cal forest of Ceará, Pernambuco, Paraíba, Minas gerais, goiás and Rio de Janeiro (Barneby & grimes 1996) .

Anadenanthera Speg. is a neotropical genus with two species (Altschul 1964) and is characterized by the glo-merulate flowers, follicular fruit that are slightly winged, and compressed seeds . The seeds of A. colubrina (Vell .) Brenan were traditionally used as an hallucinogen by

Fig. 2. A. Abarema langsdorffii. B. A. obovata. c. Albizia polycephala. d. Anadenanthera colubrina. E. Calliandra fasciculata. F. C. linearis g. Enterolobium gummiferum. H. Mimosa arenosa. i. M. calocepha-la. Scale bars: A, C, D, G, H, I - 5 cm; B, F - 5 mm; E - 10 cm. (A, B - L.M. Borges; C, G, H, I - D.C. Zappi; D - W. Miliken; F - F.U. Yamamoto).

46

South American Indians (Altschul 1972) . Both species are present in Brazil, distributed through Amazonian, Atlantic Forest, Caatinga and Cerrado domains (Morim 2014d) .

At Serra do Cipó, the genus is represented only by Anadenathera colubrina (Vell .) Brenan (Fig . 2 .D), a com-mon tree of different formations, from riverine forest to xeromorphic environments of South America . It is found in semideciduous forests and cerrado-”campo rupestre” transitions in the western slopes where this commonly vigorous tree reaches only 2--3 m tall .

Calliandra Benth. is a neotropical genus with ca. 135 species, distributed from southwest United States to Ar-gentina (Barneby 1998, Lewis & Rico-Arce 2005). The ge-nus can be distinguished from other genera of Mimosoide-ae with multiple stamens fused in a tube (tribe Ingeae) by the thick-margined, elastic dehiscent fruits and absence of extrafloral nectaries. In Brazil 74 species occur in the Am-azon, Atlantic Forest, Caatinga and Cerrado domains (Sou-za 2014), but the main centre of diversity and endemism of the genus is found in the mountains of Bahia (Queiroz 2009, Renvoize 1981) . Six species occur at Serra do Cipó .

Calliandra asplenioides (Nees) Renvoize is an endemic of the “campos rupestres” of the Espinhaço Range (Barneby 1998) and was collected in this vegeta-tion formation at Serra do Cipó .

Calliandra brevipes Benth., an species wide-ly used for gardening, is natural of Riverine Forests and rocky stream banks in Argentina, Paraguay, Uruguay and the south-east and south regions of Brazil (Barneby 1998). At Serra do Cipó it was collected only once, near a waterfall.

Calliandra dysantha Benth . is a polymorphic and widely distributed species, occurring in cerrados and “campos rupestres” from Piauí to Paraná, in Brazil, and in Paraguay (Barneby 1998) . At Serra do Cipó it occurs in cerrados and “campos rupestres”.

Calliandra fasciculata Benth . (Fig . 2 .E) is en-demic to stone outcrops and “campos rupestres” of the montaneous regions of central and northern Minas gerais (Barneby 1998), including Serra do Cipó, where it is very common in rocky stream banks .

Calliandra linearis Benth . (Fig . 2 .F) is endem-ic to the “campos rupestres”, where it is found at Serra do Cipó, and stone outcrops in the southern portion of the Espinhaço Range (Barneby 1998) .

Calliandra parvifolia (Hook . & Arn .) Speg . is distributed trough river banks, riverine forest borders, dunes and stone outcrops from the eastern Brazil, reach-ing Argentina, and disjunctly the Amazonian campina (Barneby 1998) . At Serra do Cipó it has only been col-lected in riverine forest .

Enterolobium Mart . -This genus consists of 11 spe-cies restricted to the Neotropics and occurs mainly in hu-mid forests (Lewis & Rico-Arce 2005; Mesquita 1990). The most important characteristic of the genus is the indehiscent, internally septated and recurved fruit (Que-iroz 2009) . In Brazil, there are nine species distributed through the Amazonian, Atlantic Forest, Caatinga and Cerrado domains (Mesquita 1990; Morim 2010a) .

A single species is found in the cerrados of Serra do Cipó: Enterolobium gummiferum Mart. (Fig. 2.G), which is very common and distributed throughout all of the Cer-rado Domain (Mesquita 1990) .

Inga Mill . -This genus has ca . 300 species restricted to the Neotropics, from Mexico to Uruguay (Pennington 1997). The genus attains its highest diversity in wet forest environments, being limited to riverine forests when found in other biomes. Inga is the only genus with pinnate leaves in Mimosoideae, where bipinnate leaves prevail (there are rare exceptions in other genera) . It is represented in Bra-zil by 131 species, mainly at the Amazonian and Atlantic Forest domains, also in Caatinga and Cerrado but lacking in the Pampas Domain (garcia & Fernandes 2014) . Seven species are found at Serra do Cipó . Except for three taxa mentioned below, all Inga species inhabit the semidecidu-ous forest of the eastern slopes of Serra do Cipó .

Inga cyllindrica (Vell .) Mart . occurs from Cos-ta Rica to the southern region of Brazil, in tropical rain forest, semideciduous forest and cerrado (Pennington 1997) .

Inga edulis Mart . is a very common species in tropical South America, occurring in tropical and semide-ciduous forests from Venezuela to northern Argentina . It is know from the seasonally dry tropical forest of Serra do Cipó (Pennington 1997) .

Inga sessilis (Vell .) Mart . is restricted to the southern and south regions of Brazil, occurring in river-ine forest in Cerrado and in lowland and montane for-ests (Pennington 1997). At Serra do Cipó, it was found in riverine forest within the “campos rupestres” and the semideciduous forest of the eastern slopes of Serra do Cipó .

Inga striata Benth . occurs in tropical and semideciduous forests of the guyanas, Bolivia, Peru, Ec-uador, Colombia and the Amazonian and eastern Brazil (Pennington 1997) .

Inga subnuda Salzm . ex Benth . is restricted to tropical and semideciduous forests of several coast-al states of Brazil, from Paraíba to Santa Catarina, and Minas gerais (garcia 1998; Pennington 1997) .

Inga vera Willd . is one of the most common species in the genus and occurs throughout the Neo-

47

tropics (Pennington 1997) . At Serra do Cipó it is a com-mon species found in riverine forest and in humid areas of the semideciduous forest .

Inga vulpina Mart . ex Benth . extends from Bahia to Santa Catarina through the south-east region of Brazil, occurring in the montane forests of the Cerrado in transitional areas between cerrado and “campo rupestre” (garcia 1998; Pennington 1997) . Besides the semidecidu-ous forest of the eastern slopes of Serra do Cipó, it is also locally found in “campos rupestres”.

Leucochloron Barneby & J .W . grimes - A genus endemic to the Atlantic and Planaltine Brazil, with four to five species (Lewis & Rico-Arce 2005). A peculiar characteristic of this genus is the presence of perulate axillary resting buds, a feature only shared, within the Mimosoideae of Serra do Cipó, with Pseudopiptadenia warmingii (Benth.) G.P. Lewis & M.P. Lima which is a diplostemonous species . The genus occurs in the Atlan-tic Forest, Caatinga and Cerrado domains (Barneby & grimes 1996; Morim 2014b) .

Leucochloron incuriale (Vell .) Barneby & J .W . Grimes, which occurs in “capões de mata” and cerra-do-riverine forest transitions from the southern region of Brazil and eastern Paraná (Barneby & grimes 1996), is the single species of the genus found at Serra do Cipó in both of these habitats .

Mimosa L . is one of the largest genera of Mimosoideae with more than 500 species (Luckow 2005), is almost re-stricted to the Neotropics but with a few species also oc-curring in Africa and Asia (Barneby 1991) . It can be dis-tinguished from other genera of Mimoseae by the lack of extrafloral nectaries (with the exception of a few species) and the common presence of craspedium fruit . In Brazil 361 species of Mimosa are found distributed throughout the country and its six phytogeographic domains (Dutra & Morim 2014) . The genus is especially diverse in central Brazil, in the Cerrado and Caatinga domains, which together form one of the centers of endemism of the genus. Twen-ty-seven species of Mimosa are found at Serra do Cipó:

Mimosa adenotricha Benth . is restricted to cer-rados and “campos rupestres” of the Espinhaço Range (Barneby 1991) and was collected on these formations at Serra do Cipó .

Mimosa arenosa Poir . (Fig . 2 .H) is distributed from Mexico to Rio de Janeiro (Queiroz 2004) and in cer-rado and riverine forest of Serra do Cipó is represented by M . arenosa var . lysalgica Barnby, an endemic of the cerrado of the Espinhaço Range .

Mimosa barretoi Hoehne is endemic to the hu-mid areas of “campos rupestres” and riverine forest of

Serra do Cipó . Mimosa bimucronata (DC .) Kuntze is natu-

ral from humid soils and roadsides of the coastal states of Brasil, from Alagoas to Rio grande do Sul (Barneby 1991). It was found in disturbed areas by the roadside at Serra do Cipó .

Mimosa bombycina Barneby var bombycina is endemic to cerrado and “campo rupestre” of Serra do Cipó (Barneby 1991) .

Mimosa calocephala Mart . (Fig . 2 .I) is an en-demic of the southern portion of the Espinhaço Range and occurs in “campos rupestres” (Barneby 1991), the same habitat in which it was collected at Serra do Cipó.

Mimosa diplotricha C .Wright ex Sauvalle has a discontinuous, but wide distribution, from Mexico to Argentina (Barneby 1991) . It is common in the cerrado, but also in disturbed areas where it was collected at Serra do Cipó .

Mimosa dolens Vell . (Fig . 3 .A) is distributed through Argentina, Paraguay, Bolivia and the center-west and southeast regions of Brazil. Two varieties were found in the “campo rupestre” of Serra do Cipó: M. dolens var. acerba (Benth) Barneby and M . dolens var . dolens .

Mimosa filipes Mart . (Fig . 3 .B) occurs from Maranhão, Pernabuco and Bahia to the southern Espin-haço Range in cerrado, Caatinga and “campo rupestre” (Barneby 1991). At Serra do Cipó it was collected in cer-rado and “campo rupestre”.

Mimosa foliolosa Benth . var . pachycarpa (Fig . 3.C) is endemic to cerrado and “campo rupestre” of the southern Espinhaço Range (Barneby 1991) . It is found in cerrado, “campo rupestre”and riverine forest of Serra do Cipó .

Mimosa gemmulata Barneby (Fig . 3 .D) occurs in cerrado and “campo rupestre” of Bahia, Goiás, Mato grosso, Minas gerais, and disjunct in Venezuela (Barne-by 1991). It was collected in the cerrado of Serra do Cipó.

Mimosa gracilis Benth . var capillipes (Benth .) Barneby is distributed through cerrado, humid fields and disburbed environments of Missiones in Argentina and Minas gerais, São Paulo, Paraná, Santa Catarina and the center-west region of Brazil (Barneby 1991). It is found in the cerrado of Serra do Cipó .

Mimosa hirsutissima Mart . occurs from Vene-zuela to Paraguay, in cerrado, sandy fields and “campo rupestre” (Barneby 1991). At Serra do Cipó, two of its three varieties (M. hirsutissima var . hirsutissima and M. hirsutissima var. barbigera (Benth .) Barneby) are found in “campo rupestre”, sandy fields, cerrado and by the roadsides .

Mimosa macedoana Burkart (Fig . 3 .E) is an endemic of the “campo rupestre” and stone outcrops of

48

Serra do Cipó and gouveia (Diamantina Plateau) (Barne-by 1991) .

Mimosa maguirei Barneby is endemic to the southern part of the Espinhaço Range and occurs at “cam-po rupestre” of Serra do Cipó, the Diamantina Plateau (Barneby 1991) and Serra do Cabral. It was collected in “campo rupestre” and stone outcrops at Serra do Cipó.

Mimosa nuda Benth . var . glaberrima (Chodat & Hassler) Barneby occurs in cerrados and “campo rup-estre” of central Brazil, Minas Gerais, Paraná, São Pau-lo and Argentina (Barneby 1991) . At Serra do Cipó it is found in cerrado .

Mimosa paucifolia Benth . (Fig . 3 .F) occurs in sandy fields of Minas Gerais and São Paulo. At Serra do Cipó it was collected at “campo rupestre” areas.

Mimosa pigra L . (Fig . 3 .g) is distributed throughout all tropical America in stream margins and seasonally humid cerrado (Barneby 1991) . At Serra do Cipó it was found by the roadside near a stream within “campo rupestre”.

Mimosa pithecolobioides Benth . (Fig . 3 .H) oc-curs in cerrado and “campo rupestre” of the Espinhaço Range (Barneby 1991) . It is a very common species of cerrado–“campo rupestre” transition at the northwest slope of Serra do Cipó .

Mimosa radula Benth . var . calycina (Benth .) Barneby is distributed through cerrados in the southern Espinhaço Range, Minas gerais, São Paulo and goiás . It was collected in cerrado and “campo rupestre” at Serra do Cipó .

Mimosa setistipula Benth . occurs only on mountainous areas at the Minas gerais state . At Serra do Cipó, it was collected on “campo rupestre”.

Mimosa setosa Benth . var . paludosa (Benth .) Barneby (Fig . 3 .I) is found mostly in humid areas, such as riverine forest and swamps, but is also by roadsides in Ceará, Bahia, the southern Espinhaço Range, São Paulo and Goiás (Barneby 1991). At Serra do Cipó it was found in all those habitats . M. setosa aff . var . pseudomelas Barneby, which was previously thought to be restricted to São Paulo (Barneby 1991), was also recorded in the cerrado of the northwest slope of Serra do Cipó .

Mimosa somnians Humb . & Bonpl . ex Willd . var . somnians occurs throughout the Neotropics, in cerra-do, margins of riverine forest and as weeds in plantations (Barneby 1991). At Serra do Cipó it was collected in cer-rado and by the roadside within “campo rupestre” areas.

Mimosa sordida Benth . is endemic to the cerra-do at the southern portion of Serra do Cipó and Lagoa San-ta, a municipality located close to the Espinhaço Range .

Mimosa stylosa Barneby, known for Serra do Cipó by a single collection, is a rare endemic of the south-ern Espinhaço Range cerrado .

Mimosa velloziana Mart. is an weedy oppor-tunist species distributed in open fields of the Neotropics. At Serra do Cipó it was collected in disturbed areas by roadsides .

Mimosa xanthocentra Mart . var . xanthocentra is distributed from Maranhão to Santa Catarina, Brazilian Central Plateau, Colombia, Bolivia, Argentina and Vene-zuela, in open formations, cerrado and roadsides . At Serra do Cipó it was collected in cerrado.

Piptadenia Benth. is a genus with more than 20 spe-cies (Jobson & Luckow 2007) restricted to the tropical region of South America (Queiroz 2009) . With the seg-regation of Pityrocarpa (Benth .) Britton & Rose and eventually of Piptadenia viridiflora (Kunth) Benth ., Pip-tadenia possess, as distinguishing features from other Mimoseae, the presence of prickles, spikes organized in secondary inflorescences and the ovary exserted above the corolla (Jobson & Luckow 2007). The genus is rep-

Fig. 3. A. Mimosa dolens. B. M. filipes. c. M. foliolosa var. pachycarpa. d. M. gemmulata. E. M. macedoana. F. M. paucifolia. g. M. pigra. H. M. pithecolobioides. i. M. setosa var. paludosa. Scale bars: A, B, E, F, G, I - 2 cm; D, D, H - 5 cm. (A, C, G, I - L.M. Borges; B, H. D.C. Zappi; E - M.F. Santos; D, F - W. Miliken).

49

resented in Brazil by 21 species distributed throughout the Amazonian, Atlantic Forest, Caatinga and Cerrado domains (Morim 2014e) and four occur in Serra do Cipó .

Piptadenia adiantoides (Spreng .) J .F .Macbr . is distributed thoughout Caatinga, Cerrado and Atlantic Forest from Paraíba to São Paulo and also Pará (Morim 2014e) . At Serra do Cipó, it is common in riverine forest borders .

Piptadenia gonoacantha (Spreng .) J .F . Mac-br . (Fig . 4 .A) occurs in the coastal states of Brazil, from Paraíba to Rio grande do Sul, and in Minas gerais and Mato Grosso do Sul (Morim 2014e). It was collected in semideciduous forests of Serra do Cipó .

Piptadenia macradenia Benth . is restricted to the Brazilian states of goiás and Minas gerais (Morim 2014e), occurring mainly in areas of seasonally dry trop-ical forest. At Serra do Cipó, it is know only from this vegetation formation .

Piptadenia paniculata Benth . is a rare species

of the Atlantic Forests from Bahia, Paraná, Santa Catarina and the southern region of Brazil (Morim 2014e, Burkart 1979). At Serra do Cipó, it is know only from the semide-ciduous forest of the eastern slopes .

Plathymenia Benth . - An endemic genus of South America, is considered monotypic by Warwick & Lew-is 2003 . Distinctive characters include the articulated pedicels and a flattened legume with papery endocarp separating from epicarp and breaking up in one-seeded segments . Plathymenia reticulata Benth . (Fig . 4 .B-C) is distributed througout the Amazonian, Atlantic Forest, Caatinga and Cerrado domains in Brazil (Warwick & Lewis; 2003, Morim 2014c) and is also found in Serra do Cipó in cerrado and semideciduous forest .

Pseudopiptadenia Rauschert is a genus with 11 spe-cies restricted to South America . The main centre of diversity for the genus is the Brazilian Atlantic Forest, where seven species occur (Lewis & Lima 1991, Luck-ow 2005). It is differentiated from Piptadenia mainly by lomentiform fruit (Lewis & Lima 1991). In Brazil nine species are found distributed in the Amazonian, Atlantic Forest, Caatinga and Cerrado domains (Morim 2014f) . Three species are present at Serra do Cipó . Pseudopip-tadenia contorta (DC.) G.P. Lewis & M.P. Lima occurs in the Atlantic Forest and Caatinga of Paraíba, Bahia and in the southern region of Brazil (Lewis & Lima 1991). It was collected in the semideciduous forest of the eastern slopes of Serra do Cipó .

Pseudopiptadenia leptostachya (Benth .) Raus-chert (Fig . 4 .D) is restricted to forests on mountain slopes in Minas Gerais, Rio de Janeiro and São Paulo (Lewis & Lima 1991). It was collected in the semideciduous forest of the eastern slopes at Serra do Cipó .

Pseudopiptadenia warmingii (Benth .) g .P . Lewis & M.P. Lima is common in the Atlantic Forest of Bahia, Minas gerais, Rio de Janeiro and from São Paulo to Santa Catarina (Lewis & Lima 1991). At Serra do Cipó it is restricted to the seasonally dry tropical forest .

Senegalia Raf . is a genus of pantropical distribu-tion with ca. 200 species (Seigler et al. 2006; Rico-Arce 2007) . The genus Senegalia is roughly equivalent to Aca-cia subgen . Aculeiferum but Senegalia is adopted based on its status as a monophyletic group and the retypifica-tion of Acacia based on an Australian species (McNeill et al. 2006). It has polystemonous flowers with free or shortly connate stamens and prickles, but lacks stipular spines . It is represented in Brazil by 60 species and occurs in all phytogeographic domains (Morim & Barros 2014) . At Serra do Cipó there are four species of Senegalia .

Fig. 4. A. Piptadenia gonoacantha. B-C. Plathyme-nia reticulata. d. Pseudopiptadenia leptostachya. E. Senegalia polyphylla. F. Stryphnodendron adstrin-gens. g. S. gracile. H. S. polyphyllum. A, H - 5 mm; B - 2 cm; C - 50 cm; D, E, F, G - 1 cm. (A,B,D,F, H - L.M. Borges; C,D - M.F. Santos; G - F.U. Yama-moto).

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Senegalia martiusiana (Steud .) Seigler & Ebin-ger occurs in riverine forests and less often in caatingas of Bolivia, Colombia, French guyana, Paraguay and, in Bra-zil, from Bahia to Santa Catarina. It was collected at river-ine forests within “campos rupestres” of Serra do Cipó.

Senegalia polyphylla (DC .) Britton & Killip (Fig. 4.E) is a polymorphic and widespreaded species, occurring in many vegetational types from Mexico to Ar-gentina . At Serra do Cipó it is the most common species of the genus, where it can be found in semideciduous for-est and riverine forests within the Cerrado.

Senegalia riparia (Kunth) Britton & Rose is belongs to a widespread complex of species occurring from Mexico to Argentina in seasonally dry tropical for-est, Cerrado and Caatinga. It is known only from the cer-rados of Serra do Cipó and is very common along the western slopes of the range where the floristic influence of the Cerrado domain is stronger .

Senegalia tenuifolia (L .) Britton & Rose is dis-tributed from Mexico to Paraguay in many environments . At Serra do Cipó it was collected in cerrado.

Strhyphnodendron Mart. Is a genus with Neotropical distribution and 30-35 species (Luckow 2005; Scalon 2007). Two centres of diversity to the genus are recog-nized, one at Amazonia with ca. 14 species., and another at central and southeastern Brazil with ca. 13 species (Luckow 2005). It is represented in Brazil by 21 species occur in the Amazonian, Atlantic Forest, Caatinga and Cerrado domains (Scalon 2014) . In Serra do Cipó, three species are found:

Stryphnodendron adstringens (Mart .) Coville (Fig. 4.F), a typical species of the Cerrado domain, which occurs through all its limits . As expected, at Serra do Cipó, it was collected in cerrados.

Stryphnodendron gracile Heringer & Rizzini (Fig . 4 .g) is an endemic species of the “campos rupes-tres” from Serra do Cipó.

Stryphnodendron polyphyllum Mart . (Fig . 4 .H) occurs in the semideciduous forests and cerrados of Minas Gerais and at Serra do Cipó it is known only from the first formation, at the western slopes.

Zygia P. Browne – The genus consists of ca. 60 species (including Marmaroxylon Killip as a synonym) (Barneby & grimes 1997) exclusively distributed in the Neotropics. Cauliflory and an intrastaminal disc are dis-tinctive characters of the genus . Some of the species lack one of these features but never both (Barneby & grimes 1997). Nineteen species are known in Brazil occurring in Amazonian, Atlantic Forest, Caatinga and Cerrado do-mains (garcia et al . 2014) .

A single species is found in riverine forests of Cerra-do sites along the western slopes of Serra do Cipó: Zygia latifolia (L.) Fawc. & Rendle, which occurs in riverine forests from Mexico to Paraguay .

Discussion

Of all Brazilian Mimosoideae (32 genera and 810 species), nearly 44% of the genera and 7 .6% of the spe-cies are present at Serra do Cipó, while the whole state of Minas gerais accounts for 17 genera (53%) and 235 spe-cies (29%) (Lima et al . 2014) . The occurrence of such a high diversity of mimosoid legumes in a relatively small area is probably favoured by the peculiar location of Ser-ra do Cipó, between two of the widest and most diverse Brazilian phytogeographic domains: the Cerrado and the Atlantic Forest . The vegetation types in the area possess a high diversity of mimosoid taxa, except for the season-ally dry tropical forest where only three species are found (Anadenanthera colubrina, Piptadenia macradenia and Pseudopiptadenia warmingii). A similar situation was highlighted by Forzza et al. (2010a) for the whole State of Minas gerais, the richest in the country in number of species and endemics and where the Caatinga, Cerrado and Atlantic Forest are present .

The role of the Cerrado Domain is also relevant, for its great influence in the number of species: ca. 65% of the Ser-ra do Cipó mimosoids are found in cerrado (25 spp .) and

Fig. 5. diagram depicting the number of species of Mimosoideae of Serra do Cipó distributed by Phy-togeographic Domains. CER - Cerrado Domain; ARF - Atlantic Rain Forest Domain. Species collec-ted in disturbed areas are excluded.

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“campo rupestre” (15 spp.) areas, both encompassed in that domain (Fig 5). This is in agreement with the pattern high-lighted by Shrire et al. (2005), who indicate that the majority of Mimosoideae are mostly found through grassy biomes with fire-tolerant species and with a dry season, such as the Cerrado Domain, or biomes under semi-arid conditions rich in succulents, such as the Brazilian Caatinga .

An analysis of the distribution of habits amongst the species of Serra do Cipó indicates that 33 species are trees and the remaining 29 are shrubs or subshrubs (Table 1) . Tree species are found in all genera except for Calliandra . The predominance of this habit is expected since the major-ity of mimosoid legumes from Brazil are trees. However, all the shrubby or subshrubby species here studied belong either to Calliandra and Mimosa, with Stryphnodendron gracile, an endemic subshrub with a woody xylopodium, being the only exception . Both Calliandra and Mimosa are diversified in the Cerrado Domain, which is seasonally subjected to fire. The presence of the (sub)shrubby habit in these genera may be seen as an adaptation to the fire-re-gime, such as Calliandra dysantha and especially Mimosa where many species have woody xylopodiums. It is im-portant to mention that many of the tree species from the Cerrado are also adapted to fire-regimes, commonly pos-sessing thick corky barks. A recent work suggests a strong correlation between the development of structures adapted to fire and the occupation of the cerrados by some Mimosa lineages (Simon et al . 2009) .

The Cerrado Domain, which includes both cerrados and “campos rupestres”, is also very rich in Mimosa spe-cies (Barneby 1991; Simon & Proença 2000, Simon et al . 2009) and, amongst the genera of Mimosoideae found at the Serra do Cipó, it has one of the highest number of spe-cies: 26 . The remaining genera are represented by seven or less species and some just by a single species (Fig . 6) .

Amongst the Mimosoideae of Serra do Cipó, 24 show a wide geographic distribution, eight and five are endemic to the Cerrado and Atlantic Forest domains respectively, two occur both on the Cerrado and Caatinga domains (Mi-mosa filipes and M. gemmulata), two both on the Cerrado and Atlantic Forest domains (Calliandra parvifolia and Inga vulpina), and one at the Atlantic Forest and Caatin-ga domains (Pseudopiptadenia contorta) . The particular importance of the Serra do Cipó mimosoid flora, howev-er, relies on the presence of the following occurrences: a. four endemic taxa, two of the “campos rupestres” (Mimosa barretoi and Stryphnodendron gracile), one of cerrado and “campo rupestre” (M. bombycina var . bombycina) and one of a restrict area of cerrado at the southern portion of the range (M. sordida), b . 12 endemic species of the cerrado and/or “campo rupestre” of the Espinhaço Range, ten be-ing restricted to its southern portion, c . one species exclu-

sive of the semideciduous forest of Minas gerais (Stryph-nodendron polyphyllum), d . one exclusive species of sandy fields at Minas Gerais and São Paulo (M. paucifolia), and e. two rare species of the Atlantic Forest (Abarema villo-sa and Piptadenia paniculata) . The presence of all those endemic or restricted species highlights the relevance of the southern Espinhaço Range as an important centre of endemism, as has been pointed out for other authors based on data from different families (e.g. Schefflera J.R.Forst. & g .Forst . – Fiaschi & Pirani 2008; Chamaecrista Moench . – giulietti & Pirani 1988, Rando & Pirani 2011; Asclepia-doideae – Rapini 2010) .

Of the total of 15 endemics to the southern Espinhaço Range that occur at Serra do Cipó, ten belong to Mimo-sa, the richest mimosoid genus at the area . At Catolés, a surveyed region situated in the northern portion of the Es-pinhaço Range in Bahia State, the genus is represented by nine species, of which just two are endemic to that part of the range (Zappi et al . 2003) . These data suggest that Mi-mosa may show the same pattern highlighted by Rapini (2010) for Asclepiadoideae, with a higher rate of specia-tion, or lower rate of extinction, in the southern portion. On the other hand, Calliandra, with two endemics of the southern Espinhaço occurring at Serra do Cipó, presents a higher number of species and endemics in the northern Espinhaço (Barneby 1998), and is the richest mimosoid genus at Catolés, where 21 species were found, 17 being endemic to this portion of the range . (Zappi et al . 2003) .

To date, the completion of family-wide taxonomic treatments in the Flora of Serra do Cipó series has usually shown a tendency to increase species numbers when com-pared to the previous records in the checklist by giulietti et al. (1987). Some examples were highlighted by Pirani et al. (2009): Aquifoliaceae, with four species at the initial check-list and nine after taxonomic treatment (groppo & Pirani 2005), Scrophulariaceae s .l ., from ten to 17 species (Souza & giulietti 2003) and Apocynaceae s . str ., from 15 to 27 spe-cies after a detailed treatment (Kinoshita & Simões 2005) .

The same pattern is observed in the Mimosoideae treat-ment of Serra do Cipó (Borges & Pirani 2013): 39 new re-cords were found, and the total number of species (62) has almost tripled the 23 taxa listed by giulietti et al . (1987) .

Of the 39 new records, eight of them were found in herbaria collections (BHCB, ESA, MBM, RB, SP, SPF & UEC - acronyms according to Thiers 2014) and the re-maining 31 originated from new collections by different researchers in the last five years. These facts highlight the need to intensify field efforts to properly explore the bio-logical diversity of an area, even in well-studied regions. Similar situations still prevail in most tropical regions, as pointed out by several authors who indicate the impor-tance of developing fieldwork, florulas and monographs

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to access the still underestimated diversity of Angio-sperms in the Neotropics (e .g . Mori 1989; Thomas 1999; Prance et al 2000; Sobral & Stehmann 2009) .

Both the cerrado and the semideciduous forests are the vegetation types that most contributed to increasing the species number, followed by “campos rupestres” and disturbed areas, as roadsides, scattered through “campos rupestres” (Fig. 6). The first two formations brought the major contributors to the increase in Mimosoid species numbers because of recent collections expeditions to re-gions dominated by those biomes in Serra do Cipó .

In the cerrado, Mimosa accounts for 16 out of the 39 new records for the Serra do Cipó and it is the most di-verse genus of the area . Of the 13 cerrado and 6 “campos rupestres” new records, nine and three respectively be-long to Mimosa . The remaining species of Mimosa were collected in disturbed areas .

On the other hand, the intensive sampling of ar-boreous species carried out on a floristic survey in the eastern slopes of Serra do Cipó by Santos (2011), where semideciduous forests prevail, was critical in locating new Mimosoideae distributional records. As a direct con-sequence of this specific effort, all new records from this site are trees, which belong to taxa that are either typical or even restricted to that vegetational formation .

The four species collected in disturbed areas were Mi-mosa (M. bimucronata, M. diplotricha, M. pigra and M. velloziana), a habitat that has been very well explored since the beginning of the Flora of Serra do Cipó project . Even though they have a wide distribution, they may constitute

examples of recent invasions promoted by extensive dis-turbance related to the expansion and improvement of the roads . In fact, during the last four years, an evident increase in the population size of M. velloziana has been observed along the roadsides in some parts near the National Park .

The remarkable increase in Serra do Cipó Mimo-soideae species numbers is attributed mainly to three factors: better sampling effort in semideciduous forests, a vegetation type that was formerly only accounted for studies of the patches of “capões de mata”, increase of sampling effort in already well collected areas, and final-ly, new expeditions and collections in poorly explored areas like Santana de Pirapama, Lapinha — where the highest peak of Serra do Cipó is located — and Cong-onhas do Norte .

Conclusion

The goal of obtaining a comprehensive species list in areas with high biodiversity is a task accomplished only after a systematic and extended period of research (Rapini et al . 2008) . For the Mimosoideae of Serra do Cipó, extensive herbaria, field and taxonomic work was essential to reveal its full diversity . In fact, some treat-ments already published for families in the area need to be updated, since the increase of field expeditions to the area revealed unanticipated diversity .

The account of Mimosoideae of Serra do Cipó indi-cates that even a well explored area may show a high-er diversity than the initial inventory, after a taxonomic treatment based on intensive field work is accomplished. This indicates than an even higher effort must be done in areas that were not commonly target of research, such as many sites at the Brazilian center-west and north regions (Forzza et al 2010b), their biodiversity probably being underestimated and poorly protected .

Unfortunately with the expansion of agrobusiness, natural environments in Brazil are being damaged, to-gether with unknown data about living organisms (Bran-don et al . 2005; Klink & Machado 2005) . As stressed by Funk (2006), facing the biodiversity crisis, the need to develop and sponsor Floras — as well as Faunas and tax-onomic studies of other organisms — is urgent and must be carried out together with a public agenda committed to the better control of land usage in the country .

The data resulted from this survey of the Serra do Cipó Mimosoideae also show the urgent need to develop region-al floras and expand collections in order to build for the Brazilian Flora, one of the richest in the world (Forzza et al. 2010b), a well-founded knowledge base with data about its composition, distribution, diversity and conservation .

Fig. 6. Diagram depicting the new records of Mi-mosoideae of serra do cipó (as compared to the checklist by Giullieti et al. 1987) distributed by ve-getational formation. CAM - campo rupestre; CER - Cerrado; SDTF - Seasonally Dry Tropical Forest; SF - Semideciduous Forest; DIST - disturbed areas.

53

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