BOLETIM 1)0 MUSEU PARAENSE EMÍLIO GOELDI

Série BOTÂNICA

GOVERNO DO BRASIL

Presidência da República

Presidente - Itamar Franco

Ministério da Cipncia e Tecnologia - MCT

Ministro - José Israel Vargas

Conselho Nacional de Desenvolvimento Cientifico e Tecnológico - CNPq

Presidente - Lindolpho cie Carvalho Dias

Museu Paraense Emílio Goeldi - MPEG

Diretor - José Guilherme Soares Maia

Vice-Diretor de Pesquisa - Pedro Luiz Braga Lisboa

Vice-Diretor de Difusão Científica - Denise llamú Marcos de La Penha

Comissão de Editoração - MPEG

Presidente - William L. Overal

Editor-Associado - Pedro Luiz Braga Lisboa

Equipe Editorial -Lairson Costa, Lais Zumero, Graça Overal

CONSELHO CIENTÍFICO

Consultores

Ana Maria Giulietti - USP

Carlos Toledo Rizzini - Jardim Botânico do Rio de Janeiro

Dana Griffin III - Univcrsity of Florida

Enrique Forero - New York Botanical Garden

Fernando Roberto Martins - UNICAMP

Ghillean T. Prance - Royal Botanic Garden

Hermógenes Leitão Filho - UNICAMP

João Murça Pires - Museu Paraense Emílio Goeldi - CNPq

João Peres Chimelo - IPT

Nanuza L. Menezes - Instituto de Biociências - USP

Ortrud Monika Barth - Fundação Oswaldo Cruz

Paulo B. Cavalcante - Museu Paraense Emílio Goeldi

Thcrezinha SanfAnna Melhem - Instituto de Botânica de São Paulo

Warwick E. Kerr - Universidade Federal de Uberlândia

William A. Rodrigues - Instituto Nacional de Pesquisas da Amazônia

por/by/MCT/CNPq/Museu Goeldi

V4 SEI 1995

ISSN 0077-2216

Ministério da Ciência e Tecnologia

Conselho Nacional de Desenvolvimento, ( Científico e Tecnológico

MUSEU PARAENSE EMÍLIO GOELDI

Boletim do

Museu Paraense

Emílio Goeldi

Série

BOTÂNICA

Vol. 9(2)

Belém - Pará

Dezembro 1993

MCT/CNR]

MUSEU PARAENSE EMÍLIO GOELDI

Ministério da Ciência e Tecnologia - PR

CNPq - Conselho Nacional de Desenvolvimento Científico e Tecnológico

Parque Zoobotânico - Av. Magalhães Barata, 376, São Braz

Campus de Pesquisa - Av. Perimetral, Guamá

Caixa Postal: 399. Telex: (091) 1419. Telefones: Parque, (091) 224-9233. Fax (091) 241-7384

Campus, (091) 228-2341 e 228-2162.

66.017-970. Belém-Pará-Brasil

O Boletim do Museu Paraense de História Natural e Elhnographia foi

fundado em 1894 por Emilio Goeldi e o seu Tomo I surgiu em 1896. O atual

Boletim é sucedâneo daquele.

The Boletim do Museu Paraense de História Natural e Ethnographia was

founded in 1894, by Emilio Goeldi, and the first volume was issued in 1896.

The present Boletim do Museu Paraense Emilio Goeldi is the successor to

this publication.

REVISTA FINANCIADA COM RECURSOS DO

Programa de Apoio a Publicações Científicas

MCT ÊlcNPq 0FINEP

CDD 583.309811

ASSOCIAÇÃO RIZÓBIO-LEGUMINOSAS

NA AMAZÔNIA 1

Fátima M.S. Moreira 2 3

Marlene F. da Silva 1

RESUMO - Grande parte das espécies de leguminosas tropicais ainda carece

de informações sobre sua capacidade de nodular, isto é, de se associar

simbioticamente a bactérias fixadoras de nitrogênio dos gêneros Rhizobium.

Bradyrhizobium. Azorhizobium e Sinorhizobium. Sendo assim, foi realizada

uma pesquisa extensiva sobre a ocorrência de nodulação em leguminosas,

através do projeto "Levantamento da capacidade de nodulação e/ou fixação

de nitrogênio, em espécies forestais da Região Amazônica " no período com-

preendido entre junho/ 1984 e junho/1987. Os resultados obtidos foram, então,

divididos, de acordo com as áreas pesquisadas, em 4 comunicações que serão

divulgadas sucessivamente. Nesta primeira comunicação são apresentados os

resultados relativos às coletas realizadas no Município de Manaus, incluindo

a Reserva Ducke, e no Município de Manacapuru. Das 96 espécies e 48 gêne-

ros coletados nesta etapa, 57 espécies e 8 gêneros não possuíam referências

anteriores na literatura sobre a capacidade de nodular. Foram encontrados

nódulos em espécies dos gêneros Campsiandra, Etaballia e Abarema. Por ou-

tro lado, os gêneros Aldina. Bocoa. Lecointca. Taralea e Marmoroxylon pare-

cem incapazes de nodular. A maior parte das estirpes de bactérias isoladas

dos nódulos tem características típicas do gênero Bradyrhizobium.

PALAVRAS-CHAVE: Leguminosas - Rizóbio, Leguminosas da Amazônia. As-

sociação Rizóbio-Leguminosas

ABSTRACT- A great part of tropical leguminoits species hasyet no informations

about nodulation capability, e.a., capabUily of simbiotic association with

nitrogen fxing bactéria belonging to the genera Rhizobium. Bradyrhizobium.

Azorhizobium and Sinorhizobium. Therefore, an extensive research on

nodulation occurrence in leguminoits species of the Amazon region was

performed by the project " Survey on nodulation capability and/or nitrogen

fxalion in forest species of Amazon region ” during the period june/1 984-june/

1 Projeto 54/84/0294/00, financiado pelo Convênio FINEP/INPA/CODEAMA

2 Coordcnaçâo de Pesquisas em Ciências Agronômicas-CPCA/INPA

3 Coordcnação de Pesquisas cm Botânica-CPBO/INPA; Centro de Ensino e Pesquisas Florestais-CEPEP/

UTAM

129

Boi Mus. Para. Emílio Goeldi, sér. Bot. 9(2). 1993

1987. Results of this project are divided in 4 phases according to the areas of

collection. In this paper, results of the phase I . related to collection on

Municípios of Manaus, including Reserva Florestal Ducke and Manacapuru

are introduced. From 96 species and 48 genera collecled in this phase, 57

species and 8 genera had no previous informations on the literature, about

nodulation capability. Nodules were found in species of the genera:

Campsiandra, Etaballia and Abarema. By contrary, the genera Aldina, Bocoa,

Lecointca, Taralea and Marmoroxylon seems not able to nodulate. The majority

of bactéria isolated from nodules have typical cultural charactheristics of the

gentis Bradyrhizobium.

KEY WORDS: Leguminosac - Rhizobia, Leguminosae of the Amazon,

Association Rhizobia-Leguminosae

INTRODUÇÃO

Na flora amazônica a família Leguminosae tem grande diversidade em

espécies e é uma das mais abundantes em número de indivíduos (Ducke 1 949,

1959; Black et al. 1950; Rodrigues 1961; Rodrigues 1967 a e b; Klinge &

Rodrigues 1971; Prance et al. 1976). Ducke (1949) reportou 867 espécies

distribuídas em 1 1 8 gêneros de Leguminosae na Amazônia brasileira. Desde o

trabalho de Ducke ( 1 949) novas espécies e gêneros têm sido descritos (Barneby

& Grimes 1984; Cowan 1966, 1967, 1975; Forero 1972; Hopkins 1986; Lee

& Langenheim 1975; Mesquita & Silva 1984; Rodrigues 1974, 1975; Rodrigues

& Lima 1989; Rodrigues & Matos 1980; Silva 1981,1986; Silva & Graham

1980, e outros) aumentando significativamente aqueles números. Em uma

“check-list” preliminar elaborada com base nas coleções dos principais

herbários da região (INPA, MG e EMBRAPA/CPATU (1 AN), antigo IPEAN)

e algumas revisões taxonômicas modernas. Silva et al. (1989) encontraram

um total de 1 46 gêneros, 1 .24 1 espécies e 3 1 0 táxons infra-específicos nas 3

subfamílias consideradas (Caesalpinioideae, 48 gêneros, 475 espécies, 22

subespécies e 175 variedades; Mimosoideae, 23 gêneros, 288 espécies, 3

subespécies e 18 variedades; Papilionoideae, 75 gêneros, 478 espécies, 2

subespécies, 84 variedades e 6 formas).

Bactérias aeróbicas fixadoras de nitrogênio dos gêneros Rhizobium,

Bradyrhizobium, Azorhizobium e Sinorbizobium vivem saprofiticamcnte no

solo ou em simbiose com várias espécies leguminosas, formando estruturas

nodulares em suas raízes ou, excepcionalmente, no caule como em Sesbania

rostrala (Dreyfus et al. 1 988).

130

Associação Rizóbio-leguminosas na Amazônia

A fixação biológica de nitrogênio é um processo que já vem sendo explo-

rado economicamente no cultivo de espécies agrícolas importantes como a

soja, o feijão e a ervilha, entre outros, em substituição aos onerosos adubos

nitrogenados, obtidos através de processos industriais.

O potencial de utilização de várias espécies de leguminosas nativas da

Amazônia como alimento, madeira, adubação verde, produtos químicos, etc.,

vem sendo destacado por diversos autores (Loureiro et al. 1979; National...

1979; Arkool 1984a,b; Moreira & Franco 1992) e a fixação biológica de ni-

trogênio pode desempenhar papel importante na exploração econômica

dessas espécies.

Como a associação simbiótica de bactérias fixadoras de nitrogênio com

muitas espécies tropicais ainda era desconhecida (Allen & Allen 1981 ), foi

realizado um levantamento extensivo sobre a capacidade de nodulação de es-

pécies nativas da Amazônia durante o período de junho/l 984 e junho/l 987.

Neste trabalho são fornecidas informações sobre a capacidade de nodular

de espécies encontradas no Estado do Amazonas em áreas dos Municípios de

Manaus, incluindo a Reserva Florestal Ducke, na BR 1 74, e de Manacapuru.

MATERIAL. E MÉTODOS

Foram realizadas várias excursões para coleta de material durante o pe-

ríodo de junho/1984 e junho/1987, no Município de Manaus, inclusive na

Reserva Ducke e uma excursão ao Distrito de Caviana, no Município de

Manacapuru, de 12 a 27 de março de 1985.

Espécimes de leguminosas com diferentes hábitos de crescimento,

preferentemente com flores e/ou frutos, foram coletadas ao longo das margens

de rios, estradas ou em picadas abertas dentro da mata. Para prevenir o ataque

de fungos, as amostras, em geral, foram embebidas em álcool comercial e

colocadas dentro de sacos plásticos e fechados. Em Manaus, elas foram pren-

sadas, colocadas na estufa para secar e posteriormente foram identificadas

sendo, em muitos casos, incorporadas ao Herbário do INPA.

Em plantas adultas foram procurados nódulos seguindo-se as raízes la-

terais a partir do tronco até suas ramificações mais finas. Quando havia mu-

das próximo à planta matriz seu sistema radicular era retirado intacto do solo

com auxílio de um enxadeco. Em ambos os casos, cavar o solo foi uma

131

SciELO

10 11 12 13 14 15

Associação Rizóbio-legiiminosas na Amazônia

capacidade de nodular. Foram encontrados nódulos em espécies dos gêneros

Campsiandra, Etaballia e Abarema. Os gêneros Aldina, Bocoa, Lecointea,

Marmaroxylon e Taralea parecem incapazes de nodular, principalmente os

gêneros Lecointea e Taralea que também não nodularam em condições de

viveiro.

Cassia leiandra Benth. não nodulou mesmo quando cultivada em vivei-

ro. Este resultado é conflitante com a observação feita anteriormente porNorris

( 1 969), sobre ocorrência de nodulação nesta espécie, na região de Manaus.

Sclerolobium, Hymenolobium, Pterocarpus, Derris e Calhandra , ge-

ralmente reportados como nodulíferos por outros autores (Faria et al„ 1989)

foram encontrados sem nódulos, nas áreas pesquisadas. Porém , Andira

núcrantha Ducke e Platytniscium duckei Huber, também encontradas sem

nódulos no campo, nodularam quando cultivadas em viveiro. Estes exemplos

indicam a necessidade de não se considerar somente dados de campo como

prova da incapacidade de nodular de uma espécie. No campo, condições de

equilíbrio na nutrição de nitrogênio, ausência de estirpes de rizóbio específi-

cas e outros fatores limitantes, podem impedir o estabelecimento e o desenvol-

vimento da simbiose em uma espécie nodulífera. Em viveiro, estes fatores

podem ser controlados para que a nodulação seja induzida, de modo que, se a

espécie é capaz de nodular, ela terá condições adequadas de expressar esta

característica. No caso de Pterocarpus , que também não nodulou em viveiro,

provavelmente, o período de crescimento (5 meses) não foi suficiente para a

indução da formação de nódulos. Faria (com. pessoal) relata que algumas

espécies só começam a nodular em viveiro, 7 meses após a germinação.

O cultivo de espécies em viveiro no levantamento da capacidade de

nodular, em leguminosas, também é útil no caso de indivíduos cujo sistema

radicular é de difícil acesso como nas matas muito densas e/ou em condições

alagadas, por exemplo. Outro objetivo do estudo na nodulação das espécies

em viveiro é a possibilidade de isolamento de rizóbio nativo de diferentes solos

da região amazônica.

A autenticidade dos nódulos encontrados como estruturas induzidas por

bactérias fixadoras de N2 foi confirmada pela medida da atividade de

nitrogenase (enzima responsável pela fixação de N2) através da técnica de

redução do acetileno (ARA). Em todas as espécies noduladas foram encontra-

dos nódulos com atividades da nitrogenase (ARA), positiva (+).

SciELO

10 11 12 13 14 15

Boi. Mus. Para. Emílio Goeldi. sér. Boi. 9(2). 1993

Um nódulo de aproximadamente I Ocm, e outros semelhantes, foram en-

contrados em Swartzia schomburgkii Bentli. e a ARA positiva confirmou a

autenticidade destas estruturas.

Machaeritim leiophyllum (DC). Benth. var cristacastrense (Mart. ex

Benth.) Rudd., Tachigalia plúmbea Ducke e Swartzia ulei Harms também

foram encontradas nas áreas estudadas, mas, suas capacidades de nodulação

não puderam ser pesquisadas porque seus sistemas radiculares estavam

submersos.

Outros 20 indivíduos coletados não puderam ser identificados a nível de

espécie. Esses indivíduos foram classificados nos gêneros: Sclerolobium,

Tachigalia. Cassia, Bauhinia, Pentaclethra, Acacia, Inga. Pilhecellobium,

Diplotropis, Plaiymiscium , Dalbergia e Dioclea.

Várias estirpes com características típicas de rizóbio, a maior parte com

características do gênero Bradyrhizobium, foram isoladas dos nódulos

coletados. A caracterização morfológica, bioquímica, fisiológica e genética

destas estirpes se encontra em andamento e estes resultados serão divulgados

em trabalhos posteriores.

134

Associação Rizóbio-legnminosas na Amazônia

Tabela 1 . Ocorrência dc nodulação em leguminosas da Amazônia, nos Municípios de Manaus (MAO),

incluindo a Reserva Florestal Duckc (RFD) e Manacapurú (Mc).

CoI.FMSM

Número dc

Local

Hábito

Nodulaçào

Referências anteriores

TAXON

Hcib.INPA

indivíduos

Campo/

sobre capacidade de

***

obsenados

Coleta

Planta*

Viveiro

nodular

CAFSALPINIOIDEAE

Cacsalpinieae

( 'aesalpinia ferre a Mait. cx Tul .

252

MAO

Faria et al. 1984

('. pulcherrima(L.) Sw.

Dimorplutndra pani/lora

254

MAO

Allen&Allen 1981

Spruce ex Bcnth.

( 'ampsiat ulra comosa ( B th . ) Cowan

239/133625

RFD

■

ixl

Magalhães 1 986

var. lauiifolia (Bth.) Cowan

Sclero/obium chry sopln 'Hum

10/133836

Mc. MAO

•f

Poepp.&Endl.

157/151904

RFD

Vouacapoua palliilior Duckc

153

RFD

Magalhães & Fernandes 1984

Cassicac

( ' assia leiandra Bth.

( hamaecrislaniclidans (L)

235/133623

Mc. MAO

ixl

Allen&Allen 1981

Mocnch. ssp. disadena {Slcuá.)

Invin & Bamcby

51/133854

Allen&Allen 1981

Dialiwn guianensis ( Aubl .) Sandw.

■

Magalhães et al. 1982;

Faria et al. 1984

Se una quinquagulala ( Rich . )

Invin & Bamcbv

215/133609

MAO

EJVt

S. siumea( Lam.) Invin & Bamcbv

227/133617

MAO

Allen&Allen 1981

S. ,v/7vt'.s7/*/.v(Vel.) Invin & Bamcby

60/133860

Faria et al. 1987

Detaricac

( vnomeim bauhiniifoHa Bth.

20/133839

Mc. MAO

( 'opaiferu muhiju^a Hayne

141/133870

RFD

Magalhães 1986

('rudia amazônica Spr. cx Bth.

27/124720

( puhescens Spr. cx Bth.

18/133838

Pehopyne pauiculala Bth .

I*. caiinpae Duckc ssp. x lahra

224

RFD

‘

Magalhães et al. 1982

(W.Rodr.)M.F. da Silva

173

RFD

ixl

1*. excelsa Duckc

178

RFD

íxl

P venosa Bth.

Magalhães 1 986

Amhci-sticac

A lacrolobiiim limbamm

Spruce cx Bth.

147

RFD

M microcalvx Duckc

221

RFD

ixl

A/, acaciifolinm Bth.

24/124719

Mc. MAO

Faria et al. 1987

A 1 anpitsiiftliitm ( Bth.) Cowan

46/133849

íxl

(*) A = árvore: Ah = arbusto: C -= cipó: E erva:J indivíduo jovem: M -muda

(**) ' nódulos presentes:- - ausência de nódulos: nd - nodulaçâo náo verificada

1***1 col coleção

I Icrb = herbário

135

Boi. Mus. Pura. Emílio GoeliJi. sér. Boi. 9(2). 1 993

Col.FMSM

Número de

L.ocal

I lábito

Nodulaçào

Referências anteriores

TAXON

Hat.INPA

indivíduos

Campo/

sobre capacidade de

***

observados

Coleta

Planta*

Viveiio

nodular

PAPILIONOIDEAE

Swartzieae

Alílina lati folia Spr. ex Blli.

MAO

- nd

A. helcrophylla Spr. ex Bth.

MAO

liocoa viridifolia (Ducke) Cowan

165

RFD

Lecoiníea amazônica Ducke

Swartzia reliculala Ducke

149

RR)

- nd

S. laevicarjm Amsh.

57/133858

- nd

S. scricca Vog.

36/133846

S. schomhurgkii Bth.

146

RH)

’ nd

S u lei Harms.

197/133593

RFD

S ingifolia Ducke

203/133599

RFD

S. cuspidata Bth.

148

RH)

S. polyphylla DC.

36/133845

+ nd

Sophoreae

Acosmium nilen.s ( Vog.) Y acovlcv

28/133841

- nd

Magalhães et al. 1982

Diplolropis purpura (Rich.) Amsh.

170

RFD

+ ixl

( írmosia macrocalvx Ducke

48/133851

+ +

O. smithii Rudd.

204/133600

RFD

- nd

O. discolor Spr. ex Bth.

47/133850

■

Diptcrygeae

Diplervx odoram (Aubl.) Willd.

207/133603

RFD

- nd

Váiias

IX magnifica Ducke

237/133624

RH)

•

Taraleaopposilifolia Spr. cx Bth.

175

RFD

ixl

■

Aeschynomencae

Acschynomene rudis Bth.

03/133833

Al)

+ nd

Allcn & Allcn 1981; Fana et

al. 1984

Dalbergieae

156/133872

RFD

Allcn & Allcn 1981

A. surinamensis ( Bth. ) Splitg.

151/133871

RFD

- nd

A. nnifoliolala Ducke

166/133875

RFD

- nd

Sylvester-Bradlcy ct al. 1 980

Dalbergia inundam Bth

23/124717

Mc; MAO

4- nd

■

1). riparia ( Mai1.)Bth.

54/133857

+ ixl

Jàahaliaduhia Rudd .

ffymcnolohium hclerocarpum

31/133843

- nd

Ducke

158/133873

RFD

■

H. scriccum Ducke

154

RID

ixl

A lachacriumfirocsii Rudd.

• nd

Matos.

136

SciELO

10 11 12 13 14 15

Associação Rizóbio-leguminosas na Amazônia

TAXON

Col.FMSM

Hcrb.INPA

+**

Número de

indivíduos

obseivados

Lrocal

Coleta

Hábito

Planta*

Nodulaçào

Campo/

Viveiro

Referências anteriores

sobre capacidade de

nodular

M inundatum (Mait. ex Bth.)

Ducke

23-A/124718

ixl

Magalhães 1 986

Platymiscium ducke i Huber

08/139629

McMAQRFD

AJM

Pterocarpus amazonicus I íuber

26/124721

P. amazomun (Mait. cx Bth.)

Amsli.

163/133874

Mc; RFD

Vatairea guianensis Aubl .

19/124710

Faria et al. 1987

Milletieac

Derris longifolia Bth.

21/133840

Phaseoleae

Cliloriajavitensis (H.B.K.) Bth.

448

MAO

Dioclea macrocarpa Hub.

449

MAO

MIMOSOIDEAE

Parkieae

Parkia discolor Bth.

189/133586

MAO

P. nni/iijuga Bth.

201/133597

RFD

Magalhães et al. 1982;

P. opposil ifolia Bth.

225/133615

MAO;RFD

Magalhães 1 986

P pendula Bth . ex Walp.

210/133604

RFD

Vários

Penlaclelha macroloba /Willd.)

Kunth.

226/133616

MAO

Vários

Mimoseae

Adenanlhera pavonina L.

230/133618

MAO

A. peregrina (L.) Bth.

231/133619

MAO

vários

Untada polyphylla Bth .

Campeio 1976

Mimosa pigra L.

194/133690

MAO

-f

Sylvester-Bradleyctal 1980

M. polystachya Kunth.

186/133597

MAO

Stryphnodendronpulclierriinnm

(Willd.) Hochr.

202/133598

Mc; RFD

Faria et al. 1984

S. microstachyum Poepp. & EndI.

187/133584

MAO

Ingeae

( .' alliandra tenuijlora Bth .

160

RFD

Cedrellinga catenaeformis Ducke

140

RFD

Magalhães et al. 1982

Unterohbiwn schomburgkii Bth.

242

RFD

Magalhães et al. 1982

lngacapitata Dcsv.

232/133620

MAO

Ingaedullis Mail.

vários

/. ingoides (R\ch.) Willd.

Allen & Allen 1 98 1

/. nobilis Willd.

05/133835

/. panurensis Spr. cx Bth.

200/133596

RFD

■

137

1, | SciELO

Associação Rizóhio-legwninosas na Amazónia

referências bibliográficas

ALLEN, O.N. & ALLEN, F.K. 1981 . The Leguminosae. London, University ofWisconsin Press Macmillan

Publishing Company.

ARKOLL, D. 1984a. A coniparison ofsomc fast growing specics suitable forwood lots iii lhe wetlropics.

Pesq. Agropec. liras. 19: 61-68.

ARKOLL. D. 1 984b. Sonic Icguminous trees providing uscful fruits in lhe Nortli of Brazil. Pesq. Agropec.

liras. (s/n):235-239.

BARNEBY, R.C. & G RIMES, J.W. 1984. A new specics ot Abarema Pitticr (Mimosaccae-Ingeae), from

Amazônia Brazil. Acta Amazon. I4( l/2):95-99. Suplemento.

BLACK, G.A.; DOBZI IANSKI.T 1 1.: PAVAN. C. 1950. Some attempts to estimate specics diversity and

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Associação Rizóbio-leguminosas na Amazônia

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Recebido em 09.06.92

Aprovado em 30. 10.92

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CDD: 583. 1150981

STUDIES IN ANNONACEAE. XVI. A TAXONOMIC

REVISION OF DUGUETIA A. F. C. P. DE

SAINT-HILAIRE SECT. DUGUETIA (ANNONACEAE)

IN EASTERN BRAZIL

Ping He'

P. J. M. Maas -

ABSTRACT - This paper contains a laxonomic revision of lhe species of

Duguetia Secl. Duguetia that occur in Brazil E of 50 IVL. Three species and

one variely recognized in lhe carlier trealment of lhe gentis by Fries are now

pui into synonymy. The presenl study is par I of lhe monographic sludv of lhe

whole gemis currenlly under way, lo be published in Flora Neolropica.

KEY WORDS: Annonaceae, Duguetia. laxonomic revision

RESUMO - Este trabalho é uma revisão taxonômica das espécies do gênero

Duguetia Secção Duguetia, ocorrentes no Brasil a Este de 50° JVL. Três espé-

cies e uma variedade, anleriorinente reconhecidas no tratamento do gênero

por Fries, são agora colocadas em sinonimia. Esta estudo faz parte duma

obra monográfica do gênero inteiro atualmente em andamento e a ser publicada

na Flora Neotrópica.

PALAVRAS-CHAVE: Annonaceae, Duguetia

INTRODUCTION

Being kindly invited to spend a year at the Herbarium of the Department

of Plant Ecology and Evolutionary Biology, University of Utrecht, the first

author has been able to concentrate his attention on the taxonomy of the eastern

Brazilian species of Sect. Duguetia of the genus Duguetia between February

and Deccmber, 1990.

1 Department of Biology, Southwest China Teachers University, Beibei, Chongqing, Sichuan, China

- Department of Plant Bcology and Evolutionary Biology, Herbarium Division, University of Utrecht,

Heidelberglaan 2, 3584 CS Utrecht, The Netherlands

143

Boi. Mus Para. Emílio Goeldi. sér. Boi. 9(2), 1993

TAXONOMIC HISTORY

The geiuis Duguetia was established by Saint-Hilaire (1824) in Flora

Brasiliae MeridionaliswiththetypespeciesZ). lanceolata A. F. C. P. deSaint-

Flilaire. The gentis was named after Jacobus Joseph Dugiiet, a french activist

and author of “Ouvrage des six jours” (1731). Martins (1841) enlarged the

genus with 7 new species. Afterwards, Baillon ( 1 868) regarded Duguetia as a

synonym of Aberemoa Aublet with Aberemoa guianensis as type. Aberemoa

guianensis , however, belongs to Guatteria Ruiz et Pavón. Thus the generic

liame of Duguetia was used again by Prantl (1891). In the following years

many new species were described.

Duguetia occurs throiighout the Neotropics and is characterized by the

the combination of a pseudosyncarpous fruit and stellate to entire scales and/

or stellate hairs.

In bis revision of Duguetia in 1934, R. E. Fries grouped the 67 species

then known in 1 3 sections. A new section, D. Sect. Xylopipetalum, was added

to accomodate a new species ( 1 937). The section Duguetia (“Eu- Duguetia")

was by far the largest with 25 species. D. caudata was described by Fries in

1 937 and added as the 26th member of the section.

Several of Fries’s sections are based on unique character States, and appear

quite natural-looking. Whether or not these are truly natural groups remains to

be settled yet. At any rate, these sections are easily recognisable. D. Sect.

Duguetia, on the other Iiand, is not so easily recognized as such. It is defined

rather by a combination of character States, viz., entire scales on all floral

parts, sinal I bracts, (nearly) free calyx lobes, and two whorls of petals of

about the same size.

In lhe present study, those species of D. Sect. Duguetia are treatcd th at

are found in the eastern part of Brazil, i.e., east of 50° WL (Table 1 , Figures 1-

3). Some species are restricted to small areas along the east coast (Rio de

Janeiro, São Paulo, Bahia). The distribution patterns of others, on the contrary,

are extended far more toward the north-west, (up to Colombia, Venezuela, and

Suriname), west (Matto Grosso), or south-west (as far as Paraguay or Bolivia).

Among the species of th is section, D. furfuracea (including D.

hemmendorffii and D. jonasiana) is the most widely sprcad, ranging from

Brazil (Mato Grosso. Paraná, São Paulo, Bahia, Goiás to Ceará) to Paraguay

144

Slndies In Annonaceae. A77. A Toxononiic Revisipn of Duguetia A. F. C. P. de Sanl-Hilaire

and Bolivia. Besides, it is the most variable species showing dynamic diversity

of infra-specific morphological and ecological variations. It is fonnd in dense

secondary forest, as well as in dry and open vegetations (cerrado).

Apart from some species in Annona , the last-mentioned vegetation type

is very unusual for annonaceous species, which mostly occur in primary or

secondary forest. Hovvever, within Sect. Duguetia there are some notable

exceptions, too. D. dicholepidota and D. moricandiana, botli highly similar

to D. furfuracea, are found in caatingas and restingas of Bahia. Four otlier

species, viz., D. lanceolata , D. riede liana, D. microphylla , and D. salicifolia

are also regarded as closely related. They occur in cerrados, or in semideciduous

or montane forest along the east coast of Brazil. D. glabriuscula, known from

Mato Grosso and Bolivia, is found in comparable cerrado vegetation. It is so

similar to D. lanceolata, that vve decided to include D. glabriuscula in the

present treatment, notwithstanding its slightly more westerly distribution.

Three otlier species, viz., D. echinophora, D. marcgraviana (including

D. brevipedunculata ), and D. lepidota, show a rather high resemblance to D.

furfuracea, especially in their leaf indument. Although theirmain area is more

in the north-west (map 4), they are found as far east as Maranhão or Piauí,

and are therefore included in this study. Their habitat is primary or secondary

forest (D. echinophora, D. lepidota), or wet savannas and gallery forest

(D. marcgraviana).

Of the species of Sect. Duguetia not included in the present treatment,

two show clear links to eastern Brazilian species, and therefore should be

mentioned here. D. lúcida is closely related to D. lanceolata, but occurs in

Trinidad, Tobago, Venezuela, Guyana, and the Brazilian States Acre and

Roraima. D. surinamensis comes morphologically close to D. echinophora,

D. lepidota , and D. marcgraviana, but occurs from Colombia to Pará, as far

south as Goias. D. quitarensis should also be mentioned. It is found from

Guyana to E Peru and Bolivia, but may occur as far east as Maranhão. The

species is, however, morphologically highly variable and represents perhaps a

complex of species. For this reason, it lias also been omitted from the present

study. Although D. Incida, D. surinamensis and D. quitarensis are not treated

in full, they are discussed underthe descriptions oftlie allied species to íàcilitate

their identification.

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Boi. Mus. Para. Emílio Goelcli. sér. But 9(2). 1993

MATERIAL AND METHODS

For th is stucly, material of the following Iierbaria was investigated: A,

AAU, B, BM, BR, C, CEPEC, COL, E, ECON, F, G, GB, GH, GUA, HBG,

1AN, IN PA, K, LE, M, MICH, MG, MO, NA, NY, OXF, P, R, RB, S, SP, U,

UC, US, VEN, W, WIS, and Z.

Fora better understanding of the interspecific variation, material from

the whole area of a specics was sludied. Under “Specimens examined”, however,

only Brazilian material is cited.

All quantitative data are based on mature material. Measurements were

mostly taken from many specimens.

The terminology used in the descriptions is mainly based on Radford et

al. ( 1 974) and I iickey ( 1 979). Some terms need some explanation.

Indument

Within Dugiielia , the indument and the structure of the individual

trichomes is important forthecircumscription and identi fication ofthe species.

Basically, the trichomes can be classified into two types: scales and stellate

hairs.

In this study, scales with free rays ovcr less than 1/3 of their length are

referred to as “entire scales”. “Stellate scales” applies to trichomes with rays

free over 1/3 - 2/3 of their length. The term “stellate hairs” lias been reserved

for trichomes with at least 2/3 of rays free (Figure 3).

When stellate hairs are present, tliey may have a slightly different

morphology on different plant parts.:

a. Appressed: the rays of the stellate hairs remain close to the plant surface

(may be found on both upper and lower side ofthe leaves).

b. Erect: the rays of the stellate hairs are straight, more or less perpendi-

cular to the plant surface (may be found on adaxial side of lamina, on ovary

and stigma).

c. Curviform: the rays of the hairs are irregularly curved (found on inner

side of bracts, sepals, and petals of all species, and sometimes on stigmas as

well).

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Studies lnAmonaceae. XVI. A Toxonomic Revision ofDugiietia A. F. C. R de Sant-l/ilaire

The density of the indument varies considerably, betvveen species and

with age. The following categories are used:

- “sparse": less than 5 trichomes per mm 2 ;

- “rather dense”: 5- 1 0 trichomes per mm 2 ;

- “dense”: more than 10 trichomes per mm 2 . In th is category, the

trichomes often touch each other.

Venation patterns

Venation patterns may change more or less during development of a leaf.

Therefore, only mature leaves were described. Besides, many leaves have

different venation patterns in the lower, middle, and upper part of the leaf

(Klucking 1986). In th is paper, only the middle part of the leaf was taken into

consideration.

SYSTEMATIC TREATMENT

Duguetia A. F. C. P. de Saint-I lilaire Sect. Duguetia R. E. Fries, Acta Horti

Berg. 12(1): 32. 1934 (“Eu-Diigiietia”); R. E. Fries in Engler& Prantl,

Nat. Pflanzenfam., ed. 2, 17 a II: 55. 1959.

Shrubs to trees. Twigs striate or ribbed, lenticel late, sparsely to densely

covered with yellowish fimbriate to stellate scales, becoming glabrous with

age. Leaves petiolatc. Lamina elliptic, ovate, to obovate. or narrowly so,

chartaceous to coriaceous, base acute to obtuse, apex acute, acuminate to

roimded, upper side glabrous to rather densely covcred with whitishor yellowish,

5- 1 8(-20)-rayed, appressed orerect stellate hairs. becoming glabrous with age.

lower side sparsely to densely covered with fimbriate to stel late scales, primary

vein impressed on the upper side, venation camptodromous, secondary veins

uniformly or abruptly curved, raised on both sides, between 5-22 on either

side of the primary vein, angles with primary veins between 45°-90°, intercos-

tal arca distinct, distance between the middle ofclosure and margin 1-10 mm,

tertiary veins (slightly) raised on the upper side, slightly raised to Hat on the

lower side, forming (1-16 more or less irregular nets.

Inflorescence leaf-opposed, pseudo-terminal or rarely supra-axillary,

simple, 1-3-flowered. or ramose, ( l-)3- 1 6-flowered rhipidia, subsessile oron

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Boi. Mus. Para. Emílio Goelcti, sér. Boi. 9(2), / 993

the top of a 2-5 mm long, ligneous, rudimentary twig. Pedicels straight or

recurved, to 25 mm long, to 35 mm long in fruit. Bract(s) 1-2 to the pedicel,

small, caducous, the lower bract below the articulation, amplectant, mostly

broadly to depressed ovate-triangular, the upper bract amplectant below the

articulation, broadly to depressed ovate-triangular. Indument of inflorescence

(inciuding fruits) consisting of stellate scales and/or stellate hairs; pedicels,

outer side of bracts and sepals rather densely to densely covered vvith stellate

scales, inner side of bracts, sepals, and petals rather densely to densely covered

vvith stellate hairs to glabrous, fruit sparsely to densely covered with stellate

hairs and scales or rarely glabrous.

Flower buds broadly ovoid to subglobose, vvith or without 3 ribs, apex

acute to obtuse or mucronate, to 1 0(- 1 2) mm long. Flowers green to cream,

yellow to red (-purple), usually pink to dark red at inner base (in vivo). Sepals

connate at the base, ovate-triangular to very broadly ovate-triangular, much

shorter than the petals. Outer and inner petals ovate to obovate, apex acute to

obtuse, sometimes mucronate, to 30(-45) mm long. Torus depressed to very

depressed ovoid-ellipsoid, apex concave. Stamens between 60-300, apical

prolongation of connective shallowly to very shallovvly conical, puberulous to

glabrous. Carpels between 40-380(-480), glabrous to densely covered with

erect stellate hairs, style terete, glabrous to densely covered with stellate hairs.

Fruitingreceptacleellipsoid,obovoid to subglobose, 10-35 mm long, 5-25

mm in diam. Fruit pseudosyncarpous, rarely syncarpous, green, brown to red

(in vivo), ovoid to transversely broadly ellipsoid to spheroid, to 8 cm long,

basal collar present and composed of up to 20 connate sterile carpels, mostly

broadly to shallovvly conical oroblong-ellipsoid, to 1 8 mm long, fertile carpels

30-350(-450), narrowly to broadly angular-obovoid, or obconical, tetragonous

to hexagonous, to 30(-35) mm long, apex Hat, truncate, conical to broadly

conical or uncinate, carpels fused over basal 1 /4 or fully fused, rarely completely

free. Seed íllling up3/5-6/7 ofthecarpel, narrowly to broadly ovoid orobovoid,

tawny, reddish-brovvn to dark brown, smooth, roughish or rugged.

Key to the species

1 . Lower side of lamina densely covered vvith contiguous scales even when

old 2

1 . Lower side of lamina, when old, sparsely covered vvith scales nottouching

each other or nearly glabrous 5

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Slndies In Annonaceae. XVI. A Toxonomic Revisinn ofDugiielia A. F C P de Sant-Hilaire

2. Small shrubs, usual ly no more tlian 3 m tall; lamina narrowly obovate to

obovate, bract below articulation more than 3 mm in length; carpels sborter

than 17 mm; seeds sborter than 11 mm; C, E, and SE Brazil.

1. D. furfuracea

2. Trees or rarely big shrubs, (2-)5-20(-30) m tall; lamina narrowly ovate to

ovate, narrowly elliptic to elliptic, bract below articulation sborter than 3

mm; carpels more than 1 7 mm long; seeds more than 1 I mm in length 3

3. Carpels at apical part straight (not contracted); seeds usual ly with 2-4

ribs 4

3. Carpels at apical 1/5 to 1/4 usual ly suddenly contracted and forming a

peltate top; seed without ribs; Amazonas, Pará, and Maranhao.

6. D. lepidota

4. Lamina narrowly ovate (to narrowly elliptic), upper side sparsely (to rather

densely) covered with stellate hairs; fertile carpels tetragonous, broadly to

transversely rhombic in transverse section, top smooth: seeds narrowly

ovoid to narrowly ellipsoid; C Brazil

4. D. marcgraviana

4. Lamina elliptic, oblong-elliptic (to narrowly ovate), upper side glabrous

from the very beginning; fertile carpels angular-obovoid to obconical, top

irregularly rugose; seeds obovoid; N and NE Brazil 5. D. echinophora

5. Upper side of lamina sparsely to rather densely covered with stellate hairs;

Bahia D. dicholepidota

5. Upper side of lamina glabrous, slightly shiny 6

6. Lamina rounded (-acute orapiculate) at the apex, rounded (-obtuse) at the

base 7

6. Lamina acute to acuminate at the apex, acute to attenuate at the base. . 8

7. Carpels densely covered with scales and stellate hairs, basal 1/2 to 3/5

fused; inner side of bracts densely covered with curviform stellate hairs;

apical prolongation of connective glabrous; ovary sparsely to rather densely

covered with curviform stellate hairs; Bahia and Sergipe.

3. D. moricandiana

7. Carpels glabrous, totally free from each other; inner side of bracts glabrous;

apical prolongation (4 connective puberulous, ovary glabrous; Brazil (Mato

Grosso) and Bolivia (Santa Cruz) 7. D. glabriuscula

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Boi. Mus. Para. Emílio GoehJi. sér. Boi. 9(2). 1993

8. Fertile carpels glabrous; ovary glabrous, style sparsely to densely covered

vvith erect stellate ha i rs; from Minas Gerais to Santa Catarina.

8. D. lanceolata

8. Fertile carpels sparsely to densely covered with stellate scales; ovary densely

covered with stellate scales and/or stellate liairs, upper part of style

glabrous 9

9. Lamina broadest at lhe middle, narrowing more or less equally toward

both ends; São Paulo 11. D. salicifolia

9. Lamina broadest belovv the middle, narrowing more quickly toward the

base than toward the apex 1 0

10. Lamina 2.5-4(-4.5) cm wide, index 2-3(-3.5), lower side sparsely (0-2

scale/mm2) covered with stellate scales; apical prolongation ofconnective

puberulous; top of carpels conical or uncinate, yellowish, densely covered

with stellate scales and liairs; from Ceará to Rio de Janeiro.

9. D. riede liana

1 0. Lamina ( 1 .2-) 1 .4-2 cm wide, index 4.5-6(-6.4), lower side rather densely

(3-9 scales/mm2) covered vvith stellate scales; apical prolongation of

connective glabrous; top of carpels truncate, blackencd, sparsely covered

with stellate scales; Rio de Janeiro 10. D. microphylla

I . Duguetia furfuracea (A. de St. Hilaire) Bentham et Hooker f., Gen. pl. I :

24. 1 862; R. E. Frics, Acta I lorti Berg. 12(1): 35. 1 934. Figure 4

Aimona furfuracea A. de St. flilaire, Fl. Bras. Merid. 1 : 34. t.6. 1 825. Martius,

Fl. bras. 13(1): 8. 1841. Type: Brazil. Minas Gerais: without locality,

1816-1821, A. deSaint Hilaire s.n., fl. (syntypes, P [a lectotype still lias

to bc selected]; duplicates in F, G, NY).

Duguetia coriacea Sonder, Linnaea 22: 557. 1849. Type: Brazil. Minas Ge-

rais: Caldas (fr), Regnell 11-3 (syntypc. O); Brazil. Minas Gerais: Uberaba,

“ad Formizas”, 13 Dcc 1848 (fl), Regnell 11-3 (syntype, S); Brazil. São

Paulo: Canna Verde, Apr 1 848 (tl), Regnell 11-3 (syntypes, MO, S); Brazil.

São Paulo: Batataes, Apr 1848 (tl), Regnell 11-3 (syntype, S).

Aberemoa furfuracea (A. de St. I lilaire) Baillon, 1 1 ist. pl. 1 : 204. 1 868; R. E.

Fries, Kongl. Svenska Vetenskapsakad. Handl., n.s., 34(5): 21. 1900.

150

Studies In Annonaceae. XVI. A Toxonomic Revision oj Dugneiia A. F. C. P. de Sant-Hilaire

A beremoa furfuracea (A. de St. Hilaire) Baillon var . jonasicma Barbosa

Rodrigues, Pl. mattogr. 7. t. 4. 1898. Type: Brazil. Mato Grosso: near

Rio Coxipó and Rio do Peixe, near Cuiabá, Jun I 8??, Barbosa Rodrigues

s. n. (not seen).

Aberemoa jonasiana (Barbosa Rodrigues) R. E. Fries, Ark. Bot. 4(19): 10.

t. l, f.9. t.2, f.6-8. 1905.

Duguetia jonasiana (Barbosa Rodrigues) R. E. Fries, Acta Horti Berg. 6(6):

16. 1919, 12(1): 38. 1934.

Duguetia henmendorffii R. E. Fries, Acta Horti Berg. 12(1): 37. 1934. Type:

Brazil. São Paulo: Santa Rita do Passo Quatro, Santa Albertina, 2 Feb

1900 (11), Henuuendorff278 (holotype and 3 isotypes, S).

Shrub, (0.5-) 1 - 1 ,5(-3) m tall. Young twigsdcnsely covered witli yellowish

fimbriate to stellatescales, becomingglabrous vvitli age, oldertwigs yellovvish-

brown to dark brown, irregularly striate. Petiole ( 1 .5-)2-4(-5) mm long, ( l-)l .8-

2.5 mm in diam., densely covered witli yellowish fimbriate scales. Lamina

narrowly obovate to obovate (narrowly oblong-obovate to oblong-obovate),

coriaceous, grcen to greenish-yellowabove, greenish-yellowto yellow below,

(6-)9- 1 4(- 1 7) cm long, (2-)3-5(-6) cm wide, index 2-3.5(-5.5), base narrowly

cuneate (broadly cuneate, attenuate or rarely obtiise), apex obtuse to acute

(acuminate or rarely rounded), uppcr side sparsely to densely covered vvith

yellowish. (6-)8-l 8(-20)-rayed appressed orerect stellate hairs and/or stellate

scales, becoming sparsely covered orglabrous vvith age (permanently hairy on

the base of primary vein), lower side densely covered vvith yellowish fimbriate

to stellate scales, secondary veins uniforinly curved, (9-) 1 0- 1 5(- 1 7) on either

side of primary vcin, angles vvith primary veins (50°)60 o -70 o (-85°). intercostal

area irregularly spaced, distancc betvveen the middle of closure and margin

( 1 -) 1 .5-4(-5) mm, tertiary veins slightly raiscd on upper side. flat to slightly

raised on lower side, irregularly forming ( l-)2-12(-16) nets, 4th order veins

slightly raised to flat on both sides, but nearly invisible on lower side because

ofthe rather densely covering of scales.

Inflorescence leaf-opposed, 1-2-fIowercd, subsessile, pedicel (7-)10-

1 5(- 1 7 ) mm long, rccurved, 1-2 mm in diam. at base, (3-)3.5-5(-6) mm in

diam. just below flower. fruiting pedicel (1 1-) 15-30 mm long, 2-3.5(-4)mm in

diam. at base, (3-)3.5-5(-6) mm in diam. below the fruit. Bracts2 to the pedicel,

one amplectant on basal 1/3 to l/2(-5/6) of pedicel, very broadly ordepressed

151

Boi. Mus. Paru Emílio Goeldi. súr. Boi. 9(2). 1993

ovate to subcirciilar, 3-5 mm long, 5-7 mm wide, easily broken, another one

triangular, 4-5 mm long, 2. 5-3. 5 mm wide. Pedicels and outer side of bracts

densely covered with yellowish fimbriate to stellate scales, inner side of bracts

rather densely to densely covered witli curviform stellate liairs; outer side of

sepals densely covered with yellowish fimbriate scales, inner side densely

covered with curviform stellate liairs (at basal 1/5 to 1/3 usually sparsely

covered); outer side of petals rather densely to densely covered with yellowish

fimbriate to stellate scales, inner side densely covered with curviform stellate

liairs (basal 1/5 to 2/5 of outer petals and 1/3 to 1/2 of inner petals glabrous).

Flower buds broadly ovoid to subglobose, apex obtuse to rounded, (8-)9-

12 mm long, 8-12 mm in diam. Flowers green, yellow or red, with dark red at

inner base (in vivo). Sepals connate to 1/5- 1/4 from base, ovate to broadly

ovate (triangular to broadly triangular), (7-)l 0-1 5(-22) mm long, (5.5-)9-

1 2(- 1 4) mm wide. Outer petals elliptic to rhombic-ell iptic, apex acute, ( 1 0-) 1 5-

20(-30) mm long, 6-9(- 1 2) mm wide, inner petals rhombic-el I iptic to elliptic,

apex acute, ( 1 0-) 1 6-20(-3 0) mm long, 6-9(-13) mm wide. Torus depressed

ovoid. Stamens 150-300, 0.9- 1.2 mm long, 0.5-0. 6 mm wide, apical

prolongation of connective very shallowly conical, puberulous, 0.2-0.4(-0.5)

mm long, 0. 7-0.9 mm wide. Carpels 40-150, ovary and stigma sparsely to

densely covered with erect stellate liairs.

Fruiting receptacleellipsoid (oblong-ellipsoid), 15-20 mm long, 5-8 mm

in diam. Fruit green lo greyish-green (greenish-brown) (in vivo), spheroid

(ellipsoid), (2-)3-5(-7) cm in diam., the basal collarcomposed of 1 0-1 6 connate

sterile carpels, (broadly conical) deltoid, 6-12 mm long. 6-14 mm in diam.,

covered with some scales and stellate liairs, fertile carpels (40-)70-l40.

obconical. 10-15 mm long, 1 0- 1 4(- 1 7) mm in diam., with 4-5 ri bs or without

rib, top itself flat, depressed ovoid to conical, densely covered with scales and

stellate liairs, fused in the basal (2/3-)5/6 tototally. Seed filling up to (2/3-)5/

6 of the carpel, obovoid to broadly obovoid, 9- 1 1 mm long, 7-9 mm in diam.,

ycllowish-brown to reddish-brown.

Distribution (Figure 1). C, S, and SE Brazil (Mato Grosso, Paraná,

Distrito Federal, São Paulo, Minas Gerais, Bahia, and Goiás) to NE Brazil

(Ceará), also in Paraguay and Bolivia. Typical pyrophyte. Iiighly adapted to

Central Brazilian savannas. Common, particularly in various cerrado typcs

(campos rupestres, arborcous cerrado, open grassy to shrubby campos) or in

dense secondary forest, at elevation from 200 to 1400 m. but mainly restricted

152

Studies tn Armonaceae . XVI. A Toxonomic Revision of Dugiielia A. F. C. P. de Sanl-Hilaire

to deep white or brovvn sandy soil, rarely on red clay or lateritic soil. It is a

smai I slirub with several-many ascending stems whicli sprout froin a woody,

elongate xylopodimn. Flowering and fmiting all the year aronnd, but mostly

flovvering from Novemberto May and fruiting from Septemberto June.

Specimens examined. Bahia: Espigão Mestre, 23 km W of Barreiras,

680 m, 3 Mar 1972 (fl), W. R. Anderson et al. 36544 (AAU, NY, U); 16 km

NW of Lagoinha on side road to Minas do Mimoso, 950-1000 m, 4 Mar 1 974

(fl), Harley et al. 16717 (CEPEC, K, MO, NY, P, U, US); 15-20 km from

Andarai, along the road to Itaeté wliich branches E off the road to Mucugé,

500-600 m, 13 Feb 1977 (fl & fr), Harley et al. 18634 (CEPEC, F, K, MO,

NY, U, US); lowerNE slopes of the Pico das Almas, 25 km WNW ofthe Vila

do Rio de Contas, 1400 m, 20 Mar 1977 (fl), Harley et al. 19748 (CEPEC,

M, N Y, U, US); Rio Piau, 225 km S W of Barreiras, 850 m, 1 2 Apr 1 966 (fl),

ínvia et ai 14639 (F, NY, S, SP, US); 5 km S of Rio Roda Velha, 900 m, 15

Apr 1966 (fl), Irwin et ai 14870 (F, NY, S, SP, US); road to Agua de Rega, 23

km N of Seabra, 1 000 m, 24 Feb 1 97 1 (fl), Irwin et al. 30914 (F, MO, NY, U,

US); Mun. Formosa do Rio Preto, 550 m, 8 Apr 1989 ( fl), Mendonça et ai

1374 (U); Highway-BA-265, 9 km E of Victória da Conquista, 900 m, 4 Mar

1978 (fl), Mori et ai 9437 (CEPEC, U); Mun. Caetité, 2 km S ofCaetité, 800

m, 19 Mar 1980 (11), Mori & Benton 13472 (CEPEC, U); Mun. Piatã, Piatã,

1 100 m, 3 Mar 1980 (11 & fr), Mori & Fitnch 13369 (CEPEC, NY); Mun.

Formosa do Rio Preto, 7 Apr 1989 (fl), near Rio Riachao, Walter et al. 210

(IBGE, U). Ceará: Serra do Araripe, 12 Km SE ofCrato, 800 m, 14Jun 1944

(fr), CutlerSl 16 (F, US); top of Chapada de Araripe, 10 Km S of Crato, 950

m, 1 3 Feb 1 985 (fr), Genlry et al. 50040 (G, M, U); Taboca, Serra do Araripe,

1 4 May 1 934 (fl), von Liietzelburg 26061 (M, N Y). Distrito Federal: Brasília,

Bacia do Rio São Bartolomeu, 15 Apr 1980 (fl & fr), Heringer et al 4372

(MO, U, US); Brasília, 6 km from Cachoeirinha, 14 Jan 1981 (fl), Heringer et

al 5971 (K, MO, U, US); Brasília, Convênio Florestal, 10 Feb 1962 (fl),

Heringer 8868 (NY); Brasília, Horto do Guará, 24 Apr 1974 (fl), Heringer

13250 { U); Brasília, Monumento do Candango, 12 Nov 1980, Heringer 17966

(NY); 20 km S of Brasília, on road to Belo Horizonte, 700-1000 m, 26 Aug

1964 (fl), Irwin & Soderslroin 5572 (NY, S, SP). Goiás: Serra dos Cristais,

12 km N of Cristalina, 1060 m, 3 Apr 1973 (fl), W. R. Anderson et al. 8025

(F, MO, NY, U, US); 26 Km W of Rio Verde, 7 Jul 1966 (fl), Goodland 482

(NY); 6 km Sof Cristalina, 1 175 m,5Nov 1965 (fl ), Irwin et al. 9952 (F,GH,

MO, NY, S, SP); 5 km E of Goiás boundary on road to Guarapauá, 1000 m,

16 Nov 1965 (fl), Irwin et al 10295 (F, MO, NY, S, SP); 3 km N of Cristalina,

153

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Boi. Mus. 1‘ara. Emílio Goeldi. sér. Boi. 9(2). 1993

1250 m, 2 Mar 1966 (fl), Invin et al. 13266 (F, MO, NY, SP); Rio da Prata,

6 km S of Posse, 800 m, 4 Apr 1966 (fl & fr), Invin et al. 14357 (NY, S, SP);

70 km SE of Aragarças, road to Piranhas, 700 m, 23 Jun 1966 (fr), Invin et al.

17646 (F, MO, NY, S, US); 45 km S of Caiapônia, road to Jataí, 900 m, 28

Jun 1 966 (fr), Invin et al. 7 7973(F,NY, S,SP,US); 15 km SofNiquelândia,

1000 m, 21 Jan 1972 (fl), Invin et al. 34693 (NY); Mim. Piranhas, 30 km N

of Piranhas, 26 Feb 1982 (fl). Oliveira & Anderson 475 (MICH, U). Mato

Grosso: Belo Horizonte, Cardoso, 20 Nov 1932 (fl), Barreto 420 (RB);

Aquidauna, 23 Jul 1972, de Jesus 1 730 (RB); Mim. Barra do Garças, 7 km N

of Córrego Tangúru, 12 Dec 1969 (fl), Eiten & Eiten 9890 (SP, US); Mim.

Rio Brilhante, Fazenda Bela Vista, 25 Jan 1971 (fl), Hatschbach 26118 (C,

UC); Mim. Campo Grande, road from Campo Grande to Rochedo, 12 Jul

1969 (fl & fr), Hatschbach & Guimarães 21837 (S, UC); Mim. Nova

Andradina, Douradinho, 12 Apr 1972 (st), Hatschbach 29419 (UC); Mun.

Bataguaçu, Rodovia BR-265, 6 Feb 1975 (fl), Hatschbach et al. 35904 (U);

20 km N of Amambai, Rodovia MT-642, 16 Dec 1983 (fl), Hatschbach &

Callejas 47293 (U); 20 km S of Garapú, 300-400 m, 30 Sep 1964 (fr), Invin

& Soderstrom 6491 (F, GI I, NY, P, S, SP); Rio Turvo, 2 1 0 km N of Xavantina,

500 m, 28 May 1966 (fr), Invin et al. 16206 (F, NY, SP, US); 86 km N of

Xavantina, 550 m, 3 Jun 1966(11 & fr), Invin et al. 16528(¥ , NY, S, SP. US);

80 miles out of Campo Grande on road to São Paulo, 500 m, 24 Nov 1 959 (fl),

Maguire & Maguire 44529 (NY); 370 km W of Cuiabá, 26 Sep 1963 (fr),

Maguire et al. 56853 (NY); Cuiabá, 17 Jun 1902 (st), Malme 1779a (S);

Mun. Rio Pardo, 700 m, Martinelli 402 (RB); Mun. Chapada dos Guimarães,

3 km W of Buriti, 600 m, 12 Jul 1984 (fr), Mori et al. 16717 (CEPEC, U);

Mun. Rio Brilhante, Posto Zuzu, 6 Sep 1979 (fl), P. I. Oliveira 9(U); Mun.

Santana do Riacho, Mãe d’ Agua, along road from Belo Horizonte to Conceição

do Mato Dentro, 1 2 Jan 1981 ( fl), Rossi et al. CFSC6985 (K); highway from

Campo Grande to Aquidauana, 10 km from Fazenda Leão, 14 Dec 1976 (st),

Shepherd et al. 4133 (RB); Mun. Três Lagoas, Mar 1960 (st), Válio 60 (NY,

SP). Minas Gerais: Serra do Espinhaço, 6 km N of Gouveia on road to

Diamantina, 1250 m, 10 Apr 1973 (fl), W. R. Anderson et al. 8585 (F, MO,

NY, RB, U, US); Vicinity of Mendanha, 24 Sep 1936 (fr), Archer 4069 (A,

NA, RB); between Patos and Pirapora, 17 Sep 1963 (fr), Castellanos 24190

(GUA); Serra de Lenheiro, São João Del-Rei, Jan 1960 (y 11), Duarte 5136

(NY, RB); Belo Horizonte, Leitão, 5 Feb 1919 (fl), Gehrt SP3200 (NY. S.

SP); Turvo, 24 Apr 1926 (11), Gehrt SP 17501 (S); N of Uberaba, 426.8 km

from Brasília, 9 May 1966, Goodland 326 (NY, U); Mun. Prata, Rio

Douradinho, 23 Jul 1 977 (fr), Hatschbach 40053 (NA); between Brasília and

154

Studies In Annonaceae. XVI. A Toxonomic Revision of.Duguetia A F. C. /’. de Sam-Hllaire

João Pinheiro, 30 Aug 1979 (st), Heringer A Rizzini 17419 (U); Serra do

Espinhaço, 28 km SW of Diamantina on road to Gouveia, 1300 m, 15 Jan

1961 (íl). Ir win et ai 22017 { MO, NY, U); 25 km W of Monte Claros, road to

Agua Boa, 1 000 m, 23 Feb 1 969 (fl), Irwin et ai 23699 ( AAU, NY, RB, U);

25 km NE of Patrocínio, 1 050 m, 28 Jan 1970 (fl), Irwin et al. 25488 ( F, MO,

NY, U); Serra da Anta, 2 km NW of Paracatú, 700 m, 7 Feb 1970 (fl), Irwin

et al. 26312 (F, MO, N Y, U, US); Rio Bicudo, 20 km W of Corinto, 525 m, 3

Mar 1970 (fl), Irwin et al. 268 1 7 (F, MO, NY, RB. U, US); Serra do Espinhaço,

25 Km E of Diamantina, near Rio Jequití, 800 m, 15 Mar 1970 (fl), Irwin et

al. 2756>9 (11BG, NY, U); 10 km S of São João da Chapada, 1 150 m, 27 Mar

1 970 (fr), Irwin et al. 28427 { F, MO, NY, RB, U, US); 53 km E of Araxá, Rio

Quebra Anzol, 30 Jan 1 978 (fl), Krqpovickas et al. 33360 ( WIS); Mun. Grão

Mogol. road to Francisco Sá, 10 km from Grão Mogol, 10 May 1979 (fl), II.

C. Lima et al. 969 (RB. U); 2 km from Alfenas, 26 Feb 1963 (fl), Magalhães

11 (SP, U); Mun. Prata, 2 km from Prata. 19 Mar 1963 (fl), Magalhães 52 (P,

RB, S, SP, U); Mun. Patrocínio, Fazenda Grão de Ouro, 28 Feb 1989 (fl),

Mendonça et al. 1179{\3)\ Pirapora, 5 Mar 1 962 (íl), Rizzini RBJ14058 (RB);

Lavras, 15 Jan 1914 (fr), Shatnel et al. 255/;(US). Paraná: Itararé, 16 Apr

1910 (fl), Dusén 9667 (G, MO, S); Jaguariaiva, 19 Apr 1910 (fl), Dusén

9731 (F, GH, MICH, MO, NY. S): Mun. Paraguaçu Paulista, 4 km N of

Paraguaçu Paulista, 500 m, lOFeb 1 965 (fl), Eiten et al. 6024 (MO, NY, SP,

US); Mun. Cianorte, Fazenda Lagoa, 28 Apr 1966 (fl), Hatschbach 14251

(NY, P, U, US, WIS); Campo Mourão, 8 Dec 1965 (fl), Hatschbach et al.

132 76 (U); Jaguariaiva, 19 May 1914 (11), Joensson 386a (S, Z); Fazenda

Lagoa, S of Rio 1 vai, 1 5 km E of São Tomé, 5 Apr 1 966 (fl), Lindeman A de

Haas 894 (C, F, NY, U, W); 3 km N of Campo Mourão, 26 Jan 1967 (fl),

Lindeman A de Haas 4500 (NY, U). Rondônia: Vilhena, 4 Jan 1979 (fl), M.

G. Silva A A. Pinheiro 4136 (NY, U, US). São Paulo: Loreto, Dattistella

213=SPI451 1 (S, SP); Jeriquara, 26 Mar 1967 (fl, fr), Bicalho 30 (SP);

Moóca,20 Apr 1913 (fl, fr). Brade 5882 (S, SP); Braste, Jacguariuna, Brantjes

701607 (U); Fazenda Paulistas, Matão, 24 May 1955 (st), Dedecca 5 1 4 (UC);

Mun. São Carlos, 5 km NW of São Carlos, 28 Jan 1961 (st), Eiten A Eiten

2535 (SP); Mun. Brotas, N W of the intersection of Brotas-ltirapina road vvith

thc road to Campo Alegre, 750 m, 16 Jun 1961 (fl), Eiten et al. 2962 (SP);

Barretos, Rio Pardo, Nov 1917 (st), Frazão RB8659(S)\ São Miguel, 19 Jan

1 944 (fl), Gonçalves SP5036I (SP); Moj i-Guaçú, 27 Apr 1 979 ( fl), O. Handro

2294 (SP, US); Santa Rita do Passa Quatro, Hemmendorff 279 (S); Santa

Rita do Passa Quatro, 27 Nov 1981 (fr), Kirizawa A Morretes 608 (SP);

Anhembí, Fazenda Barreiro Rico, 2 May 1959 (11), M. Kuhlmann 4524 (SP);

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Boi. Mus. Para. Emílio Goe/cli, sér. Boi. 9(2). / 993

Mun. Itapetininga, km 1 50 of road Raposo Tovares, 1 7 Jul 1 962, Labouriau

113 (SP); São Simão, 6 Feb 1 964 (fl), Labouriau & Válio 1098 (SP); Sorocaba,

Dec 1817 (fl), Martins 550 (M); Serra do Botucatú, 1 4 Mar 1 967 (fl), Mattos

& Mattos 14427 (SP); Sorocaba, 30 Jun 1 934 (fr), Mendes 1 (SP); Itú, 1 5 Feb

[noyear] (fl), Russel 286 (S, SP); Fazenda Hollambra, 35 km N of Campinas,

600 m, 25 Feb 1976 (fl), Shepherd & Gibbs 11246 (K, NY); Fazenda Graci-

osa, betvveen Sao Carlos and Descalvado, 2 Feb 1966 (fl), Siqueira 8 (SP);

Jundiaí, 3 Feb 1907 (11), Usteri 3a=SP 17878 (K, S, SP); Mun. Itirapina, 5 km

from Rodovia Washington Luiz, near road from Itirapina to Rio Claro, 1 1 Apr

1 962 (fl), Válio 269 (P, RB, SP).

Vernacular Names. Araticum, Araticum-do-campo, Araticum-rasteiro,

Araticum-vermelho, Araticunzinho/Araxicunzinho, Araticum lanceta, Ata, Ata

do Campo, Aticum, Bruto, Moroua/alathê, Orelha-de-burro, Pinha braba. Pi-

nha brava. Pinha do Campo.

This is the most widesprcad and common species of the group. It is also

the most variable one, especial ly in the shape and size ofthe leaves. The shape

of the lamina is mainly narrowly obovate to obovate, sometimes very narrovv,

the leaf index mainly falis into the range from 2 to 3.5, buton some specimens

it is much higher (5-6). Generally speaking, most collections from Paraguay

have narrowerand smaller leaves than those from Brazil. But it is hardly possible

to give thcm a formal taxonomic rank to express the infra-specific differentiation

within D. furfuracea because intermediate forms can bc found easily.

Based on the narrovv leaf shape and the glabrescent upper side ofthe

lamina, Fries distinguished D. hemmendorffii from D. furfuracea, be it vvitli

some reluctance. D. jonasiana was distinguished from D. furfuracea on the

baseof its narrovv leaves vvith the upper side glabrous from the very beginning,

and its narrovv petals.

In this paper, the three spccies are considered to be conspecific. In overall

resemblance, D. hemmendorffii is within the variation range of D. furfuracea,

and there is no reason for keeping it apart.

The inclusion of D. jonasiana may need some more explanation. The

main distinctivc feature of D. jonasiana appears to be the upper leaf surface

vvhich is glabrous from the very beginning. In typical D. furfuracea the leaves

are sparsely to densely covered vvith stellate scales. In D. jonasiana, hovvever,

the primary vein does appear to be covered with some hairs, too. In D.

156

Studies In Annonaceae. XV! ,1 Toxonomic Revisioit of Duguetia A. F. C. P deSant-llilaire

fiirfuracea, on lhe other hand, leaves may become glabrous above witli age.

The diíTerence, therefore, is only small, and certainly does not provide suffleient

reason for maintaining two species. As for other characters, we vvere unable to

find any differences that could support a specific statiis for D. jonasiami, and,

consequently, we united it with D. fitijuracea.

2. Diigiielia dicholepidota Martins, Fl. Bras. 13(1): 22. 1841; R. E. Fries,

Acta Horti Berg. 12(1): 43. 1 934. Type: Brazil. Bahia: Serra do Açurua,

Sertão de Rio San Francisco, 1 839 (fl & fr), Blanchet 2828 (lectotype,

G, selected here; isolectotypes, B, BM, F, G, K, NY, OX, P,). Figure 5

Shrub to 3 m tal I, young twigs densely covered with yellowish stellate

scales, becoming glabrous with age, older twigs greyish brown to dark brown,

irregularly ribbed. Pctiole 2-3(-4) mm long, (1.2-)1.5-2(-2.5) mm in diam.,

(sparsely to) densely covered with stellate scales. Lamina elliptic ovate to

narrowly ovate. coriaceous. dark glossy grecn above, green below, (5-)7- 1 0(- 1 2)

cm long, (2.2-)3-4 cm wide. index 2-2.5(-2.9). base rounded to obtuse (acute),

apex acute to acuminatc (acumen 5- 1 0 mm long), upper side sparsely to rather

densely covered with whitish, (6-)7- 1 5(- 1 7)-rayed appressed or erect stellate

hairs, becoming glabrous with age (primary vein permanently covered with

stellate hairs), lower side densely covered with yellowish to whitish stellate

scales, becoming sparsely covered with age, secondary veins abruptly curved,

(8-) 1 0- 1 4(- 1 6) on either side of primary vein, angles with primary veins

(60 o -)65 o -80 o (-90°), intercostal area somewhat regularly spaced, distance

between the middlc of closure and margin 2.5-4(-5) mm, tertiary veins slightly

raised on botli sides, irregularly forming 5-1 1 nets, 4th order veins slightly

raised on upper side. nearly invisible on lower side.

Inflorescence leaf-opposed. 1 -2-flowered, sessile, pedicels (5-)8- 1 2(- 1 4)

mm long, l-l.5(-1.8)mm indiam.at base, ( 1 .5-)l ,8-2.2(-3) mm indiam. just

below (lower, fruiting pedicel 11-17 mm long, 1.5-2. 5 mm in diam. at base,

2. 8-3.8 mm indiam. just below the fruit. Bracts 2 to the pedicel, one amplectant

on basal 1/3 to 1/2 of pedicel, transvcrsely broadly elliptic, apex mucronulate

with a 1-1.5 mm long tip, 3-6(-8) mm long, 3-4.5(-6) mm wide, anotherone

shallowly triangular, 2-3 mm long, 3-4 mm wide. Pedicels and outer side of

bracts densely covered with ycl lowish I imbnate to st cl late scales, i nner side ol

bracts densely covered with curviform stellate haif s, outer side of sepals densely

covered with yellowish I imbriatc to stellate scales, inner side densely covered

with curviform stellate hairs (at basal 1/5 to 1/3 usually sparsely covered).

157

Hol. Mus. Para. Emílio Goelcli. sér. Boi. 9(2). 1993

outer side of petals ratlier densely (to densely) covered witli yellowish stellate

scales, inner side densely covered with curviform stellate hairs (basal 1/5 of

outer petals and 1/3 of inner petals glabrous).

Flower buds very broadly ovoid, apex obtuse to mucronate, 5-7 mm

long, 5-6.5 mm in diam. Flowers yellowish green (in vivo). Sepals connate to

1/5- 1/4 from base, (broadly) triangular to (broadly) ovate, apex acute to

mucronate, 11-15 mm long, 8-1 1 mm wide. Outer petals rhombic-el I iptic, apex

acute (mucronate), 1 6- 1 9(-2 1 ) mm long, 10-11.5 mm wide, inner petals

rhombic-el I iptic to e 1 1 iptic, apex acute to mucronate, 1 4- 1 8(-22) mm long,

(9-) 1 0-12 mm wide. Torus depressed ovoid. Stamens 200-280, 1.1-1 ,4(-l .6)

mm long, (0.3-)0.4-0.5(-0.7) mm wide, apical prolongation of connective

shallowly to very shallowly conical, puberulous, (0. 1 -)0.2-0.3(-0.4) mm long,

0.7-0. 8 mm wide. Carpels 1 20- 1 50, stigma sparsely covered with curviform

stellate hairs and ovary sparsely to ratlier densely covered with curviform’

stellate hairs.

Fruit greyish-green (in vivo), spheroid, ea. 3 cm in diam., the basal collar

composed of 1 0-14 connate sterile carpels, deltoid, ca. 7 mm long, 5-6 mm in

diam., ratlier densely covered with scales and stellate hairs. Fertile carpels ca.

80, obovoid to angular-obovoid, 11-13 mm long, 6.5-9 mm in diam., with a

0.5-1 mm long unciform lip, densely covered with stellate scales and stellate

hairs, fused in the basal 1/4 to 1/3, apical free parts spacing 1-2.5 mm apart.

Seed filling up to 5/7 to 6/7 oftlie carpel. ellipsoid, 9-1 I mm long, 5-6 mm in

diam., smooth, tawny to reddish brown.

Distribution (Figure 3). Only known from Açuruá, E Bahia; in caatinga

vegetation.

Specimen examined. Bahia. Serra do Açuruá, 12 km E of Gentio do

Ouro on road to Boa Vista and Ibipeba, 500-700 m, 22 Feb 1977 (11 & fr),

Harley et ai 18917 (CEPEC, K, M, NY, U).

Fries's referring to “ Duguetia dolicholepidota Mart.” (Fries 1959) is

obviously an error for D. dicholepidota Mart.

Owingto the strong similarity in flowers and fruits with D. furfuracea,

D. dicholepidota sliould be placed near D. furfuracea, ratlier than near D.

marcgraviana as was suggested by R. E. Fries (1934). One might wonder if

the two collections of D. dicholepidota , known from the type locality only.

158

ShicJies In Annonaceae. XI I .1 / oxonomic Revision ofDugiietia A. /•' C. P. de Sant-HHaire

justify the maintenance of a separate species. But it can be d istingu ished from

D. furfuracea by the sparse indument on tlic lovver side of the mature lamina,

the abniptly curved secondary veins, the more regularly spaced intercostal

areas, a larger angle between secondary veins and the primary vein, and by the

friiit with carpels only fused in the basal l/4th to l/3rd part.

3. Duguetia moricandiana Martius, Fl. Bras. 13(1): 22. 1841; R. E. Fries,

Acta Horti Berg. 12(1): 44. l934.Type: Brazil. Bahia: without location,

1 834 (fl & yoimg fr), Blunchet 1678 (holotype, G (2 sheets); isotypes,

BM, F). Figure 6

Tree or shrub, (0.4-)3-5 m tal I. Yoimg twigs densely covered with

yellowish fimbriate scales becoming glabrous with age, older twigs greyish-

white, ribbed. Petiole darkened, 2-6 mm long, 1 .5-2.5 mm in diam., sparsely

covered with fimbriate scales, sometimcs nearly glabrous. Lamina elliptic to

oblong-el 1 iptic (obovate), coriaceous, green to dark green above, brownish-

green below. (4-)5.5- 1 ()(- 1 2.5) cm long, (2.5-)3-5.5(-7) cm wide, index

1.5-2(-2.2), base rounded (obtuse), apex rounded (apiculate or mucronate),

iipper side essentially glabrous, lovver side densely covered with stellate scales

when young and becoming sparsely covered (on primary vein permanently

densely covered), secondary veins abruptly curved. (5-)7- 1 0(- 1 2) on either

side of primary vein, anglcs with primary vein 50°-70°(-75°), intercostal area

somewhat regularly spaced, distance between the middle of closure and margin

4- 8 mm, tertiary veins conspicuous, raised on both sides, irregularly forming

5- 1 0(- 13) nets, 4th orderveins subconspicuous.

Inilorescencc leaf-opposed or pseudo-terminal, directly attached to the

twigs or on the top of a 5-8 mm long rudimentary twig, simple, 1 -2-flowered.

Pedicels 8-10 mm long, 1-1.5 mm in diam. at base, 2-2.5 mm in diam.just

below flower, fruiting pedicels 11-18 mm long, 2-3.5 mm in diam. Bract(s)

1-2 to the pedicel, oneon basal 1/3 to 1/2 of pedicel, (deltate) broadly ovate,

1-2 mm long, 1-1.8 mm wide, another one (if present) triangular, 0.5-1 mm

long, 0.3-0. 8 mm wide. Indument of floral parts: peduncle, pedicel and outer

side of bracts rather densely to densely covered with yellowish fimbriate scales,

inner side of bracts densely covered with curviform stellate hairs; outer side of

sepals densely covered with yellowish fimbriate scales, inner side densely

covered with whitish curviform stellate hairs; outer side of petals rather densely

to densely covered with yellowish stellate scales, inner side densely covered

with curviform stellate hairs.

159

1, | SciELO

Rol. Mus. Beira. Emílio Goelcli. sér. Rot. 9(2). 1 993

Flower buds broaclly ovoid, apex mucronate vvitb a ca. 2 mm long apicule,

8-10 mm long, 7-9 mm in diam. Flowers greenish-white to yellow (in vivo).

Sepals connate to 1/4- 1/3 from base, broadly triangular-ovate, apex acute to

acuminate with a 1.5-2. 5 mm long acumen, 9-12 mm long, 5-9 mm wide.

Outer and inner peta Is almost the same, elliptic (obovate), (7-)9-13(-l 5) mm

long, 5-6.5(-9.5) mm wide. Torus depressed ovoid, vvitb an ovoid apicule.

Stamens 200-250, 1 . 1 - 1 .3 mm long, 0.4-0. 5 mm wide, apical prolongation of

connective very sballowly triangular, glabrous, 0. 2-0.4 mm long, 0.4-0. 6 mm

wide. Carpels 40-80. stigma and ovary sparsely to rather densely covered with

curviform stellate liairs.

Fruit unknown (only loose carpels present). Fertile carpels >30, broadly

obtrulloid with 5-6 ribs, 14-16 mm long, 7-9 mm in diam., apex forming a ligneous

cap 6- 1 0 mm long, 8- 1 0 mm across, the top of the cap densely covered with scales

and stellate liairs, with a I -2 mm long curved apicule, basal 1/2 to 3/5 of carpels

fused. Seeds filling up to 5/6 of the carpcl, obovoid (broadly obovoid), 10-13 mm

long, 7-9 mm in diam., dark brovvn, smooth and slightly shiny.

Distribution (Figure 3). Only known from the Brazilian States Bahia and

Sergipe. Growing in dunesand restinga. Floweringand fruitingall theyeararound.

Specimen.s exainined. Bahia: Mun. Salvador, Alamedas da Praia, between

Itapuã and airport, 25 Jan 1977 (fr), ./. S. Araújo et al. 135 (CEPEC); Mun.

Salvador, 3 km from Salvador, W of airport, 12 Nov 1983 (fl), Callejas c£

Carvalho 1744 (CEPEC, U); Mun. Salvador, 30 km N of Salvador, near airport,

2 Jan 1 982 (fl). Carvalho et al. 1092 (CEPEC); Itapuã near Lagoa de Abaeté,

Salvador, 1 7 Jun 1985 ( fl), M. L. Guedes et al. 15826 (CEPEC); Mun. Salva-

dor, Itapuã, near airport Dois de Julho, 23 May 1981 (11), Mori et al. 14078

(CEPEC, NY); Km 4 ofthe highway from Bali ia to Rio de Janeiro, Jaguaquara,

6 Jul 1971 (fr), Pinheiro 1420 (CEPEC): Jaguaquara, km 6 of the highway

from Bahia to Rio de Janeiro, 3 Oct 1 972 (fl). Pinheiro 1961 (CEPEC); Mun.

Salvador, along road from Itapuã to airport Dois de Julho, 27 Jan 1983 (fl),

Plowman 12783 (CEPEC, F, U). Sergipe: Piramba, 19 Nov 1 974 ( 11), Fonseca

131 (RB).

Although closely related to D. furfiiracea, tliis species can easily be

distinguished from D. Jiirfuracea by the sliape ofthe leaves and their indument.

lt should be pointed out, tliat the fruit is poorly known. Only individual carpels

of one collcction (,/. S. Araújo 163) wcre available.

160

Sludies In Annonaceae. XI 7. .1 Toxonomic Revision ofDugnetia A. F. C. P. de Sant-Hilaire

4. Duguetia marcgraviana Martins, Fl. Bras. 13(1): 25. 1841; R. E. Fries,

Acta Horti Berg. 12(1): 42. 1934. Type: Brazil. Mato Grosso: location

unknown (“ in locis tempore pluvioso inundatis pr. Mato-Grosso “), Feb

1 828 (st), Riecfel 1471 (lectotype, LE, selected here a collection annotated

by Martins; isotypes, BR, LE, S). Figure 7

Aberenwa marcgraviana (Martius) R. E. Fries, Kongl. Svenska

Vetenskapsakad. Fiandl., n.s., 34(5): 20. 1900.

Duguetia sanctae-cmcis S. Moore, Trans. Linn. Soc. London, Bot. ser.2. 4:

299. 1895. Type: Brazil. Mato Grosso: Santa Cruz, Nov 1891 (fr), S.

Moore 576 (holotype, BM; isotypc, B).

Aberemoa sanctae-crucis (S. Moore) R. E. Fries, Kongl. Svenska

Vetenskapsakad. 1 landi., n.s., 34(5): 22. 1900.

Aberemoa brevipeáunculata R. E. Fries. Ark. Bot. 4(19): 8. 1. 1, f. 10. 1905.

Type: Brazil. Mato Grosso: Santa Anna da Chapada, 18 Sep 1902 (fl,

fr), Malme 11-2322 (holotype & isotype, S).

Duguetia brevipedimculata (R. E. Fries) R. E. Fries, Acta Horti Berg. 6(6):

16. 1919, 12(1): 40. 1934.

Tree (or rarely shrub), (2-)3-l 5(-25) m tall. stems 1 0-25(-30) cm in diam.

Young tvvigs densely covercd vvith yellowish flmbriate scales, becoming sparsely

covered with age, older twigs greyish-white, yellowish-brovvn to dark brown,

fissured. Pctiole (2-)3 .5-6 mm long, 1-2 mm in diam., densely covered with

yellowish fimbriate scales. Lamina narrowlyovate to narrowly elliptic (elliptic),

coriaceous, light green to dark green above, pale green below, (8-) 1 0- 1 8(-23)

cm long, 3-5.5(-6.5) cm wide, index (2.7-)3-4.5. base acute to slightly attenuate

(rounded), apex acuminate to acute, upper side sparsely to rather densely

covered with 5-8(-12)-rayed, appressed and/orerect stellate hairs becoming

glabrous and slightly shiny with age, lower side densely covered with yellowish

fimbriate scales and whitish stellate scales, secondary veins uniformly curved,

( 1 0-) 1 3- 1 8(-22) on either side of primary vein, angles with primary vein

(45-)50-65(-75), intercostal area irregularly spaced, distance between the middle

of closures and the margin 2-5(-7) mm, tertiary veins slightly raised on both

sides, irregularly forming (2-)5- 1 2(- 1 5) nets, 4th order veins slightly raised on

upper side, slightly raised to flat on lower side.

161

Boi. Mus. Paru. Emílio Gockli. ser. Boi. 9(2). 1 993

I nflorescence leaf-opposed or pseudo-terminal, 1 -3-flowered, subsessile

or on tlie top of a 2-5 mm long, ligneous rudimentary twig, pedicel

(4-)10-25(-30) mm long. straight (recurved), thickened towards flower, 1-2

mm in diam. at base, (1 ,8-)2-3.5(-4.2) mm indiam.just belovv flower, fruiting

pedicel (1 2-) 15-30 mm long, 2-3(-4) mm in diam. at base, (4.5-)5-7 mm in

diam. just belovv fruit. Bracts 2 to the pedicel, one amplectant on basal 1/2 to

3/4 of pedicel, depressed ovate to subcircular, 5-7 mm in diam., anotber one

broadly triangular, 2-3 mm long, 2. 5-3. 5 mm wide. Pedicels and outer side of

bracts denscly covered witli yellowish fimbriate scalcs, inner side densely

covered witli curviform stellate liairs; outer side of sepals densely covered

witli ycllowisb fimbriate scales, inner side (rather densely) densely covered

with curviform stellate liairs (basal ca. 1/5 sparsely covered to glabrous); outer

side of petals densely covered witli yellowish fimbriate to stellate scales, inner

side rather densely to densely covered with curviform stellate liairs (basal 1/4

to 1/3 of outer petals and basal 1/3 to 1/2 of inner petals glabrous, striate).

flower buds broadly ovoid, apex acute to rounded, 5.5-8 mm long, 6-9

mm in diam. Flower yellowish-green, pinkish-red (purple) with dark red inner

base. (in vivo). Sepals connate to 1/6- 1/4 from base, broadly to very broadly

ovate(-deltate), 1 1 - 1 5(- 1 9) mm long, 1 0- 1 4(- 16) mm wide, apex acute (obtuse).

Outer petals rhombic-obovate to spathulate (broadly rhombic-obovate), apex

acute, at basal 1/4 to 1/3 usually contracted, (1 5-)l 8-30(-45) mm long,

(8-) 10-1 5(- 1 7) mm wide, inner petals rhombic-obovate (spathulate), apex acute,

at basal ca. 1/4 to 1/3 contracted, (18-)20-35(-45) mm long, (8-)l 2- 1 5(- 1 7)

mm wide. Torus very depressed ovoid. Stamens 100-200, 0.9- 1.4 mm long,

0.3-0. 4 mm wide, apical prolongation of connective (very) shallowly conical,

(0. 1 -)0.2-0.3 mm long, 0.4-0. 5 mm wide. Carpels 200-380, stigma and ovary

densely covered with curviform and/orerect stellate liairs.

Fruiting receptacle cllipsoid, ca. 18-22 mm long, 10-14 mm in diam.

Fruit green with an red centre, broadly ovoid to globose, ca. 4-5 cm in diam.,

the basal collar composed of 14-20 connate sterile carpels, shallowly conical,

ca. 6- 1 1 mm long, 10-20 mm in diam. Fertile carpels 200-350, tetragonous,

broadly to transversely rhonibic in transverse section, ( 1 5-)20-25(-27) mm

long, 6-9 mm in diam., top itself Hat, with a 1 .5-2.5 mm long uncinate tip,

densely covered with stellate scales stellate liairs, basal 2/3 to 4/5 oftlic carpels

fused, the upper parts spacing 0-1 mm apart. Secds filling up 2/3 to 4/5 ofthe

carpcl, narrowly ovoid to narrowly cllipsoid (cllipsoid to compressed cllipsoid),

1 5-22 mm long, 6-8(- 1 0) mm in diam., tawny to dark brown, rugged, usually

with 2-3(-4) ribs.

162

Studies In Annonaceae. XVI. A Tnxonomic Revision of Duguetia A. F. C. P de Sant-Hilaire

Distribiition (Figure 2). Mato Grosso, Rondônia, Goiás, and rarely to S

Pará and SW Maranhão. In wet savannas or in gallery forest up to the altitude

of 550 m. Flowering and fruiting mainly from June through January.

Specimem examined. Goiás: Serra do Caiapó, 12 km S of Caiapônia,

840 m, 2 May 1973 (st), W. R. Anderson et ai. 9637 (U); Couto de Maga-

lhães, Rio Araguáia, 5 Jul 1953 (fl ),Fróes 30122 (S); Serra do Caiapó, 40 km

S of Caiapônia, road to Jatai, 900 m, 26 Jun 1966 (st), Irwin et al. 17764

(NY); Araguáina, Rio das Lontras, 300 m, 14 Mar 1968 (fr), Irwin et al.

21192 (NY); km l-5ol’Road from Estreita to Tocantinópolis, 9 Aug 1 964 (fl,

fr), Prance & Silva 5S630 (B, F, GH, K, MO, NY, RB, S, UC, US); Serra do

Caiapó, 66 Km of Jatai, 21 Oct 1964 (fl), Prance & Silva 59551 (F, GH, K,

NY, S, U, US): Ilha do Bananal, 2 km from Macaúba. 18 Sep 1980 (11. fr),

Ratteret al. 4438 ( E, K); Mun. Santa Izabel, Ilha do Bananal, Pargue Nacio-

nal do Araguaia, 19 Jun 1979 (st), F. C. Silva et al. 27 7 (SP); Rio Araguáia,

between Rio Caiepõs and Santana do Araguáia, 12 Aug 1978 (fl, fr), TV. T.

Silva 4801 (U). Maranhão: Ilha dos Botes, Rio Tocantins, near Carolina, 25

May 1950 (fl), Pires & Black 2107 (NY, S, U, UC). Mato Grosso: Mun.

Sinop, Fazenda Missionaria, Rio Teles Pires, 25 Sep 1985 (fl, fr), GUI et al.

6265 (U); Mun. Santa Terezinha, 21 km S of Santa Terezinha, Serra da

Cobrinha, 13 Oct 1985 (fr), GUI et al. 6439 (U); Aripuanã, near Salto Ando-

rinha, 21 Oct 1976 (11), Gomes & Miranda 356 (INPA); Royal Geographic

Society Expedition Base Camp, ca. 12°49' S. 51°46' W, 16 Oct 1968 (fr),

Harley 10665 (K, MO, NY, P. UC); Cáceres, Sep 1 908 (11), F. C. Hoehne 549

(SP); São Luis de Cáceres, Jun 1911 (fl), F. C. Hoehne 3461 (SP); São Luis

de Cáceres, Jul 191 1 (fl), F. C. Hoehne 3464 (S); Serra do Roncador, 84 km N

of Xavantina, 550 m, 1 Jun 1 966 (fl, fr), Irwin et al. 16418(MO. NY, S, SP);

Estrada Ranchão da Lagoa-Engenho Velho (Cuiabá), 22 Nov 1976 (y fr),

M. Macedo et al. 305 (INPA); Cuiabá. 30 Nov 1893 (fl), Mahne 1178* (S);

Cuiabá, 17 Jun 1902 (11). Mahne 1779 (F, G, LE, S); Mun. Santa Terezinha,

Rio Araguáia, 10 Oct 1985 (fr), Piranietal. 1193{ U); Mun. Luciara, Distr.

Porto Alegre, 10-11 km from I Iiglnvay BR 1 58, 1 6 Oct 1 985 (fr), Pirani et al.

1278(11): Gorge of Veu do Noiva, Chapada dos Guimarães, 1 7 Oct 1973 (11),

Prance et al. 19087 (K, MO. NY, S, U); 50 km N of Chavantina, Rio Vau, 8

Oct 1 964 (11), Prance & Silva 59305 (F, Gl I, K, NY. S. U, US); 8 km NE of

the Base Camp ofthe Expedition, close to Xavantina-São Fclix road, 12 Apr

1968 (st), Ratler et al. 1019 (K); Fazenda Santa Teresa, Rio Negro arca. 29

Apr 1 978 (st), Schaller 153 (NY): Mun. Vila Bela da Santíssima Trindade, 41

163

Boi. Mus. Para. Emílio Goeldi, súr. Boi. 9(2). 1993

km NNW of Potes and Lacerda on BR 364 to Vilhena, 31 Oct 1985 (fr),

Thomas et al. 4712 (U). Pará: Mun. Conceição do Araguaia, 20 km W of

Redenção, near Córrego São João and Troncamento Santa Teresa, 350-620

m, 8 Feb 1980 (fr), Plowman et al. 8532 (U) Chapada dos Parecis, 20 Jan

1 985 (fl), Venturieri & Menezes 1 8 (1NPA). Rondônia: Mun. Presidente Mediei,

BR 364, km 300 of road from Cuiabá to Porto Velho, 28 Jun 1984 (fl, fr), Cid

et al. 4874 (U); Mun. Ariquemes, Minerção Mibrasa, Setor Alto Candeias,

km 128, 19 May 1982 (11), Teixeira et al. 612 ( U).

Vernacular names. Araticum, Articum, Cunduru, Envira Biriba.

D. marcgraviana lias becn lectotypied by Fries ( 1 934) vvith Riedel 1471.

In Leningrad there wcre 2 collcctions of it, one a specimen with a labei in

Riedel’ s handwriting ("no. 1471. Anona, Arbor 20-25 ped. Foliisab squamulis

cineras crescentibus florib. flavescentis. In humidis tempore pluv. inundatis

pr. Matto Grosso. Febr. 1828") and annotated by Fries (1905) as Aberemoa

marcgraviana. The second collection has been selected by usas lectotype as it is

apparently a duplicate of Riedel 1471 as it has been annotated by Martius with

a.o.: “ Duguetia marcgraviana Mart. Icon. nomine Biriba in Libro qui dicitur

Príncipes quem Marcgravii composuit Menzelius, in biblioth. Berol. aspervat. “.

This is alinost the same phrase as is given in Martius’s original description.

D. marcgraviana appears to be a highly variable species, i.e., in the

length ofthe pedicels, leaf size, pcrsistence ofthe leaf indument, petal length,

etc. In all characters, D. brevipedunculata R. E. Fries Fits in our description

of D. marcgraviana. with the sole exception ofthe upper leaf side which is

glabrous from the very beginning. This character, however, does not seem to

be sufficient to maintain a separate specics, in particular as only the type of D.

brevipedunculata from Santa Anna da Chapada of Mato Grosso is known up

to now. Thereforc, D. brevipedunculata is brought into synonymy with D.

marcgraviana.

D. marcgraviana can easily be confused with both D. echinophora and

D. lepidota. It comes most close lo D. echinophora , from which it differs in

some features ofthe leaves, fruit and seeds (Table 2). Vegetatively, D. lepidota

comes close to D. marcgraviana and D. echinophora , but its inflorescence

may be more-llowered and its fruit is very distinctive (carpels at apical 1/5-1/

4 suddenly contracted and forminga black, decayed-ligneous, irregularly rugose

peltate top).

164

Síudies In Annonaceae. XVI. A Toxonomic Revision ofDiiguelid A. F. C P. de Sant-Hilaire

5. Duguetia echinophora R. E. Fries, Acta Horti Berg. 12(1): 40. 1934. Type:

Brazil. Pará: Bragança. 14 Jan 1923 (fl, fr), Ducke RB1 7868 (holotype,

S; isotypes, B, K, RB , US). Figure 8

Tree (rarely shrub), (3-)5-20(-25) m tal 1, stems (8-)l 5-20(-40) cm in

diam. Young twigs densely covered with yellowisli fimbriate scales, becoming

glabrous with age, older twigs greenish-brown to dark brown, striate. Petiole

(2.5-)3-4.5(-5) mm long, (1 .2-) 1 .5-1 .8 mm in diam., (ratlier densely) densely

covered vvitli yellowish fimbriate scales. Lamina elliptic, oblong-elliptic to

narrowly ovate, chartaceous to slightly coriaceous, green to dark green and

shiny above, greyish-green to green bclow, (6-)9- 1 5(- 1 7) cm long, (2.5-)3-5(-6)

cm wide, index (2.2-)2.5-3.5(-4), base attenuate, acute to obtuse, apex

acuminate (to acute). upper side glabrous from the very beginning (sometimes

covered with stellatc liairs on the base of primary vein), lower side densely

covered with yellowish fimbriate scales and whitish stellate scales, secondary

veins uniformly curved, (8-) 1 1 - 1 5(- 1 8) on either sides of primary vein, angles

with primary vein (45°-)55 0 -65 0 (-75°), intercostal area irregularly spaced,

distance between the middle ofclosures and margin (1 .5-)2-5(-8) mm, tertiary

veins slightly raised on botli sides, irregularly forming (2-)5- 1 3 (- 1 5 ) nets, 4th

order veins slightly raised on upper side, slightly raised to flat on lower side.

Inflorescence leaf-opposed, l-2(-3)-flowered, subsessile, pedicels

9- 1 5(- 18) mm long, straight (recurved), (l-)l ,2-2(-2.5) mm in diam. at base,

2.5-4 mm in diam. just bclow flower, fruiting pedicel 14-22(-25) mm long,

(2-)2.5-4.5(-6) mm in diam. at base, 3-7 mm in diam. just below fruit. Bracts

2 to the pedicel, one amplectant on basal 2/5 to 4/5 of pedicel, depressed

ovate, 3-4.5 mm long, 6-7(-8) mm wide, another one broadly ovate todeltate.

2-3 mm long, 3-4 mm wide. Pedicels and outer side of bracts densely covered

with yellowish fimbriate scales, inner side of bracts (sparsely to) densely covered

with curviform stellatc liairs; outer side of sepals densely covered with yellowish

fimbriate scales, inner side ratlier densely covered with curviform stellate liairs

(basal ca. 1/5 usual ly very sparsely covered to glabrous); outer side of petals

(ratlier densely to) densely covered with yellowish fimbriate to stellate scales,

inner side densely covered with curviform stellate liairs (basal 1/5 to 2/5 ot

outer petals and 1/5 to 1/2 of inner petals glabrous, striate).

Flower buds broadly to very broadly ovoid, apex acute (rounded). 5-8

mm long, 5-9 mm in diam. Flower green to greenish-yellow with red inner

base (in vivo). Sepals connate to 1/5- 1/4 from base, ovate to very broadly

165

SciELO

Boi. Mus. Para. Emílio Goeldi. sér. Boi. 9(2). 1993

ovate (elliptic to oblong-elliptic), 1 3- 1 6(-20) mm long, 9-14 mm wide, apex

acute. Outer petals (narrovvly) obovatc to spathulate, apex acute to obtuse, at

basal i/4 to 1/3 strikingly contracted, 15-25 mm long, 8-1 1 mm wide, inner

petals obovatc to spathulate, apex acute, at basal 1/3 to 1/2 strikingly contracted,

14-25(-30) mm long. 8-14 mm wide. Torus transversely narrowly ellipsoid.

Stamens 200-300. (0.9-) 1 . 1 - 1 .4 mm long, 0.2-0. 3 mm in diam., apical

prolongation of connective shallowly conical, (0.2-)0.2-0.3 mm long,

0.3-0.5(-0.6) mm wide. Carpels 1 50-250(-300), stigma densely covered with

curviform and/orerect stcllate hairs.

Fruiting rcceptacle subglobose to broadly ellipsoid, 1 0- 1 8(-20) mm in

diam. Fruit red at maturity with an orangc edible pulp (in vivo), transversely

broadly ellipsoid to subglobose, 3.5-6 cm long, 4.5-7 cm in diam., the basal

collar composed of 12-16 connate sterile carpels, very broadly to shallowly

conical, 9-14 mm long, 12-16 mm in diam. Fertile carpels 1 50-250(-280),

angular-obovoid to obconical, (18-)20-26 mm long, 7-1 I mm in diam., top

itself flat to conical, irregularly rugose, w ith a 2-3 mm long uncinate tip, densely

covered with stcllate scalesand stcllate hairs, basal 4/5 ofthe carpels to totally

fused. Seed filling up 3/5 to 3/4 ofthe carpel. obovoid, ( 1 1 -) 1 3- 1 8 mm long,

5-7(-9) mm in diam., lawny to dark brown, roughish, usual ly with 3-4 ribs.

Dislribiition ( Figure 2 ). Amapá, Amazonas, Maranhão, Pará, and Piauí.

In primary or secondary forest. Flowering and fruiting all the year around.

Specimens examinei!. Amapá: Rio Araguari, 5 Sep 1 96 1 (fl). Pires cl al.

50732 (G, GH, MG, NY. U). Amazonas: Colosso, Faz. Fsteio, km. 23, ZF-3,

24 Jul 1 986 (y fr), ( 'ariloso N Set: 36 (INPA); Estrada Manaus-Caracaraí, km

57, 1 5 Sep 1 976 (fr), Damiào 4/ Mola 632 (INPA); Manaus, Estrada Manaus-

Caracaraí, km 57, 24 Sep 1 976 (y fr). Nascimento 53 (U): Km 69, E of Highway

Manaus-ltacoatiara, 5 Oct 1960 ( fr), Rodrigues /#09(INPA, MG); Maranhão:

Fazenda Bacaba, 5 km S of MA I 19, from entrance 3 km NW of Lago do

Junco, 4 Oct 1980 (fr). DaJy et al. 459 (NY, IJ); 50 km from Santa Luzia on

highway to Açailândia. 24 Oct 1980 (fr). Daly et al. 7 41 (NY, U); São Luis,

Granja Barreto, 25 Oct 1948 (II), Ducke 10 (S); Mim. Lorêto, 40 km S of

Lorêto, 250 m, 25 Mar 1962 (fr), Eiten & Eiten 3832 (NY, SP. US); Rio

Maracaçumé region, 14 Sep 1932, Fróes 1905 (A, B, BM, F, G, K, MICII.

MO, NY, P, S, U, IJS): Island of São Luiz. Feb-Mar 1939 (fl), Fróes 1 1641

(A. F, G. K. LIL. MIC 1 1. MO, NY. S, U. UC, US); São Luiz, 12 May 1949,

Fróes 24246 (SP); Al/i landia. Rio Pindaró. 0- 1 00 m. 1 2 Dec 1 978 (fr), Jangoux

166

Studies In Annonaceae. XVI. A To.xonomic Revision ofDuguetia A. F. C P. de Sant-Hilaire

& Bahia 438 (N Y, U); Bom Passar, 24 km of Peritoró, 6 Jun 1 979 (st), Jangoux

& Bahia 1058( U); Buriticiipu, Reserva Florestal da CVRD, 5-30 Sep 1975 (y

fr), N. A. Rosa et ai 51 70 (MG); Mim. Tuiitum, 5 km N of Palmeirinhas, 25

Feb 1983 (fr), Schatz et ai 772 (U); Mim. Barra do Corda, Cachoeirinha, 57

km SSW of Barra do Corda, 2 Mar 1983 (fl), Schatz et ai 827 (U); Mim.

Barra do Corda, Cachoerinha, 57 km SSW of Barra do Corda, along Rio

Corda, 2 Mar 1983 (fl, fr), Schatz et ai 835 (U); Mim. Codo, 50 m, 27 Jun

1972 (fl), Sucre & Si/va 9400 ( RB, U). Pará: Curiiça, Iririteua, 14 Dec 1978

(st), Cavalcante 3359 (NY); Acará, Fazenda Borba Gato, near Rio Acará, 7

Nov 1980 (st). Da/y et ai 873 (F, M1CH, NY, U); km 90 of Highvvay from

Belém to Brasília, 21 Sep 1959 (fl, fr), M. Kuhlmann & Jimbo 257 (SP, US);

Ciiruçá, 4-10 Nov 1 985 (fr). Lobato et ai 141 (MG); 50-60 km N of Gurupk

Belém-Brasília Highvvay, 1 2 Aug 1 964 (fr), Prance <Sc Silva 58684 (F, GH, K,

NY, S, U); Vieinity of Paragominas, km 161 of Belém-Brasília Highvvay, 28

Aug 1964 (fl), Prance & Silva 58931 (F. GH, NY, S, U); Ilha de Marajó, Mim.

Salvaterra, Joanes, 25 Apr 1980. N. A. Rosa 3638 (U); Ilha de Marajó, Mun.

Salvaterra, Joanes, 1 0 Jau 1 982 ( fl), Rosário & Taylor 116 (HBG, MG). Piauí:

E of Piauí, Mun. Cocai. I 1 0 m, 2 1 Jun 1 972 (fl), Sucre et ai 9247 (RB).

Vernacular naines. Aineiju, Aineju de touceira. Ata braba / Ata brava,

Ata-meju (= Envira surucucu). Envira-preta, Envira surucucu, Envireira,

Pindaeua.

Uses. Wood of th is specics is used for frame building. The bark is used

as cure for fever. The orange flesh of thc mature fruit is cdible.

For discussion oflhe differences betvveen D. echinophora , D.lepiclota

and D. marcgraviana , see note under description oi' D. marcgraviana.

6. Duguetia lepiclota (Miquel) Pulle, Enum. Vasc. PI. Surinam 176. 1906;

R. E. Fries, Acta I lorti Berg. 12(1): 41. 1 934. Figure 9

Annona lepiclota Miquel. Linnaea 19: 129. 1847. Type: Surinam. Surinam

River(“in regionibus interioribusad 11. Surinam”), Oct 1 844 (fl), Kappler

1686 (holotype. U: isotypes, B, F: fragment, G, P, S).

Aberemoa lepiclota (Miquel) R. E. Fries, Kongl. Svenska Vetenskapsakad.

I landi., n.s., 34(5): 20. 1900.

Trec (or rarely shrub), (7-)10-20(-28) m tal 1, stems (6-)l 5-25(-65) cm in

diam. Young tvvigs densely covered vvith yellovvish fimbriate scales, becoming

167

Boi. Mus. Para. Emílio Gocldi, ser. Boi. 9(2). 1 993

glabrous wilh age, older twigs greyish-brown to dark brown, irregularly striate.

Petiole 4-6(-8) mm long, ( l-)l .5-2(-2.5) mm in diam., densely covered witli

yellowish fimbriate scales. Lamina narrowly ovate to narrowly elliptic-ovate

(narrovvly elliptic), cliartaceous to sliglitly coriaceous. dark green and slightly

shiny above, greyish-grcen below, (7-)9-20(-23) cm long, (2.2-)3-5(-6.2) cm

wide, index 3-4(-4.3), base acute, narrowly cuneate to slightly attenuate, apex

acuminateto acute, tipperside sparsely to rather densely covered with whitish

(yellowish), (5-)6- 1 0(- 1 2)-rayed appressed and/or crect stellate hairs bccoming

sparsely covered to glabrous with age. lower side densely covered with yellowish

fimbriate scales and stellate scales, secondary veins uniformly curved, (9-)l 2- 1 8

on either side ofprimary vein, angles with primary veins (50 o -)60°-70 o (-80°),

intercostal area irregularly spaced, distance between the middle of closures

and margin (l-)2-5(-6) mm, tertiary veins slightly raised on both sides,

irregularly forming 2-13 nets, 4th order veins slightly raised to flat on both

sides, but nearly invisiblc on lower side because ofthe dense indument.

Inflorescence leaf-opposed or pseudo-terminal, ramose. 1 -9(- 1 2)-

llowcred, subsessile or on the top of a 2-4 mm long, ligneous rudimentary

twig, pedicel (1 l-)15-18(-22) mm long, straight, (0. 8-) 1-1. 2 mm in diam. at

base, ( 1 .5-)2-3(-4) mm in diam. jnst below llower, fmiting pedicel (20-)25-35

mm long, (2-)3-5(-6) mm in diam. at base, (4-)5-8(-9) mm in diam.just below

fruit. Bracts 2 to the pedicel, one amplcctant on basal l/2(-3/4) of pedicel.

depressed ovate to very broadly ovate (subeireular). 2. 5-4. 5 mm long. 4-6 mm

wide, another one depressed ovate to broadly ovate (to deltatc), 1-2.5 mm

long, 2-4 mm wide. Ped iceis and outer side of bracts rather densely to densely

covered with yellowish fimbriate scales, inner side of bracts (rather densely

to) densely covered with eurviform stellate hairs; outer side of sepals rather

densely to densely covered with yellowish fimbriate scales, inner side densely

covered with eurviform stellate hairs (basal ca. 1/5 usually very sparsely covered

to glabrous); outer side of petals densely covered with yellowish fimbriate to

stellate scales (stellate hairs), inner side densely covered with eurviform stellate

hairs (basal 1/3- 1/2 of outer petals and basal I/2-2/3 of inner petals glabrous,

striate).

Flower buds broadly to very broadly ovoid, apex acute (rounded), (4-)6-8

mm long, (4 .5- )5 .5-8 mm in diam. flower green to greenish-yellow, with pink

to dark red inner base ( in vivo). Sepals connatc to 1/6- 1/4 froin base, ovate to

broadly ovate. 9- 1 3 mm long. 8- 1 3 mm wide. Outer petals obovate to slightly

168

Studies In Annonaceae. XVI. A Toxonomic Revision ofDuguetia A. F. C. P. de Sant-Hilaire

spatluilate, apexacute, at basal 1/3- 1/2 strikingly contracted, (10-)12-20(-22)

mm long, (6-)7- 1 0(- 1 2) mm wide, inner petals obovate to slightly spathulate,

apex acute to obtuse, at basal 1/3- 1/2 apparently contracted, (10-)14-20(-28)

mm long, (6-)8- 1 0(- 1 2) mm wide. Toras transversely narrowly ellipsoid.

Stamens 1 20-250, ( 1 .2-) 1 .4- 1 ,5(- 1 .7) mm long, (0.3-)0.4-0.5 mm wide, apical

prolongation of connective shallowly (to very shallowly) conical, (0.2-)0. 3-0.4

mm long, (0.4-)0.5-0.6(-0.7) mm wide. Carpels 1 50-300(-480), stigma and

ovary sparsely to densely covered with curviform stellate bairs.

Fruiting receptacle ellipsoid, ca. 30-35 mm long, 20-25(-30) mm in diam.

Fruit green when immature, outer surface becoming pink and the centre becoming

orange with age (in vivo), ovoid to very broadly ovoid (to spheroid), 5.5-8 cm

long, 5.5-7 cm in diam.. lhe basal collar composed of 12-15 connate sterile

carpels, broadly conical to deltoid, ca. 14-18 mm long, 14-16 mm in diam.,

fertile carpels 1 50-350(-450), narrowly angular-obovoid to angular-obovoid,

(15-)20-28mm long,(6-)8-12 mm in diam., at apical I/5-1/4 suddenly contracted

and forming a black. decayed-ligneous, irregularly rugose and peltate top, basal

2/3 to 3/4 of the carpels fnsed, the upper parts spacing 1-5 mm apart. Seed

filling up 2/3 to 3/4 of the carpel, ellipsoid (obovoid), ( 1 0-)l 2- 1 6(- 1 8) mm

long, (5-)6-8 mm in diam., tavvny to dark brown, roughish (smooth).

Disíribvtion (Figure 2). Colombia, Venezuela, Surinam and the Brazilian

States of Amazonas, Pará and Maranhão. Mostly in primary forest on terra

firma. Floweringand fruiting all theyeararound.

Specimens exatninecl. Amazonas: Mun. Barcelos, 3 km S of Serra Cen-

tral of Serra Aracá, 8 km F of Rio Javari, 9 Mar 1984 (fr), Amaral et al. 1702

(U). Maranhão: Mun. Santa Luzia, Fazenda AGRIPEC, 7 km W ofBuriticupu,

3 Apr 1 983 (fr), Taylor cl al. El 129 (U). Pará: 1 8 km from Tucuruí, 1 6 km S

on old Highway BR 422, 1 Nov 1981 (11), Daly et al. 1155 (U); 25 km from

Tucuruí, SW of ncw road serving Fazendas, 14 Nov 1981 (fl, Y Ir), Daly et al.

1323 (U); Breu Branco, 40 km S of Tucurui, then 5 km W on old logging road,

22 Nov 1981 (fl), Daly et al. 1535 (NY, U); Rio Tocantins, Jacundá, Rio

Cajazciras, 1 6 May 1 978 ( 11). Aí Cl. Silva A Bahia 3603 (NY), 1 ucurui, BR230,

5 Nov 1 979 (fr), M. F. F. Silva et al 39 (HBG). Roraima: Mun. Alto Alegre,

I lha de M aracá, SEM A Ecological Station, 4 Jun 1 986 ( fr), Hopkins et al 507

(U); Mun. Alto Alegre. Ilha de Maracá, 6 Jun 1986 (fr), Hopkins et al. 512

(U); Canto Galo, Rio Mucajaí bctween Pratinha and Rio Apiaú, 21 Jan 1967

(fr)' Prance et àl. 3961 (F.’ GIl, K. NY. S. U, US); Ipiranga, Rio Mucajaí,

160

Boi. Mus. Para. Emílio GoelcJi. sér. Bot. 9(2). 1 993

between Pratinha and Rio Apiaú, 27 Jan 1967 (fr), Prance et ai. 4130 (F, K,

NY, S, U, US); Base of Serra Tepequem, 1 1 Feb 1967 (fl), Prance et al. 4327

(F, GH, K, NY, S, U, US); Ilha de Maracá, SEM A Ecological Station, 29 Oct

1987 (y fr), Ratter & Milliken 5916 (U); Ilha de Maracá, SEMA Ecological

Station, 15 Feb 1979 (fr), IV. Rodrigues 10144 (MO); near Ecological Station

of Ilha de Maracá, 6 Feb 1979 (fr), N. A. Rosa 3058 (NY, U).

Vernacular names. Ameiju, Ata braba.

Uses. The wood is used for construction.

Vegetatively D. lepidota comes close to D. tnarcgraviana and D.

echinophora (see discussion imder D. tnarcgraviana), but its fruit is very

distinctive as the upper 1 /5-1/4 part of the carpels is suddenly contracted fonning

ablack, decayed-ligneons, irregularly-rugose peltate top.

7. Duguetia giabriuscula (R. E. Fries) R. E. Fries, Acta Florti Berg. 12 (1):

46. 1 934. Figure 1 0

Aberetnoa lanceolata (A. F. C. P. de Saint-Hilaire) Warming var .giabriuscula

R. E. Fries, Ark. Bot. 4(19): 6. 1 905. Type: Brazil. Mato Grosso: between

Buriti and Santa Anna da Chapada, 4 Oct 1902 (fl), Malrne 11-2440

(holotype and isotype, S).

Shrub, 0.3-2 m tall. Youngtwigs sparsely todensely covcrcd with whitish

fimbriate scales, becoming glabrous with age, older twigs dark brown,

irregularly striate. Petiole 2.5-4 mm long, 1 .5-2 mm in diam., sparsely to rather

densely covered with whitish stellate scales. Lam ina elliptic to ovate, coriaceous,

green on botli sides, 6-10 cm long, index 1.6-2(-2.3), base acute to

obtuse(-attenuate), apex obtuse(-rounded or acute), upper side glabrous, lower

side sparsely covered with whitish stellate scales, secondary veins abruptly

curved. 8-12 on either sides of primary vein, angles with primary veins

(45 o -)50 o -65 o (-75°), intercostal area irregularly spaced, distance between the

middlc ofclosure and margin 2-4 mm, tertiary veins conspicuous, raised on

botli sides, irregularly forming 7-10 nets, 4th ordcr veins subconspicuous,

slightly raised on both sides.

Inllorescence leaf-opposed, 1-2-flowered, subscssile, pcdicel 6-8(-10) mm

long, recurved, 0.6-1 mm in diam. at base. 1-2 mm in diam. just below flower.

Bracts 2, one amplectant on basal 1 13-213 of pedicel, very broadly ovate, 1 .5-2.5

mm long, 2. 5-3. 5 mm wide, another one broadly to very broadly triangular.

170

Studies In Annonaceae. ATI A Toxonomic Revision oJDuguelia A. T. C. P. de Sant-Hilaire

1.5-2 mm long, 1.5-2 mm wide. Pedicels and outer side of bracts densely

covered witli vvhitish iunbriate scales, inner side of bracts glabrous (or rarcly

sparsely covered with stellate liairs); outer side of sepals densely covered with

whitish fimbriate scales, inner side densely covered with whitish curviform

stellate liairs (basal 1/3- 1/2 sparsely covered to glabrous); outer side of petals

densely covered vvitli whitish stellate scales, inner side densely covered with

whitish curviform to erect stellate liairs (basal 1/6-1/5 of outer petals and 1/

5-1/4 of inner petals sparsely covered to glabrous).

Flower buds ovoid to broadly ovoid, apex acute to obtuse, 4-8 mm long,

4-6.5 mm in diam. Flower green to greenish salmon, with red at inner base (in

vivo). Sepals connate to 1/6- 1/5 from base, triangular-ovate to broadly ovate,

9-12 mm long, 6-9 mm wide. Outer petals rhombic-elliptic, apex acute, 15-22

mm long, 7-1 1 mm wide, inner petals rhombic-elliptic, apex acute, 14-20 mm

long, 7-10 mm wide. Torus depressed ovoid. Stamens 80-150, 0.8-1 mm long,

0.4-0. 6 mm wide, apical prolongation of connective very shallowiy conical,

puberulous, 0.2-0. 3 mm long, 0.5-0. 7 mm wide. Carpels 50-130, ovary

glabrous, upper part of stigma densely covered with whitish erect stellate liairs.

Fruiting receptacle ellipsoid to oblong-ellipsoid, 13-16 mm long, 9-12

mm in diam. Fruit subglobose, 3-4 cm in diam., the basal collarcomposed of

8-10 connate sterile carpels, angular-ovoid or conical, 8-10 mm long, 8-10

mm in diam., glabrous. fertile carpels 60-120, obtrulloid, 12-15 mm long,

7-10 mm in diam., with 4-5(-7) ribs, top itself shallowiy conical, uncinate,

glabrous, total ly free from eacli other. Seeds not seen.

Distribution (Figure 3). Brazil: Mato Grosso and Bolivia: Santa Cruz.

In cerrado and Pantanal vegetation. Flowering mainly from September to

November, and fruiting from Novemberto March.

Specimem examined. Moto Grosso: I orixoréu, J. S. Costa 75 (RB); Mt.

Chapada dos Guimarães, 3 Mar 1 983 (y I r), da C unha et al. 788 ( MG); 50 km N

oi Barra do Garças, on road to Xavantina, 300-400 m, 14 Oct 1964 (11), Irwin &

Soderstrom 6870 (N Y, S); 1 2 km of Chapada dos Guimarães, E of Cuiabá, 630 m,

21 Mar 1981 (y fr), Jangoux & Sebastião 1403 (MG); Fazenda Miranda, ncar

Rio Aquidauana, 18 Sep 1980 (11), Pires & Furtado 17164 (U); Chapada dos

Guimarães, próxime a Escola Evangélica de Buriti, 21 Nov 1982 (y II), ./. U.

Santos & Rosário 302 (MG); Chapada dos Guimarães, próximo a Escola Evan-

gélica de Buriti, 21 Nov 1 982 (fr), J. U. Santos & Rosário 406 (MG).

171

Boi. Mus. Para. Emílio Goeldi. sér. Boi. 9(2). 1993

The shape and venation of the mature leaves of tliis species most resemble

tiiat of D. moricandiana. Differences are found in the puberulous apical

prolongation of the connectives, and the glabrous ovary (vs. glabrous apical

prolongation of connective and densely hairy ovary in D. moricandiana).

D. glabrinsciila is also very close to D. lanceolata. lt can be distingnished

by its mature lamina acute to obtuse at the base, and obtuse to rounded at the

apex (vs. narrowly cuneate to attenuate at the base, and acute to acuminate at

the apex in D. lanceolata).

8. Dnguetia lanceolata A. F. C. P. de Saint-Hilaire, Fl. Bras. merid. 1 : 35.

t. 7. 1825: R. E. Fries, Acta florti Berg. 12(1): 47. 1934. Type: Brazil.

Minas Gerais: Sumidouro, near Villa do Principe, (fr), A. F. C. P. de

Saint-Hilaire s.n. (holotype & isotype, P). Figure 1 1

Aberemoa lanceolata {A.. F. C. P. de Saint-Hilaire) Warming, Vidensk. Meddel.

Dansk Naturhist. Foren. Kjobenhavn 1873: 146. 1873.

Dnguetia lanceolata A. F. C. P. de Saint-Hilaire var. parvifolia (R, E. Fries)

R. E. Fries, Acta Horti Berg. 12(1): 47. 1934.

Aberemoa lanceolata (A. F. C. P. de Saint-Hilaire) Warming var. parvifolia

R. E. Fries, Kongl. Svenska Vetenskapsakad. Handl. n.s. 34(5): 20. 1 900.

Type: Brazil. Rio de Janeiro: Pctropolis, Santo Antonio, 25 Sep 1875

(fl), Glaziou 8255 (syntypes, B, C, G, K, LE).

Tree (rarely shrub), (2.5-)8-18 m tall, ca. 25 cm in diam. Young twigs

rather densely to densely covered with yellovvish fimbriate to stellate scales,

becoming sparsely covered or glabrous with age, older twigs yellowish-brown

to greyish white, irregularly striate. Petiole 2-5 mm long, 1-2 mm in diam.,

densely covered with yellowish to whitish fimbriate to stellate scales. Lamina

narrowly elliptic to elliptic (ovate-elliptic or oblong-elliptic), coriaceous, green

above, green to greenish yellow below, (4-)6- 1 0(- 1 4) cm long, ( 1 ,5-)2.5-4(-5.5)

cm wide, index 1.9-3, base narrowly cuneate to attenuate, apex acute to

acuminate, upper side glabrous, slightly shiny, lower side (sparsely) rather

densely covered with whitish (fimbriate to) stellate scales, secondary veins

uniformly curved, (8-) 10-1 5 on eithersides of primary vein, angles with primary

veins (45°-)50 o -70 o (-80°), intercostal area irregularly (orsomewhat regularly)

spaced. distance between the middle of closure and the margin 2-4 mm, tertiary

veins raised on both sides, irregularly forming 5- 1 0(- 13) nets, 4th order veins

slightly raised on both sides.

172

Studies In Annonaceae. XVI. A Toxonomic Revision ofDuguetia A. F. C P. de Sanl-Hilaire

Inflorescence leaf-opposed or supra-axillary, ! -3(-4)-flowered, subsessile,

pedicel (5-) 10-20 mm long, 0.8-1 .2 mm in diam. at base, 1 .5-2.5 mm in diam.

just below flower, fruiting pedicel (1 1 -) 1 5-20(-22) mm long, 2-4 mm in diam.

at base, 3.5-4.5(-6) mm in diam. just below the fruit. Bracts 2 to the pedicel,

one amplectant on basal 2/3-5/Ó of pedicel, very broadly to depressed ovate,

2-4 mm long, 4-6 mm wide, easily broken, another one deltate to very broadly

ovate, 1 .5-2.5 mm long, 2. 5-3. 5 mm wide. Pedicels and outer side of bracts,

densely covered with yellowisb fimbriate to stellate scales inner side of bracts

glabrous (or rarely sparsely covered with stellate hairs); outer side of sepals

densely covered with yellowish fimbriate to stellate scales, inner side densely

covered with curviform stellate hairs (basal 1/5 usually sparsely covered);

outer side of petals densely covered with yellowish to whitish stellate scales to

stellate hairs, inner side densely covered with yellowish to whitish curviform

stellate hairs (basal 1/1 0 of outer petals and 1/5-1/4 of inner petals glabrous).

Flower buds broadly to very broadly ovoid, apex acute (to obtuse), 3.5-6

mm long, 3-6 mm in diam. Flower green, brownish pink to red, with dark red

at inner base (in vivo). Sepals connate to 1/6- 1/5 from base, triangular-ovate

(broadly triangular-ovate), 9- 1 1 mm long, 6-8 mm wide. Outer petals rhombic-

elliptic to narrowly ovate(-spathulate), apex acute to obtuse, 14-20 mm long,

5-9 mm wide, inner petals rhombic-elliptic to spathulate, apex acute(-obtuse),

1 5-20 mm long, 6-9 mm wide. Torus depressed ovoid, densely covered with

whitish erect stellate hairs. Stamens 80-150, 1.1-1 .3 mm long, 0.5-0. 7 mm wide,

apical prolongation ofconnective shallowly to very shallowly conical, compressed,

setulose, 0.2-0.4 mm long, 0.6-0. 8 mm wide. Carpels 40-100, ovary glabrous,

upper part of stigma sparsely to densely covered with erect stellate hairs.

Fruiting receptacle ellipsoid to broadly ellipsoid (or oblong-ellipsoid) or

obovoid, I 5-30 mm long, 1 5-2 1 mm in diam. Fruit green to red, subglobose,

4-6 cm in diam., the basal collar composed of 7-14 connate sterile carpels,

oblong-ellipsoid or broadly angular-ovoid, 8-17 mm long, 8-18 mm in diam.,

glabrous, fertile carpels 30-80, obtrulloid, 20-25 mm long, 10-13 mm in diam.,

with 5-7 ribs, top itself shallowly conical, glabrous, totally free from each

other. Seed filling up 5/6 of the carpel, obovoid, 1 8-22 mm long, 8-10 mm in

diam., yellowish-brown, rugged.

Dislribiiíion (Figure 3). The Brazilian States of Minas Gerais, Paraná, Rio

de Janeiro, Rio Grande do Sul, Santa Catarina, and São Paulo. In cerrado or in

semidecidous forest from nearly sea levei up to 900 m at elevation. Flowering

mainly from Septemberto March, and fruiting mainly from Octoberto May.

173

Boi Mus. Pura. Emílio Gochli, sér. Boi. 9(2), 1993

Specimens examined. Minas Gerais: São Scbestião da Campina, 22 Dec

1949 (y fr), Duarte 2289 (RB); Findai- reuna, near Felixlândia, 20 Jul 1959

(fl), A. de Mattos 121 (RB); “ ln circuitu urbis Franca”, 1 902 (fr), Wacket s.n.

(W); Lagoa Santa, 8 Mar 1865 (fl, fr), Warmings.n. (C, F, K, NY, S). Paraná:

Sangés, 29 Jun 1910 (y 11), Dusén 9939 (GH, NY, S); Patrimônio, 900 m, 1 1

Mar 1915 (st), Dusén 167S5 (GH, MO). Rio de Janeiro: Ypanema, Jan 1826

(y fr), Riedel 174 (BM, LE. M, S); Itaipava, 1940 (fl), Stickney RB42422

(RB). Rio Grande do Sul: Torres, Limoeiro, 21 Apr 1978 (fr), Waechter &

Baptista ICNS26 (ICN); Torres, Lagcadi, 1 8 Oct 1 980 (fl). Waechter ICN1 735

(ICN); Três Cachoeiras, Pôrto Alegre, 26 Mar 1977 (yfl), Waechter ICN 42 185

(F); Torres, Três Cachoeiras, Sep 1978 (fl) Waechter & Baptista ICN47652

(ICN). Santa Catarina: Reserva Florestal des Pilões, 250 m, 3 Dec 1950 (fl),

Duarte & Frctzão 3208 (BM, F, M, NY, RB, Z); Brusque, near Azambuja, 50

m, 25 Nov 1 949 (11), Klein 182 (NY, S, US); Morro da Fazenda, Itajaí, 200 m,

30 Nov 1954 (11), Klein 886 (B, NY, S, UC, US); Horto Florestal I.N.P.,

Ibirama, 200 m, 27 Dec 1954 (fl), Klein 936 (B, NY, P, S, US); Mun. Mata-

dor, Matador, 300 m, 17 Dec 1967 (fl), Lourteig 2312 (C, P); Horto Florestal

I.N.P., Ibirama, 300 m, 2 Mar 1954 (fr), Reitz & Klein 1647 (NY, S, UC);

Pilões, Palhoça, 250 m, 5 Apr 1956 (fr), Reitz & Klein 3044 (B, GH, NY, S,

U, UC, WIS); Azambuja, Brusque, 50 m, 15 Nov 1949 (st), Reitz 3204 (S);

Azambuja, Brusque, 50 m, 15 Nov 1949 (fl), Reitz 3585 (B, F, G, NY, P, U,

UC,'W); Ibirama, 100 m, 2 1 Sep 1956 ( 11), Reitz & Klein 3750 (S); Matador,

Rio do Sul, 350 m, 1 8 Oct 1 958 (11), Reitz & Klein 7342 (G, M, S, Z); Vargem

Grande, Lauro Miiller, 350 m, 24 Oct 1 958 (11), Reitz & Klein 7483 (B, K, S);

Pirão Frio, Sombrio, 10 m, 29 Oct 1959 (fl), Reitz & Klein 9248 (B, G, K, M,

S, Z); Itajaí, 1 1 Nov 1 953 (11), Schultz 1103 (ICN). São Paulo: Mun. Anhembi,

Barreiro Rico, 19 Sep 1979 (fl), Assumpção 7548 (E); Paccas, Oct 1945 (fl),

Edwall 145 (S); Jardim Botânico, 26 Sep 1932 (11), Hoelme SP29784 (NY);

São Paulo, 5 Nov 1937 (fl), Hoehne SP3S975 (SP, U, US); Jardinopolis,

margins of Rio Pardo, Parque Rio Pardo, 19 Nov 1947 (y fr), M. Kuhhnann

1589 (SP); Bairro das Três Fazendas, Fazenda Santa Tereza, 1 5 Nov 1 947 (y

fr), M. Kuhhnann & Gonçalves 1667 { SP); Anhembi, Fazenda Barreiro Rico,

4 Oct 1956 (II), M. Kuhhnann 3951 (K, SP, US); Mun. Teodoro Sampaio,

Reserva Morro do Diabo, 23 May 1986 (fr). Leite & Klein 58 (RB); Campo

Pasto, São José dos Campos, 12 Mar 1909 (y 11), Loefgren 244 (RB, S); São

José dos Campos, 24 Sep 1909 (11), Loefgren 4008 (S); Queluz Sorocabana,

25 Jun 1899 (st), Loefgren 5986 (S); Pindorama, property of F. Santos, 14

Mar 1939 (11), Mendes SP44191 (SP); Campinas, Novaes 4576 (S); “ Interior

do Estado de São Paulo”, 18 Mar 1932 (fl), A. Pereira SP29284 (SP, US);

174

Studies In Annonaceae. XVI. A Toxonomic Revision ofDuguetia A. F. C. P. de Scmt-Hila ire

Mugi, Nov 1833 (y fr), Riedel 1686 (K, LE, S); Loreto, Unknown collector

SP30620 (SP); Campinas, 10 Oct 1938 (y fr), Valentim 3401 (LIL).

Vernacular names. Cabreuva vermelho, Pindabuna, Pindahiba, Pindaíba/

Pindaíba, Pindaibuna, Pindaúva.

This species resembles most D. lúcida, a species also placed in Sect.

Duguetia by Fries, but excluded from this treatment because of its distribution

(Trinidad, Venezuela, Guyana and the Brazilian State of Acre). The species

differ in the size of the lamina: usually no more than 10 cm long vs. lamina

usually more than 10 cm long, and in the indument of ovary and carpel:

essentially glabrous vs. sparscly covered with stellate hairs at upper part in D.

lúcida). See also under D. glabriuscula.

9. Duguetia riedelianaW. E. Fries, Buli. Herb. Boissier sér. 2. 7: 1002. 1907;

R. E. Fries, Acta Horti Berg. 12(1): 45. 1934. Type. Brazil. Rio de Ja-

neiro: without exact location, 1836 (fr), Riedel s.n. (lectotype, S:

designated by Fries, 1934; isolectotype, LE). Figure 12

Tree or shrub, (0.5-)4-8(-25) m tall. Young twigs sparsely to densely

covered with yellowish stellate scales, becoming glabrous with age, older twigs

greyish white to yellowish-brown, irregularly striate. Petiole 3-5 mm long,

0.8- 1.2 mm in diam., sparsely covered with fimbriate to stellate scales. Lamina

narrowly ovate to ovate (narrowly oblong-el liptic to oblong-elliptic),

chartaceous to siightly coriaceous, green on both sides, (4-)5- 1 0(- 1 2) cm long,

2.5-4(-4.5) cm wide, index 2-3(-3.5), base attenuate, apex acuminate(-acute),

upper side glabrous, lower side sparsely covered with yellowish stellate scales,

secondary veins abruptly curved, 12-16 on either sides of primary vein, angles

with primary veins (50 o -)60°-75 o (-85°), intercostal area somewhat regularly

spaced, distance between the middle of closure and margin 3-5 mm, tertiary

veins raised on both sides, irregularly forming 7- 1 5 nets, 4th order veins siightly

raised to flat on upper side, siightly raised on lower side.

Inflorescence leaf-opposed, 1 -2-fIowered, subsessile, pedicel 4-6 mm long,

straight(-recurved), 0.8-1 mm in diam. at base, 1-1.5 mm in diam. just below

the flower, fruiting pedicel 1 0-20 mm long, 1 .5-5 mm in diam. at base, 2.5-6

mm in diam. just below the fruit. Bracts 2 to the pedicel, one amplectant on

basal 1/2-2/3 of pedicel, very broadly ovate, 1-2 mm long, 1-2 mm wide,

another one shallowly triangular, ca. 1 mm long, 1 .5-2 mm wide. Pedicels and

outer side of bracts densely covered with yellowish fimbriate to stellate scales.

175

Rol Mus. Rara. IímilioGoelili. ser. Boi. 9(2). 1993

inner side of bracts glabrous; outer side of sepals densely covered with yel lovvisb

fimbriate to stcllate scales, inner side densely covered witli yellowisb curviform

stellate hairs (basal ca. 1/6 sparsely covered); outer side of petals densely

covered with yellowisb stellate scales. inner side sparsely covered with curviform

to erect stellate hairs (basal 1 /6- 1/4 of outer petals and 1/5- 1/4 of inner petals

nearly glabrous).

Flower buds broadly ovoid, apex acutc to apiculate, 4-9 mm long, 5-7

mm in diam. Flower cream-coloured, with dark red inner base (in vivo). Sepals

connate to ca. 1/5 from base, triangular ovate to ovate, 10-12 mm long, 6-7.5

mm wide. Outer petals rhombic-clliptic to rhombic-obovate, apexacute, 14-18

mm long, 6-9 mm wide, inner petals rhombic-elliptic (to rhombic-obovate),

apex acute, 14-17 mm long, 6-8 mm wide. Torus depressed ovoid, densely

covered with yellowisb erect stellate hairs. Stamens 60-100, 0.7-1 mm long,

0.3-0. 4 mm wide, apical prolongation of connective very shallowly conical,

puberulous, 0.1 -0.2 mm long, 0. 5-0.6 mm wide. Carpels 50-100, ovary and

base of style densely covered with yellowisb erect stellate hairs, upper part of

style glabrous.

Fruit ellipsoid, ca. 7 cm long, 5 cm in diam., the basal collar composed of

10- 14 connate sterile carpels. broadly conical to deltoid, 8-14 mm long, 10-15

mm in diam., densely covered with yellowisb stellate scales and stellate hairs,

fertile carpels 40- 1 00, narrowly angular-obovoid to angular-obovoid, with 4-6

ribs, 15-20 mm long, 9- 12 mm in diam., top conical oruncinate, densely covered

with yellowisb stellate scales and stellate hairs, basal 1/2-3/5 of carpel fused.

Sced filling up 5A3-4/5 of the carpel, (broadly) obovoid, compressed, apex

apiculate, 10-14 mm long, 8-10 mm in diam., dark brown, smooth.

Distribution (Figure 3). Brazilian States of Bahia, Ceará,

Maranhão, and Rio de Janeiro. Flowering mainly from November to

June, fruiting in April.

Specimens examined. Bahia: Mun. Uruçuca, 30 km from Uruçuca, 15

Apr 1978 (Ir), Movi Kallunki 9920( CEPEC, U). Ceará: Serra do Bezouro,

23 Jan 1958 (fl), T. N. Guedes 484 (MG). Maranhão: 10 km W from

Barreirinhas road to Sobradinho, 30 Nov 1985 (y fl), Prance 29924 (U). Rio

de Janeiro: Mun. Cabo Frio, Búzios, near Praia Brava, 14 Feb 1985 (fl),

Araújo & Plowman 6680 (U); Rio Comprido, Lagoinha, 18 Feb 1871 (st),

Glaziou 4746 (C, P); without exact location, Glaziou 6464 (C, P); without

176

Studies In Annonaceae. XVI. A Toxonomic Revision ofDuguetia A. F. C l‘ de Scmt-IMaire

exact location, Glaziou 10230 (B, C, K, P); Gávea, Chácara do Lage, 6 Jun

1940 (fl), J. G. Kuhlmann 6037 (RB, U); Fundão, 8 Mar 1965 (fl), Lobão &

Almir RB194252 (RB, U).

This species is very similar to D. salicifolia. Vegetatively it can be

recognized by its ovate lamina vs. ellipsoid lamina in D. salicifolia), the lower

side sparsely covered with stellate scales (0-1 /mm2, 0.1 -0.2 mm in diam.) vs.

lower side more densely covered with fimbriate scales (3- 1 1 scales/mm2, 0.3-0.4

mm in diam.) in D. salicifolia (Table 3).

D. microphylla, a species with the same distribution area and often

confused, has much narrower leaves with 3-9 scales per mm2. Furthermore,

the ápices of the fertile blackened, broadly obconical carpels of D. microphylla

are truncate and sparsely covered with stellate scales, whereas in D. riedeliana

one finds carpels with conical or uncinate ápices, densely covered with yellowish

stellate scales and stellate hairs.

10. Duguetia microphylla (R. E. Fries) R. E. Fries, Acta Horti Berg. 6(6):

16. 1919; R. E. Fries, Acta Horti Berg. 12(1): 44. 1934. Figure 13

Aberemoa microphylla R. E. Fries, Kongl. Svenska Vetenskapsakad. Handl.

n.s. 34(5): 23. t. 3, f. 4-6. 1900. Type. Brazil. Rio de Janeiro: Serra do

Alto Macahé, Novo Friburgo, 1 888 (fl, fr), Glaziou 16689 (holotype, B;

isotypes, C, F: fragment, S: fragment).

Tree, 5-22 m tall. Young twigs slender, somewhat regularly striate, densely

covered with yellowish fimbriate scales, becoming glabrous with age, oldertwigs

greyish white to yellowish-brown, irregularly fissured. Petiole 2-3(-4) mm long,

0.8- 1 (- 1 .2) mm in diam., densely covered with yellowish fimbriate scales. Lamina

narrowly ovate, chartaceous, greenish on both sides, 6- 1 2 cm long, ( 1 .2-) 1 .4-2 cm

wide, index 4.5-6(-6.4), base attenuate, apex acuminate, upper side glabrous, lower

side sparsely to rather densely covered with yellowish fimbriate to stellate scales,

secondary veins (uni forni ly) abruptly curved, 1 4- 1 8(-20) on either sides of primary

vein, angles with primary veins (50 o -)55°-70 o (-85°), intercostal area irregularly

spaced, distancc between the middle of closure and margin 2-3 mm, tertiary veins

slightly raiscd on both sides, irregularly forming 5-12 nets, 4th orderveins slightly

raised to Hat on both sides.

Inflorescence leaf-opposed, 1 -2-Ilowered, subsessile, pedicel 6-8 mm long,

straight, 0.8-1 mm in diam. at base, 1 .2- 1 .5 mm in diam. just below flower,

177

Boi Mus. Para. Emílio Goeldi. sér. Boi. 9(2). 1993

fruiting pedicel 1 0-35 mm long, 2.5-4 mm in diam. at base, 4-7 mm in diam.

just belowtlie fruit. Bracts2 to lhe pedicel, one amplectanton basal 1/3- 1/2 of

pedicel. very broadly triangular-ovate, 0.5- 1 mm long, 1-1.5 mm wide, another

one broadly to very broadly triangular, ca. 0.5 mm long, 0.5-0. 8 mm wide,

caducous. Pedicels and outer side of bracts densely covered with yellowish

fimbriate(-stellate) scales, inner side of bracts glabrous or sparsely covered

with stcllate hairs; outer side of sepals densely covered with yellowish

fimbriate(-stellate) scales, inner side densely covered with yellowish curviform

stellate hairs (basal ca. 1/5 usual ly sparsely covered); outer side of petals

densely covered with yellowish stellate scales and stellate hairs, inner side

densely covered with yellowish to whitish curviform stellate hairs (basal 1/61/

5 of outer petals and 1/5- 1/4 of inner petals sparsely covered to glabrous).

Flower buds broadly ovoid, apex acute, 3-7 mm long, 3-6.5 mm in diam.

Flowercream, with dark red at inner base (in vivo). Sepals connate to 1/6- 1/5

from base, broadly triangular-ovate to broadly elliptic-ovate, apex acute, 6-10

mm long, 5-8 mm wide. Outer petals rhombic-obovate to oblong-rhombic,

apex acute, 12-18 mm long, 6-8 mm wide, inner petals rhombic-obovate to

oblong-rhombic, apex acute, 11-17 mm long, 6-8 mm wide. Torus depressed

ovoid, glabrous. Stamens 1 00- 1 80, 0.8- 1 .2 mm long, 0.3-0. 4 mm wide, apical

prolongation ofconnective deltoid to shallowly conical. glabrous, 0.4-0. 6 mm

long, 0.5-0. 7 mm wide. Carpels 60-100, ovary and baseof style densely covered

with whitish stellate scales and stellate hairs, upper part of style glabrous,

blackened.

Fruit spheroid, 4-5 cm in diam., the basal collar composed of 8- 1 0 connate

sterile carpels, very broadly conical, 8- 10 mm long, 12-14mm in diam., sparsely

to densely covered with yellowish stellate scales and stellate hairs, fertile carpels

ca. 40, broadly obconical, pentagonous, 15-20 mm long, 13-15 mm in diam.,

top truncate, blackened, sparsely covered with yellowish stellate scales, basal

4/5 of carpel fused. Seed filling up 3/5-3M of the carpel, obovoid(to broadly

obovoid), 9- 1 2 mm long, 7-9 mm in diam., dark brown, smooth, slightly shiny.

Distribution (Figure 2). Only known from the State of Rio de Janeiro,

Brazil. Flowering from Septemberto November, and fruiting in April.

Specimens examined. Rio de Janeiro: Mun. Macaé, Pico do Frade de

Macaé, 900-1000 m, 22 Oct 1985 (fl), Leitmam et ai 17 (U); Mun. Macaé,

Pico do Frade de Macaé, 700-900 m. 1 8 Apr 1985 (fr), Martinelliet al. 10734

178

Studies In Annonaceae. XVI A Toxonomic Revision of Ditgiielia A. F. C P de Sant-Hilaire

(U); Mun. Novo Friburgo, Macaé de Cima, 1 100 m, 26 Nov 1986 (fl),

Martinelli et al. 11934 (U); Petrópolis, Quitandinha, 15 Sep 1963 (fl),

E. Pereira 7660 (B, MO, RB).

Th is species comes very close to D. riedeliana and D. salicifolia , from

which it can be distinguished by its narrowly ovate lamina (Iess than 2 cm

wide) and blackened, broadly obconical carpels with truncate apex, sparsely

covered with stellate scalcs Table 3.

1 1. D. salicifolia R. E. Fries, Acta Horti Berg. 12(1): 48. 1934. Type. Brazil.

São Paulo: Serra do Mar, Fazenda Campo Grande, 1 8 Nov 1 892 (fl, fr),

Edwall 1988 (holotype, S; isotype, B). Figure 14

Big tree, height unknown. Young twigs densely (15-25 scales/mm2)

covcred with yellowish fimbriate to stellate scales, becoming glabrous with

age, older twigs greyish vvliite to dark brown, irregularly fissured. Petiole 3-5

mm long, 1-1.5 mm in diam., densely covered with yellowish fimbriate to

stellate scales. Lamina narrowly elliptic, chartaceous to slightly coriaceous,

green above, yellowish green below, (6-)8-l 1 cm long, (2-)2. 5-3.4 cm wide,

index (2.7-)3-3.7(-4), base attcnuate, apex acuminate, upper side glabrous,

slightly shiny, lower side sparsely to densely covered with yellowish

fimbriate(-stellate) scales, secondary veins uniformly curved, (1 1 -) 1 3- 1 6(- 1 8)

on either sides of primary vein, angles with primary veins (50 o -)55°-65 o (-75 o ),

intercostal area somewhat regularly shaped, distance between the middle of

closure and margin 2-5 mm, tertiary veins raised on both sides, irregularly

forming 6-12 nets, 4th order veins slightly raised on both sides.

Inflorescence leaf-opposed, l-2-llowered, subsessile, pedicel 6-10 mm

long, straight, 0.6-1 mm in diam. at base, ( 1 -) 1 .5-2 mm in diam. just below

flower. Bracts 2 to the pedicel, one amplectant on basal 1/5-1/2 of pedicel,

very broadly to depressed triangular-ovate, 0.8- 1 .5 mm long, 1 .5-2.5 mm wide,

anothcr one broadly to very broadly triangular, 0.5-1 mm long, 0.5-1 mm

wide, caducous. Pedicels and outer side of bracts densely covered with yellowish

(fimbriate to) stellate scales, inner side of bracts glabrous; outer side of sepals

densely covered with yellowish fimbriate to stellate scales, inner side of sepals

densely covered with yellowish ctirviform stellate hairs (basal 1/5- 1/4 sparsely

covered to glabrous); outer side of petals densely covered with yellowish stellate

scales, inner side densely covered with yellowish curviform stellate hairs (basal

1/6- 1/5 of outer petals and 1/5- 1/4 of inner petals glabrous).

179

Boi. Mus. Para. Emílio Goeldi. sér. Bot. 9(2), 1993

Flower buds broadly to very broadly ovoid, apex acute, 6-8 mm long,

7-8 mm in diam. Sepals connate to 1/6-1/5 from base, triangular-ovate to

broadly triangular-ovate, 9- 1 1 mm long, 7-8 mm wide. Outer petals rhombic-

obovate, apex apiculate to acute, 1 7-20 mm long, 8- 1 1 mm wide, inner petals

rhombic-obovate, apex apiculate (or acute), 1 6-20 mm long, 7-1 1 mm wide.

Torus depressed ovoid. Stamens 120-200, 0.8-1 .1 mm long, 0.3-0.4 mm wide,

apical prolongation of connective shallowly conical, glabrous, 0.3-0.4 mm

long, 0.5-0. 7 mm wide. Carpels 80-120, ovary densely covered with whitish

stellate scales and stellate hairs, stigma blackened, glabrous.

Fruit unknown (only loose carpels present). Fertile carpels narrowly an-

gular-obovoid to angular-obovoid, 30-35 mm long, 1 0- 1 6 mm in diam., with

5-6 ribs, top itself conical, densely covered with yellowish stellate scales, basal

2/3 of carpel fused. Seed filling up 2/3-3/4 of the carpel, ellipsoid, 1 8-22 mm

long, 14-16 mm in diam., yellowish-brown, roughish.

Distribution (Figure 2). Only known from São Paulo in Brazil. Flowering

mainly from September to December. The only record of fruiting time is in

November.

Speciniens examined. São Paulo: Alto da Serra, 28 Oct 1 936 (fl), Hoehne

& Gehrt s.n. (F, K, NA, NY, US, UC); Estação Biologica, Alto da Serra, 29

Sep 1 922 (fl), ./. G. Kuhlmann RB 19670 (RB, S); Estação Biologica, Alto da

Serra, 30 Sep 1922 (fl), .7. G. Kuhlmann RB 1967 6 (MO, RB, S); Estação

Biologica, Alto da Serra, 2 Dec 1 93 1 (fl), Lemos SP28313 (NY, S).

This species is very similar to D. microphylla and D. riedeliana , see

discussion under those species and Table 3.

ACKNOWLEDGEMENTS

The authors arc much indebted to the curators of all herbaria that sent us

material on loan. Thanks also go to J. C. Lindeman (U), who provided a

Portuguese abstract, and to H. Rypkema for his assistance in preparing the

drawings.

180

Studies In Annonaceae. XVI. A Toxonomic Revision of Dtignelia A. F. C. P. de Sanl-Hilaire

BIBLIOGRAPHICS REFERENCES

BAILLON, H. 1 868. Mémoire sur la famille cies Anonacccs. Adansonia 8: 327-336.

FRIES. R. E. 1934. Revision der Arten ciniger Anonaceen- Gattimgen - III. Acta Horti Berg. 12(1 ): 1 -44.

FRIES, R. E. 1937. id.- IV. Acta Horti Berg. 12(2): 221-228.

FRIES, R. E. 1959. Annonaceae, In: ENGLER. A.& PRANTL, K. Die Natiirlichen PJlanzenfamilien, 2.

ed. Berlin, Duncker & Humblot. 1 7A II: 1-171.

HICKEY, L. J. 1979. A reviscd classiflcation of thc architccture ofdicotyledonous leavcs. 2 ed. Oxford,

Clarendon Press.

KLUCKING, E. P. 1986. Leafvenation pattems(l). Annonaceae.: Berlin, J. Cramer, p. 1-256.

MARTIUS, C. F. Ph. von 1841. In: MARTIUS - Flora Brasiliensis. V. 13 Leipzig, Frid. Fleischerp. 1-64.

PRANTL, K. 1891. Dic natUrlichen Pflanzenfamilien. III. V. 2. Abteilung: 23-39.

RADFORD. A. E. et al. 1974. Vascular plant systematics New York., FIarpcr& Rovv, p. 85-166.

SAINT-HILAIRE, A. F. C. P. de. 1824. Flora Brasil iac Meridionalis. V. I : Paris, p. 34-36.

Recebido em 26 10.92

Aprovado em 19 0.1.93

NUMERICAL LIST OF TAXA

1. Dugueíia furfuracea (St. Ililairc) Bcntham et Flookcr f.

2. Dugueíia dicholepidota Martius

3. Dugueíia moricandiana Martius

4. Dugueíia marcgraviana Martius

5. Dugueíia echinophora R. E. Fries

6. Dugueíia lepidota (Miqucl) 1’tille

7. Dugueíia glabriuscula (R. E. Fries) R. E. Fries

8. Dugueíia lanceolala A. de Saint-I lilaire

9. Dugueíia riedeliana R. E. Fries

10. Dugueíia microphylla (R. E. Fries) R. E. Fries

1 1. Dugueíia salicifoUa R. E. Fries

LIST OF EXSICCATAE

Alemão e Cysneiros, F. F., 25. R60662 ( 1).

Almeida. E. de F„ 13(1).

Almeida, S. P. et al.. 852 (1).

Amaral. I. L. et al.. 1702 (6)

181

SciELO

10 11 12 13 14 15

Boi. Mus. Para. Emílio Goeldi, sér. Boi. 9(2). 1 993

Andcrson, W. R. etal., 8025, 8585, 8737. 9031, 9493 (1); 9637 (4); 10072, 35139, 35640,

36544, 36831 (1).

Andersson, L. & Ilagberg, M, 1635 (1).

Andrade, A. G. & Emmerich, M, 448 (1),

Andrade, N. de, SP25004 (1); R60903 (8).

Araújo, J. S. et al., 135 (3).

Araújo, D. & Plowman, T. C., 6680 (9).

Archer, W. A., 128.4069 (1).

Argent, G. ct al., 6525 (1).

Assumpção, C. T. de, 7548 (8).

Ayala, F„ 221 (1).

Azevedo. M. L. M. & Lopes. R. D., 787 (1).

13amps, P„ 5489 (1).

Barçante, I., 190 (I).

Barreto, H. L. M. [etal.], 420. 422, 424, 425, 1331. 3284, 4418, 8693 (1).

Battistclla, P„ 213=SP1451 1 (1).

Becker, R„ 37 (1).

Belém, R. P. [etal.]. 131, 3653 (1).

Bernardi, A. L.. 18926, 18930. 19460 (1).

Bicalho, H. D., 30 (I).

Black, G. A., 387 (1).

Blanchet, .1. S„ 1678 (3): 2828 (2); 3578 (1).

Bockermann, W., SP1 92940(1).

Bogner, J., 1 193 (I ).

Bordo, A., 42 (I ).

Brade, A. C„ 5882, 1 1 889, 1 3473 ( 1 ).

Brantjcs, N. B. M., 701607 (I).

Braza, J. E. M.. 38 (1).

Burchell, W. .1., 4974, 5669 (1).

Callcjas, R. & Carvalho, A. M. V. de, 1744 (3).

Carauta, .1. P. P„ 771 (I).

Cardoso, R. M. & Setz, E., 36 (5).

Carlos Filho, L. C.. 72 (1).

182

Stuclies In Annonaceae. XVI. A Toxonomic Revision ofDugiietiaA. F. C. P. de Sant-Hilaire

Carvalho, A. M. V. de et al., 1092 (3): 1716 (1).

Castellanos, A. [et al.J, 22526, 24190, 24288 (I).

Cavalcante, P. B., 3359 (5).

Cid, C. A. et al„ 4873. 4874. 6265. 6439 (4).

Claussen. M„ 36, 138. 331. 482. 1091. s.n. (1).

Cobra. L. Q. & Oliveira, J„ 233 (1).

Costa, E. & Mautonc, L., 60 (1).

Costa, J. S„ 75 (7); 111 (1).

Cunha, C. N. da et al., 788 (7).

Curran, H. M.. 178. 184 (1).

Cutler, H. C„ 8116(1).

Daly, D. C. et al., 459, 741, 873 (5); 1 155. 1323. 1535 (6).

Damião. C. & Mota. A.. 632 (5).

Davidse, G. et al., 121 14(1).

Dcdecca, D. M.. 514 (1).

Dias. A. A., 33 (1).

Duarte. A. P. [et al.|, 1389 (I); 2289 (8): 2790 (1); 3208 (8); 5136 (I).

Ducke, A.. 10. 432, 691 (5); RB7142 (1); RB 17868 (5).

Dusén, P. K. H„ 9667. 9730. 9731 (1); 9939 (8); 1 1751, 16198. 16442 (1); 16785 (8).

Eiten, G. et al., 2535. 2920. 2962 (1); 3832 (5); 5925, 6024. 9625. 9757. 9890 (1).

Edwall, G„ 145 (8); 1988 (11).

Feliciano. J. B.. 22 (1).

Eclippc, G. M.. 136 (1).

Ferreira. C. A. Cid et al.. see Cid, C. A. et al.

Ferreira. V. F. et al.. 694 (1).

Fonseca, M., 131 (3).

F razão. A.. RB8659 ( 1 ).

Frócs, R. L.. 1905, 1 1641. 24246 (5); 30122 (4); 30691. 34223 (5).

Gehrt, A.. SP3200. SP 17501 (I).

Gentry. A. 1 1. et al., 50040 ( 1 ).

Gibbs. P. E. et al.. 5102 (I).

GilTord. D. R.. 79 (1).

Glaziou, A. F. M.. 4746, 6464 (9); 8255 (8); 10230 (9); 10234 (1); 16689 (10); s.n. (1.8).

183

SciELO

10 11 12 13 14 15 16

Boi. Mus. Para. Emílio Goeldi, sér. Boi. 9(2). / 993

Gomes, M & Miranda, S., 356 (4).

Gomes, ?, 1 128 (1).

Gonçalves, D., SP50361 (1).

Goodland, R„ 324, 326, 482 (1).

Gregory, W. C. et al., 10587 (1).

Guedes, M. L. et al., 15826 (3).

Guedes, T. N„ 435 (I); 484 (10).

Guimarães, J. G., 164. 1122, 1513 (1).

Gumièrc, G„ RB 199 176 (1).

Handro, 0„ 2294 (1).

Handro, W„ 39(1).

Hanke, W„ 28 (1).

Harley, R. M. [et al.], CFCR6424=SPF36024 ( 1 ); 10665(4); 15041. 16717. 18634(1); 18917

(2); 1 9748, 20960, 2 1 1 1 4, 2 1 742, 2 1 856 ( I ); 24 1 02 (3); 25964, 27161, 27607 ( 1 ).

Hatschbach, G. et al., 13276. 14251, 21837. 26118, 29419, 31873. 35904. 40053, 42806,

43774, 47293 (1).

Heiner, A., s.n. ( I ).

I lemmendorff, E„ 278, 279. 280 (1).

Ilenschen, S., 11-3 (1).

1 leringer, E. P. [et al.], 4372, 5971,8868, 13250, 17419, 17580. 17966. 18400 (1),

llochne, F. C. [et al.], 549 (4); 1946 (I); 3461. 3462, 3463, 3464 (4); 4478. 4479. 4532.

4565, 4566, 4567, 4602. 4754. SP 17501 (1); SP29784 (8); SP35768 (I); SP38975

(8); s.n. (12).

Hopkins, M. J. G. et al., 507, 512 (6).

Irwin, H. S. et al.. 5460, 5572,6491 (1); 6870 (8); 7431, 9127, 9952, 10295, 11108, 13266,

13675, 14357, 14639. 14870. 14894. 16028. 16206(1): 16418(4); 16528. 17646(1);

1 7764 (4); 1 791 3 ( 1 ); 2 1 1 92 (4); 220 1 7. 22572. 22954. 23699. 25488, 25868. 263 1 2.

26817, 27569, 28427. 30914, 31496. 32471. 34693 (I).

Jangoux, .1. et al.. 438, 1058 (5); 1403 (7); 1410(1).

Jesus, .1. A. de, 1730 (1).

Joensson, G., 386a (1).

Kappler, A.. 1 686 (6).

Kirizawa, M. et al., 608, 1220 (I).

Klein, R. M„ 182. 886 (8); 936 (8).

Klug, G„ 1587 (7).

184

Studies In Annonaceae. XVI A Toxonomic Revision ofDuguetiaA. F. C. P. de Sant-Hilaire

Krapovickas, A. et al„ 32694, 33360 (1).

Kuhlmann, J. G., 6037 (9); RB 1 9670 ( 1 1 ); RB 1 9676 (11).

Kuhlmann, M. [et ah], 257 (5); 1589, 1667, 3951 (8); 4524 (1).

Labouriau, L. & Válio, 1. M., 1098 (I).

Labouriau, M. S„ 41. 86, 113(1).

Leitão, H. F. et al„ 13263 (1).

Leite, P. F. & Klein, R. M„ 58 (8).

Leitmann, M. et al„ 17 (10).

Lemos, C„ SP28313 (1 1).

Lima, A., & Magalhães, M., 53-1 138 (1).

Lima, H. C. et al„ 969 (1).

Lindeman, J. C. & Haas, J. H. de, 894, 4500 (I).

Lisboa, P. et at„ 2696 (4); 3059 (1); 3271 (4).

Lobão, J. & Almir, ?, RB 194252 (9).

Lobato, L. C. B. et al„ 141 (5).

Loefgren, A., 244 (8); 523. 596 (1): 4008, 5986 (8).

Lopez. M. L. S., 43 (1).

Loureiro, A. et al„ INPA35808 (5).

Lourteig, A., 2312 (8).

Luetzelburg, Ph. von, 26061, 26135 (1).

Lund. P. W., s.n.. ( I ).

Macedo, A., 965. 1593, 2395 (1).

Macedo, M. et al„ 305 (4).

Machado. 0„ 264. 284 (5).

Maciel, A.. 31 (1).

Maciel, U. N. et al„ 382 (5); 1597 (4).

Magalhães, F. M. R., 1 1 , 52 ( 1 ).

Magalhães, M„ 259, 1917. 4658. 14065, 17331 (1).

Magnago, I I. , 345 (I).

Maguire, B. et al„ 44529. 56853 (1).

Malmc, G. O. A.. 1 1 78. 1 1 78* (4); 1 756 ( I ); 1 779 (4); 1 779a, 2006, 2006a ( 1 ); 11-2322 (4);

11-2440(7).

Martinel li, G. [et al.], 402 (1); 10734, 11934 (10).

185

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Martius, C. F. P. dc, 214. 550, s.n. (1).

Mattos, A. de, 121 (8).

Mattos, J. & Mattos, N., 14427 (I).

Mello f„ L. E. de, 245 (1).

Mendes. L. O. T„ 1(1); SP44191 (8).

Mendonça, R. C. et al., 23, 1179, 1335, 1 374 ( 1 ).

Mileski, E„ 101 (I).

Miranda. F. E. et al., 384 (5).

Miranda, R.. 30 ( I ).

Moore, S. M.. 163 (1); 576 (4),

Moraes, E. C. C. et al., 124 (I).

Moreira, M. E. & Almeida, E. F.. 13 (1).

Mori. S. A. [et al.], 9437 (1); 9920 (9); 13369. 13472 (1); 14078 (3); 16717, 16902 (1).

Mosén, C. W. Ii„ 1098 (I).

Nascimento, .1. R„ 53 (5).

Neto, G. G, et al.. 813 (1).

Noberto. F. et al.. 40 ( 1).

Novaes, C„ 687 (1); 4576 (8).

Oliveira, E.. 170 (5).

Oliveira, J. E„ R1367977. RB71557 (1).

Oliveira. P. 1. [et al.], 9, 475 (1).

Orlandi, R. P„ 314. 319 (I).

Pacheco, M. ct al., 168 (5).

Pedersen. T. M„ 12052 (1).

Peixoto, A. E. & Peixoto, O. L.. 1012(1).

Pereira, A., SP29284 (9).

Pereira, E„ 73 1 3, 7466 ( 1 ); 7660 ( 1 0); 9 1 08 ( 1 ).

Philcox, D. et al.. 3065. 3484, 3822 (1).

Pinheiro, R. S.. 1420. 1961 (3).

Pirani, J. R ct al.. 1 193. 1278 (4); 1593, SPF22847=CFCR920, SPF36550=CFCR7550 ( 1 ).

Pires. .1. M. et al„ 2107 (4); 9079 (1); 12549 (6); 16506 (I); 17164 (7); 50732 (5).

Plowman. T. C. |ctal.|, 8532 (4); 12723AO); 12783 (3).

Polil. .1. 13. E.. 450. 937 (1).

186

Stndies In Annonaceae. XVI. A Toxonomic Revision ofDiiguetia A. F. C. I’. de Sanl-IIilaire

Prance.G. T. [et al.], 3961, 4130, 4327 (6); 6899, 7178 (4); 11467(5); 18891 (1); 19087(4);

25599 (5); 26272 (I); 29924 (9); 58630 (4); 58684, 58931 (5); 59305 (4), 59551 (4).

Puiggari, .1. I., 523=SP17876 (1).

Ramamoorthy, T, P., 499, 623 ( 1 ).

Ramos, J. F. & Sousa, R., 7(1).

Rankin, J. et al., 42 (5).

Ratter. J. A. et al., 384, 852, 918 (1); 1019 (4); 2097 (1); 4438 (4); 5916 (6).

Regnell, A. F„ II-3, s.n. (1).

Reitz, R. [et al.], 1647, 3044, 3204, 3585. 3750. 7342, 7483. 9248 (8).

Riedel, L„ 174 (8); 1471 (4); 1626, 1686 (8); 1889 (1); s.n. (1,9).

Rizzini, C. T., RB1 14058 (1).

Robert, A., 432b ( 1 ).

Rodrigues, 1. A. et al., 329 (1).

Rodrigues, W. A., 1809 (5); 10144 (6).

Romariz, D., 103 (1).

Rosa, N. A. | et al.], 3058 (6); 3638, 4903, 5170 (5).

Rosário, C. S. et al., 1 16 (5); 665 (6).

Rossi, L. et al., CFSC6985 (1).

Russel, A., 286 (1).

Saint-IIilaire, A. F. C. P. de, s.n. (1, 8).

Sali. A„ SP184741 (1).

Sampaio, A. J., 6507, 6624, 6925, s.n. (1).

Santos, E. et al., 1801.2509 (1).

Santos, .1. U. & Rosário, C. S.. 392, 406 (7).

Santos, M. R„ 570 (5).

Santos, N„ R60848 (1).

Santos, R. S. & Castellanos, A.. 24288, 25190 (1).

Santos, T. S. dos, 2522 (1).

Schallcr, G„ 153 (4).

Schatz, G. E. et al., 772, 827, 835 (5).

Schwacke, C. A. W., 14057 (1).

Schultz, ?, 1 103 (8).

Sellow, F., s.n. (9).

187

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Shamel. ? et al.. 235b. 303b (1).

Shepherd, G. J. et al.. 4133, 7267, 1 1246 (1).

Silva, A. S. L. da et al., 1617 (6).

Silva, F. C. da et al., 217 (4).

Silva. M. F. F. da et al., 39 (6).

Silva, M. G. et al.. 2545 (5); 3603 (6); 4136, 4609 (1); 5459 (6); 5636 (5); 6029 (1).

Silva, N. T. fct al.], 3596 (6); 4801 (4).

Silva, P. E. N. et al., 175 (1).

Siqueira, O. P., 8(1).

Souza, O., 3 1 38=R1 12827=SP40614 (I).

Souza, S. A. M. da et al., 252 (6).

Stannard, B. L. et al., SPF36269CFCR6669 (1).

Stickney, F„ RB42422 (8).

Sucre, D. [et al ], 586 (1); 9247. 9400 (5); 10438 (1).

Tamberlik, ?, s.n. (1).

Taylor, E. L. et al„ EI 129 (6).

Teixeira, L. O. A. et al.. 612 (4).

Thomas, W. W. et al., 4550 (1); 4712 (4); 5578 (1).

Toledo, J. F. & Gehrt, A., SP43 1 94 ( 1 ).

Unknown collector, SP30620 (8).

Usteri, A., 3a=SPI7878 (I).

Valentim, A„ 3401 (8).

Válio, I. M„ 60, 239, 269, 270 (1).

Vanni, R. et al., 245 (1).

Vauthier, ?, 578 (1).

Vavrek, M., 158 (1).

Vecchi, O., 221 (8).

Venturieri, G. A. & Menezes, J. M. T., 18 (4).

Vidal, J„ V-228 (1).

Viegas, A. P. [et al.], 5386, 5551 (1 ).

Vieira, H. C. W„ 588 (8).

Violatti, L. G. et al., 15 (1).

Wacket, M., s.n., (1, 8).

188