New Phytol. (1992), 121, 563-570

Occurrence of nodulation in legume species in the Amazon region of Brazil

BY FATIMA MARIA DE SOUZA MOREIRA\ MARLENE FREITAS DA SILVA^AND SERGIO MIANA DE EARIA^ ^ Coordenagdo de Pesquisas em Ciencias Agronomicas (CPCA), Instituto Nacional de Pesquisas da Amazonia (INPA), Alameda Cosme Ferreira, 1756 Caixa Postal 478, Manaus, AM, Brazil CEP 69083 ^Centro de Ensino e Pesquisas Florestais (CEPEF)/ Universidade de Tecnologia da Amazonia (UTAM), Av. Darcy Vargas, 1200 Parque Dez, Manaus, AM ^Centro Nacional de Pesquisa de Biologia do Solo (CNPBS), Empresa Brasileira de Pesquisa Agropecudria (EMBRAPA), Estrada Rio, Sao Paulo, km 47, Seropedica, Itaguai, Rio de Janeiro, Brazil CEP 23850 {Received 23 October 1991 \ accepted 8 April 1992)

SUMMARY This paper reports the nodulation status of 172 legume species from different ecosystems within the Amazon region of Brazil. The nodulation ability of 98 species and eight genera was observed for the first time. Occurrence of nodulation is discussed in the context of the taxonomy of the family and the different ecosystems. The frequency of nodulation among tribes and genera was found to be higher in flooded {varzea and igapo) than in non-flooded sites {terra firme); moreover, more plants had nodulated in nursery beds consisting of soil from flooded areas. The symbiosis may therefore be favoured in flooded areas.

Key words: Nodulation, Leguminosae, taxonomy, Amazon region.

INTRODUCTION MATERIAL AND METHODS The Brazilian Amazon region comprises an area of The location of the six different sites examined is about 5 X 10® km^, with non-uniform ecosystems shown in Figure 1. Details of each site are shown in (Sioli, 1967; Braga, 1979; Falesi, 1986). Almost all of Table 1. Field observations and sampling of nodules this area is covered by forests with a very diverse from seedlings and/or adult plants were made as flora, among which the Leguminosae is well repre- described earlier (Faria et al., 1984; Magalhaes, sented, being the largest family in diversity and the 1986). Nitrogenase activity (acetylene reduction), fifth in density (Ducke, 1949; Black, Dobzhansky & was measured on individual nodules from each Pavan, 1950; Rodrigues, 1961, 1967a, b; Klinge & species. Some nodules were desiccated in vials Rodrigues, 1971 ; Prance, Rodrigues & Silva, 1976). containing CaClj or silica gel, others were preserved Many legume species have been used by local in aqueous 70 % ethanol for subsequent structural farmers for timber and firewood production studies. Rhizobia were isolated from nodules show- (Loureiro, Silva & Alencar, 1979; Arckoll, 1984a), ing the highest rates of acetylene reduction following in agro-forestry systems (Escalante, Herrera & the procedure of Vincent (1970). Plant material was Aranguren, 1984), in reafforestation of abandoned collected for identification and cataloguing at the sites and also for human food (Arckoll, 19846). The Herbarium of the National Amazon Research In- capacity of some of these species to fix Nj is regarded stitute (INPA). Whenever seeds were available, they as an important feature in tree establishment on were collected for confirmation of the nodulation nutrient-deficient soils, and is also taken into account status found in field conditions. They were sown in in the choice of species for the various purposes plastic bags containing sand, fertilized with nitrogen- listed above. free nutrient solution and inoculated with more than This paper reports the nodulation status of 172 a hundred rhizobial strains. Alternatively seeds were legume species from different ecosystems in the germinated in representative soils of the Amazon Amazon region. region: gley humic soil and podzolic soil (ultisol)

38 ANP 121 564 F. M. De Souza Moreira, M. F. Da Silva and S. M. De Faria

Figure 1. Location of tbe sites wbere tbe nodulation status of plants was observed. For details of eacb site see Table L , Roads; , state limit.

respectively from flooded and non-fiooded sites. Table 1. Details of sampling sites shown in Figure 1 Occurrence of nodulation was recorded after 5 months. Vegetation type wbere nodulation status of plants Sites was observed RESULTS (1) Manacapuru Forests seasonally flooded Municipality by black rivers Unless specified, the classification of the genera Terra firme forests follows Polhill & Raven (1981). The nodulation Disturbed areas in terra status of 172 legume species was examined: 49 firme Caesalpinioideae, 60 Mimosoideae and 63 Papilio- Forests seasonally flooded by wbite rivers noideae. Of the 98 species examined for the first (2) Careiro Municipality Terra firme forests time, 43 exhibited nodulation while 55 were non- Disturbed areas in terra nodulating (Table 2). In the Caesalpinioideae only firme eight species were found to be nodulated out of the Forests seasonally flooded 49 examined. The status of three genera was by black rivers (3) Anavilbanas Arquipelago Forests seasonally flooded observed for the first time: two nodulating {Camp- by black rivers siandra and Dicorynia), and one non-nodulating (4) Manaus City Terra firme forests {Heterostemon). The 45 original reports on the status Disturbed areas in terra of the nodulation in the Papilionoideae include five firme new genera: one nodulating (Etaballia) and four Planting in terra firme Forests seasonally flooded non-nodulating {Bocoa, Lecointea, Monopteryx and by black rivers Aldina). Twenty-nine mimosoid species are also Forests seasonally flooded reported for the first time as being able to nodulate. by wbite rivers The observations were made basically in two (5) Road Manaus-Caracarai, Terra firme forests different types of vegetation: igapo and vdrzea, Br. 174 Disturbed areas in terra where the vegetation is flooded during part of the firme (6) Br. 369, Porto Acre City, Terra firme forests year; and terra firme, where flooding does not occur. Placido de Castro City Disturbed areas in terra Sites five and six (see Fig. 1 and Table 1) are firme exclusively on terra firme. Nodulated legumes of the Brazilian Amazon 565

Table 2. Occurrence of nodulation in legume species in the Amazon region of Brazil

Occurrence of nodulation^ No. of SUB-FAMILY Herbarium plants Habit Terra Igapo and Tribe reference or exam- of the firme vdrzeas Nursery Species collector no. ined species^ (non-flooded) (flooded) bed

CAESALPINIOIDEAE Caesalpinieae Caesalpinia ferrea Mart, ex Tul. FMSM 252 6 A/S ne Caesalpinia pulcherrima (L.) Sw. FMSM 254 6 S ne ne Dimorphandra caudata Ducke*'' FMSM 415 1 Y — ne ne Dimorphandra parviflora INPA 133.638 10 A - ne 4- Spruce ex Benth. Campsiandra comosa Bth, var. INPA 133.836 47 A/S ne 4- 4- laurifolia (Benth.) Cowan* Mora paraensis Ducke* INPA 133.640 28 A/S ne Sehizolobium amazonicum Hub. ex Ducke 3 A/S — ne ne Sclerolobium melanocarpum Ducke INPA 133.613 1 A — ne ne Sclerolobium chrysophyllum INPA 151904 1 A — ne ne Poepp. of Endl.* Sclerolobium hypoleucum Benth.* INPA 138.806 3 S ne ne -1- Sclerolobium odoratissimum FMSM 80 1 A ne — ne Spruce ex Benth.* Tachigali paniculata Aubl. INPA 138.801 32 A/S ne 4- 4- var. paniculata Vouacapoua pallidior Ducke FMSM 153 3 A + ne ne Cassieae Apuleia molaris Spruce ex Benth.* FMSM 543 1 A ne ne Cassia fastuosa Willd, ex Benth. 1 A — ne ne Cassia grandis L. 2 A — ne ne Cassia leiandra Benth. INPA 133.623 12 S ne ne — Chamaecrista negrensis (Irwin) INPA 124.723 17 A/S — 4- 4- Irwin & Barneby* Chamaecrista nictitans (L.) Moench. INPA 133.854 7 H/S 4- 4- ssp. disadena (Steud.) Irwin & Barneby Dialium guianensis (Aubl.) Sandw. FMSM 37 4 A ne Dicorynia paraensis Benth.* FMSM 409 1 A ne 4- ne Senna iatifolia (G.F.W. Mey) 1 A/S — ne ne Irwin & Barneby Senna multijuga (Rich.) INPA 133.622 14 S/A - ne — Irwin & Barneby Senna quinquangulata (Rich.) INPA 133.609 7 L/S — ne — Irwin & Barneby* Senna siamea (Lam.) Irwin & Barneby INPA 133.617 6 A ne ne Senna silvestris (Vel.) INPA 133.860 2 A/S/A — ne ne Irwin & Barneby* Detarieae Copaifera multijuga Hayne INPA 133.870 22 A/S ne Copatfera reticulata Ducke 2 S ne — ne Crudia amazonica Spruce ex Benth.* INPA 124.720 16 A/S ne — — Crudia pubescens Spruce ex Benth.* INPA 133.838 18 A ne — — Cynometra bauhiniifolia Benth.* INPA 133.839 25 A/S ne — — Eperua bijuga Mart, ex Benth. INPA 133.614 14 A/Y/S — ne — Heterostemon mimosoides Desf.* INPA 138.793 28 A/S ne — — Elizabetha bicolor Ducke* FMSM 62 1 A — ne ne Hymenaea courbaril L. FMSM 2T 5 S ne ne — Hymenaea parviflora Huber FMSM 106 13 A — ne Peltogyne paniculata Benth. INPA 133.633 15 A/S — — — Peltogyne prancei M.F. da Silva* INPA 133.861 6 A/S — ne — Peltogyne catingae Ducke ssp. glabra FMSM 173 1 A — ne ne (W\ Rodr.) M.F. Silva* Peltogyne excelsa Ducke* FMSM 178 1 A ne ne Peltogyne venosa Benth. FMSM 56 2 A ne — ne Amherstieae Macrolobium limbatum Spruce ex Benth.* FMSM 147 2 A ne ne Macrolobium microculyx Ducke* FMSM 221 1 A — ne ne Macrolobium acaciifolium Benth. INPA 124.719 67 A/Y/S ne — — Macrolobium angustifolium* INPA 133.849 1 A ne ne (Benth.) Cowan 38-2 566 F. M. De Souza Moreira, M. F. Da Silva and S. M. De Faria

Table 2. (cont.)

Occurrence of nodulation^ No. of SUB-FAMILY Herbarium plants Habit Terra Igapo and Tribe reference or exam- of the firme vdrzeas Nursery Species collector no. ined species' (non-flooded) (flooded) bed Macrolobium gracile Spruce ex Benth.* INPA 133.630 1 A — ne ne Cercideae Bauhinia acreana Harms. 1 A — ne Bauhinia angulata L.* FMSM 444 1 Y — ne ne Bauhinia longicuspis Benth. FMSM 67 1 L/S — ne ne PAPILIONOIDEAE Swartzieae Aldina latifolia Spruce ex Benth.* 2 A ne _ ne Atdina heterophylta Spruce ex Benth.* 2 A ne - ne Bocoa viridifolia (Ducke) Cowan* FMSM 165 2 A — ne ne Lecointea amazonica Ducke* FSMS 9 1 A/S ne - — Swartzia auriculata (Poepp.) Endl.* INPA 133.639 1 A ne + ne Swartzia bannia Sandw.* FMSM 397 1 A ne + ne Swartzia reticutata Ducke* FMSM 149 1 A — ne ne Swartzia laevicarpa Amsh.* INPA 138.784 16 A/S ne - + Swartzia panacoco (Aubl.) Cowan* INPA 133.862 1 A - ne ne Swartzia sericea Vog.* INPA 133.846 1 S — ne ne Swartzia schomburgkii Benth.* FSMS 146 2 A -1- ne ne Siuartzia ulei Harms* INPA 133.593 6 A — ne ne Swartzia ingifolia Ducke* INPA 133.599 4 A/Y — — ne Swartzia cuspidata Benth.* FSMS 148 3 A — ne ne Swartzia potyphylla DC* INPA 133.845 24 A/S + -1- + Sophoreae Amburana acreana (Ducke) A.C. Smith 6 A/S — ne — Acosmium nitens (Vog.) Yakovlev* INPA 133.841 2 A ne - ne Bowdichia virgitioides Kunth FSMS 445 1 A — ne ne Ctathrotropis nitida (Benth.) Harms* INPA 133.643 3 A/Y ne -1- ne Diplotropis purpurea (Rich.) Amsh. FSMS 170 2 A -1- ne ne Ormosia excelsa Benth.* INPA 138.791 21 A/S ne -1- + Ormosia macrocalyx Ducke* INPA 138.851 19 A/S ne + 4- Ormosia smithii Rudd* INPA 133.600 1 A — ne ne Ormosia paraensis Ducke* INPA 133.866 2 A ne - ne Ormosia discolor Spruce ex Benth.* INPA 133.850 19 A/S — ne -1- Monopteryx inpae W. Rodr.* FSMS 63 1 A — ne ne Dipterygeae Dipteryx odorata (Aubl.) Willd. INPA 133.603 27 A/S — ne — Dipteryx pimctata (Blake) Amsh. FMSM 61 1 A ne — ne Dipteryx magnifica Ducke* INPA 133.624 1 A — ne ne Taralea oppositifolia Aubl.* FMSM 175 1 A — ne ne Aeschynomeneae Aeschynomene rudis Benth. INPA 133.833 1 Y ne + ne Dalbergieae Andira micrantha Ducke* INPA 133.872 10 A/S — ne + Andira parviflora Ducke* INPA 133.867 1 A ne — ne Andira surinamensis Benth. INPA 133.871 1 A — ne ne Andira unifoliotata Ducke INPA 133.875 2 A — ne ne Dalbergia inundata Benth.* INPA 124.717 32 L/S ne -1- + Dalbergia riparia (Mart.) Benth.* INPA 133.857 20 L/S ne -1- -1- Dalbergia riedelii (Radlk.) Sandw.* FSMS 403 1 L ne + ne Etaballia dubia Rudd* INPA 133.843 4 A ne + ne Hymenotobium heterocarpum Ducke* INPA 133.873 2 A + ne ne Hvmenolobium modestum Ducke 9 S + — + Hymenolobium petraeum Ducke* FSMS 130 1 A — ne ne Hymenolobium sericeum Ducke* FSMS 154 1 A — ne ne Machaerium hoeneanum Ducke* FSMS 539 1 A - ne ne Machaerium ferox (Mart. FSMS 124 13 L ne 4- ne ex Benth.) Ducke* Machaerium froesii Rudd FSMS 15 1 A ne 4- ne Machaerium inundatum (Mart, ex INPA 124.718 54 L/S 4- 4- + Benth.) Ducke Machaerium madeirense Pittier 2 A + ne ne Machaerium quinata (Aubl.) Sandw.* INPA 138.792 30 L/S + ne Nodulated legumes of the Brazilian Amazon 567

Table 2. {cont.)

Occurrence of nodulation^ No. of SUB-FAMILY Herbarium plants Habit Terra Igapo and Tribe reference or exam- of tbe firme vdrzeas Nurserv Species collector no. ined species' (non-flooded) (flooded) bed

Platymiscium duckei Huber* INPA 139.629 21 A/S ne Pterocarpus amazonicus Huber* INPA 124.721 27 A/S ne Vatairea guianensis Aubl. INPA 124.716 42 A/S ne — Millettieae Derris amazonica Killip. INPA 138.795 2 L ^ ne Derris negrensis Bentb.* INPA 138.790 52 L ne ne Derris longifolia Bentb.* INPA 133.840 1 L ne ne Pbaseoleae Clitoria pinnata (Pers.) Baill 5 S ne ne Clitoria amazonum Mart, ex Bentb.* INPA 133.837 28 A/Y ne 4- -1- Clitoria javitensis (Kuntb) Bentb.* FMS 448 1 L ne 4. ne Clitoria fairchildiana Howard FSMS 264 12 A ne ne + Dioclea malacocarpa Ducke* FSMS 431 1 L -t- ne ne Dioclea macrocarpa Huber* FMS 449 1 L + ne ne Erythrina fusca Loureiro INPA 133.832 1 A ne -1- ne Mucuna urens (L.) Medic. INPA 138.789 6 L ne ne MIMOSOIDEAE Parkieae Parkia decussata Ducke FSMS 260 14 S ne Parkia discolor Bentb.* INPA 133.586 21 A/S ne Parkia multijuga Bentb. INPA 133.597 13 A/S — — — Parkia oppositifolia Bentb.* INPA 133.615 27 A/S — ne — Parkia pectinata (Kuntb) Bentb.* FSMS 70 1 A — ne ne Parkia pendula Bentb. INPA 133.604 13 A/S — ne — Parkia ulei (Harms) Kubl.* FSMS 427 1 A ne — ne Pentaclethra macroloba (Willd.) Kuntze INPA 133.616 23 A/S ne -1- -1- Acacieae Acacia multipinnata Ducke* FSMS 505 1 L — ne ne Acacia paniculata Willd.* FSMS 524 1 Y — ne ne Acacia polyphylla DC. FSMS 424 2 A -1- ne ne Mimoseae Adenanthera pavonina L.* INPA 133.612 6 S ne ne — Anadenanthera peregrina (L.) Speg. INPA 133.619 7 A/S -1- ne Dinizia excelsa Ducke FSMS 261 12 S _ ne Entada polyphylla Bentb. INPA 133.621 32 L -1- -1- -1- Mimosa pigra L. INPA 133.590 13 A ne -1- -1- Mimosa polystachya Kuntb* INPA 133.597 6 H ne ne -f. Mimosa spruceana Bentb. 1 A — ne ne Stryphnodendron guianense 20 S — ne -1- (Aubl.) Bentb. Stryphnodendron pulcherrimum INPA 133.646 12 s -1- -1- -1- (Willd.) Hocbr. Stryphnodendron microstachyum INPA 133.584 6 s ne ne — Poep. & Endl.* Ingeae Abarema jupunba (Willd.) FSMS 176 8 A -1- -1- ne Britton & Killip Abarema adenophora (Ducke) FMS 450 1 A + ne ne Barneby & Grimes* Albizia corymbosa (L.C. Ricb) FSMS 422 1 A H- ne ne G.P. Lewis & P.E. Owen Albizia polyantha (Spreng) G.P. Lewis* INPA 124.726 5 S ne ne + Albizia saman (Jacq.) F.V.M. FSMS 267 13 A/S -1- ne + Calliandra tenuiflora Bentb.* FSMS 437 25 A/S — ne Calliandra surinamensis Bentb. FSMS 256 12 S ne ne + Cedrelinga catenaeformis Ducke FSMS 126 13 A/S -1- + -1- Enter olobium schomburgkii Bentb. FSMS 105 16 A/S + ne -1- Inga acreanum Harms* INPA 138.786 25 S ne ne 4- Inga calantha Ducke* FSMS 538 26 A/S -1- ne Inga capitata Desv.* INPA 133.869 1 A ne -H ne Inga cinnamomea Spruce ex Benth. INPA 133.620 7 S ne ne .^- Inga disticha Bentb.* INPA 138.805 25 s ne ne -1- 568 F. M. De Souza Moreira, M. F. Da Silva and S. M. De Faria

Table 2. {cont.)

Occurrence of nodulation^

N1 ^ nU . oKJif SUB-FAMILY Herbarium plants Habit Terra Igapo and Tribe reference or exam- of the firme vdrzeas Nurserv Species collector no. ined species^ (non-flooded) (flooded) bed

Inga edulis Mart. FSMS 41 1 A -1- ne ne Inga heterophylla Willd. INPA 133.627 12 A/S -1- ne -1- Inga ingoides (Rich.) Willd. FSMS 7 44 A ne + -1- Inga laterifiora Miq. FSMS 100 1 S + ne ne Inga macrophylla Kunth INPA 133.834 14 A/S — -1- -1- Inga meissneriana Miq.* FSMS 389 1 A ne + ne Inga nobilis Willd.* INPA 133.835 1 A ne + ne Inga panurensis Spruce ex Benth.* INPA 133.596 1 A — ne ne Inga rubiginosa (Rich.) DC* INPA 133.848 1 A -1- ne ne Inga speciosa Spruce ex Benth.* FSMS 295 1 Y ne — ne Inga thibaudiana DC. INPA 133.606 1 A — ne ne Inga uraguensis Hook & Arn. 10 S + ne -1- Marmaroxylon racemosum (Ducke) Killip INPA 133.595 2 A — ne ne Pithecetlobium (Macrosamanea) INPA 138.809 3 S ne ne + adiantifolium Benth.* Pithecellobium (Abarema) INPA 133.591 41 A/S + ne + arenarium Ducke* Pithecellobium {Abarema) INPA 133.592 1 A ne — ne auriculatum Benth.* Pithecellobium (Marmaroxylon) INPA 133.847 1 A — ne ne claviflorum Benth.* Pithecellobium (Macrosamanea) FSMS 112 1 Y — ne ne duckei Huber* Pithecellobium (Macrosamanea) INPA 133.588 3 A/S ne 4- ne lindsaefolium Spruce ex Benth.* Pithecetlobium (Albizia) pediceltare INPA 133.594 1 A 4- ne ne (DC.) Benth. Pithecellobium {Zygia) INPA 138.799 25 S ne ne + sanguineum Benth.* Zygia caulifiora (Willd.) Killip INPA 133.853 26 A ne 4- + Zygia glomerata (DC.) Pittier* INPA 133.855 1 A ne + ne Zygia inaequalis (Humb. INPA 133.844 1 A ne 4- ne & Bonpl.) Pittier Zvgia latifoha INPA 138.800 30 S ne ne -t- (L.) Fawc. & Rendle

'- A, adult plant; Y, young plant; S, seedling; H, herbaceous; L, liana. ' —, plant without nodules; -I-, plant with nodules; ne, not examined. ^ Species designated by an asterisk are new records.

With the exception of Etaballia nodules, nitro- nodulated species to total number of species analyzed genase activity was detected in at least some samples grown on (1) sand inoculated with rhizobia, (2) gley of nodules from all of the species studied. In 52 humic soil, (3) podzolic soil (ultisol) with N-free species, from which seeds were harvested and fertilizer and (4) podzolic soil without fertilizer were, inoculated with a large number of rhizobial strains in respectively, 95, 85, 73 and 65%. plastic bags, nodulation in the field was confirmed in nursery beds. Six other species {Dimorphandra parviflora, Swartzia laevicarpa, Ormosia discolor, DISCUSSION Andira micrantha, Platymiscium duckei, Calliandra The fact that 56 % of the species examined for their tenuiflora), though lacking nodules in the field, were ability to nodulate are new reports, shows how little shown to be capable of nodulation in nursery beds. is known about the occurrence of nodulation in the These were all found as adult plants in the field and, Leguminosae. with the exception of S. laeviearpa, were observed in No nodules have been found on species of the terra firme. Three species lacking nodules in the field tribes Amherstieae, Detarieae and Cercideae, con- and not analyzed in nursery beds are already well firming early reports that nodulation in the Caesal- known as nodulating species: Inga thibaudiana, pinioideae is largely restricted to the Caesalpinieae Bozvdichia virgilioides and Andira surinamensis (Allen and Cassieae (Allen & Allen, 1981; Faria et al., & Allen, 1981; Faria et al., 1984). The ratios of 1989). In the Caesalpinieae, nodules have been Nodulated legumes of the Brazilian Amazon 569 found in species that belong to the groups Pelto- nutrients are well known in the Amazonian oxisols phorum, Dimorphandra and Sclerolobium. Sclero- and ultisols, the most representative soils of lobium chrysophyllum was found without nodules Amazonia terra firme. Sites fiooded by 'white water during this survey, but its ability to nodulate has rivers', rich in nutrients, such as the Amazon itself been confirmed in the state of Bahia (Faria, un- (Sioli, 1967), exhibit high levels of nutrients (except published results). Reports of the nodulation of for N, because of leaching). Sites fiooded by black Mora species are disputed (Allen & Allen, 1981): and clear rivers show lower levels of nutrients (Sioli, here the inability to nodulate was confirmed in the 1967). At both sites, nodulation may be favoured by field and also after massive rhizobial inoculation. In nitrogen limitation. Furthermore, higher numbers of the Cassieae, only species of Chamaecrista have been free-living Ng-fixing bacteria have been found on found to nodulate, though the first genus of the fiooded sites (Sylvester-Bradley, Oliveira & Podesta Dialineae {Dicorynia) group is now reported as Filho, 1980; Magalhaes & Dobereiner, 1984). nodulating. Nodulation in the Swartzieae is far from being a general feature of the tribe: only species of the genus ACKNOWLEDGEMENTS Swartzia were found with nodules. The nearest The authors would like to thank G. P. Lewis, H. C. Lima relatives of Swartzia, Aldina and the other genera and J. I. Sprent for helpful discussion, and FINEP and have been observed not to nodulate. CNPq (Brazil) for financial support. So far no nodule has been reported on any genus within the Dipterygeae, which once more confirms this to be the only entire tribe of the Papilionoideae REFERENCES where nodulation does not occur (Faria et al., 1989). ALLEN, O. N. & ALLEN, E. K. (1981). The Leguminosae. Uni- The new report of nodulation of Etaballia, along versity of Wisconsin Press. MacMillan Publishing Company, with its nodule morphology and anatomy (results not Madison and London. ARKCOLL, D. B. (1984a). Uma compara^ao de algumas especies shown), confirms the actual position of this genus in de crescinfiento rapido adequadas para produ^ao de lenha em the Dalbergieae (Polhill, 1981). Nodules oi Etaballia tropicos umidos. Pesquisa Agropecudria Brasileira 19, 61-68. are of the aechynomenoid type according to Corby ARKCOLL, D. B. (19846). Algumas arvores leguminosas que produzem frutos uteis no norte do Brasil. Pesquisa Agropecudria (1981). Faria et al. (1984) and Sprent, Sutherland & Brasileira 19, 235-240. Faria (1989). This type of nodule is found only in Bu-^CK, G. A., DoBZHANSKY, T. H. & PAVAN, C. (1950). Some the Aeschynomeneae, Adesmieae and part of the attempts to estimate species diversity and population density of trees in amazonian forest. Botanical Gazette 3 (4), 413-^25. Dalbergieae. BRAGA, P. I. S. (1979). Subdivisao fitogeografica, tipos de vege- Nodulation in the Mimosoideae has been shown to tapao conservaijao e inventario Horistieo da floresta amaz6nica. be a general feature: 90 % of observed species Acta Amazonica 9 (4), Suplemento, 53-80. CORBY, H. D. L. (1981). The systematic value of leguminous root nodulate (Faria et aL, 1989). In this survey the lower nodules. In: Advances in Legume Systematics, part 2 (Ed by frequency of nodulation among mimosoid species R. M. Polhill & P. H. Raven), pp. 657-669. Royal Botanic (63 %) was mainly due to the non-occurrence of Gardens, Kew. DUCKE, A. (1949). Notas sobre a Flora Neotropiea. II. As nodules in Parkia spp. leguminosas da amaz6nia brasileira. Boletim Tecnico do Instituto The fact that five adult, potentially nodulating Agronomico do Norte 18, Belem. species had been observed without nodules on terra EscALANTE, G., HERRERA, R. & ARANGUREN, J. (1984). Nitrogen fixation in shade trees {Erythrina poeppigiana) in cacao planta- firme sites indicates the difficulty of finding nodules tions in North Venezuela. Pesquisa Agropecudria Brasileira 19, in dense forests and/or on adult plants. In most 223-230. tribes that were examined at fiooded sites, more FALESI, I. C. (1986). Present knowledge of the soils of the Brazilian Amazonia. In : Proceedings of the 1st Symposium on the species were nodulated (37 out of 69 examined), as Humid Tropics vol. I, pp. 168-191. EMBRAPA, CPATU, compared to the situation at non-flooded sites Belem, PA. (31/100). At fiooded sites, more abundant nodulation FARIA, S. M. DE. FRANCO, A. A., JESUS, R. M. DE, MENANDRO, M. DE S., BAITELLO, J. B., MUCCI, E. S. F., DOBEREINER, J. & was also observed. SPRENT, J. I. (1984). New nodulating legume trees from south These observations were confirmed in the nursery East Brazil. New Phytologist 98, 317-327. bed experiment, where 85 % of the species from FARIA, S. M. DE, LEWIS, G. P., SPRENT, J. I. & SUTHERLAND, J. M. (1989). Occurrence of nodulation in the Leguminosae. different tribes had nodulated when grown on soil New Phytologist 111, 607-619. from flooded sites while only 65 % had nodulated on KLINGE, H. & RoDRiGUES, W. A. R. (1971). Materia organica e non-fiooded soil as a substrate. This proportion nutrientes na mata de terra-firme perto de Manaus. Acta Amazonica 1 (1), 69-72. was increased to 73 % when the latter treatment LouREiRO, A. A., SILVA, M. F. & ALENCAR, J. C. (1979). Essencias was chemically fertilized with all nutrients except madeireiras da Amazonia, vols I and II (Ed. by Conselho nitrogen. This indicates that the symbiosis is Naeional de Desenvolvimento Cientifico e Tecnologico, Insti- tuto Nacional de Pesquisa da Amaz6nica and Superintendencia limited by nutrients on non-flooded sites, but not da Zona Franca de Manaus). Imprensa oficial do Estado do on gley humic soils (Ranzani, 1979; Magalhaes & Amazonas. Dobereiner, 1984). Flooded sites may therefore be MAGALHAES, F. M. M. (1986). Present state of knowledge on biological nitrogen fixation in Amazonia. In: Proceedings of the more favourable areas for the legume-rhizobia sym- 1st Symposium on the Humid Tropics, vol. I, pp. 499-512. biosis. Deficiencies of P, K, Ca, Mg and some micro- EMBRAPA, CPATU, Belem, PA. 570 F. M. De Souza Moreira, M. F. Da Silva and S. M. De Faria

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