AMAZONIANA XIV (3/4): 33s-341 Kiel, Dezember 1997
On the vegetation of a seasonal mixedwal¿r inundation forest near Manaus, Brazilian Amazonia
by
I.L. do Amaral, J. Adis & G.T. Prance
M.Sc. Iêda Leão do Alnaral, Instituto Nacional de Pesquisas da Amazônia (INPA)' Caixa Postal 478,6901l-970 Manaus, AM, Brazil. Priv.-Doz. Dr. Joachim Adis, Max-Planck-lnstitut für Limnologie, AG Tropenökologie, Postfach 165, D-24302 Plön, Germany' Prof. Dr. Ghillean Prance, Royal Botanic Gardens, Kew, Richmond, Suney TWg 3AB' England. (Accepted for publication: September 1997).
Abstract
The species composition of an area of 0.5 hectare of seasonal inundation forest located between the blackwater Negro River and the whitewater Solimões River at Lago Janauari near Manaus. Brazil. was basal area studie Keywords: lnundation forest, igapó, vtirzea, floodplains, ecotone, Amazon, Neotropics. ISSN 0065-67551199713351 o MPI für Limnologie, Ac Tropenökologie, Plön; INPA, Manaus 335 Introduction Results and discussion In Central Amazonia, there are seasonat changes in the water level of the Rio Floristic composition Solimões-Amazonas and the Rio Negro averaging about l0 m. Forests beside the rivers Thirty-one plant families were recorded on the study site (Table l). Leguminosae are flooded to a depth of several meters for five to seven months each year. These sensu latu was richest in species: eighteen species belonged to the Fabaceae, seventeen natural events have occurred for at least 2.4 years million (lRIoN et al. 1997). The to the Mimosaceae, and ten species to the Caesalpiniaceae. The farnily Euphorbiaceae, pronounced seasonal periodicity of the "monomodal floodpulse,'(JUNK et al. l9g9) has with seven species, ranked highest for Family ltnportance Value (FIV) equal to 35 (cf. brought about long-term adaptations and the evolution of endemic species of plant and MORI et al. 1983). It was followed (in decreasing order of FIV; see Table l) by the animals (cf. JUNK 1997). They represent organisms characteristic either of "seasonal Sapotaceae, Clusiaceae, Mimosaceae, Fabaceae and Moraceae. Caesalpiniaceae, Lecythi- igapó", (forest annually flooded by black- and/or clearwater rivers) or of "seasonal daceae, Myristicaceae and Lauraceae ranked intermediate, with a FIV front l9 to I l. varzea" (forest flooded by whitewater rivers), pRANCE definitions accor 336 JJI Vegetation type CAMPBELL, D.G.. DALY. D.C.. PRANCE, G.T. & U.N. AMCIEL (1986): Quantitative ecologicat According to the floristic categories of Amazonian forest types subject to inundation inventory of terra firme and várzea tropical forest of the Rio Xingú. Brazilian Amazon. - Brittonia (PRANCE 1979, 1980), the vegetation in the study area contains tree species charac- 38(4):369-393. teristic the seasonal ofboth várzea, and the seasonal igapó. and also trees which occur ERWIN. T.L. (1983): Beetles and other insects of tropical forest canopies at Manaus, Brazil, sampled by in both types of inundation forest (Table 4). Therefore the forest investigated is a insecticidal fogging. - In: SUTTON, S.L.. WHITMORE. T.C. & A.C. CHADWICK (eds.): Tropical mixture of seasonal varzea & igapo an is better termed a"mixedw,ater" inundation forest rain forest: Ecology and management: 59-75. - Proc. Trop. Rain Forest Symp. (Leeds 1982). Blackwell because of the influence of both whitewater and black/clear waters. One of the reasons Scientific. Publ., Oxford: 498 pp. is the dispersal of diaspores by water and fish (hydrochory, ichthyochory) during the lRION, C., JUNK, W.J. & J.A.S.N. DE MELLO (1997): The large Central Amazonian river floodplains near Manaus: Geological, high water period (cf. KUBITZKI & ZIBURSKI l9S4). The mixedwater inundarion hydrological, and geomorphological aspects. - ln: JUNK, W.J. (ed.): The Central Amazon floodplain. Ecology ofa pulsing system:23-4ó. - Ecological Studies l2ó, Springer, forest should therefore be considered an ecotone with respect to both flora and terrestri- Berlin: 525 pp. al invertebrate fauna (cf. ADIS 1992; MORAIS 1995; MORAIS et al. 1997a, b). IRMLER, U. (1975): Ecological studies ofthe aquatic soil invertebrates in three inundation forests of Another example of this occurs in the Anavilhanas Archipelago in the Rio Negro, where Central Amazonia. - Amazoniana 5(3): 337-409. the large whitewater river Rio Branco influences the igapó vegetation of the Rio Negro IRMLER, U. (1976): Zusammensetzung, Besiedlungsdichte und Biomasse der Makrofauna des Bodens in (cf. RODRIGUES l96l). Further floristic and faunistic inventories of this area are der emersen und submersen Phase zentralamazonischer Úberschwemmungswälder. - Biogeographica currently in progress. 7: 79-99. IRMLER. U. (1977): lnundation-forest types in the vicinity of Manaus. - BiogeographicaS: 17-29. JUNK, W.J. (ed.) (1997): The Central Amazon floodplain. Ecology of a pulsing system. - Ecological Resumo Studies 126, Springer, Berlin: 525 pp. JUNK, W.J.. BAYLEY, P.B. & R.E. SPARKS (1989): The flood pulse concept in river-flooplain sy- Estudou-se a conrposiçào dc espécies numa área de 0.5 hectare de unra floresla inundável sazonal, stems. - Can. Spec. Publ. Fish. Aquat. Sci. t0ó: I l0-127. situada no Lago Janauarí perto de Manaus, Brasil, entre o Rio Negro. cle água preta, e o Rio Solir¡òes, de KEEL. S.H. & C.T. PRANCE (1979): Studies of the vegetation of a white-sand black-water lgapó (Rio água branca. Registrou-se 66 espécies arbóreas, representando 55 gêneros e 3 I lamílias. A área basal total Negro, Brazil). - Acta Amazonica 9(4): 645-655. foi de l5 mr e o volutne do luste 125 mr. As espécies mais abundantes îoran Poutct.la sp. (Sapotaceae) KLINGE. H., ADIS, J. & M. WORBES (1995): The vegetation of a seasonal várzea forest in the lower e Plentcurpus Qtttlzonum (Fabaccae). A maior área basal por espécic foi dc 2.15 r¡t parn Culoph.tllunt Solimòes River. Brazilian Amazonia. - Acta Amazonica 25(314): 201 -220. brttsiliense (Clusiaceae). A vegetação na área de estudo representou urn ecótono com espécies dc árvores KUBITZKI, K. & A. ZIBURSKI (1984): Seed dispersal in flood plain forests of Amazonia. - Biotropica características para a floresta sazonal na regiào ila água branca (véúzeal ou/e da água preta (igapó) da 26( I ): 30-43. Amazônia Central. Seguindo a tertninologia para os tipos de florestas amazônicas sujeitas à inundação LAMPRECHT. H. (1972): Einige Strukturmerkmale natürlicher Tropenwaldtypen und ihre waldbauliche (PRANCE 1979) denomina-se, desta forma. "várzea & igapó sazonal" ou floresta inundável de "água Bedeutung. - Forstwiss. Centralblatt 9l: 270-277. nlista". MORAIS, J.W. DE (1995): Abundância, distribuição vertical e fenologia da fauna de Arthropoda de uma região de água mista, próxima de Manaus. AM. - Ph.D.-thesis, ESALQ/Univ. São Paulo, Piracicaba, Brasil: 226 p. Acknowledgments MORAIS, J.W. DE, ADIS, J., MAHNERT, V. & E. BERTI-FILHO (1991a): Abundance and phenology of Pseudoscorpiones (Archnida) from a mixedwater inundation forest in Central Amazonia, Brazil. - We thank the technicians of Botany at INPA, Manaus/Brazil. in particular Luiz Coelho. for help Revue suisse Zool.: 104(3): 475-4835. received in the fìeld and herbarium. Berit Hansen. Max-Planck-lnstitute for Linrnology, plön/Germany, MORAIS, J.W. DE, ADIS, J,, BERTI.FILHO. E., PEREIRA, L.A., MINELLI, A. & F. BARBIERI kindly rnade the drawings. B.Sc. Francisca Dionízia de Alrneida Matos, INpA. Manaus/Brazil. helped (1997b): On abundance, phenology and natural history ofGeophilomorpha (Chilopoda. Myriapoda) computing the plant inventory data. Dr. Danielle Waldhoff, University of Kicl/Gerrnany. Dr. Maria T.F. from a mixedwater inundation forest in Central Amazonia. - Ent. scand. Suppl.: in press. Piedade and Dr. Lúcia Maia. INPA, Manaus/Brazil as well as PD Dr. Martin Worbes. University of MOREIRA, E. (1970): Os lgapós e o seu aproveitamento. - lmprensa Universitária, Belém: l-33. MORI, Göttingen/Germany made valuable comments o the manuscript. Special thanks to Sr. Dió, supervisor of the S.4., BOOM, 8.M., CARVALHO, A.M. DE & T.S. DOS SANTOS (1983): Southern Bahian moist Ecological Park ofSelvatur at Lago JanauarilBrazil. forest. - Bot. Rev. 49: l-232. PIRES, J.M. (1984): The Amazon forest. - ln: SIOLI, H. (ed.): The Amazon. Limnology and landscape ecology of a mighty tropical river: 581-602. - Monographiae Biologicae, Junk Publ., Dordrecht: References 763 pp. PRANCE, G.T. (1979): Notes on the vegetation of Amazonia III. The terminology of Amazonian forest (1992): types subject to inundation. - Brittonia 3l(l): 26-38. ADIS, J. Überlebensstrategien terrestrischer Invertebratcn in überschwemmungswäldern Zen- PRANCE, G.T. (1980): A terminologia dos tipos de florestas amazônicas sujeitas a inundação. - Acta tralanrazoniens. - Verh. naturwiss. Ver. Hamburg (NF) 33: 2l-l 14. Amazonica l0(3): 495-504. ADIS' J. (1997): Terrestrial invertebrates: Survival strategies, group spectrum, dominance and activity patterns. REVILLA, J.D. (1981): Aspectos florísticos e fitosociológicos da floresta inundávei (lgapó) Praia Crande, - In: JUNK, w.J. (ed.): The central Amazon floodplain. Ecology of a pulsing system: 299- Rio Negro, Amazonas, Brasil. - M.Sc.-thesis, 129 p. 317. - Ecological Studies 126, Springer, Berlin:525 pp. INPA, Manaus: 338 339 -o RODRIGUES. W.A. (1960): Relatório preliminar da viagem de reconhecimento pelos Rios Uatumã e o\o !+o o Urubu. - Bol. Geogr. l8(159): 1045-1072. z(o C\¡ o RODRIGUES, W.A. ( l96l ): Estudo preliminar de mata de várzea alta de uma ilha do baixo Rio Negro de o, solo argiloso e úmido. - CNPq/INPA, Sêrie Botânica l0: l-50. o; TAKEUCHI, M. (1962): The structure of the Amazonian vegetation. Vl. lgapó. - J. Fac. Sci. Tokyo. Sect. Ât o) 3, Bot 8(7): 297-304. WORBES, M. (1983): Vegetationskundliche Untersuchungen zweier Überschwemmungswälder in Zen- tralamazonien - vorläufige Ergebnisse. - Amazoniana 8(l):47-65. oì ø WORBES, M. ( 198ó): Lebcnsbedingungen und Holzwachstum in zentralanrazonischen ÜbcrschwcrÌtnungs- o) d o) d wäldern. - Scripta Geobotanica 17: l-112. I ¿ (1997): o WORBES, M. The forest ecosystem of the floodplains. - In: JUNK. W.J. (ed.): The Central o) o Amazon floodplain. Ecology ofa pulsing system: 223-265. - Ecological Studies 126. Springer. Berlin: 525 pp. o? d O) WORBES, M., KLINGE, H., REVILLA, J.D. & C. MARTIUS (1992): On the dynamics. floristic sub- @ 6 I division and geographical distribution of várzea forests in Central Amazonia. - J. Veg. Sci. 3: 553-564. U) o d O J q d o) ! (o |\ oI o f.- 6 oìÊ q)() (o \ t- s tU) o \ õE P (d o?3 d o)o o roõ 6 (f) rc o o: d ro .!9 E o oì o) : \f $ F I o o o $ Þ. 6 ol o) llho N (r) do Coreiro F I 0 o (f) ^t D .o q o o) o rO C\I (f) æ oI o' C\I o q o o) o Map l: € I a Location of the mixedwater inundation forest Lago Janauarí (03'2'S.60"17'W). at o o F "o Ë ooooo òöd =F o@(o$c\t o üi5 340 341 N o N% ro 79 80 40 \to 70 35 60 (Ðâo 30 È Ci {¿ ct gD cr) c É 50 25 _o 6 R! (t H¿ o 40 35 20 I a É 29 28 o 30 .t G 15 (t 19 o þo 20 10 d o -J o o o d 5 ôl 10 5 o roo € 0 0 o 6 0-4.9 5-9.9 10 - 14.9 15 - 19.9 20 - 24.9 >25 o 13 tr $ É height classes (m) ù o a3 N G (Ðão \ 25 È ÑP a t¿ oil c È _o Oñ CL' a a (\t o 20 G (É o o o o ø o o o q. E 5 o o o o o. .o o E o o o o c: 0 ô.t o o rO õ I o .0) $o a 0 o I (Ðã o 2468 10 12 14 16 18 20 22 24 26 ct È o l¿ height (m) ¡sS -c s _-o -o oil ñ! ! Fis.2: a o Dbh-class distribution of 198 trees (dbh > l0 cmi top) and frequency oftree height (bottom) sampled on a. dôt) 0,5 ha of a seasonal mixedwater inundation forest at Lago Janauarí near Manaus. I o 'Úo o o. 6s o _è0 o o o Þoõ C\I [Å (u) ¡q6¡eq 342 343 Table l: Plant families in 0.5 ha ofthe seasonal mixedwqter inundation forest at Lago Janauari near Table 2: Species composition in 0.5 ha of the seasonal mixedwater inundation forest at Lago Janauarí Manaus. FIV Family = Importance Value. near Manaus. SIV = Species Importance Value. Plant family Number Number Number Relative Relative FIV Planl sp€c¡ss Family Number Basal stv of trees area (m2) of genera of species of trees basal area tree density 1 Pouteria sp. Sapotacoae o.73 '17.53 2 Câlophyilumbras,Tlense Camb. Clusiaceae 1 Euphorbiaceae 7 7 23 12.97 11.62 35.19 2.15 17.19 3 Pterocarpus amæonum (Matl.ex Bth.) Amsh. Fabaceae 2 Sapotaceae 4 0.76 15.89 4 24 7.61 12.12 25.79 4 Pseudolmedia sp. Moraceae 1.24 12.63 3 Clusiaceae 3 3 I 16.29 4.55 25.38 5 Hura crop¡tans l. Euphorbiaceae 0.9'1 12.53 6 Leco¡ntea amazon,bá Ducko Câêsalpiniaceae 4 Mimosaceae 4 5 17 7.46 0.60 10.53 8.59 23.62 7 /Veeá sp. Nyctaginaceae 't8 o.20 9.32 5 Fabaceae 3 4 8.14 9.09 23.29 I Eschwe¡leru sp. Lecylhidacêae 0.31 9.09 6 Moraceae 2 4 10 9.32 5.05 20.44 I Heveaspruceana Muell. Arg. Euphorbiaceae 0.29 8.95 10 Ttích¡l¡a sp. Meliaceae 7 Caesalpiniaceae 3 3 10 9.57 0.34 8.60 5.05 19.16 '11 Vircla suñnamans¡s (Rol.) Warb Myrislicaceae 0.50 7.49 8 Lecythidaceae 2 4 10 2,59 5.0s 13.70 12 P¡thecellob¡um sp- M¡mosaceae o.22 7.25 '13 Rheedia sp. 9 Myristicaceae 1 2 I 3.88 4.55 11.45 Clusiaceae 0.20 7.10 14 Painari excelsa Sab. Chrysobalanacêae 10 Lauraceae o 6.98 2 3 1.79 4.55 10.89 15 ¡ndêt. Laurac€ae 0.23 6.63 11 Anonaceae 2 3 7 0.67 3.54 8.75 16 Unonopsis sp. Annonac6ae 0.07 6.37 17' Acac¡a sp. M¡mosaceae 0.54 6.1'1 12 Nyctaginaceae 1 I 1.40 4.55 7.46 '18 I S/oanaa sp. Elaeocarpaceaê o.47 5.58 13 Meliaceae 1 1 7 2.37 3.54 7.42 19 Cecropia sp. C6cropiaceae 0.13 5.42 'l Microphol¡s sp. 14 Chrysobalanaceae 1 2 4.63 1.01 7.16 20 Sapolaceae 0.19 5.38 21 Camp¡s¡andra sp. Caêsalpiniaceae 15 Olacaceae 2 2 2 2.29 0.42 5.27 1.0'l 6.33 22 Eugenia sp. Myrtaceae 0.07 5.19 16 Elaeocarpaceae 1 2 2 3.22 1.01 5.75 29 So/ocaa sp. Moracêa€ 0.04 5.00 24 V¡rola elongata (Bth.) Warb. Myrist¡cacêae 17 Combretaceae 2 1 2 1.15 1.01 5.19 0.06 4.98 25 Labat¡a sp. Sapotaceae 18 Rubiaceae ,| 0.13 4.95 2 3 0.41 1.52 4.95 26 Swanzia sp. Caesalpinlacsae 0.36 4.87 19 Cecropiaceae 1 1 5 0.87 2.53 4.91 27 Vata¡rca sp. Fabacêae 0.19 4.83 28 Leon¡a sp. Violacêae 20 Apocynaceae 2 2 2 0.37 1.01 4.41 0.17 4.79 29 Mabea sp. Ëuphorb¡acêae 0.05 4.41 2'l Violaceae 1 1 3 1.19 1.52 4.22 30 Phanheatñlol¡ata Baill. Euphorbiacaae 0.46 4.35 Ha¡steia sp. 22 Myrlaceae 1 1 4 0.47 2.02 4.01 31 Olacaceae 0.32 3.38 32 lnga sp.2 Mimosacoaê o.17 3.37 23 Polygonaceae 1 1 2 0.45 1.01 2.98 33 /nga sp.1 Mimosacoae 0,08 3.25 24 Sterculiaceae 1 1 2 0.24 1.01 2.76 34 Cæoloba sp. Polygonacêae 0.07 2.81 35 /nga sp. Mimosaceae 0.06 25 Efihroxylaceae 1 1 1 0.13 0.51 2.15 2.81 36 indêt Rubiacoae 0.05 2.70 26 Ochnaceae 1 1 1 0.12 0.51 2.14 37 Neoxythecê elogans (A.DC.) Aubr. Sapolaceae 0.05 . 2.67 27 Rutaceae 1 1 1 0.11 0.51 2.13 38 indet. Sterculiaceaê 0.03 2.60 'l 39 Nectandra sp. Lauraceae o.o2 2.44 28 Bombacaceae 1 1 0.09 0.51 2.'t1 40 Buchenav¡amacrophyla Eichl Combretaceao 0.16 2.26 29 Hippocrateaceae 1 1 1 0.09 0.51 2.11 4'l Vataircagu¡anens¡s Aubl. Fabaceae 0.14 2.17 42 Alchom¡a sp. Euphorbiaceaê 1 '1.97 30 Ebenaceae 1 1 1 0.08 0.51 2.10 0.1 43 Drypetês sp. Euphorbiaceae o.o2 1.81 31 Flacourtiaceae 1 1 1 0.05 0.51 2.07 44 Etabail¡a sp. Fabaceaê 0.08 1.76 45 Pseudolmedia ct. laev¡gata Trcc. Moraceaê 0.06 1.58 46 Asp¡dospema sp. Apocynacêao Total 55 66 198 100.00 100.00 300.00 0.04 1.47 47 Gustav¡a hexapsfala (Aubl.) Smith Lêcyth¡dacêae 0.03 1.41 48 Sap¡um sp. Euphorb¡aceaê 0.03 1.36 49 Xylop¡a sp. Annonac€ae 0.02 1.31 50 Etythroxylum sp Erythroxylacêae 0.02 1.31 51 Duro¡a sp. Ochnacêaê 0.02 1.30 52 Zanthoxylum compactum (Hubsr êx Albuq.) Wat. Rutaceae 0.02 1.29 5g Eschwe¡lèra albillora (4. DC.) Miere Lecythidaceae 0.01 1.28 54 Endl¡cheda sp. Lauraceae 0.01 1.28 55 Ce¡ba pentandra Gaed.. Bombacaceae 0.01 1.27 56 Sa/ac,a sp. H¡ppocrateacôaè 0.01 1.27 57 M¡nquartia sp. Olacaceaê 0.01 1.27 58 Himathanthue sp. Apocynaceae 0.01 1.26 59 D¡ospyrce sp. Ebenac€ae 0.01 1.26 60 Gustav¡a augusta l. Lecylhidacoaê 0.01 1.26 61 indet. Annonacoae 0.01 - 1.25 62 Th¡loa paragua¡lensls Eichler Combretâc6ae 0.01 1.25 69 S¡ckingia t¡nctoña (H.B.K.) K.Sch. Rub¡acêae 0.01 1.25 64 Cara¡pa sp. Clusiaceae 0.01 1.24 65 Caseaila aculeata Jacq. Flacourl¡aceae 0.01 66 ¡ndel. Moraceae 0.01 1.22 Total 14.48 300 '= Lianas 344 34s --l õ- xf o DJ Oo Jo 1t l+ õi ða-!o) =E 5O=.@ Ð! oa õ' Iu ) O¿ iIUJJ[\)JJIUIUIU - l+E (¡)\¡!(o(J)cl)(o('|OIu Ît =@ (tl tto o o \Þ o À(JJ ::Þìo o\ @ è' !t ø ìâ ir g ' -ò, -N--SDPgtù99 Ð l+L o(o(¡)tuoo)(oJo)@ =o @('rcDÀo(Jl(oÍ\)JJ o ÞÀ -@ !, o 53^-= 3 N ao Ø3ôÐ E o o o- ñ- J LN N) l\) fU o :=9 l;1 (rrroo!Àæ(tr('torf¡l\)lgof¡!¡Psosrg, SOö oo(/)G)ofÐrJsæo' 3 :m o) o 3 o Þ= ?= Table 4: Tree species of the mixedwa¡er inundation forest at Lago Janauarí near Manaus, representing the characteristic vegetation of seasonal varzea orland seasonal igapó. (Assignment according to KEEL & PRANCE 1979; KLINGE et al. 1976; KUBITZKI & ZIBURSKI 1994; MOREIRA 1970; PRANCE 1979, 1980; REVILLA l98l: RODRIGUES 1960; TAKEUCHI 1962; WORBES 1983, 1986. 1997; WORBES et al. 1992). + = recorded on the study site of 0.5 ha. vatzea ¡qapo várzea & iqapó * Buchenavia macrophylla E¡chl. Neoxythece elegans (A.DC.) Aubr. Astrocaryum jauari Mart. (Combretaceae) (Sapotaceae) (Arecaceae) * Caesaria aculeata Jacq. Th i loa paragu arien s is Eichl. Calophyllum brasiliense Camp. (Flacourtiaceae) (Combretaceae) (Clusiaceae) Ceiba pentandra Gaert. Virola elongata (Bth.) Warb. Eschweilera albiflora (A.DC.) Miers (Bombacaceae) (Myristicaceae) (Lecythidaceae) Crataeva benthamii Eichl. " Gustavia augusta L. (Capparidaceae) (Lecythidaceae) Gustavia hexapetala (Aubl.) Smith * Hevea spruceana Muell. Arg. (Lecythidaceae) (Euphorbiaceae) sUJ Heisteria spruceana Engl. Parinari excelsa Sabine -J (Olacaceae) (Chrysobalanaceae) Hura crepitans L. * Vatairea guianensis Aubl. (Euphorbiaceae) (Fabaceae) Lecointea amazonica Ducke (Caesalpiniaceae) Piranhea trifoliata Baill. (Euphorbiaceae) Pseudoxandra polyphleba R. E. F ries (Annonaceae) Rheedia brasiliensis (Maft.) Tr. & Pl. (Clusiaceae) Triplari s su ri namen sis Ch am. (Polygonaceae) Virola surinamensis (Rol.) Warb. (Myristicaceae) Vitex cymosa Bth. ex Spreng. (Violaceae) Zanthorylum compactum (Huber. ex Albuq.) Wat. (Rutaceae)