Allpahuayo: Floristics, Structure, and Dynamics of a High-Diversity Forest in Amazonian Author(s): Rodolfo Vasquez Martinez and Oliver L. Phillips Reviewed work(s): Source: Annals of the Missouri Botanical Garden, Vol. 87, No. 4 (Autumn, 2000), pp. 499-527 Published by: Missouri Botanical Garden Press Stable URL: http://www.jstor.org/stable/2666143 . Accessed: 12/07/2012 11:15

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http://www.jstor.org ALLPAHUAYO: FLORISTICS, Rodolfo Vdsquez Martznez2and STRUCTURE, AND Oliver L. Phillips3 DYNAMICS OF A HIGH-DIVERSITY FOREST IN AMAZONIAN PERU'

ABSTRACT

This paper describes the results of a floristicinventory at the Allpahuayo Reserve, near Iquitos in Amazonian Peru. Two long-termone-hectare plots were established using a pre-determinedsampling grid, with each individual and liana over 10 cm diameter collected at least once, except forpalms. The plots were re-censused after5 years to quantify forestdynamics. Floristic analysis shows that the Allpahuayo forestis among the most diverse site yet inventoried,with 281 to 311 per hectare, and at least 466 species and 61 families in the 1277-stem two-hectare sample, confirmingthat upper Amazonia is a world center of tree biodiversity.The ecologically most dominant and speciose in the plots is sensu lato, with 231 stems and 89 species; no other family represents more than 7% of the species or 10% of the stems. In contrast to the exceptional floristicdiversity, both the structureand the dynamics of the Allpahuayo forest are similar to those recorded fromother old-growthneotropical forests. Many tree and liana canopy species were previously unknown to both the Iquitos area and to Amazonian Peru, which demonstrates the significance of Amazon ecological studies to systematic botany.

RESUMEN

Este documento describe el estado actual de los resultados de un inventario floristicoa largo plazo, en la Reserva Biol6gica de Allpahuayo, cerca de Iquitos, en la Amazonfa Peruana. Alli se establecieron dos parcelas de una hect. cada una, usando un muestreo pre-determinado, cada Arbol, palmera o liana, mayores o iguales que 10 cm de diAim. fu6 marcado y colectado, excepto las palmeras que fueron colectadas solo una vez por especie. Las parcelas fueron re-censadas despues de 5 afios para cuantificar el proceso de la dinamica del bosque. El analisis de los resultados nos muestra que el bosque de Allpahuayo, esta entre los bosques mas diversos hasta ahora inventariados, con 281 a 311 especies por hect., y al menos 466 especies y 61 familias en 1277 individuos en dos hect. muestreadas; 6stos datos apoyan la idea de que el oeste de la Amazonia es un centro mundial de diversidad de arboles. En las parcelas la familia Fabaceae s.l., es la mas dominante ecologicamente y en numero de especies, contiene 231 tallos y 89 especies; ninguna otra familia representa mas que el 7 por ciento de las especies o el 10 por ciento de los tallos. En contraste a la excepcional diversidad floristica, tanto la estructura del bosque como la tasa de mortalidad en Allpahuayo, estan dentro los rangos de otros sitios neotropicales. El valor de los estudios ecol6gicos para la botanica sistematica, se demuestra en tanto que, antes de que empezaramos a tomar datos ecol6gicos e instalar parcelas permanentes, muchas especies de arboles y lianas eran desconocidas tanto en el area de Iquitos, como en la Amazonia Peruana. Key words: Amazonia, floristics,inventoiy, Neotropics, species richness, species turnover.

' We gratefullyacknowledge the help of institutionalsupport fromInstituto de Investigaciones de la Amazonia Peruana (IIAP) for permission to work in Allpahuayo Reserve and for logistical support forour work; the John D. and Catherine T. MacArthur Foundation for long-termsupport of the Missouri Botanical Garden's floristicand ecological research at Allpahuayo and elsewhere in Peruvian Amazonia; the National Geographic Society for support for forest dynamics research in Amazonian Peru (grant #5472-95), and the U.K. Natural Environment Research Council for a Research Fellowship (OP). Melchor Aguilar, C6sar Grandez, Rosa Ortiz de Genthy,Nestor Jaramillo, Dennis Milanowski, JohnJ. Pipoly III, Peggy Stern, Henk van der Werff,and Arturo Vasquez Martinez assisted in establishing, inventoiying,and re-censusing the Allpahuayo plots. Herbarium determinationswere provided by numerous systematistsat MO and other herbaria, and we are especially grateful to: C. C. Berg (BG); T. Pennington, Sir G. Prance (K); A. Chanderbali, R. Gereau, R. Liesner, J. J. Pipoly III, G. Schatz, C. Taylor, M. Timana, H. van der Werff,and the late A. H. Gentry(MO); D. Daly, S. Mori, and M. Stern (NY). The maps were drawn by A. Manson at the School of Geography Graphics Unit, University of Leeds. We thank V. Hollowell, Sir G. Prance, and H. van der Werfffor their constructive suggestions to improve our manuscript. 2 Missouri Botanical Garden, Apartado 020, Ja6n, Cajamarca, Peru. 3Centre for Biodiversity and Conservation, School of Geography, Universityof Leeds, Leeds LS2 9JT, U.K.

ANN. MISSOURI BOT. GARD. 87: 499-527. 2000. 500 Annals of the MissouriBotanical Garden

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10Km B1 Figure 1. Map showinglocation of studyregion in relationto Iquitos and Peru. -A. Peru. -B. Iquitos region, showinglocation of Allpahuayo.Modified from Kalliola et al. (1998).

Biodiversityis at riskin muchof the tropics. Yet floristicunderstanding by generatinglarge numbers biologicalresearch and conservationinitiatives are ofnew collectionsthat are associatedwith site-spe- hampered by inadequate baseline information. cific ecological information.Botanical institutions Thus, sufficientdata on the numbers,kinds, and play a vital role in initiatingand supportingthis abundanceof most major and animaltaxa are work,and contributeessential expertise for ensur- not available. Moreover,knowledge of species as- ing accuratevoucher determination. Equally, eco- sociationsand biogeographicdistributions of taxa logical inventoriesgenerate much biogeographical remainmeager (McNeely et al., 1990; NationalSci- data ofrelevance to monographicand phylogenetic ence Board, 1990; National Research Council, studies. Such eco-floristicresearch has recently 1992; Phillips& Raven, 1996). been initiatedin severaltropical countries, includ- In theNeotropics, the extraordinary species rich- ing Madagascar(Lowry et al., 1997; Rakotomalaza ness, ourlimited taxonomic understanding, and the & Messmer,1999), (Rudas, 1996), and physicalinaccessibility of mostareas are formida- Peru (Gentry,1988a, b). This paper reportsthe re- ble barriersto phytogeographicinvestigation. Tra- sults of a similarresearch program at the Allpa- ditionalbiological inventory efforts that rely mostly huayoReserve, near Iquitosin northernPeru. on ad hoc collectionefforts and lengthymonograph studies cannot address the knowledgegap alone. STUDY SITE Alternativemethods are criticalto improveour un- derstandingof the factors determining species com- The AllpahuayoReserve (3O57'S, 73026'W) lies positionand theecological dynamics of tropical for- southwestof Iquitos in AmazonianPeru, between est ecosystems.In recent years there has been theblackwater Rio Nanayon thenorthwest and the growingscientific interest in moreecological, plot- Iquitos-Nautaroad to the southeast(Fig. 1). This based workas a means of understandingtropical 2750-ha reserveis administeredby the Peruvian forests(e.g., Gentry,1988a, b; Phillips & Gentry, Institutefor Amazonian Research (IIAP). The cli- 1994; Phillipset al., 1994; Dallmeier& Comiskey, mateis humidand hot (withthe mean annual pre- 1998a, b). This approachcan contributegreatly to cipitationabout 3000 mmand an averagetemper- 0 0 ~~0m500m 25 B20 24...... 24 I 23 ~~~~~~~~23m I5 229....Z 22 2m c

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4-lO~~~~m-~~~ 500x 300m x mm e-1O ...... Fiur 2 Dara sowngte rrngmetofpltswthnerannt507 50 00m amliggrd t lpauao.Th21mplt7(1"an octe t heede ofthgid Adiioalprale(rasets"E.ad.U,.haedinth.fgue).Frcopaisn.e tmpray amle f 00X mwee stblsed10 minfrm heede "".aels.so th rlaiv ostinso adiioal50 2 mpot tatinopoat tasets"E ad"U a ohr its n eu CucoA aznio.ndScuar/AER)bu.nt.tAlpauao 502 Annals of the Missouri Botanical Garden

atureof 260C; Marengo,1998). AllpahuayoReserve posited in two Peruvian herbaria(AMAZ, USM) lies between110 m and 180 m above sea level. and at MO, withpartial collections existing at HAP Edaphicconditions are variable,representing a mo- and the UniversidadAgraria La Molina (MOL), as saic of patches rangingin texturefrom clayey to well as distributionsto familyspecialists world- almost pure sand, and in drainage fromwater- wide. loggedswamps to well-drainedhill tops.Vegetation Fertilecollections were made on occasionwhen- is mature,old-growth forest, although some palm everflowers or fruitsof plot species wereseen, but (Oenocarpusbataua Mart.)have been cut to in commonwith other tropical plot inventories most harvesttheir . Allpahuayo and neighboring species (90%) are representedonly by sterilema- Mishanacomprise one ofthree locations within 100 terial. At MO every collectionnumber was first kmof Iquitos where the MissouriBotanical Garden identifiedto familyby the firstauthor and Ron has been conductinga long-termfloristic inventory Liesner,with each familyset then distributedto effortwith the institutionalcollaboration of HAP familyspecialists (, Apocynaceae, and ExploramaTours S.A. (Vasquez, 1997). The Bignoniaceae,Burseraceae, Chrysobalanaceae, Dil- other two sites, Reserva Ecologica Explorama leniaceae, Fabaceae, Lauraceae, Lecythidaceae, Lodge (Yanamono)and ReservaEcologica Sucusari Malpighiaceae,Melastomataceae, , Mor- (ACEER), have ratherdifferent soil conditionsand aceae, Myrtaceae,Rubiaceae, Sabiaceae, Sapinda- floristiccomposition. The 3750 ha of Allpahuayo- ceae, Sapotaceae, Simaroubaceae,Violaceae). For Mishanais extremelyspecies-rich, with more than mostspecies at least threefertile Peruvian collec- 10% of the plant species reportedfrom the whole tionswere notedfor verification (MO), mostderiv- of Peru (Brako & Zarucchi,1993) recordedhere ing fromthe Iquitos Florula project (Vasquez, (Vasquez, 1997). 1997). Re-census of both plots was done in February METHODS 1996, withthe re-measurementof all treesand li- anas. Plantsthat had died in theintervening period Samplingtook place withina 500 X 500 m grid of 5.25 years were noted as such, with evident in theAllpahuayo Reserve, sufficiently far from the cause of death. Each new recruitinto the 10 cm Iquitos-Nautaroad (> 2 km at its nearestpoint) to DBH class was measured,tagged, and given a minimizehuman disturbance, yet close enoughto unique number.New recruitswere collected and the researchstation (3 km) to allow us to commute vouchersdistributed to herbariaas forthe original daily.Within the sampling grid, two long-term plots collections. wereestablished in a pre-determinedlocation with- In order to maximizepotential comparability in the grid (Fig. 2) to eliminatesubjective bias. with otherforests, diversity values for each plot Each plot is 20 X 500 m, witheach plant - 10 were expressedin termsof both species richness cm DBH (diameterat breastheight, 1.3 m) marked and Fisher's Alpha, using various subsets of our withsequentially numbered aluminum tags. Addi- data. We estimated"species richness"-the sumof tional narrowersamples of 2 X 500 m with all thenumber of tree species-as thismeasure is eas- plants2 2.5 cm DBH collectedbut not tagged were ily understoodand widelyreported from other for- made at pre-determinedpoints in the grid:the re- ests. We also estimatedFisher's Alpha, where: sults of these inventorieswill be reportedin a fu- species richness= Alpha * ln(I + stems/Alpha) turepublication. The linearnature of all our plots meansthat they each traversea mixtureof edaphic Fisher'sAlpha values fromsmall tropicalforest formations.Soil conditionswithin the gridranged treesamples providegood estimatesof the overall frompoorly to adequatelydrained clay soils ("Sha- diversityof each forest(Condit et al., 1998). Both paja") to well-drainedwhite sand soils at topo- species richnessand Fisher's Alpha values were graphichigh points ("Varillal"). calculated on a per-areabasis (i.e., foreach one- In the long-termplots, every tagged plant's hectare plot), and on an area-independentbasis heightwas visuallyestimated and its diametermea- (i.e., forthe first500 stems encounteredin each suredat 1.3 m withdiameter tape. Herbariumcol- plot) in orderto removethe complicatingeffect of lections (Vdsquez 14592-15829) were made in varyingstem density. Finally, each diversityindex 1990 and 1991 fromevery individual plant (except was also estimatedseparately for all woodystems, forpalm tree species, whichwere only collected and fortrees alone. once each) using extendablealuminum collecting In orderto comparestem densities of individual poles and, wherenecessary, spiked climbingirons species and families,average per-hectarevalues forclimbing trees. A full set of duplicatesis de- werecalculated. The mostspeciose familiesacross Volume87, Number4 VAsquezMartinez & Phillips 503 2000 High-DiversityPeruvian Forest

Table 1. Allpahuayoforest diversity in 1990, forstems 2 10 cm diameterin 1-ha plots. The values givenfor species richnessand Fisher'sAlpha are forthe minimumand the mostlikely (underlined) estimates, respectively. The diversityvalues are calculatedfor (A) all woodystems (i.e., treesand lianas),and (B) fortrees only. See textfor further details.

Species per Fisher's Fisher's first500 Alpha,first Stems Species Alpha stems 500 stems (A) All woodystems Plot 1 643 281/293 190/208 233/242 170/185 Plot 2 634 306/311 233/242 250/255 199/208 Plots 1 and 2 combined 1277 466/480 264/280 N/A N/A (B) Treesonly Plot 1 616 264/275 175/191 228/237 162/176 Plot 2 608 290/295 218/226 243/248 186/195 Plots 1 and 2 combined 1224 433/444 239/250 N/A N/A

our whole sample were tabulatedon the basis of beit unnamed),but for a feweven thiswas notpos- thetotal species recordedin bothhectare plots. The sible. degree of departurefrom randomness in species' distributionwith respect to soil typewas explored (B) SPECIES RICHNESS AND DIVERSITY using a binomialtest based on the relativefre- quency with which individualswere recordedin Because of the difficultieswith identification clay soil and sandysoil forests. thereis some uncertaintyabout the exact number Standardmeasures of forest structure (density of of species in our samples. We estimateda "most stemsand totalbasal area ofstems - 10 cm DBH) likelyvalue" forspecies richness(S) bymultiplying werecomputed for each hectareplot. Annual mor- the numberof unidentifiedplants per sample that talityand recruitmentrates were estimatedusing could notbe allocatedto morphospecies(U) by the standardprocedures that use logarithmicmodels ratio of species:individualsfor the in that thatassume a constantprobability of mortality and sample thatwere identifiedto species or morphos- recruitmentthrough each inventoryperiod (Phillips pecies (R), and addingthis value to the numberof et al., 1994; Swaine& Lieberman,1987), and com- morphospeciesand species conceptsactually dis- puted separatelyboth for woody stems and for tinguished(D). woodybasal area. Turnoverrates foreach period Thus, (S = (U * R) + D. were representedby the mean of recruitmentand mortalityrates (Phillips, 1996). This estimateprobably errs on the conservative side because the most difficultplants to identify RESULTS AND DISCUSSION tendto be the rarest,and thereforeit is likelythat the ratio of species to individualswill be greater (A) TAXON IDENTIFICATION for non-identifiedplants than it is for identified Of the 1277 plants in the plots in 1991, 100% plants. were identifiedto family,1249 (97.8%) to , Diversityvalues foreach plot and forboth plots and 1168 (91.5%) to species (Appendix 1). We combinedare presentedin Table 1, withresults made 1160 collectionsfrom the two plots, of which given separatelyfor all stemsand fortrees alone. 1053 (90%) were sterile,22 (2%) were in , 33 These revealthat the Allpahuayoplots are some of (3%) werein ,and 52 (5%) werein .The the richest1-ha samples ever reported.The best highproportion of sterilematerial complicated the estimateof Fisher'sAlpha fortrees in our plot 2 identificationprocess. Near-complete identification (226) is the greatestvalue ever recordedfor trees was possible onlybecause of the large numberof 2 10 cm DBH in 1-ha plots,comparing with 221 fertilecollections from Allpahuayo and nearbylo- at Yanamonoin Peru (Conditet al., 1998), 211 at calities available forcomparison at MO. For most Cuyabenoin Ecuador(Condit et al., 1998), and 191 of the unidentifiedspecies, it was possibleto allo- in Allpahuayoplot 1 (thisstudy), all of whichare cate collectionsto morphospecies(i.e., to allocate upper Amazoniansites. In termsof publishedre- collectionsto a morphologicalspecies concept,al- sultsof numbers of species per 500 stems,the All- 504 Annals of the Missouri Botanical Garden

pahuayo plots are surpassed only by Yanamono dominateother Amazonian forests (Gentry, 1988a). (Phillips et al., 1994). There are also some pub- Fabaceae are byfar the most species-rich, and even lished inventoryresults of, variously, lianas, hemi- whentreated individually the three legume subfam- epiphytes,trees, and saplings fromthis part of ilies each have aboutas manyspecies as the other Amazonia(e.g., Gentry,1988a, b, 1992; Valencia top-fivelargest families (, Lauraceae, An- et al., 1994; Clinebellet al., 1995; Duivenvoorden nonaceae,Burseraceae). & Lipps, 1995). These confirmthat the region's un- The plots each traversetwo contrastingsoil floodedforests generally have globallyexceptional types,richer clay soils (the "Shapaja" series) and levels of diversityfor woody plants. poorwhite sand soils (the"Varillal" series), and the Within-community("alpha-") diversityin neo- stronglocal edaphiccontrasts might be expectedto tropicalforests is closely correlatedwith climatic result in some floristicdifferentiation within the conditions,with the richest forests found in thehot plots. Because of the high diversity,few species and aseasonal equatoriallowlands (Clinebell et al., weresampled frequently enough in thetwo plots to 1995). High levels of local diversitymay also be a allow us to testthe extentto whichthey are habitat consequenceof close juxtapositionof differentto- specialistsor generalists.However, among the 56 pographicaland soil formationsin westernAma- species thatwere relativelycommon, with five or zonia (Tuomistoet al., 1995) with "mass-effect" more individualplants in the two hectares,a bi- processescausing species to spill over fromadja- nomialtest reveals that a greaternumber are con- cent edaphicallydefined communities. Each All- finedto one soil or anotherthan would be pahuayo plot traversescontrasting substrates, of expectedunder a null modelof random distribution sandy and clayey soils, which contributesto the (Table 3). This analysisreveals a large numberof high diversityvalues, as some species are appar- species that are apparentspecialists on the poor entlyspecialized on each particularsoil type (cf. whitesand soils (forexample, Macrolobium micro- (c) below). However,the Allpahuayoplots were calyx,Ocotea aciphylla,Tachigali ptycophysca, Ta- placed withoutregard to foresttype, so these di- ble 3A), and a smallergroup mostly confined to the versityvalues are presumablytypical for most 1-ha relativelyrich clay soils (forexample, Leonia gly- patcheswithin the Allpahuayoforest. Data are also cycarpa,Lindackeria paludosa, Senefelderaskutch- available fromone 0.1-ha sample of plants 2 2.5 iana, Table 3B). Note that these resultsare not cm DBH, purposivelylaid out at Allpahuayoby A. proofof specialization,since apparentspecialists Gentryand the firstauthor so as to only sample could arguablybe constrainedto one patchof forest clay soil forest (Clinebell et al., 1995; http.// by chance alone, especiallyif theyhave poor dis- www.mobot. org/MOBOT/research/appliedLresearch/ persal mechanisms.However, a parallel studyat gentry.html).Here 275 species oftrees, lianas, and Allpahuayo(Vormisto et al., 2000) demonstrated hemiepiphyteswere found in 401 individuals,giv- that the local-scale distributionalpattern of soil ing a Fisher'sAlpha value of386, thehighest spe- typesis closelyrelated to plant distributionalpat- cies richnessand Fisher'sAlpha value of all Gen- terns,not onlyfor tree species but forother plant try's227 0.1-ha samples worldwide(Phillips & groupsas well, and these spatial patternsare con- Miller,in prep.). gruentamong different groups. This researchcon- firmsthat the local distributionalpatterns of many

(C) FLORISTIC COMPOSITION plants withinthe Allpahuayoplots are influenced by edaphic conditions.The high diversityof the The Allpahuayoforest is not onlyvery species- Allpahuayoplots is thereforenot onlydue to high rich,but also has a verylow degreeof dominance intra-communitydiversity, but also derivesfrom the by any one species. The ten mostcommon species contrastingedaphic conditionsevident within each togetherrepresent less than20% ofall stems(Table plot. 2A). Most of the familiesthat physically dominate the forest(Table 2B), such as Fabaceae, Myristi- (D) STRUCTURE AND DYNAMICS caceae, Euphorbiaceae,Sapotaceae, and Moraceae, do so by virtueof havingmany species almostall Althoughthe Allpahuayoplots are exceptionally ofwhich are presentat a verylow density. The palm species-rich,they do not appear to be particularly familyis atypicalin thatits statusas the second- remarkablein termsof their physical structure. moststem-dense family is due mostlyto the high This is indicatedby the factthat both theirstem densityof one species (Oenocarpusbataua Mart.). densityand treebasal area appearto be wellwithin The mostspeciose familiesin our entireAllpa- thetypical ranges for most lowland Amazonian for- huayosample (Table 2C) are thesame familiesthat ests (Table 4). However,we could findno published Volume87, Number4 VWsquezMartinez & Phillips 505 2000 High-DiversityPeruvian Forest

Table 2. Dominantwoody plant taxa at Allpahuayo.

A. The 10 species withthe highestdensity of stems2 10 cm diameterper hectare,on averageand by soil type. The twoplots included 1.16 ha of clay soil forest,and 0.84 ha of sandysoil forest. Stemdensity, Stemdensity, clay soil sandysoil Family Genus Stemdensity forest forest Palmae Oenocarpusbataua 39.0 26.3 55.2 subsp. bataua Myristicaceae Iryantheraulei 14.5 8.8 20.8 Myristicaceae Virolapavonis 11.5 11.3 14.6 Euphorbiaceae Hevea guianensis 10.0 1.3 18.8 Fabaceae Macrolobiurninicrocalyx 9.5 0 19.8 Annonaceae Diclinanonatessmannii 7.5 0 15.6 Euphorbiaceae Micrandaelata 7.5 0 15.6 Euphorbiaceae Senefelderaskutchiana 7.5 17.5 1.0 Apocynaceae Aspidospermaexcelsurn 7.0 1.3 12.5 Palmae Astrocaryurnmacrocalyx 7.0 16.3 1.0 Top 10 species 121.0 82.8 175.9 All species 638.5 637 640

B. The 10 familieswith the highest density of stems2 10 cm diameterper hectare. Family Stemdensity Fabaceae: 115.5 Caesalpiniaceae 53.0 Mimnosaceae 23.5 Papilionaceae 36.5 Indet.Fabaceae 2.5 Palmae 61.5 Myristicaceae 59.0 Euphorbiaceae 56.5 Sapotaceae 33.0 Moraceae 30.5 Annonaceae 27.0 Lauraceae 26.5 Burseraceae 21.5 Lecythidaceae 16.0 Top 10 families 420.0 All families 638.5

C. The 10 familieswith the highest number of species 2 10 cm diametersampled throughout our 2-ha sampledarea. The values givenare forthe minimumand mostlikely (underlined) number of species per family. Totalspecies Family inventoried Fabaceae: 82/89 Caesalpiniaceae 27/29 Mimosaceae 25/28 Papilionaceae 28/29 Indet.Fabaceae 2/3 Moraceae 31/31 Lauraceae 29/29 Annonaceae 27/27 Burseraceae 23/24 Euphorbiaceae 23/23 Sapotaceae 22/23 Myristicaceae 21/21 Lecythidaceae 14/14 -Myrtaceae 10/12 Top 10 families 262/273 All families 433/444 506 Annals of the MissouriBotanical Garden

Table 3. Edaphic specialistsin the Allpahuayoplots. For all species withfive or moreindividuals, a binomialtest was conductedto testthe null hypothesisof random distribution with respect to soil type.The hypothesiswas rejected forhalf the 56 species tested(* (P < 0.05); ** (P < 0.01); *** (P < 0.001)). See textfor further details.

A. Specialistson whitesand ("Varillal"soils). Probability Total Shapaja Varillal of random Family Taxon stems stems sterns distribution Annonaceae Diclinanonatessmnannii 15 0 15 Apocynaceae Aspidospermaexcelsum, 14 1 12 ** Bignoniaceae Jacarandainacrocarpa 5 0 5 * Euphorbiaceae Hevea guianensis 20 1 18 Euphorbiaceae Micrandaelata 15 0 15 Fabaceae (Caes.) Macrolobiurnbifolium 8 0 8 ** Fabaceae (Caes.) Macrolobiuminicrocalyx 19 0 19 Fabaceae (Caes.) Sclerolobiumbracteosum 5 0 5 * Fabaceae (Caes.) Tachigaliptychophysca 10 0 10 Fabaceae (Caes.) Tachigalitesmnannii cf. 13 0 13 Fabaceae (Pap.) Swartziaracemosa var. klugii 11 1 10 ** Icacinaceae Emmoturnfloribundumr 5 0 5 * Lauraceae Ocoteaaciphylla 8 0 8 ** Malpighiaceae Byrsonimastipulina 8 0 8 ** Myristicaceae Iryantheraulei 29 7 20 * Nyctaginaceae Neea floribunda 8 1 7 * Palmae Oenocarpus bataua subsp. bataua 78 21 53 Rosaceae Prunusdetrita vel sp. aff. 5 0 5 * Rubiaceae Ferdinandutsachlorantha 5 0 5 * Sapotaceae Chrysophyllurnbombycinurn 9 0 9 ** B. Specialistson clay ("Shapaja" soils). Euphorbiaceae Conceveibarhytidocarpa 7 6 1 * Euphorbiaceae Nealchorneayaputrensis 9 8 1 ** Euphorbiaceae Senefelderaskutchiana 15 14 1 Flacourtiaceae Lindackeriapaludosa 6 5 0 * Moniimiaceae Siparutnadecipiens 5 5 0 * Palmae Astrocaryumrnacrocalyx 14 13 1 Palmae Iriarteadeltoidea 13 12 1 Violaceae Leonia glycycarpavar. glycycarpa 8 6 0 *

Table 4. Stemdensity and basal area of somelowland Amazonian forests, for stems ?10 cm diameterper ha. Data are fromthis study(Allpahuayo, 1990 census) or othersources (Mishana,Tambopata, Yanamono: Gentiy, 1988b; Afiangu,Jatun Sacha, Bel6m,San Carlos de Rio Negro:Phillips et al., 1994; Cusco Amaz6nico:Phillips et al., un- publisheddata). Stemdensity values are per ha. Basal area values are m2per ha.

Liana and Liana and Tree stem strangler Tree basal strangler Site density stemdensity area basal area Bel6m,Brazil 572 27.7 JatunSacha, Ecuador 724 30.5 Afiangu,Ecuador 734 23.1 AllpahuayoPlot 1, Peru 616 27 26.77 0.42 AllpahuayoPlot 2, Peru 608 26 27.33 0.27 Cusco AmazonicoPlot 1, Peru 489 45 25.9 Mishana,Peru 842 16 28.7 TambopataPlot 1, Peru 585 17 26.9 Yanamono,Peru 580 26 32.7 San Car-losde Rio Negro,Venezuela 744 23.0 Volume87, Number4 VAsquezMartinez & Phillips 507 2000 High-DiversityPeruvian Forest

Table 5. Annual rates of naturalstem and basal area mortality,recruitment, and increment,for stems 2 10 cm diameter,between 1990 and 1996. All resultsare based on 0.84 ha (plot 1) and 0.96 ha (plot2).

A. Stemdynamics. Annualturnover Annual Annual (mean of loss and mortality recruitment gain) Plot 1: Tree stems 1.89% 1.57% 1.73% Plot 2: Tree stems 1.84% 2.04% 1.94% Plot 1: Liana and stranglerstems 4.41% 1.89% 3.15% Plot 2: Liana and stranglerstems 4.87% 8.02% 6.44% B. Basal area dynamics.Note that the annual gain in basal area is thesum of new recruitment into the 10 cm diameter size-class,plus the growthof pre-existingstems 2 10 cm diameter,and annual turnoveris the mean of mortality and gain rates. Annual Annual mortality recruitment Annualgain Annualturnover Plot 1: Tree basal area 1.76% 0.48% 2.08% 1.92% Plot 2: Tree basal area 2.21% 0.57% 2.32% 2.26% Plot 1: Liana and stranglerbasal area 5.69% 1.03% 5.72% 5.71% Plot 2: Liana and stranglerbasal area 8.29% 7.24% 8.57% 8.43%

data forliana basal area (lianas are rarelysystem- averaged close to 2% per year in the firsthalf of aticallycensused in ecologicalplots, and liana bas- the 1990s. This rate is not unusual by the standards al area is even morerarely reported), so it is not of western Amazonian forests,but it is higher than possible to compareAllpahuayo with other forests the average turnover rate for other tropical moist in thisrespect. Elsewhere (Phillips et al., 1998) we forests (- 1.5% per year based on studies at 40 reportedthat tree basal area values have been in- differentsites; Phillips, 1996). Based on our small creasingin the majorityof Amazonianplots cen- sample of lianas and stranglers, large climbing sused since the mid-1970s,which we interpreted plants appear to turn over notably fasterthan trees, as being a possible effectof long-termfertilization about 5% a year, which indicates that these organ- by risingatmospheric concentrations of carbondi- isms may have shortercanopy residence times than oxide. In this contextit is interestingto notethat most trees. Whether this pattern is repeated else- by 1996, bothplots at Allpahuayohad experienced where remains to be seen, but if faster turnoverof small net increases in basal area (by 0.1% and lianas and stranglers is a general propertyof Am- 1.9%) over the 1990 values shownin Table 4, in azon forests it would have implications for plot spite of some illegal fellingof palm treeswithin a studies, most of which still ignore climbers. We may fewof the sub-plots.Clearly a longercensus inter- be overlooking a component of the forest that is val will be needed to confirmwhether the small more significantecologically than has been appre- change in foreststructure is part of a long-term ciated. trendat Allpahuayo,or simplypart of a patternof randomfluctuation around a long-termstable state. CONCLUSIONS To estimateannual naturalmortality and growth rateswe excluded the sub-plotswhere palm trees Our floristicand ecological results at Allpahuayo were cut (Table 5). Stem turnoverand basal area well demonstrate the ecological and systematic turnoverfunctions measure slightly different attri- benefits that can result when we concentrate our butesof the standdynamics-stem turnover is con- joint effortson inventories in fixed plots. For the cernedwith population dynamics (i.e., the meanof ecologist the benefitsare clear-without the collab- populationmortality and recruitmentrates), while oration of botanists in the field and the herbarium basal area turnoveris concernedwith basal area it is impossible to characterize patternsof diversity dynamics(i.e., themean of basal area mortalityand and floristic composition in most tropical forests, recruitmentplus growthof existingtrees). Over the let alone explore the factors that determine these. long-terman old-growthforest would be expected As forests become more vulnerable to widespread to have similarvalues of each, and at Allpahuayo environmental stresses such as fragmentationand theturnover rates of tree stems and basal area both climate change (Laurance et al., 1997; Phillips, 508 Annals of the MissouriBotanical Garden

1997), plots are also needed to monitorthese im- sityand floristiccomposition on environmentaland geo- pacts on biodiversity.Systematists at botanicalgar- graphicalgradients. Ann. Missouri Bot. Gard. 75: 1-34. . 1988b. Tree species richnessof upper Amazo- dens thereforeoffer essential expertise for under- nianforests. Proc. Natl. Acad. Sci. U.S.A. 85: 156-159. standingthe biologicaleffects of global change. . 1992. Diversityand floristiccomposition of An- For the systematistor floristicmonographer, es- dean forestsof Peru and adjacent countries:Implica- tablishingpermanent sample plots can help in un- tions for their conservation.Mem. Mus. Hist. Nat. (Lima) 21: 11-19. derstandingthe local flora.The preciserigor of eco- Kalliola, R., K. Ruokolainen,H. Tuomisto,A. Linna & S. logical samplingforces researchers to look equally Maki. 1998. Mapa geoecol6gicode la zona de Iquitosy at everyplant thatmeets pre-determined criteria. variaci6nambiental. Pp. 443-457 in R. Kalliola & S. In contrast,peripatetic botanizing may catalog the Flores Paitdn(editors), Geoecologfa y DesarrolloAma- weedy,common, obvious, and accessible species, z6nico.Turku, Finland. Kallunki,J. A. 1994. Revisionof Raputia Aubl. (Cuspa- whilemissing rarer or largerplants, especially can- riinae,Rutaceae). Brittonia,46: 279-295. opy trees,lianas, and epiphytes.Rigorous plot in- Laurance,W. F., S. F. Laurance,L. V. Ferreira,J. M. Ran- ventoriescan thereforereduce the spatial, taxonom- kin de Merona,C. Gascon & T E. Lovejoy.1997. Bio- ic, life-form,and even seasonal biases prevalentin mass collapse in Amazonianforest fragments. Science 278: 1117-1118. herbaria(e.g., Nelson et al., 1990), especiallywhen Lowry,P. P., II, G. E. Schatz & P. B. Phillipson.1997. integratedinto a largerintensive collecting effort. The classificationof naturaland anthropogenicvegeta- Thus, a comparisonof taxa in Vdsquez (1997) with tionin Madagascar.Pp. 93-123 in S. M. Goodman& those listed in Brako and Zarucchi (1993) shows B. D. Patterson(editors), Natural Change and Human that the intensivecollecting effort in permanent Impact in Madagascar.Smithsonian Institution Press, Washington,D.C. plotsand surroundingforest at severalIquitos sites Marengo,J. 1998. Climatologfade la zona de Iquitos, has yieldednearly 250 taxa new to Peru. At All- Peru. Pp. 35-57 in R. Kalliola & S. FloresPaitin (ed- pahuayo,new tree taxa have been recognizedin itors),Geoecologfa y DesarrolloAmaz6nico. Turku, Fin- Annonaceae (2 species; Chatrou,1998) and in land. McNeely,J. A, K. R. Miller,W. V. Reid, R. A. Mittermeier Lauraceae (3 species includinga new genus; van & T. B. Werner.1990. Conservingthe World'sBiodi- der Werff,1993, 1997), whilethe repetitionof plot versity.IUCN, Gland, Switzerland,and Island Press, visitshas also allowedus to collectfertile material Covelo,California. confirmingnew herb and shrubtaxa (e.g.,Kallunki, National Research Council. 1992. ConservingBiodiver- 1994; Vdsquez, 1997). sity: A Research Agenda for DevelopmentAgencies. NationalAcademy of Sciences Press,Washington, D.C. NationalScience Board. 1990. Loss of BiologicalDiver- LiteratureCited sity:A Global CrisisRequiring International Solutions. Brako,L. & J. L. Zarucchi.1993. Catalogueof the Flow- NationalScience Board,Washington, D.C. eringPlants and Gymnospermsof Peru. Monogr.Syst. Nelson,B. W., C. A. C. Ferreira,M. F. da Silva & M. L. Bot. MissouriBot. Gard. 45. Kawasaki.1990. Endemismcentres, refugia and botan- Chatrou,L. W. 1998. Changinggenera: Systematic studies ical collectingdensity in BrazilianAmazonia. Nature in Neotropicaland WestAfrican Annonaceae. Herbar- 345: 714-716. ium Division,Utrecht University. Phillips,0. L 1996. Long-termenvironmental change in Clinebell,R. C., II, 0. L. Phillips,A. H. Gentry,N. Stark tropical forests:Increasing tree turnover.Environm. & H. Zuuring.1995. Predictionof neotropicaltree and Conservation23: 235-248. liana species richnessfrom soil and climaticdata. Bio- 1997. The changingecology of tropicalforests. diversityand Conservation4: 56-90. Biodiversityand Conservation6: 291-311. Condit,R., R. B. Foster,S. P. Hubbell, R. Sukumar,E. & A. H. Gentry.1994. Increasing turnover G. Leigh,N. Manokaran,S. Loo de Lao, J.V. LaFrankie throughtime in tropicalforests. Science 263: 954-958. & P. S. Ashton.1998. Assessing forestdiversity on & J. S. Miller.In prep. Global Patternsof Forest smallplots: Calibration using species-individual curves Diversity:The datasetof AlwynGentry. Monogr. Syst. from50-ha plots. Pp. 247-267 in F. Dallmeier& J. Bot. MissouriBot. Gard. Comiskey(editors), Long-Term Monitoring of Biological & P. H. Raven. 1996. A strategyfor sampling Diversityin TropicalForest Areas. UNESCO, Paris. neotropicalforests. Pp. 141-165 in A. C. Gibson(edi- Dallmeier,F. & J. Comiskey(editors). 1998a. ForestBio- tor),Neotropical Biodiversity and Conservation.Occas. diversityResearch, Monitoring, and Modeling:Concep- Publ. MildredE. MathiasBotanical Garden 1. Mildred tual Backgroundand Old WorldCase Studies. MAB E. MathiasBotanical Garden, Los Angeles. Series Vol. 20. UNESCO, Paris. , P. Hall, A. H. Gentry,S. A. Sawyer& R. VAs- & (Editors).1998b. ForestBiodiversity in quez. 1994. Dynamicsand species richnessof tropical North,Central and SouthAmerica, and the Caribbean: rain forests.Proc. Natl. Acad. Sci. U.S.A. 91: 2805- Research and Monitoring.MAB Series Vol. 21. 2809. UNESCO, Paris. ,Y. Malhi,N. Higuchi,W. F. Laurance,P. Nufiez Duivenvoorden,J. F. & H. Lipps. 1995. A land-ecological V., R. Vdsquez M., S. G. Laurance,L. V. Ferriera,M. studyof soils, vegetation,and plant diversityin Col- Stern,S. Brown& J. Grace. 1998. Changesin the car- ombianAmazonia. Tropenbos Series 12: 1-438. bon balance of tropicalforest: Evidence from long-term Gentry,A. H. 1988a. Changesin plantcommunity diver- plots.Science 282: 439-442. Volume 87, Number 4 VWsquez Martfnez & Phillips 509 2000 High-Diversity Peruvian Forest

Rakotomalaza,P. J. & N. Messmer.1999. Structureand treealpha-diversity in AmazonianEcuador. Biodiversity floristiccomposition of the vegetationin the R6serve and Conservation3: 21-28. NaturelleIntegrale d'Andohahela, Madagascar. Fieldi- VAsquezM., R. 1997. Fl6rulade las ReservasBiol6gicas ana: Zoology,n.s., 94: 51-96. de Iquitos. Monogr.Syst. Bot. MissouriBot. Gard.63. Rudas L., A. 1996. EstudioFloristico y de la Vegetaci6ndel Vormisto,J., 0. L. Phillips,K. Ruokolainen,H. Tuomisto ParqueNacional Natural Amacayacu Amazonas-Colombia. & R. VAsquez M. 2000. A comparisonof small-scale M.Sc. Thesis,Universidad Nacional de Colombia,Bogota. distributionpatterns of fourplant groupsin an Ama- Swaine, M. D & D. Lieberman.1987. Note on the cal- zonianrainforest. Ecography 23: 349-359. culationof mortalityrates. J. TropicalEcol. 3: ii-iii. Werff,H., van der. 1993. A revisionof the genus Pleu- Tuomisto,H., K. Ruokolainen,R. Kalliola, A. Linna, W. rothyrium(Lauraceae). Ann. MissouriBot. Gard. 80: Danjoy & Z. Rodriguez.1995. DissectingAmazonian 39-1 18. biodiversity.Science 269: 63-66. . 1997. Sextonia,a new genus of Lauraceae from Valencia, R., H. Balslev & G. Paz y Mifio.1994. High SouthAmerica. Novon 7: 436-439. Appendix1. Woodyplant taxa 210 cm DBH recordedfrom two 1-ha plotsat Allpahuayo.All taxa are treesunless indicatedas *** (liana). When a voucheris fertile,we indicateits statusby thefollowing code: bd (),fl (),ft (fruits). See textfor further discussion.

No. plants in Vasquez collections Family Taxonand authority 2-ha (onlythe firstsix are listedper taxon) Anacardiaceae Tapirirapeckoltiana Engl. 2 14,608 14,614 bd Tapiriraretusa Ducke 1 15,185 Tapirirasp. 1 15,794 Annonaceae Anaxagoreabrevipes Benth. 1 14,815 fl Anaxagoreaphaeocarpa Mart. 2 14,655 bd 14,757 Annonaduckei Diels 1 15,614 Annonamontana Macfad. 2 14,788 15,332 Annonapoepipigii (Mart.) Maas & Westra 1 14,695 Bocageopsismattogrossensis (R. E. Fr.) R. E. Fr. 1 15,324 Diclinanonatessmannii Diels 15 14,819 14,840 14,899 14,907 14,919 14,928 Duguetiacaulifiora R. E. Fr. 1 15,035 ft Duguetiasp. 1 15,770 Fusaea longifolia(Aubl.) Saf. 2 14,764 bd 15,788 fl Fusaea sp. 1 15,730 o Guatteriaelata R. E. Fr. 1 15,566 | I Guatteriamegalophylla Diels 1 15,039 Guatteriaolivacea R. E. Fr. 2 14,773 14,797 ft Guatteriarugosa R. E. Fr. 1 14,943 Guatteriaschomburgkiana Mart. 2 14,770 14,812 fl Guatteriasessilis Fries 2 14,844 15,204 0 Oxandraxylopioides Diels 2 15,489 fl 15,802 a Rolliniacuspidata Mart. 1 14,877 Rolliniaperuviana Diels 4 14,776 fl 14,869 15,203 fl 15,671 Rolliniapittieri Saff. 1 14,643 fl Unonopsisfloribunda Diels 2 14,793 15,042 Unonopsisspectabilis Diels 1 15,569 Xylopiabarbata Mart. 2 15,259 15,268 Xylopiabenthamii R. E. Fr. 1 15,622 ft Xylopiaexcellens R. E. Fr. 1 14,897 Xylopianitida Dunal 1 15,536 bd Xylopiasericea A. St.-Hil. 1 15,799 ft Appendix1. Continued.

No. CD plants in Vasquez collections Family Taxonand authority 2-ha (onlythe firstsix are listedper taxon) Apocynaceae Ambelaniaoccidentalis Zarucchi 2 15,107 ft Aspidospermadesmanthum Benth. ex Mull.Arg. 2 15,251 15,425 Aspidospermaexcelsum Benth. 14 14,823 14,825 14,831 14,839 14,852 Aspidospermaspruceanum Benth. ex Mull.Arg. 1 14,818 Aspidospermaschultesii Woodson 1 15,341 Couma macrocarpaBarb. Rodr. 1 15,017 Himatanthussuccuba (Spruce ex Mull. Arg.) Woodson 1 15,676 Malouettiakillipii Woodson 2 14,601 14,622 Malouettiatamaquirina var. tamaquirina (Aubl.) A. DC.*** 1 14,882 Odontadeniapuncticulosa (Richard) Pulle*** 2 15,407 15,504 Parahancorniaperuviana Monachin 3 14,985 15,478 15,762 Aquifoliaceae flexandarensis vel sp. aff.Loes. 4 14,941 15,256 fl 15,427 15,435 < Araliaceae Dendropanaxumbellatus (Ruiz & Pav.) Deene. 2 14,908 15,440 Scheffleramorototoni (Aubl.) Maguire, Steyerm.& Frodin 1 14,858 15,009 Bignoniaceae Jacarandamacrocarpa Bureau & K. Schum. 5 14,862 14,878 14,998 15,019 15,434 Jacarandaobtusifolia subsp. obtusifolia Humb.& Bonpl. 1 15,015 (1)CD Memoracladotricha Sandwith 1 15,485 Bombacaceae Eriothecamacrophylla subsp. sclerophylla (Ducke) Robyns 2 15,514 Matisia bracteolosaDucke 2 15,620 15,662 Matisia hirtaCuatrec. 1 15,060 Matisia malacocalyx(A. Robyns& S. Nilsson)W. S. Alverson 2 15,199 15,499 Matisia soegengiiCuatrec. 1 15,733 Pachira insignis(Sw.) Sw. ex Savigny 5 14,881 15,113 15,182 15,598 15,702 Phragmothecamammosa subsp. amazonica W. S. Alverson 1 15,513 Appendix1. Continued.

No. plants in Vasquez collections Family Taxonand authority 2-ha (onlythe firstsix are listedper taxon) Boraginaceae Cordiamexiana I. M. Johnst. 1 15,135 Cordianodosa Lam. 2 14,624 15,660 Burseraceae Crepidospermumgoudotianum (Tul.) Triana& Planch. 2 15,607 15,653 Crepidospermumprancei Daly 1 15,771 Dacryodesperuviana (Loes.) J. F. Macbr. 1 15,766 Protiumaracouchini (Aubl.) Marchand 1 15,500 Protiumcrassipetalum Cuatrec. 1 15,822 Protiumdivaricatum Engl. 1 15,785 Protiumferrugineum(Engl.) Engl. 1 15,045 Protiumfimbriatum Swart 3 15,176 15,172 15,684 Protiumgallosum Daly 1 14,744 Protiumgrandifolium Engl. 6 15,179 15,308 15,492 15,574 15,599 15,778 Protiumhebetatum Daly 5 14,634 14,895 fl 14,898 15,563 15,567 Protiummacrophylla (HBK) Engl. 1 15,642 Protiumnitidifolium Daly 2 14,787 15,760 Protiumnodulosum Swart 1 15,650 Protiumopacum Swart subsp. opacum 2 14,599 14,612 15,521 15,797 Protiumopacum vel sp. aff.Swart 2 15,521 15,797 Protiumsubserratum (Engl.) A. DC. & C. DC. 1 14,975 Protiumtenuifolium vel sp. aff.(Engl.) Engl. 1 15,720 Protiumtrifoliatum Engl. 1 14,724 0 Protiumsp. aff.robustum 1 15,764 Protiumsp. 4 15,074 15,094 15,125 15,171 Tetragastrispanamensis (Engl.) Kuntze 1 15,136 Trattinickiaperuviana Diels 2 14,775 15,442 Capparidaceae Capparisschunkei J. F. Macbr. 1 15,806 fl Caryocaraceae Anthodiscusklugii Standl. ex Prance 1 15,525 Anthodiscuspilosus Ducke 1 15,395 Caryocarglabrum (Aubl.) Pers. 1 15,801 Caryocarharlingii Prance & Encarn. 2 14,802 15,201 Celastraceae Maytenusamazonica Mart. 1 14,843 Appendix1. Continued.

No. CD plants o in Vasquez collections Family Taxonand authority 2-ha (onlythe first six are listedper taxon) z Chrysobalanaceae Couepiadolichopoda Prance 1 15,284 Couepia guianensisAubl. subsp. guianensis 1 14,659 Couepiaracemosa Benth. ex Hook. f. 1 14,783 Licania arachnoideaFanshawe 1 14,777 Licania blackiiPrance 1 15,080 Licania canescensBenoist 1 15,079 Licania heteromorphavar. heteromorpha Benth. 2 14,979 14,984 Parinariklugii Prance 1 15,206 Parinariparilis J. F. Macbr. 1 15,606 Caraipa punctulataDucke 2 14,731 14,808 fl Chrysochlamysweberbaueri Engl. 1 15,488 ft Clusia laxiflora(Poepp.) D. Dietr. 1 15,625 I< Moronobeacoccinea Aubl. 1 15,198 Symphoniaglobulifera L. f. 1 15,800 Tovomitaspeciosa Ducke 3 14,732 15,148 ft 15,640 2N Tovomitastigmatosa Planch. & Triana 4 14,656 15,659 15,693 15,765 Combretaceae Buchenaviaamazonica Alwan & Stace 2 15,243 15,739 Convolvulaceae Dicranostylesscandens Benth.*** 1 15,093 Dichapetalaceae Tapuraamazonica Poepp. 1 14,673 Tapuracoriacea J. F. Macbr. 1 15,084 Dilleniaceae Doliocarpusamazonicus Sleumer subsp. | amazonicus*** 2 15,063 15,103 Doliocarpusnovogranatensis Kubitzki*** 1 14,763 Pinzonacoriacea Mart. & Zuccagni*** 1 14,680 Ebenaceae Diospyrossubrotata Hiern 1 15,561 fl Lissocarpastenocarpa Steyerm. 2 15,447 15,470 01 Appendix1. Continued.

No. plants in Vdsquezcollections Family Taxonand authority 2-ha ~~~~~~~~~~~~~~(onlythefirst six are listed per taxon) Elaeocarpaceae Sloanea durissimaSpruce ex Benth. 3 14,609 15,078 15,152 SloaneafloribundaSpruce ex Benth. 3 14,920 15,083 15,306 Sloanea guianensis(Aubl.) Benth. 2 14,888 15,631 Sloanea latifolia(Rich.) K. Schum. 1 15,753 Sloanea laxifloraSpruce ex Benth. 1 14,894 fi Sloanea robustaUittien 5 14,632 14,721 15,143 15,147 15,748 Sloanea sinemariensisAubl. 1 15,307 Sloanea tuerckheimiiDonn. Sm. 1 15,803 Sloanea sp. 1 15,304 Euphorbiaceae genus 1 14,880 Alchorneaschomburgkii Klotzsch 1 15,593 Alchorneatriplinervia (Spreng.) Muell. Arg. 3 14,729 14,771 ft 15,011 Aichorneopsisfioribunda(Benth.) Muell. Arg. 1 15,302 Conceveibamartiana Baill. 1 15,741 Conceveibarhytidocarpa Muell. Arg. 7 14,611 14,630 14,738 15,560 15,600 15,617 Conceveibasp. nov. 3 15,633 fi 15,690 15,757 Dodecostigmaamazonicum Ducke 1 14,700 Glycidendronamazonicum Ducke 1 15,048 Hevea guianensisAubl. 20 14,704 14,817 14,821 14,822 14,824 14,850 -o~ Heveapauci/1ora (Spruce ex Benth.)Muell. Arg. 6 14,667 ft 14,887 ft 15,004 15,298 15,323 15,327 Hyeronimaalchorneoides Allemao 3 14,613 bd 14,6i16bd 14,681 Hyeronimaoblonga Cuatrec. 7 14,642 14,683 14,684 14,734 14,747 15,196 fi Mabea maynensisMuell. Arg. 3 15,071 15,480 15,334 Micrandaelata Muell Arg. 15 15,250 ft 15,271 15,273 15,333 15,340 15,342 Micrandaspruceana (Baill.) R. E. Schult. 11 15,005 15,026 15,047 15,065 15,275 bd 15,265 fi Nealchorneayapurensis Huber 9 14,794 15,575 15,592 15,618 15,673 ft 15,678 Pera benensisRushy 1 15,274 Pera bicolor(Klotzsch) Muell. Arg. 1 15,424 Pera leandriye1 sp. aff.Baill. 1 15,647 Plukenetiapolyadenia Muell. Arg.** 1 15,651 Richeriagrandis Vahl 3 14,678 14,913 15,501 Senefelderaskutchiana Croizat 15 14,596 14,646 14,657 14,660 ft 14,662 14,663 Appendix1. Continued. 2 No. CD plants 0o in Vasquez collections Family Taxonand authority 2-ha (onlythe firstsix are listedper taxon) Fabaceae genus 1 15,825 2 genus 1 15,681 genus 1 15,515 genus 1 15,538 genus 1 15,624 Fabaceae: Bauhinia guianensisAubl. subsp. Caesalpinaceae guianensis*** 4 14,636 14,710 15,594 14,728 Bauhinia guianensissubsp. kunthiana(Vogel)*** 3 14,785 14,918 14,933 15,002 Cynometrabauhiniifolia Benth. 4 15,551 15,573 ft 15,700 ft 15,745 Cynometraspruceana var. spruceana (Hayne) J. F. Macbr. 5 14,708 15,123 15,156 15,635 15,682 Dialium guianense(Aubl.) Sandwith 4 14,730 15,103Aft 15,486 15,818 Dimorphandramacrostachya Benth. 1 14,845 Hymenaeaoblongifolia Huber 3 14,718 15,162 15,602 I< Macrolobiumacaciifolium (Benth.) Benth. 1 14,931 Macrolobiumangustifolium (Benth.) R. S. Cowan 2 14,689 14,837 2N Macrolobiumarenarium Ducke 1 15,197 Macrolobiumbifolium (Aubl.) Pers. 8 14,860 14,963 14,974 15,252 15,352 15,388 Macrolobiumlimbatum Spruce ex Benth.var. - > limbatum 3 15,608 15,680 15,704 15,299 Macroloboiumlimbatum var. propinquum R. S. Cowan 1 15,299 Macrolobiummicrocalyx Ducke 19 15,244 ft 15,254 15,345 15,350 15,353 15,357 Macrolobiumsp. 1 15,301 Macrolobiumsp. 1 15,339 Sclerolobiumbracteosum Harms 5 15,305 15,310 15,363 15,391 15,461 Sclerolobiummicropetalum Ducke 1 15,773 Sclerolobiumvasquezii Pipoly 1 15,634 Senna sp. 1 15,564 Tachigalicavipes (Spruce ex Benth.)J. F. Macbr. 1 15,338 fl Appendix1. Continued.

No. plants in Vasquez collections Family Taxonand authority 2-ha (onlythe firstsix are listedper taxon) Fabaceae: Tachigalichrysophylla (Poepp.) Zarucchi& Caesalpinaceae Herend. 1 14,799 Tachigalipaniculata Aubl. 1 15,717 Tachigalipoeppigiana Tul. 1 14,804 Tachigaliptychophysca Spruce ex Benth. 10 15,246 15,354 15,356 15,362 15,373 15,377 Tachigalirusbyi Harms 1 15,611 Tachigalitessmannii cf. Harms 13 14,917 14,921 14,942 14,944 15,346 15,349 Tachigalisp. 1 14,996 Tachigalisp. 1 15,030 Tachigalisp. 1 14,953 Tachigalisp. 1 14,957 Tachigalisp. 1 14,965 Tachigalisp. 1 14,961 Tachigalisp. 1 14,966 Tachigalisp. 1 14,856 Tachigalisp. 1 14,859 Tachigalisp. 1 14,871 a c(/) Tachigalisp. 1 14,872 Fabaceae: Albiziasp. 1 15,664 Mimosaceae Enterolobiumbarnebeyanum Mesquita & M. F. Silva 1 15,562 Inga acrocephalaSteud. 1 15,558 O Inga bourgonii(Aubl.) DC. 1 15,619 a Inga capitataDesv. 1 15,109 Inga coruscansHumb. & Bonpl. ex Willd. 1 14,604 Inga densiJforaBenth. 1 14,772 Inga laurina (Sw.) Willd. 2 15,311 15,312 Inga multijugaBenth. 1 15,719 Inga oerstedianaBenth. 1 15,559 Inga peziziferaBenth. 2 14,810 15,493 Appendix1. Continued. 0O

No. plants in Vasquez collections Family Taxonand authority 2-ha (onlythe firstsix are listedper taxon) Z Fabaceae: Inga poeppigianaBenth. 1 15,811 Mimosaceae Inga semialata(Vell.) Mart. 1 15,537 Inga spectabilisvar. schimpfii (Harms) Little 1 15,545 Inga velutinaWilld. 1 15,698 Inga sp. 1 15,755 Inga sp. 1 15,523 Inga sp. 1 notcollected Inga sp. 1 14,598 Marmaroxylonbasijugum (Ducke) L. Rico 4 15,024 15,776 15,792 15,804 Parkia igneiJforaDucke 3 14,991 15,124 15,138 Parkia multijugaBenth. 2 14,970 15,604 Parkia nitidaMiq. 5 14,605 14,610 14,832 15,247 15,476 Parkia panurensisBenth. ex H. C. Hopkins 1 voucherlost Parkia velutinaBenoist 2 15,260 15,335 Pithecellobiumsp. 1 14,595 < Pithecellobiumsp. 1 14,650 Pithecellobiumsp. 1 14,651 Pithecellobiumsp. 1 14,692 | N Pithecellobiumsp. 1 15,131 Pithecellobiumsp. 1 15,502 Pithecellobiumsp. 1 14,652 Stryphnodendronpolystachyum(Miq.) Kleinhoonte 2 14,972 15,568 Zygia latifolia(L.) Fawcett 1 14,626 | Fabaceae: Andiramacrothyrsa Ducke 2 15,548 15,627 Papilionaceae BatesiaftoribundaSpruce ex Benth. 2 15,044 15,705 cI Dalbergiamonetaria var. monetaria L. f.*** 5 14,676A 14,756A 15,401 15,402 15,813 Dalbergiariedelii (Benth.) Sandwith*** 1 15,761 Dalbergiasp.*** 1 15,064 Diplotropispurpurea var. leptophylla (Rich.) Amshoff 3 14,654 15,399 15,404 Dipteryxmicrantha Harms 2 14,762 15,166 Hymenolobiumnitidum Benth. 3 14,967 14,989 15,322 Lonchocarpusspiciflorus C. Mart.ex Benth. 1 15,375 0 Machaeriumsp. cf. multifoliatum*** 1 15,498 Machaeriumcuspidatum Kuhlm. & Hoehne*** 1 15,657 Appendix1. Continued.

No. plants in Vasquez collections Family Taxonand authority 2-ha (onlythe firstsix are listedper taxon) Fabaceae: Machaeriumftoribundumvar.parviflorum Benth.*** 1 14,627 Papilionaceae Machaeriumkegelli Meisn.*** 1 15,810 Machaeriummultifoliatum Ducke*** 3 14,600 15,294 14,988 Machaeriumsp.*** 1 15,073 Machaeriumsp.*** 1 15,025 Machaeriumsp.*** 1 15,095 Ormosiaarborea (Vell.) Harms 1 14,937 Pterocarpusamazonum (C. Mart.ex Benth.)Amshoff 1 14,929 Pterocarpusrohrii M. Vahl. 1 15,013 Swartziabenthamiana Miq. var.benthamiana 8 14,768 15,032 15,261 fl 15,292 15,628 bd 15,812 fl Swartziabrachyrachis var. peruviana R. S. Cowan 1 15,722 Swartziacardiosperma Spruce ex Benth. 4 14,784 15,151 15,257 fl 15,347 Swartzialaevicarpa Amshoff 1 15,116 Swartziapolyphylla DC. 2 15,281 15,713 Swartziaracemosa var. klugii R. S. Cowan 11 14,765 bd 14,834 14,841 14,993 14,994 15,258 fl . Swartziatessmannii Harms 4 14,670 14,707 14,774 15,285 CD Swartziasp. 1 15,061 Swartziasp. 1 14,875 Taralea oppositifoliaAubl. 3 14,874 14,940 14,977 o Vantaneaparviflora Lam. 3 15,772 15,789 15,824 )s Flacourtiaceae Casearia acuminataDC. 1 15,177 Casearia combaymensisTul. 1 14,751 Caseariafasciculata(Ruiz & Pav.) Sleumer 4 14,769 15,540 ft 15,552 15,686 n Caseariajavitensis Kunth 1 14,699 ft CD Casearia sylvestrisSw. var.sylvestris 1 15,508 ft Casearia ulmifoliaM. Vahl ex Vent. 1 15,645 Lindackeriapaludosa (Benth.)Gilg. 6 14,691 14,693 ft 15,104 15,694 15,715 ft 15,738 Mayna grandifolia(H. Karst.)Warb. 1 15,056 Tetracylaciummacrophyllum Poepp. 1 15,509 Tetrastylidiumperuvianum Sleumer 7 14,798 14,827 15,150 ft 15,528 15,595 bd 15,605 Hippocrateaceae Salacia sp.*** 1 15,629 Tonteleaovalifolia subsp. richardii(Miers) A. C. Sm.*** 1 14,676 Appendix1. Continued.

No. CD plants in Vasquez collections Family Taxonand authority 2-ha (onlythe first six are listedper taxon) Z

Humiriaceae Sacoglottisceratocarpa Ducke 1 14,615 CD Sacoglottissp. 1 15,511 - Icacinaceae Dendrobangiaboliviana Rusby 1 15,145 Discophoraguianensis Miers 1 15,422 EmmotumfloribundumHoward 5 14,950 14,951 15,336 15,348 15,437 Lacistemataceae Lacistemaaggregatum (Bergius) Rusby 4 14,594 ft 14,597 15,531 15,736 ft Lauraceae genus 1 15,547 genus 1 15,663 genus 1 15,674 genus 1 15,643 Anaueriabrasiliensis Kosterm. 2 14,742 14,786 Anibaperutilis Hemsl. 4 14,959 15,253 15,413 15,454 Aniba taubertianaMez 1 15,139 ft I< Endlicheriacitriodora van der Werff 2 14,922 14,949 Licaria brasiliensis(Nees) Kosterm. 1 15,496 Licaria canella (Meisn.)Kosterm. 2 14,838 15,768 Licaria sp. cf. dolichanthaKurz 1 15,280 Licaria sp. cf. rodriguesiiKurz 1 15,570 Mezilaurusopaca Kubitzki& van der Werff 2 15,746 15,759 ft Nectandraacuminata (Nees) J. F. Macbr. 1 15,621 Nectandraglobosa (Aubl.) Mez 2 15,099 15,313 . o Nectandrapearcei Mez 1 15,731 Ocoteaaciphylla (Nees) Mez 8 15,245 15,290 15,416 15,417 15,432 15,436 = Ocoteaargyrophylla Ducke 4 14,932 14,925 14,936 14,939 0? Ocoteabofo Kunth 4 15,059 15,117 15,729 15,815 Ocoteabracteosa (Meisn.) Mez 1 15,455 Ocotealeucoxylon (Sw.) Laness. 1 15,418 Ocotealongifolia Kunth 1 15,632 ft Ocoteamyriantha (Meisn.) Mez 2 14,833 15,325 Ocoteapuberula (Rich.) Nees 2 15,370 15,361 Ocoteawhitei Woodson 1 15,140 0 i Ocoteasp. 1 15,821 01 Appendix1. Continued.

No. plants in Vasquez collections Family Taxonand authority 2-ha (onlythe first six are listedper taxon) Lauraceae Ocoteasp. 1 15,751 Persea sp. cf. boliviensis 1 15,467 Pleurothyriumvasquezii van der Werff 1 15,649 Lecythidaceae Carinianasp. cf. multifloraDucke 1 14,753 Eschweilerachartaceifolia S. A. Mori 3 14,750 14,906 15,112 Eschweileracoriacea (DC.) S. A. Mon' 7 14,641 15,057 15,062 15,517 15,679 15,703 Eschweileragigantea (Knuth) J. F. Macbr. 1 15,721 Eschweileramicrantha (Berg) Miers 1 15,724 Eschweilerarufifolia S. A. Mori 9 14,761 14,690 15,089 15,088 15,110 15,128 Eschweileratessmannii Knuth 2 15,102 15,134 Eschweilerasp. 1 14,638 Eschweilerasp. 1 14,661 Gustaviahexapetala (Aubl.) Sm. 1 15,160 Gustavialongifolia Poepp. ex Berg 2 14,625 14,628 Lecythispisonis aff. Cambess. 1 15,097 Lecythissp. cf chartaceaBerg 1 15,066 Ruptiliocarponsp. 3 14,820 15,543 15,740 Linaceae Hebepetalumhumiriifolium (Planch.) Benth. 1 15,018 15,126 Raucheriapunctata (Ducke) Ducke 1 15,791 ft Loganiaceae Strychnosramentifera Ducke*** 1 15,054 Strychnossandwithiana Krukoff & Barneby*** 1 15,775 G Malpighiaceae Byrsonimacowanii W. R. Anderson 1 15,714 | Brysonimapoeppigiana A. Juss. 2 14,809 15,001 Byrsonimastipulina A. Juss. 8 14,902 14,927 15,031 15,049 15,431 fl 15,448 bd Dicellajulianii (J. F. Macbr.)W. R. Anderson*** 1 15,549 Tetrapterysmucronata Cav.*** 1 15,669 Marcgraviaceae Noranteaguianensis subsp. japurensis Aubl.*** 1 14,892 fl Melastomataceae Miconiatetragona Cogn. 1 14,969 Appendix1. Continued.

No. plants in Vasquez collections Family Taxonand authority 2-ha (onlythe firstsix are listedper taxon) Meliaceae proceraDC. 6 14,629 14,647 14,653 14,679 14,863 14,891 Guareasilvatica DC. 1 15,790 Guareasp. 1 14,759 Guareasp. 1 14,645 Trichiliasp. cf. micranthaBenth. 3 14,780 14,813 14,893 Menispermaceae Abutagrandifolia (Mart.) Sandwith 3 15,518 15,623 15,654 Abutasp.*** 1 15,264 Abutasp. cf. imene*** 1 15,058 Curareatecunarum Barneby & Krukoff*** 1 15,101 Telitoxicumkrukovii Moldenke*** 1 15,277 Telitoxicumminutifolium (Diels) Moldenke*** 1 15,466 Telitoxicumperuvianum Moldenke*** 1 15,282 Monimiaceae Siparunacristata (Peopp. & Endl.) A. DC. 1 15,527 Siparunacuspidata (Tul.) A. DC. 1 15,781 I< Siparunadecipiens (Tul.) A. DC. 5 14,674 bd 14,766 bd 15,505 15,565 15,597 ft Siparunaguianensis Aubl. 3 14,709 14,723 14,782 Moraceae genuscf. 1 15,203A Brosimumacutifolium subsp. obovatum (Ducke) C. C. Berg 1 15,289 Brosimumguianense (Aubl.) Huber 3 15,487 15,616 15,709 bd Borsimumlactescens (S. Moore)C. C. Berg 1 15,630 Brosimumpotabile Ducke 1 14,741 Brosimumrubescens Taub. 3 14,854 14,964 15,127 Brosimumutile subsp. ovatifolium(Kunth) Pittier 5 14,836 15,165 ft 15,184 15,554 15,571 o Castilla ulei Warb. 1 15,652 co Cecropiadistachya Huber 3 14,725 fl 15,021 15,288 Cecropiaengleriana Snethl. 2 15,707 fl 15,708 Cecropiasciadophylla Mart. 1 15,615 fl amazonica(Miq.) Miq. 1 15,163 Ficus subapiculata(Miq.) Miq. 1 15,626 Helicostylistomentosa (Poepp. & Endl.) Rusby 4 14,637 14,778 15,130 15,191 calophylla(Poepp. & Endl.) C. C. Berg 3 14,705 15,609 15,695 Appendix1. Continued.

No. plants in Vdsquez collections Family Taxonand authority 2-ha (onlythe first six are listedper taxon) Moraceae Naucleopsisimitans (Ducke) C. C. Berg 1 15,798 Naucleopsismello-barretoi (Standl.) C. C. Berg 1 15,668 NaucleopsisternstroemiiJfora (Mildbr.) C. C. Berg 1 15,376 Naucleopsisulei (Warb.)Ducke 1 15,716 Perebeaglabrifolia C. C. Berg 2 14,791 15,170 Perebeaguianensis Aubl. subsp.guianensis 4 14,733 14,735 14,740 15,735 Pouroumabicolor Mart. subsp. bicolor 1 15,189 Pouroumabicolor subsp. scobina (Benoist) C. C. Berg 1 15,068 15,189 PouroumaferrugineaStandl. 1 15,683 Pouroumaguianensis Aubl. subsp. guianensis 2 14,805 15,726 Pouroumaherrerensis C. C. Berg 1 15,085 Pouroumaminor Benoist 1 15,532 Pouroumasp. 1 14,631 Pseudolmedialaevis (Ruiz & Pav.) J. F. Macbr. 3 14,602 14,685 15,732 Pseudolmedialaevigata Trecul 6 14,842 14,903 bd 14,916 14,946 15,309 15,481 Trymatococcusamazonicus Poepp. & Endl. 2 14,698 ft 15,699 Myristicaceae Campsoneuracapitellata (A. DC.) Warb. 1 15,809 0 Iryantheracrassifolia A. C. Sm. 4 15,070 15,082 15,193 15,808 a Iryantheraelliptica Ducke 1 15,146 Iryantherajuruensis Warb. 13 14,618 14,640 14,712 14,748 14,800 15,503 Iryantheralaevis Markgr. 3 14,745 14,978 15,546 Iryantheralancifolia Ducke 4 14,639 14,803 15,040 15,780 Iryantheramacrophylla (Benth.) Warb. 7 14,706 14,760 14,889 15,330 ft 15,752 15,786 Iryantheraparadoxa (Schwacke)Warb. 9 14,739 14,846 14,853 14,865 14,890 14,901 Iryantheraparaensis Huber 3 14,668 15,067 15,656 Iryantherapolyneura Ducke 2 14,855 15,482 Iryantheraulei Warb. 29 14,603 14,722 14,743 15,287 ft 15,291 ft 15,510 ft Appendix1. Continued. 0o

No. CD plants in Vasquez collections Family Taxonand authority 2-ha (onlythe firstsix are listedper taxon) Z C Myristicaceae Iryantherasp. 1 15,262 Iryantherasp. 1 15,519 Osteophloeumplatyspermum (A. DC.) Warb. 4 14,701 15,484 15,561 15,750 Otobaglycicarpa W. Rodr. 2 15,560 15,641 Virolacalophylla Warb. 4 15,050 15,149 15,419 bd 15,465 ft Viroladivergens Ducke 1 15,823 Viroladuckei A. C. Sm. 1 15,591 Virolaelongata (Benth.) Warb. 2 14,767 ft 15,098 Virolamollisima (A. DC.) Warb. 1 15,725 Virolamultinervia Ducke 3 14,795 15,174 15,644 Virolapavonis (A. DC.) A. C. Smith 23 14,619 14,649 14,669 14,682 14,789 15,272 ft Myrsinaceae Cybianthusperuvianus Miq. 1 15,016 Myrtaceae genus 1 15,675 genus 1 15,687 I< genus 1 15,390 genus 1 15,769 genus 1 14,715 genus 1 14,816 genus 1 14,857 - genus 1 14,995 genus 1 15,033 genus 2 15,408 15,410 genus 1 15,400 genus 1 15,471 0 a genus 1 voucherlost Calyptranthescrebra McVaugh 1 14,956 Calyptranthescuspidata DC. 1 14,971 Calyptranthessimulata McVaugh 1 14,954 Eugeniapatrisii Vahl 1 15,534 Marlieracaudata McVaugh 3 15,266 15,343 15,443 Marlieraimperfecta McVaugh 1 14,955 Appendix1. Continued.

No. plants in Vasquez collections Family Taxonand authority 2-ha (onlythe firstsix are listedper taxon) Nyctaginaceae genus 1 15,194 ft genus 1 15,557 genus 1 14,696 genus 1 voucherlost Neea divaricataPoepp. & Endl. 2 14,671 15,820 NeeafioribundaPoepp. & Endl. 8 15,383 15,387 15,412 15,415 15,430 15,439 Neea parvifloraPoepp. & Endl. 1 15,795 Neea spruceanaHeimerl 3 15,655 15,710 15,711 Neea verticellataRuiz & Pav. 1 14,737 Neea sp. cf. macrophyllaPoepp. & Endl. 1 15,403 Ochnaceae Cespedesiaspathulata (Ruiz & Pav.) Planch. 1 14,911 14,912 Ourateasp. 1 15,248 Olacaceae Aptandratubicina (Poepp.) Benth.ex Miers 1 15,087 ft Cathedraacuminata (Benth.) Miers 2 14,909 bd 15,192 Minquartiaguianensis Aubl. 4 14,811 15,670 15,712 15,763 cJ)> Schoepfialucida Pulle 1 15,603 Opiliaceae Agonandrabrasiliensis Benth. 1 15,737 Agonandrasilvatica Ducke 1 15,530 Agonandrasp. 1 15,807 Palmae Astrocaryumchambira Burret 6 14,801 fl Astrocaryummacrocalyx Burret 14 14,592 0 Euterpeprecatoria Mart. 7 14,623 14,828 ft Q Iriarteadeltoidea Ruiz & Pav. 13 14,593 fl Oenocarpusbataua Mart. 78 14,633 15,020 Scheeleamoorei Glassman 3 no voucher Socrateaexorrhiza (Mart.) H. Wendl. 2 14,617 ft Polygonaceae Coccolobasp.*** 1 14,960 15,613 Coccolobasp. 1 15,613 Moutabeasp.*** 1 15,550 Quiinaceae Lacunariajenmanii (Oliver) Ducke 1 14,864 Quiina obovataTul. 3 14,687 15,142 15,167 Quiina sp. 1 15,749 Appendix1. Continued. |<

No. plants in Vasquez collections Family Taxonand authority 2-ha (onlythe firstsix are listedper taxon) Z C Rhizophoraceae Cassipoureaperuviana Alston 1 15,601 3 Rosaceae Prunusdetrita vel sp. aff.J. F. Macbr. 5 14,958 15,367 15,397 15,446 15,451 Rubiaceae Alseismicroarpa Standl. & Steyerm. 1 15,661 Borojoaclavifora (K. Schum.)Cuatrec. 2 15,180 15,636 Ferdinandusachlorantha (Wedd.) Standl. 5 15,242 15,344 15,372 15,374 15,452 Kotchubaeasericantha Standl. 2 14,999 15,743 Pagamea coriaceaSpruce ex Benth. 1 15,423 Rudgeafissistipula Muell. Arg. 1 14,915 ft Rutaceae Zanthoxylumhuberi Waterman 1 15,090 Sabiaceae Meliosmaherbertii Rolfe 2 15,283 ft 15,286 Ophiocaryonheterophyllum (Benth.) Urb. 2 15,497 15,541 Ophiocaryonklugii Barneby 3 15,610 15,646 15,696 Ophiocaryonmanausense (W. A. Rodrigues) I< Barneby 1 14,978 Sapindaceae Cupania diphyllaVahl 1 15,111 Cupania sp. 1 14,697 Mataybaarborescens (Aubl.) Radlk. 1 15,029 ft Mataybaguianensis Aubl. 1 15,828 Mataybasp. 3 14,935 15,000 15,023 CDCD Talisiajapurensis Radlk. 1 15,175 Talisia sp. 1 15,132 Sapotaceae Genus cf. Ecclinusa 1 21,365 Chrysophyllumamazonicum T. D. Penn. 3 15,118 15,520 15,596 1 j Chrysophyllumbombycinum T. D. Penn. 9 14,866 14,930 14,976 15,034 15,072 15,249 Chrysophyllumcuneifolium (Rudge) A. DC. 2 15,137 15,202 Chrysophyllumsanguinolentum (Pierre) Baehni 4 14,814 14,848 14,997 15,053 Ecclinusalanceolata (Mart. & Eichler)Pierre 4 14,807 15,012 15,326 15,328 Ecclinusaramiflora Mart. 1 14,884 Micropholisacutangula (Ducke) Eyma 1 14,713 Micropholiscasiquiarensis Aubrev. 4 14,675 fl 14,806 15,096 15,524 cn Micropholisegensis (A. DC.) Pierre 3 14,835 bd 14,883 15,133 Appendix1. Continued.

No. plants in Vasquez collections Family Taxonand authority 2-ha (onlythe first six are listedper taxon) Micropholisguyanensis subsp. duckeana (A. DC.) Pierre 4 14,938 14,986 15,409 15,590 Micropholisguyanensis (A. DC.) Pierresubsp. guyanensis 2 15,092 15,369 Micropholisvenulosa (Mart. & Eichler)Pierre 1 15,052 Pouteriabangii (Rusby) T. D. Penn. 5 14,849 15,037 15,076 15,129 15,187 Pouteriacaimito (Ruiz & Pav.) Radlk. 4 14,755 15,100 15,535 15,697 Pouteriacuspidata (A. DC.) Baehni 5 15,276 15,359 15,396 15,475 15,572 Pouteriaeugeniifolia vel. sp. aff.(Pierre) Baehni 1 15,119 Pouteriamultiflora vel sp. aff.(A. DC.) Eyma 2 14,727 15,086 Pouteriareticulata (Engl.) Eymasubsp. reticulata 1 15,723 Pouteriarostrata (Huber) Baehni 1 14,702 Pouteriasubrotata Cronquist 1 14,781 Pouteriatorta subsp. glabra T. D. Penn. 3 15,153 15,153 15,154 15,779 Pouteriatorta subsp. tuberculata(Sleumer) T. D. Penn. 2 14,720 fl 14,746 fl Pouteriasp. 1 14,790 Pouteriasp. 1 14,717 Simaroubaceae Simaba guianensisAubl. 1 14,607 Simabapolyphylla (Cavalc.) W. Thomas 1 15,688 Simaroubaamara Aubl. 6 14,606 14,952 15,178 15,495 15,576 15,677 0

Sterculiaceae Sterculiaapeibophylla Ducke 1 15,639 n Sterculiapruriens (Aubl.) K. Schum. 3 14,686 15,172 15,529 Sterculiatessmannii Mildbraed 2 15,186 15,507 Sterculiasp. 1 15,144 Sterculiasp. 1 15,539 Theobromaglaucum H. Karst. 1 15,734 Theobromaobovatum Klotzsch ex Bernoulli 1 15,556 Theobromasubincanum Mart. 10 14,635 14,716 14,792 15,008 15,051 15,114 (D 00 z

(D Appendix1. Continued.

No. plants in Vasquez collections Family Taxonand authority 2-ha (onlythe firstsix are listedper taxon) Tiliaceae Apeibaaspera Aubl. 3 15,014 bd 14,752 bd 15,672 fl Luehea sp. 1 14,714 fl Luehea sp. 1 14,648 Luehea sp. 1 14,658 Luehea sp. 1 15,685 Luehea sp. 1 15,754 I< Violaceae Leonia glycycarpaRuiz & Pav. var.glycycarpa 8 14,644 14,672 14,796 15,106 15,168 15,483 Paypayrolasp. 1 15,205 Rinoreaflavescens(Aubl.) Kuntze 1 15,190 Rinorearacemosa (Mart.) Kuntze 9 14,711 14,754 15,105 15,115 15,161 ft 15,200 Rinoreasp. 1 15,796 Rinoreasp. 1 15,718 ft Rinoreasp. 1 15,706 Vochysiaceae Qualea paraensisDucke 2 15,784 15,829 Vochysiadensifolia Spruce 1 14,666 0 -n ~cn