incidence of extra-floral nectaries and their effect on the growth

Incidence of Extra-Floral Nectaries and their Effect on the Growth and Survival of Lowland Tropical RainForest Trees

Honors Research Thesis

Presented in Partial Fulfillment of the Requirements for graduation “with Honors Research Distinction inEvolution and Ecology” in the undergraduate colleges of The Ohio State University

byAndrew Muehleisen

The Ohio State UniversityMay 2013

Project Advisor: Dr. Simon Queenborough, Department of Evolution, Ecology and Organismal Biology

Incidence of Extra-Floral Nectaries and their Effect onthe Growth and Survival of Lowland Tropical Rain ForestTreesAndrew Muehleisen

Evolution, Ecology & Organismal Biology, The Ohio State University, OH 43210, USA

SummaryMutualistic relationships between organisms have long captivated biologists, and extra-floral nectaries

(EFNs), or nectar-producing glands, found on many plants are a good example. The nectar produced fromthese glands serves as food for ants which attack intruders that may threaten their free meal, preventingherbivory. However, relatively little is known about their impact on the long-term growth and survival ofplants. To better understand the ecological significance of EFNs, I examined their incidence on lowlandtropical rain forest trees in Yasuni National Park in Amazonian Ecuador.

Of those 896 species that were observed in the field, EFNs were found on 96 species (11.2%), widelydistributed between different angiosperm families. This rate of incidence is high but consistent with otherlocations in tropical regions. Furthermore, this study adds 13 new genera and 2 new families (Urticaceae andCaricaceae) to the list of taxa exhibiting EFNs.

Using demographic data from a long-term forest dynamics plot at the same site, I compared the growth andsurvival rates of species that have EFNs with those that do not. This same analysis was also done with datafrom two other sites with EFN surveys, Barro Colorado Island, Panama and Pasoh Forest Reserve, Malaysia.Results showed that while species with EFNs have generally higher diameter growth rates, they also havehigher mortality rates than species without, suggesting a cost to this ecological strategy.

Keywords: extra-floral nectaries, tropical forest, growth rate, mortality rate

Introduction

Tropical forests represent a fascinating yet incrediblycomplex web of interactions, the ecology of which, inmany cases, is still largely enigmatic, and the mech-anisms that generate and maintain the remarkablediversity of plants and animals found within themremain a fundamental question in biology (Palmer1994, Hubbell 2001, Wright 2002). While one suiteof mechanisms are purely stochastic in nature (e.g.Hubbell 2001) many other mechanisms depend onniche differences between species to permit coexist-ence (Chesson 2000, Silvertown 2004). Niche differ-ences are driven primarily in response to selectionpressures, which in tropical forests include compet-ition with neighbors for (often low levels of) light,nutrients and water (Chapin et al. 1986, Denslowet al. 1987, Chazdon & Pearcy 1988), as well as in-tense predation pressure from pests, pathogens andherbivores (Barone 2000, Novotny et al. 2010).

Herbivory represents a particularly selective force,

as up to 20% of plant net primary production maybe consumed each year (Agrawal 2011). In response,tropical rain forest trees have developed a myriadof defense mechanisms, from physical (e.g. spines,hairs; Hanley et al. 2007) to chemical (e.g. low nu-trition, toxic compounds; Feeny 1976, Levin & York1978, Coley & Barone 1996). Further, many plantshave evolved mutualistic relationships with animalsin an effort to deter herbivores. A common mutual-ism is with ants and such ant-plant relationships offera considerable measure of defense from herbivory,and can have a positive impact on plant performance(Beattie 1985).

One such example of ant-plant mutualisms areextra-floral nectaries (EFNs), which are nectar-producing glands found outside of a plant’s flower,typically at the base of the leaf or on the petiole, al-though their location can vary considerably on theplant (Figure 1). EFNs vary in morphology, rangingfrom raised bowls or bulbs to very small hairs andtissues (Elias 1983). The nectar produced by these

A. Muehleisen Incidence and Ecological Significances of Extra-Floral Nectaries 2

glands serves as a food source, primarily for ants,which are believed to provide protection to the plantin return, by way of aggression toward intruding or-ganisms including herbivores (Bentley 1977a, Keeler1977, 1989, Koptur 1992). This form of ant protection-ism can result in reduced damage to both vegetativeand reproductive parts, improving plant perform-ance and fitness (Koptur 1992, Oliveira 1997). How-ever, relatively little is known about their overallecological impact at the population and communitylevels as well as on the long-term performance ofindividual plants.

Fig. 1 Example structure and locations ofextra-floral nectaries on plants. Source: Aus-tralia Biological Resources Study (environ-ment.gov.au/biodiversity/abrs)

Previous intensive surveys have determined theincidence of EFNs on Barro Colorado Island, Panama(Schupp & Feener 1991) and in the Pasoh Forest Re-serve, Malaysia (Fiala & Linsenmair 1995). Thesestudies provided an excellent picture of the distribu-tion of EFNs at these sites, and until last year were thebest data available on the phylogenetic distributionof EFNs. However, new work has drawn togetherall available data on EFN incidence currently known,to examine the phylogenetic distribution of EFNsthroughout the plant phylogeny (Weber & Keeler2012). This study found 1.0-1.8% of plant species hadEFNs, distributed in 108 families, although the au-thors suggest that the unknown incidence of EFNs

may be as great as the their currently known incid-ence (Weber & Keeler 2012), requiring further in-depth studies of EFN incidence within and betweenplant communities.

In this study, I expand upon our prior understand-ing by undertaking an intensive survey of EFN in-cidence of tree species in an old growth Neotropicalaseasonal lowland rain forest, an environment thathas not yet been studied for EFNs. I analyze the long-term demographic rates of trees with and withoutEFNs to elucidate the ecological significance of thisdefensive strategy. In a large permanent forest plot inYasuni National Park, Ecuador, I examined 896 spe-cies of tree for the presence or absence of EFNs. I usedpublished census data to compare abundance, andgrowth and mortality rates of trees with and withoutEFNs. Finally, I also used the results of surveys in BCIand Pasoh to examine how plant performance is re-lated to EFN incidence there, such that a comparisonof the phylogenetic distribution and demographicrates related to EFN can be made between three studysites. If mutualism with ants, and EFNs in particular,provide a benefit, I predict higher abundance andgreater performance in species with EFNs.

QUESTIONS

1. What is the incidence and phylogenetic distribu-tion of extra-floral nectaries on trees in a lowlandNeotropical rain forest?

2. Do trees with extra-floral nectaries have differ-ent abundances, and growth and mortality ratesthan trees without extra-floral nectaries?

3. How do findings from our study site comparewith other locations in which the incidence ofextra-floral nectaries have been studied?

Methods

STUDY SITES

I carried out fieldwork in Yasuni National Park,Ecuador, and used published data from Barro Col-orado Island, Panama (Feener & Schupp 1991) and

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Pasoh Forest Reserve, Malaysia (Fiala & Linsenmair1995) on the incidence of EFN (Figure 2).

Yasuni National Park and adjacent Huaorani ter-ritory comprise 1,600,000 ha of largely pristine trop-ical lowland aseasonal rain forest in eastern Ecuador(Finer et al. 2009, Bass et al. 2010). Yasuni ScientificResearch Station, established and maintained by thePontificia Universidad Catolica del Ecuador, is loc-ated in the north-western corner of the park, in terra-firme, mature forest bordering the Tiputini River. Theresearch station maintains a 25 ha Forest DynamicsPlot (FDP, 0◦41’S, 76◦24’W), which lies along twosmaller ridges dominated by red clays and separatedby a valley characterized by brown or grey alluvium(Valencia et al. 2004). The plot is extremely biologic-ally diverse, with a described tree species count of1,104 (Valencia et al. 2004). The climate at Yasuni isaseasonal, with an average annual rainfall of 2,826mm, with no month receiving less than 100 mm ofrainfall (Valencia et al. 2004).

Barro Colorado Island (BCI), Panama is a 1,560ha island located in Gatun Lake, formed when thePanama Canal was developed. The 50 ha ForestDynamics Plot was established in 1980 and is main-tained by the Smithsonian Tropical Research Institute(STRI). The FDP is located near the center of BCI(9◦9’S, 79◦50’W) and consists primarily of lowlandmoist tropical forest, about half of which is maturegrowth. There is a relatively high diversity of treesat the FDP, with 321 different species of tree recor-ded. The climate at BCI is seasonal, with a dry seasonlasting roughly from December to April or May andan average annual rainfall of 2,551 mm (Leigh et al.2004).

Pasoh Forest Reserve, Malaysia is a 11,000 ha re-serve situated in peninsular Malaysia. The 50 haForest Dynamics Plot situated within the reserve(2◦58’N, 102◦18’E) was established in 1986 and is andmaintained by the Forest Research Institute Malaysia.The forest consists primarily of lowland mixed dip-terocarp forest, and is surrounded by roughly 1,000ha of previously logged forest. The FDP at Pasohhas a tree diversity of around 824 species. The cli-mate at Pasoh is seasonal, with dips in precipitation

in January-February and June-July, and an averageannual rainfall of 1,571mm (Manokaran et al. 2004).

Fig. 2 Locations of Yasuni National Park, BarroColorado Island and Pasoh Forest Reserve

FIELD SURVEYS

I undertook a survey for incidence of extra-floral nec-taries on woody species at Yasuni in June-August2012. Species were censused in three ways. In thefield, I searched along trails within and around theFDP and found 787 species. A further 109 rare spe-cies were found by searching for specific individualswithin the FDP. In this way, I examined a total of896 species in the field (81% of the total 1,104 speciesin the FDP). The remaining 208 species that I couldnot find in the field were checked from dried spe-cimens in the field station herbarium. This methodwas effective for those plants with obvious nectarystructures (e.g. Fabaceae), although dried structuresare much more difficult to identify than living struc-tures. Therefore, species with EFNs determined inthe herbarium were included only in demographicanalysis.

Data on the incidence of extra-floral nectaries forspecies from the other two sites were obtained fromSchupp & Feener (1991, BCI) and Fiala & Linsenmair(1995, Pasoh). I added to these data with new datafrom Croat (1978) and Garwood (2009), for specieslocated in the BCI FDP.

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DEMOGRAPHIC DATA

At all three sites, identical methodology was followedto establish large forest plots. All plots were pro-fessionally surveyed, and within them every shruband tree stem >1 cm diameter at breast height (DBH,1.3m) are mapped, marked, measured and identi-fied every 5 years (Condit 1998). To date, threecensuses have been carried out at Yasuni, four atBCI and three at Pasoh. All demographic data canbe found at the Center for Tropical Forest Sciencewebsite (www.ctfs.si.edu, for a summary, see Table1).

From these census data, demographic rates havebeen calculated (Condit et al. 2006). Annual mortal-ity (survival from one census to the next) and growthrates (diameter increment) were determined usingBayesian hierarchical models. Abundance and demo-graphic rates were calculated for each species forindividuals in two size classes: 1-10 cm DBH and>10 cm DBH. For consistency, census years leadingup to or closest to the year 2000 were used. For eachspecies, I also assigned growth form (shrub, treelet,understory tree, canopy tree or emergent tree). Fi-nally, the higher-level taxonomy for each site wasupdated to reflect the Angiosperm Phylogeny GroupIII (APG III) system (The Angiosperm PhylogenyGroup 2009).

DATA ANALYSIS

To examine the taxonomic distribution of extra-floralnectaries, I compared the proportions of individuals,species, genera, families, and orders with EFNs ateach site using a proportion test. To test whetherspecies with EFNs were more abundant than spe-cies without, and also whether species with EFN hadhigher growth rates and higher mortality rates, I usedlinear regression. All data analysis was completed inthe statistics package R version 2.15.1.

Results

I surveyed shrub and tree species at three tropicalforest sites for extra-floral nectaries. At Yasuni, I

censused 896 species out of 1,104 species on the FDP.At BCI, Schupp & Feener (1991) surveyed 173 spe-cies, though only 150 of these are present on the FDP(of 321 total). Using additional references, I addedanother 24 species with EFNs. At Pasoh, Fiala & Lin-senmair (1995) surveyed 741 out of 824 species. Thus,I have a good sample of the species at each site, andmost of the unsurveyed species are rare and thus non-representative of the community as a whole. Detailsof species from each site with EFN can be found inAppendix 2.

TAXONOMIC DISTRIBUTION

At Yasuni, I found 96 species with extra-floral nectar-ies (11.2% of the total 896 species, Figure 3a). Thesewere distributed among 41 genera and 17 families.Over half (58) of the species with EFNs were in thefamily Fabaceae, largely thanks to the diversity ofInga (44 species) at Yasuni, all of which have EFNs.Seventy-nine percent of all species with EFNs werefound within either the Fabales or Malpighiales or-ders. In addition, I documented 13 new genera and 2new families (Caricaeae and Urticaceae) to the globallist of taxon exhibiting EFNs (Keeler 2013).

At BCI, 49 (32.7%) of 150 species of tree were foundto have EFNs (Schupp & Feener 1991, Figure 3b).They were distributed among 31 genera and 19 fam-ilies (Figure 2b). Eighteen species with EFNs were inthe family Fabaceae, also due largely to the diversityof Inga (15 species). Similar to Yasuni, 61% percent ofall species with EFNs were found within either theFabales or Malpighiales orders.

At Pasoh, 80 (9.7%) of 824 species were found tohave EFNs (Fiala & Linsenmair 1995, Figure 3c). Theywere distributed among 47 genera and 16 families.Unlike Yasuni and BCI, Pasoh exhibited a more evendistribution of EFN bearing trees across different taxa.Euphorbiaceae, rather than Fabaceae, exhibited themost species with EFNs (21 species). Forty percent ofall species with EFNs were in the order Malpighiales,while the next most important order was Malvalescontaining 17.5% of species with EFNs.

Across all study sites, Yasuni and Pasoh were

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Table 1 Demographic rates and abundances for all tree species per hectare across all three sites. Growthis measured in mm per year, mortality in % per year and abundance in individuals per ha.

1-10 cm DBH >10 cm DBH

Yasuni Pasoh BCI Yasuni Pasoh BCI

Growth

Min 0.79 1.02 0.90 0.22 0.27 0.27Max 6.25 3.13 8.69 3.01 1.66 4.22Mean 1.76 1.57 2.53 0.74 0.60 1.04SD 0.62 0.26 1.30 0.39 0.20 0.60

Mortality

Min 0.27 0.43 0.23 0.36 0.51 0.33Max 20.41 19.21 30.90 16.68 7.58 17.70Mean 2.04 1.99 4.31 1.49 1.87 2.72SD 2.27 1.67 4.66 1.02 0.75 2.20

Abundance

Min 0.00 0.00 0.00 0.00 0.00 0.00Max 184.60 159.88 638.56 71.96 11.16 36.56Mean 5.64 6.51 21.10 0.72 0.69 1.99SD 13.10 14.71 70.45 2.91 1.33 4.31

most similar in their proportion of species with EFNs(11.2% in Yasuni, 32.7% in BCI, and 9.7% in Pasoh).BCI exhibited an incidence of species with EFNs upto three times greater than Yasuni and Pasoh, and atall taxon levels, BCI showed greater proportions ofplants with EFNs than both Yasuni and Pasoh. How-ever, BCI exhibited a much lower incidence of EFNsat the individual level compared with other studysites. The distribution of species with EFNs in eachfamily for each site can be seen in Figure 4.

ABUNDANCE, AND GROWTH AND MORTALITYRATES

Species varied widely in their abundances, growthand mortality rates (Table 1). Species at BCI were onaverage three times more abundant than species fromYasuni and Pasoh, which reflects the lower speciesrichness found at BCI. Both growth and mortalityrates were also much higher at BCI than those foundat Yasuni or Pasoh.

In accordance with my prediction, species withEFN had higher mean abundance than specieswithout EFNs at Yasuni and Pasoh, but this was not

the case at BCI (Figure 5 a, b, c). At Yasuni, speciesabundances for trees with EFNs were 25-30% greaterthan those without. At Pasoh, these differences wereeven greater, where those species with EFNs were114-117% more abundant than those without. In con-trast, at BCI trees in the small size class without EFNswere almost 170% more abundant, and abundanceswere lower for those individuals >10 cm DBH.

Significantly greater growth rates were found intrees with EFNs in each plot, although this differedwith size class between sites. At Yasuni, trees withEFNs in both size classes had higher growth rates,growing about 0.28 mm extra per year than specieswithout EFNs. At BCI only trees in the small sizeclass had significantly higher growth rates, and atPasoh only trees in the large size class (Figure 5e, f).

Significantly higher mortality rates were found forspecies with EFN in the large size class (trees >10 cmDBH) at all three sites (Figure 5g, h, i), and in Pasohtrees 1-10 cm DBH also had significantly higher mor-tality rates. In all cases growth and mortality rateswere found to be either greater or the same in treeswith EFNs.

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Fig. 3 Proportion of trees that exhibit EFNs at the individual and varying taxonomic levels, by location.

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Fig. 4 Number of species examined in each family at each site, with dark bars representing species withextra-floral nectaries.

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Fig. 5 Abundance (a,b,c), and growth (d,e,f) and mortality (g,h,i) rates as a function of EFN presence inshrub and tree species in three tropical forest sites. Points are mean values per EFN group, split into two sizeclasses: trees ≤10 cm DBH (red circles),trees >10 cm DBH (black squares). Points connected by dashed linesthat are filled indicate statistical difference at P < 0.05. Error bars show 95% confidence intervals.

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Discussion

From an intensive field survey of 896 tree and shrubspecies in an Amazonian lowland tropical rain forest,I documented 96 species with extra-floral nectaries,64 of which had not previously been recorded as pos-sessing EFNs. Comparing Yasuni with two otherintensive survey sites, I found that the distribution ofEFNs across taxa was consistent between Yasuni andPasoh, and broader at BCI. The reverse was foundto be true in terms of total EFN presence on indi-viduals at each plot. Species with EFNs appeared tobe more successful ecologically at Yasuni and Pasoh,having higher abundance than species without EFNs,while at BCI the opposite was true. Further, I founda significant effect of EFN incidence on long-termplant performance. Tree species with EFNs showedhigher growth and mortality rates compared to thosewithout EFNs at all three sites in at least one sizeclass.

TAXONOMIC DISTRIBUTION AT YASUNI

This study adds 13 new genera and 2 new families(Caricaeae and Urticaceae, Appendix 1) to the listof taxon exhibiting EFNs. This increases the globalnumber of familes with EFNs to 110, 17 of whichare found at Yasuni. The family with the most num-ber of EFNs at Yasuni was Fabaceae, which is alsotrue globally. However, the family Euphorbiaceaehad the second highest incidence of EFNs at Yasuni,which stands in contrast to global patterns whichshow Passifloraceae and Malvaceae as second andthird, respectively. Only one species from Malvaceaehad nectaries at Yasuni, while there were none fromPassifloraceae, although I did not survey any vinesand lianas, the predominant growth form of Passi-floraceae. The presence of EFNs at Yasuni was muchgreater than the currently known worldwide incid-ence (11.2% at Yasuni, to 1.5% worldwide, Weber &Keeler 2012).

TAXONOMIC DISTRIBUTION BETWEEN SITES

All three locations exhibited fairly equivalent dis-tribution of EFNs across taxon, with BCI represent-ing the greatest breadth of distribution. Oddly, BCIalso exhibited the lowest total number of individualswith EFNs, despite the wide taxonomic distributionand greater number of species relative to Pasoh andYasuni. Pasoh, which overall had the smallest phylo-genetic distribution and species count of those treeswith EFNs, had the greatest number of individualswith nectaries. It is not certain what might cause thistrend, though care should be taken labeling this atrend from only three plots. This relationship couldbe examined in other plots to determine whether atrend truly exists.

In Yasuni, BCI and Pasoh the orders Fabales andMalpighiales are well represented by species withEFNs, with at least 10 species being found in eachorder with EFNs. Unlike in BCI and Yasuni, the or-ders Ericales and Malvales were also found to haveat least 10 species with EFNs in Malaysia, accordingto Fiala & Linsenmair (1995). EFNs were found inboth of these families in BCI and Yasuni, but not tothe extent that they were found in Pasoh. As such,despite an overall smaller distribution of EFNs acrossorders in Pasoh (27% in Pasoh, as opposed to 33%and 36% in Yasuni and BCI, respectively), more famil-ies were well represented by species with EFNs. BCIand Yasuni, then, have a thinner distribution of EFNsacross orders. This is generally the case for famil-ies as well, as those orders with many EFN bearingspecies in Pasoh are this way due to particularly well-represented families (Dipterocarpaceae, Ebenaceae,Euphorbiaceae and Fabaceae). This, in large part, re-flects the different floristic composition of Paleo vsNeotropical forests (Gentry 1993).

GEOGRAPHIC DISTRIBUTION

An increase in EFN presence as latitude decreaseshas been noted previously (Pemberton 1998), but itis also informative to examine how EFN distributionchanges across different habitat types at similar lat-itudes. Yasuni and Pasoh, which are two lowland

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tropical rainforests at comparable latitudes, are verysimilar in their incidence of EFNs (11.2% and 9.7%, re-spectively) suggesting little difference in distributionbetween the Neotropics and Paleotropics, as represen-ted by these two sites. The slightly higher incidencein the Neotropics may be attributed to the greaterdiversity of Fabaceae found there (Gentry 1993).

Within the Neotropics, the Brazilian cerrado hasalso been surveyed for the presence of EFNs. Anincidence of about 17% of woody plants with EFNswas found in the cerrados, which are considerablydrier than rain forest (Oliveira & Leitao-Filho 1987).Perhaps the greater presence of EFNs found in the cer-rado indicates that ant-plant interactions are strongerin this type of habitat. Further work in understandingthe differences in ant diversity and presence betweenthese two habitats may better inform our understand-ing of this mutualism across geographically similarhabitats.

ECOLOGICAL SIGNIFICANCE OF EXTRA-FLORAL NECTARIES

Differences found in growth, mortality and abund-ance rates in those trees with EFNs and those withoutare surprising, given the large and significant dif-ferences in performance I found contingent on thepresence of a single character. Nonetheless, greatergrowth and mortality rates were characteristic of treespecies with EFNs across all three sites. This consist-ency suggests that species with EFNs grow faster anddie younger than species without EFNs. Why, then, isa higher mortality rate associated with a defense traitthat is assumed to improve plant performance? Thisgeneral lifestyle is informative in that it may suggestthat many of those trees with EFNs are pioneer spe-cies. Pioneer species generally exhibit faster growthrates and higher mortality rates compared with moreslower growing but persistent shade-tolerant spe-cies (Brokaw 1985). Additionally, pioneer speciesundergo intense competition following the formationand colonization of a gap (Denslow 1980, Denslow etal. 1985), lending increased importance to adaptivestrategies such as EFNs. In this sense, pioneer spe-cies may exhibit EFNs more commonly than shade-

tolerant species, and explain this observed differencein demographic rates.

Abundances were also significantly different,though those at BCI showed the reverse trend inabundance compared with Pasoh and Yasuni. Thehigher abundances observed at Yasuni and Pasohsuggest that this defensive strategy has a positiveimpact on the plant’s ability to out-perform specieswithout this adaptation, and also counteracts the sug-gestion that differences in performance are driven bypioneer species, because these light-demanders tendto be rare (Wright 2002).

Conclusions

I found similar phylogenetic distributions of extra-floral nectaries in comprehensive surveys of tree spe-cies of an aseasonal lowland rain forest in Ecuador,and then compared to the previously determined dis-tribution of trees with EFNs lowland semi-deciduousmoist forest in Panama and lowland rain forest inMalaysia. Additionally, I found evidence of a signific-ant effect of EFN presence on individual performance.Trees with EFNs had higher growth but also highermortality rates than those without EFNs, consistentacross all three sites. These results suggest a signific-ant role for extra-floral nectaries and plant defensemechanisms in general for determining forest struc-ture and composition.

Acknowledgements

I would like to thank the Pontificia UniversidadCatolica del Ecuador and Yasuni Scientific ResearchStation for providing logistical support. I also thankPablo Alvia for aiding in the identification of treesin the Yasuni Forest Dynamics Plot. I thank the Min-isterio del Medioambiente of Ecuador for allowingthis field work to happen at Yasuni National Park. Ithank the Smithsonian Tropical Research Institute forproviding demographic data for each plot. The Un-dergraduate Research Office supported this researchthrough funding (URO Summer Research Fellow-ship) and other resources.

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Keeler, K. H. (2013). World List of Angiosperm Spe-cies with Extrafloral Nectaries. Retrieved fromhttp://bioscilabs.unl.edu/Emeriti/keeler/ ex-trafloral/worldlistfamilies.htm

Koptur, S. (1992). Extrafloral nectary-mediated inter-actions between insects and plants. Insect PlantInteractions, 4, 81-129.

Leigh Jr, E. G., Loo de Lao, S., Condit, R., Hubbell,S. P., Foster, R. B., & Perez, R. (2004). BarroColorado island forest dynamics plot, Panama.Pp: 451-463. In: Losos, E. C., & Leigh, E. G.(Eds.). Tropical Forest Diversity and Dynamism:Findings from a large-scale plot network. Universityof Chicago Press.

Levin, D. A., & York Jr, B. M. (1978). The toxicity ofplant alkaloids: an ecogeographic perspective.Biochemical Systematics and Ecology, 6, 61-76.

Manokaran, N., Seng, Q. E., Ashton, P. S., LaFrankie,J. V., Noor, N. S. M., Ahmad, W. M. S. W., &Okuda, T. (2004). Pasoh forest dynamics plot,Peninsular Malaysia. Pp. 585-598. In: Losos, E.C., & Leigh, E. G. (Eds.). Tropical Forest Diversityand Dynamism: Findings from a large-scale plotnetwork. University of Chicago Press.

Novotny, V., Miller, S. E., Baje, L., Balagawi, S., Basset,Y., Cizek, L., Craft, K. J., Dem, F., Drew, R. A.,Hulcr, J., Leps, J., Lewis, O. T., Pokon R., Stewart,A. J., Samuelson, G. A., & Weiblen, G. D. (2010).Guild specific patterns of species richness andhost specialization in plant–herbivore food websfrom a tropical forest. Journal of Animal Ecology,79, 1193-1203.

Oliveira, P. S., & Leitão-Filho, H. F. (1987). Extra-floral nectaries: their taxonomic distributionand abundance in the woody flora of cerradovegetation in Southeast Brazil. Biotropica, 19,140-148.

Oliveira, P. S. (1997). The ecological function of extra-floral nectaries: herbivore deterrence by visitingants and reproductive output in Caryocar bra-siliense (Caryocaraceae). Functional Ecology, 11,323-330.

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Pemberton, R. W. (1998). The occurrence and abund-ance of plants with extrafloral nectaries, thebasis for antiherbivore defensive mutualisms,along a latitudinal gradient in east Asia. Journalof Biogeography, 25, 661-668.

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13


Appendix 1: List of taxa from Yasuni not previously recorded with EFN

Genus Species Family Order

Lauraceae miniopacangulo Lauraceae LauralesAbarema laeta Fabaceae FabalesAlchornea glandulosa Euphorbiaceae MalpighialesAlchornea schomburkii_cf. Euphorbiaceae MalpighialesAparisthmium cordatum Euphorbiaceae MalpighialesBuchenavia congesta Combretaceae MyrtalesBuchenavia grandis Combretaceae MyrtalesBunchosia argentea Malpighiaceae MalpighialesBunchosia argentea(granpanosa) Malpighiaceae MalpighialesCaryodendron orinocense Euphorbiaceae MalpighialesCedrelinga catenaeformis Fabaceae FabalesColubrina arbol Rhamnaceae RosalesConceveiba rhytidocarpa Euphorbiaceae MalpighialesEnterolobium barnebianum Fabaceae FabalesGlycydendron amazonicum Euphorbiaceae MalpighialesHirtella excelsa Chrysobalanaceae MalpighialesInga 3crasa Fabaceae FabalesInga 3oscura Fabaceae FabalesInga 4alitarco Fabaceae FabalesInga 6cuadra Fabaceae FabalesInga acreana Fabaceae FabalesInga alata Fabaceae FabalesInga alba Fabaceae FabalesInga auristellae Fabaceae FabalesInga bourgonii Fabaceae FabalesInga brachyrhachis Fabaceae FabalesInga cayennensis Fabaceae FabalesInga chartacea Fabaceae FabalesInga ciliata_ssp.subcapita Fabaceae FabalesInga cinnamomea Fabaceae FabalesInga cordatoalata Fabaceae FabalesInga falsacre Fabaceae FabalesInga leiocalycina Fabaceae FabalesInga microcoma Fabaceae FabalesInga multinervis Fabaceae FabalesInga poeppigiana Fabaceae FabalesInga rusbyi Fabaceae FabalesInga sarayacuensis Fabaceae FabalesInga spectabilis Fabaceae FabalesInga stellaeglabra Fabaceae FabalesInga stipulacea Fabaceae FabalesInga striata Fabaceae Fabales

14


Inga tenuistipula Fabaceae FabalesInga thibaudiana Fabaceae FabalesInga thibaudiana_ssp.peltadenia Fabaceae FabalesInga tocacheana Fabaceae FabalesInga umbratica Fabaceae FabalesInga velutina Fabaceae FabalesInga vismiifolia Fabaceae FabalesInga yacoana Fabaceae FabalesJacaratia digitata Caricaceae BrassicalesLicania caudata Chrysobalanaceae MalpighialesLicania longistyla Chrysobalanaceae MalpighialesLicania nervifina Chrysobalanaceae MalpighialesMarmaroxylon basijugum Fabaceae FabalesPausandra trianae Euphorbiaceae MalpighialesPourouma minor Urticaceae RosalesPrunus debilis Rosaceae RosalesQualea paraensis Vochysiaceae MyrtalesSapium largident Euphorbiaceae MalpighialesTerminalia ob Combretaceae MyrtalesTetrorchidium macrophyllum Euphorbiaceae MalpighialesZygia heteroneura Fabaceae FabalesZygia mediana Fabaceae Fabales

15


Appendix 2: Species list and EFN incidence for each plot

Yasuní BCI Pasoh

Genus species EFN Source Genus species EFN Source Genus species EFN Source(combretaceae) ovni Abarema macradenia n Acronychia porteri n study(fabaceae) 20-25oblong Acacia melanoceras y keeler Actinodaphne macrophylla n study(fabaceae) brillafuzzy n Field Acalypha diversifolia n study Actinodaphne pruinosa n study(fabaceae) diploglauco Acalypha macrostachya n Actinodaphne sesquipedalis n study(hippocrateaceae) atenumembra n Field Adelia triloba n Adenanthera bicolor n study(hippocrateaceae) cheiloancho Aegiphila panamensis y keeler Aglaia aspera n study(hippocrateaceae) ovalo n Field Alchornea costaricensis y study Aglaia cordata n study(lauraceae) bals n Field Alchornea latifolia y study Aglaia exstipulata n study(lauraceae) chiquita n Field Alibertia edulis n study Aglaia forbesii n study(lauraceae) chorongo Allophylus psilospermus n study Aglaia ganggo n study(lauraceae) furrycanela n Field Alseis blackiana n study Aglaia glabriflora n study(lauraceae) furrymen Amaioua corymbosa n Aglaia grandis n study(lauraceae) granbrillacuspi n Field Anacardium excelsum n study Aglaia meliosmoides n study(lauraceae) granverde n Field Anaxagorea panamensis n Aglaia odoratissima n study(lauraceae) impresofalso n Field Andira inermis n study Aglaia oligocarpa n study(lauraceae) largoarco Annona acuminata n study Aglaia oligophylla n study(lauraceae) licar n Field Annona hayesii n study Aglaia palembanica n study(lauraceae) lisagroovy n Field Annona spraguei n study Aglaia ridleyi n study(lauraceae) medpubinervi n Field Apeiba hybrid n Aglaia rubescens n study(lauraceae) miniopacangulo y Field Apeiba membranacea n study Aglaia rufa n study(lauraceae) pelonegro Apeiba tibourbou n study Aglaia species_1 n study(lauraceae) subopo n Field Aphelandra sinclairiana y keeler Aglaia species_2 n study(lauraceae) tallorojo Ardisia bartlettii n study Aglaia species_3 n study(malpighiaceae) bulondu Ardisia fendleri n Aglaia species_4 n study(malpighiaceae) granoscura Ardisia guianensis n Aglaia species_5 n study(malpighiaceae) myrtosco Aspidosperma spruceanum n study Aglaia species_7 n study(melastomataceae) dienteblanca Astrocaryum standleyanum n Aglaia tenuicaulis n study(meliaceae) alarganervi Astronium graveolens n Aidia wallichiana n study(moraceae) pequecilio n Field Attalea butyracea n Alangium ebenaceum n study(myrtaceae) cortezapeq n Field Bactris barronis n Alangium griffithii n study(myrtaceae) falsasalas Bactris coloniata n Alangium nobile n study(myrtaceae) membrapelos Bactris coloradonis n Alangium ridleyi n study(myrtaceae) platatomen Bactris major n Albizia pedicellata y study(myrtaceae) prominsuave n Field Banara guianensis y keeler Alchornea rugosa y study(myrtaceae) smedcheilo Beilschmiedia pendula n study Allophylus cobbe n study(myrtaceae) smedcomun n Seed Bertiera guianensis n Alphonsea elliptica n study(myrtaceae) smedhipocrat n Field Borojoa panamensis n Alphonsea maingayi n study(myrtaceae) smedpubicost n Field Brosimum alicastrum n study Alseodaphne nigrescens n study(picramniaceae) sp.nov. n Field Brosimum guianense n Alseodaphne peduncularis n study(rubiaceae) amarillapub Calophyllum longifolium n study Alseodaphne perakensis n study(rubiaceae) ampliovada n Fieldh Capparis frondosa n study Alseodaphne species_1 n study(rubiaceae) neoide Casearia aculeata n Alseodaphne species_2 n study(rubiaceae) renato n Field Casearia arborea n study Alstonia angustiloba n study(rubiaceae) retidomatia Casearia commersoniana n Anacolosa heptandra n study(rubiaceae) viveca Casearia guianensis n Anaxagorea javanica n study(rubiaceae) wilson n Field Casearia sylvestris n Anisophyllea corneri n study(sapindaceae) amarillasper Cassipourea elliptica n study Anisoptera costata n study(sapindaceae) rua n Field Cavanillesia platanifolia n study Anisoptera laevis n study(solanaceae) plata Cecropia insignis n Anisoptera megistocarpa n studyAbarema jupunba y Field/Keeler Cecropia longipes n Antidesma coriaceum n studyAbarema laeta y Field Cecropia obtusifolia n Antidesma cuspidata n studyAbuta grandifolia n Field Cedrela odorata y study Antidesma pendulum n studyAcalypha cuneata n Field Ceiba pentandra y study Antidesma velutinosum n studyAcalypha pub Celtis schippii n study Aphanamixis polystachya n studyAcalypha sharpdent Cespedesia spathulata n Aphanamixis sumatrana n studyAcalypha sharpub Cestrum megalophyllum n Aporusa aurea y studyAcanthosyris annonagustata n Field Chamaedorea tepejilote n Aporusa bracteosa n studyAcidoton nicaraguensis n Field Chamguava schippii n Aporusa confusa n studyAegiphila cordifolia_var.villos n Field Chimarrhis parviflora n Aporusa falcifera n studyAegiphila elegans Chrysochlamys eclipes n Aporusa globifera n studyAegiphila haughtii n Field Chrysophyllum argenteum n Aporusa lunata n studyAegiphila integrifolia Chrysophyllum cainito n study Aporusa microstachya n studyAegiphila memboscu n Field Cinnamomum triplinerve n Aporusa miqueliana n studyAgonandra peruviana n Field Clidemia dentata n Aporusa nervosa n studyAgonandra silvatica n Field Clidemia octona n Aporusa nigricans n studyAiouea grandifolia_aff. Clidemia septuplinervia n Aporusa nigropunctata n studyAiouea sp.nov. n Field Coccoloba coronata n study Aporusa prainiana n studyAiphanes ulei n Coccoloba manzinellensis n study Aporusa symplocoides n studyAlbizia niopoides Cojoba rufescens n Aquilaria malaccensis n studyAlchornea glandulosa y Field Colubrina glandulosa n Aralidium pinnatifidum n studyAlchornea schomburkii_cf. y Field Conostegia bracteata n study Archidendron bubalinum y studyAlchornea triplinervia y Field/Keeler Conostegia cinnamomea n Archidendron clypearia y studyAlchorneopsis floribunda y Keeler Cordia alliodora n Archidendron contortum y studyAlibertia isernii n Field Cordia bicolor n study Archidendron globosum y studyAlibertia jorge Cordia lasiocalyx n study Archidendron microcarpum y studyAlibertia lance n Field Coussarea curvigemmia n study Ardisia colorata n studyAlibertia pelitos n Field Coutarea hexandra n Ardisia crassa n studyAlibertia pilosa Croton billbergianus y study Ardisia kunstleri n studyAllophylus amazonicus n Field Cupania cinerea n Ardisia lanceolata n studyAllophylus divaricatus n Field Cupania latifolia n Ardisia pachysandra n studyAllophylus glabra n Field Cupania rufescens n study Ardisia ridleyi n studyAllophylus pilosus n Field Cupania seemannii n Ardisia species_2 n studyAllophylus puctatus Cyathea petiolata n Aromadendron elegans n studyAlseis lugonis Dendropanax arboreus n study Arthrophyllum diversifolium n study

16


Alseis lugonis_cf. n Field Desmopsis panamensis n study Artocarpus anisophyllus n studyAlsophila cuspidata Diospyros artanthifolia y study Artocarpus dadak n studyAmpelocera edentula n Field Dipteryx oleifera n Artocarpus fulvicortex n studyAmpelocera longissima n Field Drypetes standleyi n study Artocarpus integer n studyAmyris macrocarpa n Field Elaeis oleifera n Artocarpus kemando n studyAnaxagorea brevipes Enterolobium schomburgkii y keeler Artocarpus lowii n studyAndira inermis n Field Erythrina costaricensis n study Artocarpus maingayi n studyAndira macrothyrsa Erythroxylum macrophyllum n Artocarpus nitidus_var.griffithii n studyAndira multistipula Erythroxylum panamense n study Artocarpus rigida n studyAndira sp.nov. n Field Eugenia coloradoensis n study Artocarpus scortechinii n studyAniba angulopepper n Field Eugenia galalonensis n Atuna elata n studyAniba guianensis n Field Eugenia nesiotica n Atuna excelsa n studyAniba hostmanniana n Field Eugenia oerstediana n study Austrobuxus nitidus n studyAniba riparia n Field Faramea occidentalis n study Baccaurea griffithii n studyAniba taubertiana Ficus bullenei n Baccaurea kunstleri n studyAnnona ambotay_aff. n Field Ficus citrifolia n Baccaurea maingayi n studyAnnona duckei n Field Ficus colubrinae n Baccaurea minor n studyAnnona mosaic Ficus costaricana n Baccaurea parviflora n studyAnthodiscus amazonicus n Field Ficus insipida n study Baccaurea pyriformis n studyAparisthmium cordatum y Field Ficus maxima n Baccaurea racemosa n studyApeiba membranacea n Field Ficus obtusifolia n Baccaurea ramiflora y keelerApeiba tibourbou Ficus pertusa n Baccaurea reticulata n studyAphelandra crispata n Field Ficus popenoei n Baccaurea species_1 n studyAptandra tubicina Ficus tonduzii n Baccaurea sumatrana y studyApuleia leiocacarpa n Field Ficus trigonata n Barringtonia fusiformis n studyArdisia densapunta n Field Ficus yoponensis n Barringtonia macrostachya n studyArdisia semibulada n Field Garcinia intermedia n Barringtonia pendula n studyAspidosperma blancimpreso n Field Garcinia madruno n Beilschmiedia dictyoneura n studyAspidosperma megalocarpum n Field Genipa americana n Beilschmiedia kunstleri n studyAspidosperma rigidum Geonoma interrupta n Beilschmiedia lucidula n studyAspidosperma spruceanum Guapira standleyana n Beilschmiedia madang n studyAstrocaryum chambira n Guarea fuzzy n Beilschmiedia palembanica n studyAstrocaryum murumuru n Guarea grandifolia n Beilschmiedia species_1 n studyAstronium graveolens n Field Guarea guidonia n Beilschmiedia species_2 n studyAttalea maripa n Guatteria dumetorum n study Bhesa paniculata n studyBactris corossilla n Guazuma ulmifolia n study Blumeodendron calophyllum y studyBactris marajas_sp.juruensis n Guettarda foliacea n Blumeodendron subrotundifolium n studyBactris marajas_sp.maraja n Gustavia superba y study Blumeodendron tokbrai n studyBactris simplicifrons n Hamelia axillaris n Bouea macrophylla n studyBanara nitida Hamelia patens y keeler Bouea oppositifolia n studyBatocarpus amazonica n Field Hampea appendiculata y study Brackenridgea hookeri n studyBatocarpus costaricensis n Field Hasseltia floribunda y study Bridelia pustulata n studyBatocarpus orinocensis n Field Heisteria acuminata n Buchanania sessilifolia n studyBauhinia brachycalyx n Field Heisteria concinna n study Callicarpa maingayi n studyBauhinia lisagroovy Herrania purpurea n study Calophyllum depressinervosum n studyBeilschmiedia pendula Hieronyma alchorneoides n Calophyllum dioscurii n studyBellucia pentamera n Field Hirtella americana y study Calophyllum macrocarpum n studyBertiera guianensis n Field Hirtella triandra y study Calophyllum rupicola n studyBesleria quadrangulata n Field Hura crepitans y study Calophyllum soulattri n studyBesleria stricta n Field Hybanthus prunifolius n study Calophyllum tetrapterum n studyBlakea puberula Inga acuminata y keeler Calophyllum wallichianum n studyBlakea rosea n Field Inga cocleensis y keeler Calophyllum wallichianum_var.incrassatum n studyBorojoa axiglab n Field Inga goldmanii y keeler Campnosperma auriculata n studyBorojoa claviflora n Field Inga laurina y keeler Canarium apertum n studyBrosimum acutifolium n Field Inga marginata y study Canarium littorale_var.littorale n studyBrosimum guianense n Field Inga mucuna y keeler Canarium littorale_var.purpurescens n studyBrosimum lactescens n Field Inga nobilis y keeler Canarium littorale_var.rufum n studyBrosimum potabile Inga oerstediana y keeler Canarium littorale_var.tomentosum n studyBrosimum utile n Field Inga pezizifera y keeler Canarium megalanthum n studyBrownea grandiceps y Keeler Inga punctata y keeler Canarium patentinervium n studyBrownea rosada Inga ruiziana y keeler Canarium pilosum n studyBrownea sp.nov. n Field Inga sapindoides y study Carallia brachiata n studyBrunfelsia chiricaspi Inga spectabilis y study Caryota mitis n studyBuchenavia congesta y Field Inga thibaudiana y keeler Casearia clarkei n studyBuchenavia grandis y Field Inga umbellifera y keeler Casearia species_2 n studyBuchenavia macrophylla Jacaranda copaia y study Cassia nodosa n studyBuchenavia parvifolia n Field Koanophyllon wetmorei n Castanopsis curtisii n studyBuchenavia punctata Lacistema aggregatum n study Castanopsis inermis n studyBunchosia argentea y Field Lacmellea panamensis n study Castanopsis megacarpa n studyBunchosia argentea(granpanosa) y Field Laetia procera n Castanopsis nephelioides n studyBunchosia blanquita Laetia thamnia n study Castanopsis schefferiana n studyBunchosia myrt n Field Lafoensia punicifolia n Celtis rigescens n studyByrsonima juanito Leandra dichotoma n Champereia manillana n studyByrsonima putumayensis n Field Licania hypoleuca n Chasalia longifolia n studyCabralea canjerana n Field Licania platypus y study Chassalia curviflora n studyCalliandra carbonaria n Field Lindackeria laurina n study Cheilosa malayana n studyCalophyllum brasiliense n Field Lonchocarpus heptaphyllus n Chionanthus callophylla n studyCalycophyllum megistocaulum n Field Lozania pittieri n Chionanthus macrocarpa n studyCalyptranthes bipennis n Field Luehea seemannii n study Chionanthus ramiflorus n studyCalyptranthes gigante Lycianthes maxonii n Chionanthus species_1 n studyCalyptranthes grancauli Maclura tinctoria n Chisocheton ceramicus n studyCalyptranthes graneschweil Macrocnemum roseum n study Chisocheton erythrocarpus n studyCalyptranthes loraine n Field Malpighia romeroana y keeler Chisocheton glomeratus n studyCalyptranthes pelopalida n Field Maquira guianensis n study Chisocheton patens n studyCalyptranthes plicata n Field Margaritaria nobilis n study Chisocheton sarawakanus n studyCalyptranthes pseudospeciosa Marila laxiflora n Chisocheton tomentosus n studyCalyptranthes punctote n Field Maytenus schippii n study Chrysophyllum lanceolatum n studyCalyptranthes punteada Miconia affinis n Cinnamomum iners n studyCalyptranthes ruiziana n Field Miconia argentea n study Cinnamomum javanicum n studyCalyptranthes sedosa n Field Miconia dorsiloba n Cinnamomum mollissimum n studyCalyptranthes speciosa n Field Miconia elata n Cinnamomum porrectum n study

17


Campomanesia lineatifolia n Field Miconia hondurensis n Cinnamomum sintoc n studyCapirona decorticans n Field Miconia impetiolaris n study Cleistanthus maingayi n studyCapparis detonsa n Field Miconia nervosa n Cleistanthus malaccensis n studyCapparis osmantha n Field Miconia prasina n study Cleistanthus myrianthus n studyCapparis sola n Field Mosannona garwoodii n Cleistanthus sumatranus n studyCaraipa myricoides_aff. n Field Mouriri myrtilloides n study Clerodendrum deflexum n studyCarica brillante Myrcia gatunensis n study Clerodendrum laevifolium n studyCarica microcarpa n Field Myrospermum frutescens n Clerodendrum nutans n studyCarpotroche longifolia n Field Nectandra cissiflora n Coelostegia griffithii n studyCaryocar glabrum n Field Nectandra fuzzy n Crateva religiosa n studyCaryodaphnopsis chica n Field Nectandra lineata n Cratoxylum arborescens n studyCaryodaphnopsis fosteri Nectandra purpurea n study Cratoxylum cochinchinense n studyCaryodaphnopsis tomentosa n Field Nectandra sp.4_(tiny_leaf) n Cratoxylum formosum n studyCaryodendron orinocense y Field Neea amplifolia n study Cratoxylum maingayi n studyCasearia aculeata n Field Ochroma pyramidale y study Croton argyratus y studyCasearia arborea n Field Ocotea cernua n Croton laevifolius y studyCasearia argut Ocotea oblonga n Crudia curtisii n studyCasearia bracteifera n Field Ocotea puberula n Crypteronia griffithii n studyCasearia javitensis n Field Ocotea whitei n study Cryptocarya ferrea n studyCasearia nigricans n Field Oenocarpus mapora n Cryptocarya griffithiana n studyCasearia pitumba n Field Ormosia amazonica n Cryptocarya infectoria n studyCasearia prunicerob n Field Ormosia coccinea n Cryptocarya kurzii n studyCasearia prunifolia n Field Ormosia macrocalyx n Cryptocarya rugulosa n studyCasearia sp.nov. Ouratea lucens y study Cryptocarya scortechinii n studyCasearia sylvepub n Field Pachira quinata n Ctenolophon parvifolius n studyCasearia sylvestris n Field Pachira sessilis n Cyathocalyx pruniferus n studyCasearia ulmifolia n Field Palicourea guianensis n study Cyathocalyx ramuliflorus n studyCassia cowanii n Field Pavonia dasypetala n Cynometra malaccensis n studyCastilla ulei n Field Pentagonia macrophylla n study Dacryodes costata n studyCathedra acuminata n Field Perebea xanthochyma n Dacryodes incurvata n studyCecropia acer Picramnia latifolia y keeler Dacryodes laxa n studyCecropia engleriana n Field Piper aequale n study Dacryodes longifolia n studyCecropia ficifolia n Field Piper arboreum n Dacryodes nervosa n studyCecropia herthae Piper colonense n Dacryodes puberula n studyCecropia marginalis Piper cordulatum n study Dacryodes rostrata n studyCecropia membranacea Piper imperialis n Dacryodes rubiginosa n studyCecropia putumayonis Piper perlasense n Dacryodes rugosa n studyCecropia sciadophylla n Field Piper reticulatum n Decaspermum fruticosum n studyCedrela fissilis n Field Piper schiedeanum n Dehaasia cuneata n studyCedrela odorata y Keeler Platymiscium pinnatum n Dehaasia incrassata n studyCedrelinga catenaeformis y Field Platypodium elegans n study Dehaasia longipetiolata n studyCeiba pentandra y Keeler Posoqueria latifolia n study Dehaasia polyneura n studyCeltis schippii n Seed/Field Poulsenia armata n study Deplanchea bancana y studyCentropogon loretensis Pourouma bicolor n study Dialium maingayi n studyCestrum megalophyllum n Field Pouteria fossicola n Dialium platysepalum n studyCestrum silvaticum n Field Pouteria reticulata n study Dialium procerum n studyCestrum tomentosum Pouteria stipitata n Dialium wallichii n studyChamaedorea pauciflora n Prioria copaifera n study Dichapetalum gelonioides n studyChamaedorea pinnatifrons n Protium confusum n Dillenia grandifolia n studyCheiloclinium cognatum n Seed/Field Protium costaricense n Dillenia reticulata n studyChimarrhis glabriflora Protium panamense n study Dillenia sumatrana n studyChimarrhis jacob n Field Protium tenuifolium n study Diospyros adenophora y studyChionanthus opipulv n Field Pseudobombax septenatum y study Diospyros andamanica y studyChlorocardium 2subopo Psidium friedrichsthalianum n Diospyros apiculata y studyChomelia comun n Field Psychotria acuminata n Diospyros areolata n studyChrysochlamys fragil Psychotria brachiata n Diospyros argentea y studyChrysochlamys hugo n Field Psychotria chagrensis n Diospyros buxifolia y keelerChrysochlamys membranacea_cf. n Field Psychotria deflexa n study Diospyros cauliflora n studyChrysochlamys tenuifolia Psychotria graciliflora n Diospyros demona n studyChrysophyllum amazonicum n Field Psychotria grandis n Diospyros diepenhorstii y studyChrysophyllum argenteum_ssp.argente n Field Psychotria hoffmannseggiana n Diospyros lancifolia n studyChrysophyllum baeo Psychotria horizontalis n study Diospyros latisepala n studyChrysophyllum cuneifolium n Field Psychotria limonensis n Diospyros maingayi n studyChrysophyllum manaosense n Field Psychotria marginata n Diospyros nutans y studyChrysophyllum minor Psychotria pittieri n Diospyros penangiana n studyChrysophyllum ovale n Field Psychotria psychotriifolia n Diospyros pendula y studyChrysophyllum tremi n Field Psychotria racemosa n Diospyros pyrrhocarpa y studyChrysophyllum venezuelanense Psychotria tenuifolia n Diospyros rufa n studyCinnamomum napoense n Field Pterocarpus belizensis n Diospyros scortechinii y studyCinnamomum oppretic n Field Pterocarpus rohrii n study Diospyros singaporensis n studyCinnamomum peloimpreso Quararibea asterolepis n study Diospyros species_1 n studyCinnamomum triplinerve n Field Quassia amara y study Diospyros sumatrana n studyCitharexylum poeppigii Randia armata n Diospyros venosa n studyClarisia biflora n Field Rauvolfia littoralis n Diospyros wallichii y studyClarisia racemosa n Field Rinorea sylvatica n study Diplospora lasiantha n studyClavija delgada Rosenbergiodendron formosum n Diplospora malaccense n studyClavija procera n Field Sapium broadleaf n Dipterocarpus cornutus n studyClavija weberbaueri n Field Sapium glandulosum n Dipterocarpus costulatus n studyClidemia dimorphica n Field Schefflera morototoni n Dipterocarpus crinitus n studyCoccoloba cordi n Field Schizolobium parahyba n Dipterocarpus kunstleri n studyCoccoloba densifrons n Field Senna dariensis y keeler Dipterocarpus sublamellatus n studyCoccoloba gigante n Field Simarouba amara y study Dracaena brachystachys n studyCoccoloba jill Siparuna guianensis n Dracaena elliptica n studyCoccoloba lancifuzz n Field Siparuna pauciflora n study Dracaena tetrastachys n studyCoccoloba mollis Sloanea terniflora n Dracontomelon dao n studyCoccoloba ninfi n Field Socratea exorrhiza n Drimycarpus luridus n studyCoccoloba papel n Field Solanum arboreum n Drypetes kikir n studyCoccoloba puntoblanco n Field Solanum asperum n Drypetes laevis n studyCoccoloba puntonegro n Field Solanum circinatum n Drypetes longifolia n studyCoccoloba subscab Solanum hayesii n study Drypetes microphylla n studyColubrina arbol y Field Solanum steyermarkii n Drypetes pendula n study

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Compsoneura capitellata n Field Sorocea affinis n study Drypetes polyneura n studyCompsoneura ulei n Field Spachea membranacea y keeler Drypetes rhakodiskos n studyConceveiba rhytidocarpa y Field Spondias mombin n study Drypetes species_1 n studyCordia buladent Spondias radlkoferi n study Durio griffithii n studyCordia chamissoniana n Field Stemmadenia grandiflora n Durio oxleyanus n studyCordia collococca n Field Sterculia apetala n study Durio singaporensis n studyCordia hebeclada n Field Stylogyne turbacensis n Dyera costulata n studyCordia joli Swartzia simplex_var.grandiflora n study Dysoxylum acutangulum n studyCordia kingstonina n Field Swartzia simplex_var.ochnacea n study Dysoxylum alliaceum n studyCordia nodosa n Field Symphonia globulifera n study Dysoxylum carolinae n studyCordia ucayalensis n Field Tabebuia guayacan n Dysoxylum cauliflorum n studyCostus chica Tabebuia rosea y study Dysoxylum costulatum n studyCostus gigante Tabernaemontana arborea n study Dysoxylum flavescens n studyCostus glabra Tachigali versicolor n study Dysoxylum species_1 n studyCostus hairy Talisia nervosa n study Ehretia timorensis n studyCostus mediana Talisia princeps n study Elaeocarpus ferrugineus n studyCostus pablo Terminalia amazonia y study Elaeocarpus griffithii n studyCouepia obovata n Field Terminalia oblonga n Elaeocarpus nitidus n studyCouepia parillo n Field Ternstroemia tepezapote n Elaeocarpus palembanicus n studyCouratari guianensis n Field Tetragastris panamensis n study Elaeocarpus petiolatus n studyCouroupita guianensis n Field Tetrathylacium johansenii n Elaeocarpus rugosus n studyCoussapoa orthoneura n Field Theobroma cacao n Elaeocarpus stipularis n studyCoussarea brevi n Field Thevetia ahouai n study Elateriospermum tapos y studyCoussarea cephaeloide n Field Tocoyena pittieri n Endiandra kingiana n studyCoussarea dulcifolia n Field Trattinnickia aspera n study Endiandra maingayi n studyCoussarea klugii n Field Trema micrantha n study Endocomia canarioides n studyCoussarea multiflora Trichanthera gigantea n Endospermum malaccense y studyCremastosperma cauliflorum n Field Trichilia pallida n Engelhardtia serrata n studyCremastosperma gracilipes n Field Trichilia tuberculata n Enicosanthum fuscum n studyCrepidospermum goudotianum n Field Trichospermum galeottii n study Epiprinus malayanus y studyCrepidospermum rhoifolium n Field Triplaris cumingiana n Erycibe albida n studyCryptocarya aschersoniana_cf. n Field Trophis caucana n Erythroxylum cuneatum n studyCuatresia glabra n Field Trophis racemosa n Eugenia castanea n studyCupania cinerea n Field Turpinia occidentalis n Eugenia ceraina n studyCupania livida n Seed/Field Unonopsis pittieri n Eugenia cerasiformis n studyCupania verde Urera baccifera n study Eugenia chlorantha n studyCyathea lasiosora n Field Verbesina gigantea n Eugenia claviflora n studyCyathea pungens n Field Virola multiflora n Eugenia cumingiana n studyCybianthus perseon n Field Virola sebifera n study Eugenia densiflora n studyCymbopetalum coriaceum n Field Virola surinamensis n study Eugenia duthieana n studyCyphomandra glabra n Field Vismia baccifera n Eugenia dyeriana n studyCyphomandra membra Vismia billbergiana n Eugenia fastigiata n studyCyphomandra pilosa Vismia macrophylla n Eugenia filiformis n studyDacryodes gorda n Field Vochysia ferruginea n study Eugenia flosculifera n studyDacryodes peruviana n Field Xylopia macrantha n study Eugenia glauca_var.pseudoglauca n studyDalbergia nigrescens_cf. n Field Xylosma chlorantha n Eugenia griffithii n studyDendrobangia boliviana n Field Xylosma oligandra n Eugenia inophylla n studyDendropanax arboreus n Field Zanthoxylum acuminatum n Eugenia inophylla_var.barnardi n studyDendropanax caucanus_cf. n Field Zanthoxylum ekmanii n Eugenia koordersiana n studyDendropanax caucanus_cf.(grande) Zanthoxylum panamense y study Eugenia leptostemon n studyDendropanax querceti n Field Zanthoxylum setulosum y study Eugenia napiformis n studyDialium guianense n Field Zuelania guidonia n study Eugenia nigricans n studyDilkea parviflora n Field Eugenia oblongifolia n studyDilkea pasillo Eugenia pachyphylla n studyDiospyros artanthifolia y Keeler Eugenia polita n studyDiospyros crapeifolia Eugenia polyantha n studyDiospyros ekodul Eugenia prainiana n studyDiospyros pseudoxylopia n Field Eugenia pseudocrenulata n studyDiospyros subrotata n Field Eugenia pseudosubtilis n studyDiploon cuspidatum Eugenia pustulata n studyDiplotropis pterochic n Field Eugenia ridleyi n studyDiplotropis purpurea_cf. n Field Eugenia rugosa n studyDiscophora guianensis n Field Eugenia scortechinii n studyDracontium longipes Eugenia species_10 n studyDrypetes amazonica n Field Eugenia species_16 n studyDrypetes papilosa Eugenia species_4 n studyDrypetes variabilis n Field Eugenia species_8 n studyDuguetia cortaten n Field Eugenia species_a n studyDuguetia hadrantha n Field Eugenia species_b n studyDuguetia quitarensis n Field Eugenia species_d n studyDuguetia spixiana n Field Eugenia spicata n studyDuguetia surinamensis Eugenia subdecussata n studyDulacia candida n Field Eugenia syzygioides n studyDuroia eriopila n Field Eugenia tumida n studyDuroia hirsuta n Field Eugenia valdevenosa n studyDussia delgada n Fieldl Eugenia variolosa n studyDussia pelosblancos Eugenia virens n studyDussia tessmannii Euodia glabra n studyEcclinusa angostaestipul n Field Euodia roxburghiana n studyEcclinusa guianensis n Field Euonymus javanicus n studyElaeoluma glabrescens Eurycoma longifolia n studyEndlicheria bracteata n Field Fagraea racemosa n studyEndlicheria burbujas n Field Fahrenheitia pendula y studyEndlicheria canescens Ficus chartacea n studyEndlicheria falsadyso n Field Ficus fistulosa n studyEndlicheria formosa n Field Ficus fulva n studyEndlicheria formosa n Field Ficus glandulifera n studyEndlicheria krukovii Ficus grossularioides n studyEndlicheria metalica n Field Ficus lamponga n studyEndlicheria sericea Ficus obscura y studyEndlicheria sericea_aff. n Field Ficus schwarzii n study

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Endlicheria sp.nov. n Field Ficus scortechinii n studyEndlicheria taubertiana Ficus sinuaata n studyEndlicheria tessmannii n Field Ficus variegata n studyEndlicheria tschudyana n Seed/Field Ficus vasculosa n studyEnterolobium barnebianum y Field Flacourtia rukam n studyEriotheca globosa Gaertnera oblanceolata n studyErisma uncinatum Galearia fulva n studyErythrina amazonica n Field Galearia maingayi n studyErythrochiton margot Ganua mottleyana n studyErythroxylum gracilipes n Field Ganua species_1 n studyErythroxylum macrophyllum_var.ecuadorense n Field Garcinia atroviridis n studyErythroxylum macrophyllum_var.macr n Field Garcinia bancana n studyErythroxylum squamatum Garcinia eugenifolia n studyEschweilera andina n Field Garcinia forbesii n studyEschweilera bracteosa n Field Garcinia griffithii n studyEschweilera coriacea n Field Garcinia malaccensis n studyEschweilera giga n Field Garcinia nervosa n studyEschweilera gigandin n Field Garcinia nigrolineata n studyEschweilera gigantea n Field Garcinia parvifolia n studyEschweilera gigarco n Field Garcinia prainiana n studyEschweilera juruensis Garcinia pyrifera n studyEschweilera rufifolia n Seed/Field Garcinia rostrata n studyEsenbeckia amazonica n Field Garcinia scortechinii n studyEsenbeckia lisa Garcinia species_1 n studyEugenia coffeifolia_aff. Garcinia species_2 n studyEugenia deltocrespis_aff. n Field Garcinia species_4 n studyEugenia egensis Garcinia species_5 n studyEugenia feijoi n Field Gardenia tubifera n studyEugenia florida n Field Gardeniopsis longifolia n studyEugenia graneschweil n Field Gironniera nervosa n studyEugenia granvariable n Field Gironniera parvifolia n studyEugenia lambertiana n Seed Gironniera subaequalis n studyEugenia leo Glochidion hypoleucum n studyEugenia macrocalyx n Field Glochidion obscurum n studyEugenia margot n Field Glochidion sericeum n studyEugenia membranegra n Field Glochidion species_1 n studyEugenia mimus_aff._rojo n Field Glochidion superbum n studyEugenia mimus_aff._verde n Field Glochidion wallichianum n studyEugenia minicomun n Field Gluta malayana n studyEugenia multirimosa n Field Glycosmis chlorosperma n studyEugenia myrobalana_aff. n Field Glycosmis sapindoides n studyEugenia panosadorada n Field Glyptopetalum quadrangulare n studyEugenia puntote Gnetum gnemon n studyEugenia ramamarilla n Field Gomphandra capitulata n studyEugenia schunkei n Field Gomphandra quadrifida n studyEugenia smedcomun n Field Gomphandra species_1 n studyEugenia smedcostacrasa n Field Gomphia serrata n studyEugenia smedcostadorada n Field Goniothalamus macrophyllus n studyEuplassa occidentalis_cf. n Field Goniothalamus tortilipetalus n studyEuterpe precatoria n Gonocaryum gracile n studyExostema maynense n Field Gonystylus maingayi n studyFaramea capillipes n Field Gordonia singaporiana n studyFaramea crassa n Field Grewia antidesmaefolia n studyFaramea glandulosa n Field Grewia blattifolia n studyFaramea multiflora n Field Grewia fibrocarpa n studyFaramea quinqueflora n Field Grewia laurifolia n studyFaramea torquata n Field Grewia miqueliana n studyFaramea uncinata n Field Guioa species_1 n studyFaramea vaina Gymnacranthera eugeniifolia n studyFicus bajio n Gymnacranthera forbesii n studyFicus brevibracteata n Gynotroches axillaris n studyFicus casapi n Harmandia kunstleri n studyFicus gomelleira n Harpullia cupanioides n studyFicus maxima n Hedyotis species n studyFicus minimax n Helicia attenuata n studyFicus nymphaeifolia_cf. n Field Heliciopsis velutina n studyFicus oveja n Heritiera elata n studyFicus perez-arbelaezii n Heritiera javanica n studyFicus piresiana n Heritiera simplicifolia n studyFicus tonduzii n Homalium caryophyllaceum y studyFicus trigona n Homalium dictyoneurum y studyFicus trigonata_cf. n Homalium longifolium y studyFicus trigonhirsuta n Hopea dryobalanoides n studyFicus ursina n Hopea mengarawan y studyFroesia diffusa n Field Hopea sangal n studyGarcinia brasiliensis n Field Horsfieldia brachiata n studyGarcinia macrophylla Horsfieldia crassifolia n studyGarcinia madruno n Field Horsfieldia flocculosa n studyGen.nov. subopo n Field Horsfieldia fulva n studyGenipa americana Horsfieldia polyspherula n studyGeonoma aspidifolia_cf. n Horsfieldia polyspherula_var.sumatrana n studyGeonoma maxima n Horsfieldia punctatifolia n studyGeonoma stricta_var.piscicauda n Horsfieldia sucosa n studyGeonoma stricta_var.stricta n Horsfieldia superba n studyGeonoma triglochin n Horsfieldia tomentosa n studyGloeospermum equatoriense n Field Horsfieldia wallichii n studyGloeospermum longifolium n Field Hunteria zeylanica n studyGloeospermum sphaerocarpum_cf. Hypobathrium venulosum n studyGlycydendron amazonicum y Field Hypobathrum racemosum n studyGordonia fruticosa Iguanura wallichiana n studyGrias neuberthii n Field Ilex macrophylla n study

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Guapira clasica n Field Ilex species_1 n studyGuapira granclasica Intsia palembanica n studyGuarea carinata n Field Irvingia malayana n studyGuarea ecuadoriensis Ixonanthes icosandra y studyGuarea falsakunth n Field Ixonanthes reticulata n studyGuarea fistulosa n Field Ixora concinna n studyGuarea gigakunth n Field Ixora congesta n studyGuarea glabra y Keeler Ixora grandifolia_var.grandifolia n studyGuarea gomma n Field Ixora grandifolia_var.lanceolata n studyGuarea grandifolia n Seed/Field Ixora kingstoni n studyGuarea guentheri n Seed Ixora lobbii n studyGuarea guenthfuzzy n Field Ixora nigricans n studyGuarea guidonia Ixora pendula n studyGuarea kunthiana n Field Jackiopsis ornata n studyGuarea macrophylla_2 n Field Kibara coriacea n studyGuarea macrophylla_3 Kibatalia maingayi n studyGuarea macrophylla_ssp._pac n Field Kingstonia nervosa n studyGuarea patricio Knema conferta n studyGuarea pterorhachis n Field Knema curtisii n studyGuarea pubescens n Field Knema furfuracea n studyGuarea purusana n Field Knema hookeriana n studyGuarea silvatica n Field Knema intermedia n studyGuatteria asplundiana_cf. n Field Knema kunstleri n studyGuatteria brevicuspis_cf. n Field Knema laurina n studyGuatteria citriodora n Field Knema malayana n studyGuatteria gigante n Field Knema patentinervia n studyGuatteria glaberrima n Field Knema pseudolaurina n studyGuatteria gransmoothie n Field Knema scortechinii n studyGuatteria multivenia n Field Knema stenophylla n studyGuatteria planerdorita n Field Knema sumatrana n studyGuatteria punctomarron n Field Koilodepas longifolium y studyGuatteria recurvisepala n Field Kokoona reflexa n studyGuettarda acreana n Field Koompassia malaccensis n studyGustavia hexapetala n Field Lansium domesticum n studyGustavia longifolia n Field Lasianthus lowianus n studyHasseltia floribunda y Keeler Lasianthus species_1 n studyHasseltia hasseltomen Lepisanthes fruticosa n studyHeisteria acuminata n Field Lepisanthes senegalensis n studyHeisteria flacarco n Field Lepisanthes tetraphylla n studyHeisteria grande n Field Lepisanthes tetraphylla_var.hirta n studyHeisteria multiglan Leptonychia glabra n studyHeisteria nitida n Field Licania splendens y studyHelicostylis tomentosa n Field Lindera oxyphylla n studyHerrania cuatrecasana n Field Lithocarpus conocarpa n studyHerrania nitida n Field Lithocarpus curtisii n studyHieronyma alchorneoides_var.sti y Keeler Lithocarpus cyclophorus n studyHieronyma oblonga n Field Lithocarpus ewyckii n studyHimatanthus sucuuba n Field Lithocarpus lucida n studyHirtella excelsa y Field Lithocarpus rassa n studyHirtella racemosa_var.hexandra y Keeler Lithocarpus wallichiana n studyHuertea glandulosa y? Field Lithocarpus wrayi n studyHymenaea oblongifolia n Field Litsea castanea n studyHymenolobium stipsericea Litsea costalis n studyHyospathe elegans n Litsea elliptica n studyInga 3crasa y Herb Litsea erectinervia n studyInga 3oscura y Herb Litsea ferruginea n studyInga 4alitarco y Herb Litsea firma n studyInga 6cuadra y Herb Litsea grandis n studyInga acreana y Herb Litsea machilifolia n studyInga alata y Herb Litsea magnifica n studyInga alba y Herb Litsea nidularis n studyInga auristellae y Herb Litsea resinosa n studyInga bourgonii y Herb Litsea tomentosa n studyInga brachyrhachis y Herb Litsea umbellata_var.fuscotomento n studyInga capitata y Herb/Keeler Litsea wrayi n studyInga cayennensis y Herb Lophopetalum floribundum n studyInga chartacea y Herb Macaranga conifera y studyInga ciliata_ssp.subcapita y Herb Macaranga gigantea y studyInga cinnamomea y Herb Macaranga hosei y studyInga cordatoalata y Herb Macaranga hypoleuca y keelerInga falsacre y Herb Macaranga lowii y studyInga heterophylla y Herb/Keeler Macaranga recurvata y studyInga leiocalycina y Herb Madhuca laurifolia n studyInga marginata y Herb/Keeler Madhuca malaccensis n studyInga microcoma y Herb Mallotus griffithianus n studyInga multinervis y Herb Mallotus leucodermis y studyInga nobilis y Herb/Keeler Mallotus penangensis y studyInga oerstediana y Herb/Keeler Mangifera foetida n studyInga poeppigiana y Herb Mangifera gracilipes n studyInga punctata y Herb/Keeler Mangifera griffithii n studyInga ruiziana y Herb/Keeler Mangifera indica n studyInga rusbyi y Herb Mangifera lagenifera n studyInga sapindoides y Herb/Keeler Mangifera macrocarpa n studyInga sarayacuensis y Herb Mangifera magnifica n studyInga sertulifera_ssp.lepto y Herb/Keeler Mangifera quadrifida n studyInga spectabilis y Herb Mangifera quadrifida_var.longipetiolata n studyInga stellaeglabra y Herb Mangifera rufocostata n studyInga stipulacea y Herb Mangifera species_1 n studyInga striata y Herb Mangifera superba n studyInga tenuistipula y Herb Mangifera swintonioides n studyInga thibaudiana y Herb Mastixia pentandra n study

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Inga thibaudiana_ssp.peltadenia y Herb Mastixia trichotoma n studyInga tocacheana y Herb Medusanthera gracilis n studyInga umbellifera y Herb/Keeler Melanochyla angustifolia n studyInga umbratica y Herb Melanochyla auriculata n studyInga velutina y Herb Melanochyla caesia n studyInga vismiifolia y Herb Melanochyla fulvinervia n studyInga yacoana y Herb Melanochyla species_1 n studyIriartea deltoidea n Melanochyla tomentosa n studyIryanthera grandis n Field Melastoma malabathricum n studyIryanthera hostmanniana n Field Melientha suavis n studyIryanthera hostmannii Memecylon amplexicaule n studyIryanthera juruensis n Field Memecylon cantleyi n studyIryanthera paraensis n Field Memecylon dichotomum n studyIxora acuminatissima n Field Memecylon excelsum n studyIxora killipii n Field Memecylon lilacinum n studyIxora panurensis n Field Memecylon megacarpum n studyJacaranda copaia y Keeler Memecylon minutiflorum n studyJacaranda glabra n Field Memecylon oleifolium n studyJacaratia digitata y Field Memecylon oligoneurum n studyJusticia sanchezioides n Field Memecylon paniculatum n studyKlarobelia napoensis Memecylon pubescens n studyKotchubaea semisericea Memecylon wallichii n studyLacistema aggregatum n Field Mesua cornerii n studyLacistema med n Field Mesua ferrea n studyLacmellea lactescens n Field Mesua grandis n studyLacmellea oblongata n Field Mesua kunstleri n studyLacunaria crenada n Field Mesua lepidota n studyLacunaria jenmanii n Field Mesua racemosa n studyLaetia procera n Field Metadina trichotoma n studyLaxoplumeria tessmannii Mezzettia leptopoda n studyLeandra aristigera Microdesmis caseariifolia n studyLeandra blanca n Field Micromelum minutum n studyLecointea peruviana n Field Microtropis valida n studyLeonia crassa n Field Miliusa longipes n studyLeonia glycycarpa_var.glycyc n Field Millettia atropurpurea n studyLeonia glycycarpa_var.racemo n Field Millettia species_1 n studyLeonia occidentalis Mischocarpus pentapetalus n studyLicania arborea n Field Mitrephora maingayi n studyLicania aubreuvillei_cf. Monocarpia marginalis n studyLicania caudata y Field Mussaendopsis beccariana n studyLicania harlingii n Field Myristica cinnamomea n studyLicania hipofuzzy n Field Myristica maingayi n studyLicania longipedicellata n Field Myristica malaccensis n studyLicania longistyla y Field Myristica maxima n studyLicania macrocarpa n Field Nauclea officinalis n studyLicania nervifina y Field Neesia synandra n studyLicania reticulata n Field Neobalanocarpus heimii y studyLicania silvae_cf. n Field Neolamarckia cadamba n studyLicania urseolaris n Field Neolitsea zeylanica n studyLicania zigzag n Field Neoscortechinia kingii n studyLicaria brillacuspi Neoscortechinia nicobarica y studyLicaria cannella n Field Neoscortechinia sumatrensis y studyLicaria guianensis Neouvaria foetida n studyLindackeria paludosa n Field Nephelium costatum n studyLonchocarpus seorsus_cf. n Field Nephelium eriopetalum n studyLoreya spruceana Nephelium hamulatum n studyLozania klugii n Field Nephelium maingayi n studyLozania mediana n Field Nephelium ophiodes n studyLunania parviflora Nephelium pallens n studyMabea comun n Field Nothaphoebe umbelliflora n studyMabea superbrondu n Field Ochanostachys amentacea n studyMachaerium aristulatum n Field Oncodostigma monosperma n studyMachaerium finiparalel n Field Oncosperma horridum n studyMacrocnemum roseum Orania sylvicola n studyMacrolobium angustifolium n Field Ormosia penangensis n studyMacrolobium colombianum_cf. Ormosia venosa n studyMacrolobium sp.nov. n Field Osmelia maingayi n studyMacrolobium stenocladum n Field Palaquium clarkeanum n studyMaquira calophylla n Field Palaquium gutta n studyMaquira guianensis n Field Palaquium hexandrum n studyMargaritaria nobilis n Field Palaquium maingayi n studyMargaritaria sp.nov. n Field Palaquium obovatum n studyMarila pluricostata_cf. n Field Palaquium stellatum n studyMarila puntorojo n Field Pandanus monotheca n studyMarmaroxylon basijugum y Field Pandanus yvanii n studyMatayba ocho n Field Paranephelium xestophyllum n studyMatisia bracteolosa n Field Parartocarpus bracteata n studyMatisia cordata n Field Parashorea densiflora n studyMatisia longiflora n Field Parastemon urophyllus y studyMatisia malacocalyx n Field Parinari costata y studyMatisia obliquifolia n Field Parinari elmeri y studyMatisia oblongifolia n Field Parinari oblongifolia y keelerMayna anelio n Field Parishia insignis n studyMayna odorata n Field Parishia paucijuga n studyMaytenus ala n Field Parkia speciosa y studyMaytenus ebenifolia_cf. n Field Paropsia vareciformis y studyMaytenus macrocarpa_s.l. n Field Pavetta graciliflora n studyMeliicoccus novagranatensis n Field Pavetta species_1 n studyMeliosma doly n Field Payena lucida n studyMeliosma vasquezii n Field Payena maingayi n studyMemora cladotricha n Field Pentace strychnoidea n study

22


Mezilaurus extendido Pentace triptera n studyMezilaurus triunca n Field Pentaspadon motleyi n studyMiconia abbreviata n Field Pentaspadon velutinus n studyMiconia acutipetala n Field Pertusadina eurhyncha n studyMiconia ampla Phaeanthus crassipetalus n studyMiconia atenunodu Phaeanthus ophthalmicus n studyMiconia aurea_cf. n Field Pholidocarpus macrocarpus n studyMiconia bubalina n Field Phyllanthus emblica n studyMiconia cazaletii Pimelodendron griffithianum y studyMiconia centrodesma Pimelodendron macrocarpum n studyMiconia chocofres n Field Pinanga malaiana n studyMiconia corine n Field Pinanga riparia n studyMiconia crasarb Pithecellobium splendens y studyMiconia decurrens n Field Planchonella maingayi n studyMiconia elata n Field Planchonia grandis n studyMiconia elatita n Seed/Field Platea latifolia n studyMiconia falsarug Platea species_1 n studyMiconia fosteri n Field Podocarpus motleyi n studyMiconia grancordata n Field Polyalthia cinnamomea n studyMiconia grandifolia n Field Polyalthia clavigera n studyMiconia juanito Polyalthia glauca n studyMiconia karina n Field Polyalthia hypoleuca n studyMiconia klugii n Field Polyalthia jenkinsii n studyMiconia lamprophylla n Field Polyalthia lateriflora n studyMiconia lugonis Polyalthia obliqua n studyMiconia medglauca n Field Polyalthia rumphii n studyMiconia multispicata n Field Polyalthia sclerophylla n studyMiconia napoana n Field Polyalthia stenopetala n studyMiconia nerviblanco n Field Polyalthia sumatrana n studyMiconia nervosa n Field Polyosma laete-virens n studyMiconia pablo Pometia pinnata_var.alnifolia y studyMiconia peqdorada n Field Popowia pisocarpa n studyMiconia pilgeriana n Field Popowia tomentosa n studyMiconia poeppigii n Field Porterandia anisophylla n studyMiconia prasina n Field Pouteria malaccensis n studyMiconia pterocaulon n Seed Prainea limpato n studyMiconia punctata Prismatomeris glabra n studyMiconia purpono n Field Prismatomeris species_1 n studyMiconia roselegante Prunus arborea y studyMiconia rugosa Prunus arborea_var.stipulaceum y studyMiconia sachapurp n Field Prunus grisea y studyMiconia schunkei n Field Pseudoeugenia singaporensis n studyMiconia smaragdina n Field Pseuduvaria macrophylla n studyMiconia tipica n Field Psychotria griffithii n studyMiconia tomentosa n Field Psychotria rostrata n studyMiconia trinervia n Field Psydrax maingayi n studyMiconia triplinervis n Field Psydrax species_10 n studyMiconia zubenetana n Field Psydrax species_8 n studyMicropholis brochidodroma n Field Pteleocarpa lamponga n studyMicropholis dorada n Field Pternandra coerulescens n studyMicropholis egensis n Field Pternandra echinata n studyMicropholis guyanensis_ssp.duckea n Field Pterocymbium tubulatum n studyMicropholis venulosa n Field Ptychopyxis caput-medusae n studyMinquartia guianensis n Field Ptychopyxis costata y studyMollia gracilis Pyrenaria acuminata n studyMollinedia killipii n Field Quercus argentata n studyMollinedia rojimpreso Quercus gemelliflora n studyMollinedia spinilarga n? Field Radermachera pinnata y studyMollinedia tomentosa n Field Reinwardtiodendron cinereum n studyMollinedia tridentata n? Field Rhodamnia cinerea n studyMoronobea pablo Rinorea anguifera n studyMouriri acutiflora_cf. Rinorea horneri n studyMouriri grancoala n Field Rinorea sclerocarpa n studyMouriri grandiflora n Field Rothmannia macrophylla n studyMouriri intermedia n Field Ryparosa acuminata n studyMouriri myrtilloides n Field Ryparosa fasciculata n studyMyrcia blancanueva n Field Ryparosa kunstleri n studyMyrcia nitida n Field Sandoricum beccarianum n studyMyrcia pielizard n Field Sandoricum koetjape n studyMyrcia platatomen n Field Santiria apiculata n studyMyrcia pseudonitida n Field Santiria conferta n studyMyrcia splendens n Field Santiria griffithii n studyMyrcia vertipub n Field Santiria laevigata n studyMyrciaria amazonica n Field Santiria oblongifolia n studyMyrciaria floribunda n Field Santiria rubiginosa n studyMyrciaria intermed n Field Santiria tomentosa n studyMyroxylon balsamum n Field Sapium baccatum y studyNaucleopsis glabra n Field Sapium discolor y studyNaucleopsis imitans_cf. n Field Saprosma scortechinii n studyNaucleopsis krukovii n Field Saraca declinata y studyNaucleopsis krukovii_cf. n Seed Saraca thaipingensis y studyNaucleopsis ulei n Field Sarcotheca griffithii n studyNectandra largibrachi n Field Sarcotheca monophylla n studyNectandra lineata n Field Sauropus androgynus n studyNectandra membranacea n Field Scaphium linearicarpum n studyNectandra microcarpa n Field Scaphium macropodum n studyNectandra oppositifolia n Seed/Field Scaphocalyx spathacea n studyNectandra skinny2 Schoutenia accrescens n studyNectandra skinnyret Scleropyrum wallichianum n studyNectandra viburnoides n Field Scolopia spinosa y studyNectandra yarinensis_cf. n Field Semecarpus curtisii n study

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Neea altomini Semecarpus rufovelutinus n studyNeea angostintersec n Field Shorea acuminata y studyNeea aniboid n Field Shorea bracteolata y studyNeea bajio n Field Shorea dasyphylla y studyNeea claudia n Field Shorea guiso y studyNeea comun n Field Shorea hopeifolia n studyNeea daniela Shorea lepidota y studyNeea fuzzy n Field Shorea leprosula y studyNeea garci n Field Shorea macroptera y studyNeea gigante n Field Shorea maxwelliana n studyNeea granredonda n Field Shorea multiflora n studyNeea micro n Seed Shorea ochrophloia y studyNeea mini n Field Shorea ovalis y studyNeea paty n Field Shorea parvifolia n studyNeea popular n Field Shorea pauciflora y studyNeea supercrasa n Field Sindora coriacea n studyNeea tela n Field Sindora echinocalyx n studyNeea verdeclara n Field Sindora velutina n studyNeea verdeseca n Field Sindora wallichii n studyNeosprucea grandiflora y? Field Sloanea javanica n studyOchroma pyramidale n Field Stelechocarpus cauliflorus n studyOcotea alamembra n Field Stemonurus malaccensis n studyOcotea argyrophylla n Field Stemonurus umbellatus n studyOcotea bayelshmi n Field Sterculia coccinea n studyOcotea bofo_cf. n Field Sterculia cordata n studyOcotea cernua n Field Sterculia hispidissima n studyOcotea cujumari_cf. Sterculia macrophylla n studyOcotea floribunda n Field Sterculia parviflora n studyOcotea javitensis n Field Sterculia rubiginosa n studyOcotea laurita n Field Sterculia species_1 n studyOcotea leucoxylon Streblus elongatus n studyOcotea longifolia n Field Strombosia javanica n studyOcotea luis n Field Strombosia maingayi n studyOcotea nervijens n Field Styrax benzoin n studyOcotea oblonga n Field Suregada multiflora n studyOcotea scalariformis Symplocos cerasifolia n studyOcotea tessmannii_cf. n Field Symplocos cochinchinensis n studyOcotea ucayalensis Symplocos crassipes n studyOenocarpus bataua n Symplocos ophirensis n studyOenocarpus mapora n Symplocos rubiginosa n studyOphiocaryon heterophyllum n Field Tabernaemontana corymbosa n studyOrmosia amazonica Tabernaemontana malaccensis n studyOrmosia elata n Field Talauma candollii n studyOrmosia paraensis n Field Tarenna costata n studyOssaea boliviensis Tarenna maingayi n studyOtoba glycycarpa n Field Tarenna mollis n studyOuratea flaquita n Field Tarenna species_11 n studyOxandra mediocris n Field Tarenna species_8 n studyOxandra riedeliana_aff. n Field Teijsmanniodendron coriaceum n studyPachira insignis n Field Teijsmanniodendron simplicifolium n studyPachira punga-schunkei n Field Terminalia bellirica n studyPalicourea grandiflora n Field Terminalia citrina y studyPalicourea guianensis n Field Terminalia phellocarpa y studyPalicourea lasiantha Terminalia subspathulata n studyPalicourea nigricans n Field Ternstroemia corneri n studyParadrypetes subintegrifolia n Field Tetrardisia porosa n studyParinari klugii n Field Thottea grandiflora n studyParkia balslevii Timonius species_1 n studyParkia multijuga y Timonius wallichianus n studyParkia nitida y Keeler Trema tomentosa n studyParkia velutina y Trigonachras acuta n studyPatinoa paraensis n Field Trigoniastrum hypoleucum n studyPaullinia xestophylla n Field Trigonochras species_1 n studyPausandra trianae y Field Trigonopleura malayana n studyPentagonia parvifolia n Field Trigonostemon laevigatus n studyPentagonia williamsii_cf. n Field Trigonostemon longifolius n studyPentagonia wurdackii n Field Trigonostemon malaccanus n studyPentaplaris huaoranica Triomma malaccensis n studyPera bicolor Trivalvaria macrophylla n studyPera duguet n Field Trivalvaria nervosa n studyPerebea angustifolia n Field Trivalvaria pumila n studyPerebea guianensis_cf. n Field Turpinia ovalifolia y studyPerebea guianensis_ssp._acan Unknown sp. n studyPerebea mollis n Field Urophyllum glabrum n studyPerebea tessmannii n Field Urophyllum hirsutum n studyPerebea xanthochyma n Field Vatica bella y studyPersea areolatocostae n Field Vatica maingayi n studyPersea persemed n Field Vatica pauciflora y studyPersea pseudofasciculata n Field Vernonia arborea n studyPhyllanthus attenuatus n Field Vitex pinnata n studyPhyllanthus micro Vitex quinata n studyPhytelephas tenuicaulis n Walsura chrysogyne n studyPicramnia juniniana n Field Walsura pinnata n studyPicramnia magnifolia n Field Xanthophyllum affine y studyPicramnia mini n Field Xanthophyllum amoenum n studyPicramnia pubibul n Field Xanthophyllum chartaceum n studyPicramnia pubirecta Xanthophyllum ellipticum n studyPicramnia sellowii_ssp.sprucean n Field Xanthophyllum eurhynchum n studyPiper aequale Xanthophyllum griffithii n studyPiper albertsmithii Xanthophyllum rufum n studyPiper arboreum n Field Xanthophyllum scortechinii n study

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Piper augustum n Field Xanthophyllum stipitatum n studyPiper bellidi n Field Xanthophyllum wrayi n studyPiper bellidifolium n Field Xerospermum species_1 n studyPiper bulada n Field Xerospermum noronhianum y studyPiper cordipub n Field Xylopia caudata n studyPiper crassinervium n Field Xylopia elliptica n studyPiper darkcrasa n Field Xylopia ferruginea_var.ferruginea n studyPiper falsafuzzy n Field Xylopia ferruginea_var.oxyantha n studyPiper fuzicort n Field Xylopia fusca n studyPiper granmini Xylopia magna n studyPiper macrophyllum Xylopia malayana n studyPiper maranyonense n FieldPiper minibroqui n FieldPiper minicordPiper nerviPiper obchic n FieldPiper obglab n FieldPiper obnervi n FieldPiper obtomen n FieldPiper obvil n FieldPiper peltatumPiper pubescens n FieldPiper redondaPiper renato n FieldPiper reticulatum n FieldPiper scab n FieldPiper sesivil n FieldPiptadenia pterocladaPlatymiscium pinnatumPlatymiscium stipularePleurothyrium bifidum n FieldPleurothyrium cinereumPleurothyrium cuneifolium n FieldPleurothyrium glabrifolium n FieldPleurothyrium insigne n FieldPleurothyrium willamsii_cf. n FieldPlinia caulimpreso n Seed/FieldPlinia cortezablanca n FieldPlinia pseudomouriri n FieldPlinia unop n FieldPosoqueria latifolia n FieldPosoqueria longiflora n FieldPotalia resinifera n FieldPoulsenia armata n FieldPourouma bicolor n FieldPourouma deeplob n FieldPourouma guianensis_ssp.guiane n FieldPourouma medioarco n FieldPourouma minor y FieldPourouma napoensisPourouma persecta n FieldPourouma tomentosa n FieldPouteria anchalisa n FieldPouteria angostaloopy n FieldPouteria baehniana n FieldPouteria bilocularis n FieldPouteria caimitoPouteria cuspidata_ssp.duraPouteria cuspidata_ssp.robustaPouteria doradagrande n FieldPouteria durlandii n FieldPouteria durlandii_ssp.pubicar n FieldPouteria glomerata n FieldPouteria gracilis n FieldPouteria granopaca n FieldPouteria guianensis n FieldPouteria hispida_cf. n FieldPouteria krukovii n FieldPouteria largamembra n FieldPouteria multiflora n FieldPouteria nudipetala n FieldPouteria peciolote n FieldPouteria platyphylla n FieldPouteria procera n FieldPouteria redondita n FieldPouteria reticulata n FieldPouteria rostrata n Seed/FieldPouteria smedobov n FieldPouteria torta_ssp.glabra n FieldPouteria torta_ssp.tuberculata n FieldPouteria tortachica n FieldPouteria tortachicordenPouteria trilocularis n FieldPouteria vernicosa n FieldPradosia atroviolacea n FieldPrestoea schultzeana nProtium amazonicum n Seed/FieldProtium aracouchini n FieldProtium brillanodu n FieldProtium glabrescens n FieldProtium grannodu n Field

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Protium guianense n FieldProtium nodulosum n FieldProtium sagotianum n FieldProtium trifoliolatumProtium unifoliolatumPrunus debilis y FieldPseudolmedia laevigata n FieldPseudolmedia laevis n FieldPseudolmedia macrophylla n FieldPseudolmedia rigida_ssp.eggersii n FieldPseudomalmea diclina n FieldPseudopiptadenia suaveolens y KeelerPsychotria borjensis n FieldPsychotria brachybotryaPsychotria caerulea n FieldPsychotria deflexa n FieldPsychotria dracula n FieldPsychotria huampamiensis n FieldPsychotria membradomat n FieldPsychotria officinalisPsychotria ondulada n FieldPsychotria ostreophora n FieldPsychotria poeppigiana n FieldPsychotria remota n FieldPsychotria robin n SeedPsychotria stenostachya n FieldPsychotria viridis n FieldPterocarpus rhorii_cf. n FieldQualea paraensis y FieldQuararibea amazonica n FieldQuararibea bilobata_cf. n FieldQuararibea wittii n FieldQuiina amazonicaQuiina florida n FieldQuiina grandifolia_cf. n FieldQuiina macrophylla_cf. n FieldQuiina mediana n FieldRandia bigfuzzy n FieldRandia gorky n FieldRandia manolo n FieldRauvolfia praecoxRhamnidium elaeocarpum n FieldRhodostemonodaphne grandisRhodostemonodaphne juruensis n FieldRhodostemonodaphne kunthiana n FieldRhodostemonodaphne licanoidesRhodostemonodaphne sordidaRicheria racemosa n FieldRinorea apiculata n FieldRinorea lindeniana n FieldRinorea viridifolia n FieldRollinia chrysocarpa n FieldRollinia cuspidata n FieldRollinia dolichopetala n FieldRollinia flacaglabra n FieldRollinia glomerulifera n FieldRollinia palidaRollinia pittieri n FieldRoupala montanaRuagea insignis n FieldRudgea bracteata n FieldRudgea fina n FieldRudgea japurensisRudgea nodincho n FieldRuizodendron ovale n FieldRyania speciosa n FieldRyania speciosa_var._toment n FieldSagotia racemosa n FieldSalacia atenucrasa n FieldSalacia macrantha_cf. n FieldSapium glandulosum_cf. y Field/KeelerSapium largident y FieldSapium redondaSarcaulus brasiliensis n FieldSarcaulus peloscostimp n FieldSarcaulus romolerouxii n FieldSarcaulus vestitus n FieldSchefflera morototoni n FieldSchizolobium parahybaSchoenobiblus perubianus_cf. n FieldSchoepfia lucida n FieldSenna bacillarisSenna macrophylla_var.gigan y KeelerSenna trolliiflora y KeelerSimaba orinocensis n FieldSimaba paraensis n FieldSimaba polyphylla n FieldSimarouba amara y KeelerSimira cordifolia n FieldSimira rubescens_cf. n FieldSiparuna angostadiente n Field

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Siparuna bigll3 n FieldSiparuna cervicornis n FieldSiparuna cuspidata n FieldSiparuna decipiens n FieldSiparuna gesSiparuna macrotepala n FieldSiparuna poeppigii n FieldSiparuna thecaphora(1) n FieldSiparuna thecaphora(2) n FieldSiparuna thecaphora(3) n FieldSloanea fragans n FieldSloanea gigapulvi n FieldSloanea granredonda n FieldSloanea guia n FieldSloanea membraminiSloanea nerviSloanea oak n FieldSloanea obtusidolia n FieldSloanea oppdSloanea pequrva n FieldSloanea polvorojo n FieldSloanea pubescens_cf. n FieldSloanea robusta_cf. n FieldSloanea robustipeq n FieldSloanea rufisessi n FieldSloanea synandra n FieldSmilax nerviprominSocratea exorrhiza nSolanum altissimum n FieldSolanum grandiflorum n FieldSolanum granmini n FieldSolanum lepidotumSolanum leptopodumSolanum malletii n FieldSolanum scabrosaSolanum sessileSolanum silvaticumSorocea muriculata n FieldSorocea pubivena_ssp.hirtella n FieldSorocea pubivena_ssp.oligotricha n FieldSorocea sarcocarpa_cf.Sorocea steinbachii n FieldSpondias mombin n FieldSterculia apeibophylla n FieldSterculia apetalaSterculia colombiana n FieldSterculia frondosa n FieldSterculia tessmannii n FieldStrychnos darienensis_cf.Stryphnodendron porcatumStylogyne caulifloraStylogyne longifolia n FieldStyrax cordatusStyrax guyanensisSwartzia arborescens n FieldSwartzia benthamiana n FieldSwartzia bombycina n FieldSwartzia cardiosperma n FieldSwartzia multijuga n FieldSwartzia simplex n FieldSymphonia globulifera n FieldSymplocos arechea n? FieldTabebuia ochracea y KeelerTabebuia serratifolia y KeelerTabernaemontana pequea n FieldTabernaemontana sananho n FieldTachigali formicarum n FieldTachigali paniculataTachigali paraensisTalauma ovata_cf. n FieldTalauma tyana n FieldTalisia 2-reticTalisia cerasinaTalisia gigapulvi n FieldTalisia pulvinoteTapirira guianensis n FieldTapirira myriantha_cf. n FieldTapirira obtusa n FieldTapura juruana n FieldTapura peruviana n FieldTerminalia amazonia y KeelerTerminalia axilpub n FieldTerminalia ob y FieldTerminalia oblonga n FieldTessmannianthus heterostemon n FieldTetragastris panamensis n FieldTetrathylacium macrophyllum n FieldTetrorchidium macrophyllum y FieldTheobroma speciosum n FieldTheobroma subincanum n FieldThyrsodium paraense_cf.

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Tococa guianensisTocoyena burnham n FieldTovomita alargada n FieldTovomita arbolTovomita grancrasa n FieldTovomita grande n FieldTovomita tyana n FieldToxiphon trifoliolatumTrattinnickia glaziovii_cf.Trattinnickia lancifoliaTrattinnickia lawrencei_ssp._bolivTrema micrantha n FieldTrichilia adolfiiTrichilia cip y KeelerTrichilia densapuntaTrichilia elsae n FieldTrichilia laxipaniculataTrichilia maynasiana n FieldTrichilia micrantha n FieldTrichilia obovata n FieldTrichilia pallida n FieldTrichilia pleeana_cf. n FieldTrichilia poeppigii n FieldTrichilia quadrijuga n FieldTrichilia rubra n FieldTrichilia septentrionalis n FieldTrichilia solitudinis n FieldTrigynaea triplinervia n FieldTriplaris dugandiiTrymatococcus amazonicus n FieldTurpinia occidentalis n FieldUnonopsis floribunda n FieldUnonopsis veneficiorum n FieldUrera baccifera n FieldVantanea guianensis n FieldVirola dixonii n FieldVirola duckei n FieldVirola elongata n FieldVirola flexuosa n FieldVirola fuzzyVirola microfuzzy n FieldVirola mollis n FieldVirola multinervia n FieldVirola obovata n FieldVirola pavo n FieldVirola surinamensisVirola theiodoraVismia baccifera n FieldVismia bosque nVismia floribunda n FieldVismia macrophyllaVismia spruceiVitex schunkei n FieldVitex triflora n FieldVochysia braceliniae n FieldVourana anomala n FieldWarszewiczia cordata n FieldWettinia maynensis nWittmackanthus stanleyanus n FieldXylopia aromatica_cf. n FieldXylopia cuspidata n FieldXylopia hienaXylosma tessmannii_cf.Zanthoxylum glanredonda n FieldZanthoxylum margland n FieldZanthoxylum nerviZanthoxylum pendienteZanthoxylum perpZanthoxylum setulosum y KeelerZanthoxylum sprucei_aff. n FieldZanthoxylum suaveZizyphus cinnamomumZygia heteroneura y FieldZygia mediana y Fieldzz zz

28

incidence of extra-floral nectaries and their effect on the growth

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