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Geographic variation and environmental correlates of functional trait distributions in palms (Arecaceae) across the New World Bastian Göldel 1 , W. Daniel Kissling 2 and Jens-Christian Svenning 1 1 Section for Ecoinformatics and Biodiversity, Department of Bioscience, Aarhus University, Ny Munkegade 114, DK-8000 Aarhus C, Denmark 2 Institute for Biodiversity and Ecosystem Dynamics (IBED), University of Amsterdam, P.O. Box 94248, 1090 GE Amsterdam, The Netherlands Correspondence: Bastian Göldel; Department of Bioscience, Aarhus University, Ny Munkegade 114, 8000 Aarhus C, Denmark. E-mail: [email protected] Running title: Functional traits in New World palms (Arecaceae) Word count: Manuscript text (including title, abstract, keywords, acknowledgements and references, but excluding legends and tables): 9205 ABSTRACT Functional traits play a key role in driving biodiversity effects on ecosystem functioning. Here, we examine the geographic distributions of three key functional traits in New World palms (Arecaceae), an ecologically important plant group, and their relationships with current climate, soil and glacial-interglacial climate change. We combined palm range maps for the New World (n = 541 species) with data on traits (leaf size, stem height and fruit size) representing the Leaf-Height-Seed (LHS) plant strategy scheme of Westoby (1998)to estimate median trait values for palm species assemblages in 110×110-km grid cells. We used the Akaike Information Criterion to identify minimum adequate models and then applied spatial autoregressive models to account for spatial autocorrelation. Seasonality in temperature and precipitation played a major role in explaining geographic variation of all traits. Mean annual temperature and annual precipitation were important for median leaf and fruit size, while glacial-interglacial temperature change and present-day 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32

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Page 1: pure.au.dkpure.au.dk/portal/files/121438275/goeldel_BoJLS_2015oe.pdf.docx  · Web viewGeographic variation and environmental correlates of functional trait distributions in palms

Geographic variation and environmental correlates of functional trait distributions in palms (Arecaceae) across the New World

Bastian Göldel1, W. Daniel Kissling2 and Jens-Christian Svenning1

1Section for Ecoinformatics and Biodiversity, Department of Bioscience, Aarhus University, Ny Munkegade 114, DK-8000 Aarhus C, Denmark2Institute for Biodiversity and Ecosystem Dynamics (IBED), University of Amsterdam, P.O. Box 94248, 1090 GE Amsterdam, The Netherlands

Correspondence: Bastian Göldel; Department of Bioscience, Aarhus University, Ny Munkegade 114, 8000 Aarhus C, Denmark. E-mail: [email protected]

Running title: Functional traits in New World palms (Arecaceae)

Word count: Manuscript text (including title, abstract, keywords, acknowledgements and references, but excluding legends and tables): 9205

ABSTRACT

Functional traits play a key role in driving biodiversity effects on ecosystem functioning. Here, we examine

the geographic distributions of three key functional traits in New World palms (Arecaceae), an ecologically

important plant group, and their relationships with current climate, soil and glacial-interglacial climate

change. We combined palm range maps for the New World (n = 541 species) with data on traits (leaf size,

stem height and fruit size) —representing the Leaf-Height-Seed (LHS) plant strategy scheme of Westoby

(1998)— to estimate median trait values for palm species assemblages in 110×110-km grid cells. We used

the Akaike Information Criterion to identify minimum adequate models and then applied spatial

autoregressive models to account for spatial autocorrelation. Seasonality in temperature and precipitation

played a major role in explaining geographic variation of all traits. Mean annual temperature and annual

precipitation were important for median leaf and fruit size, while glacial-interglacial temperature change and

present-day precipitation of the driest month were especially important for median fruit size, but also for

median stem height. Our results suggest that both current climate (notably seasonality) and glacial-

interglacial temperature change are important drivers for functional trait distributions of palms across the

New World, with soil playing a minor role.

Keywords

Biogeography, climate change, functional diversity, geographical ecology, Neotropics, palaeoclimate, Palmae,

Quaternary climate oscillations

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INTRODUCTION

Functional traits play a key role in driving biodiversity effects on ecosystem functioning (Mason et al., 2005;

Swenson et al., 2012). They are defined as traits which impact fitness indirectly through effects on growth,

reproduction and survival (Diaz & Cabido, 2001). Furthermore, functional traits can be divided into effect

traits and response traits (Violle et al., 2007). While the former reflect impacts on ecosystem properties and

services, the latter reflect species responses to environmental conditions and changes (Diaz et al., 2013). In

recent years, a few studies have started to explore the relationships between geographic patterns of plant

functional trait distributions and their underlying environmental drivers. Geographic variation of plant

functional traits can be strongly correlated with current environment. For instance, median seed mass of plant

assemblages across Germany correlates with soil pH and soil moisture (Tautenhahn et al., 2008), different

morphological and physiological variables of North American trees (e.g. mean tree height, seed mass)

correlate with climate (e.g. precipitation and temperature) (Swenson & Weiser, 2010), and mean leaf area of

tropical forest trees in Panama and China are mainly related to soil fertility and acidity (Liu et al., 2012).

Maximum tree height across species was shown to be related to mean annual temperature and precipitation

along the Bolivian Andes (Kessler, Böhner & Kluge, 2007), mean leaf size related to mean annual

precipitation across species in southeastern Australia (McDonald et al., 2003) and Amazonia (Malhado et al.,

2009), while abundance weighted community average of seed size was related to soil in tropical forests of

the Guiana Shield (ter Steege and Hammond, 2001) and Amazonia (ter Steege et al., 2006{ter Steege, 2006

#43}). Assemblage means of plant traits (e.g. leaf size) can further change along elevational (Gurevitch,

1988), latitudinal (Hulshof et al., 2013), and soil gradients (Liu et al., 2012).

Palms (Arecaceae) are an important plant family in tropical and subtropical regions, with high species

richness, various growth forms and keystone ecological importance in many areas (Dransfield, Uhl & Royal

Botanic Gardens, 2008; Balslev et al., 2011). The palm family occurs across the warmer parts of the world

and constitutes —with c. 2400 species worldwide (Govaerts & Dransfield, 2005)— a major canopy and

understory element in many tropical and subtropical forests (Gentry, 1988). Palms play an important role in

biogeographic theory and represent a suitable model organism for understanding the drivers of high tropical

biodiversity and its geographic variation (Eiserhardt et al., 2011). Their global distribution and diversity is

strongly linked to temperature and precipitation as well as historical regional drivers (Kissling et al., 2012a),

and palms are considered indicators for warm and humid climates in paleo-ecological reconstructions

(Greenwood & Wing, 1995; Morley, 2000). Here, we focus on New World palms which are diverse and

ecologically important in this region (Dransfield et al., 2008). For the palm family, there is evidence that

species diversity is driven by current and paleoclimatic factors (Blach-Overgaard et al., 2010; Eiserhardt et

al., 2011; Kissling et al., 2012a,b; Blach-Overgaard et al., 2013), but no studies have so far have focused on

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large-scale patterns of functional trait distributions and how they may be constrained by present-day

environment and long-term historical constraints, e.g. glacial-interglacial climate oscillations.

The Leaf-Height-Seed (LHS) plant ecology strategy scheme (Westoby, 1998) suggests that variables

related to leaf size (specific leaf area), stem height and seed size capture the main trait axes of a plant species

response to competition, stress, disturbance and variation in responsiveness to opportunities for rapid growth.

We focus here on maximum leaf size, stem height and fruit size, the first as a proxy for specific leaf area,

because it is commonly used in analyses of leaf traits (e.g. McDonald et al., 2003). Furthermore, fruit size

was used as a proxy for seed size because data availability on palm seed sizes is limited. In the LHS scheme,

the leaf component is regarded to be a representative of the light-catching area, which is responsible for

photosynthetic capacity, hence energy production of the plant, and therefore also directly involved in

competition for light with other competitors (Westoby, 1998). In palms, leaf size might be relevant to tall

species to reach canopy gaps, e.g. in disturbed forests (de Granville, 1992), but on the other hand also for

understory palms because reduced light availability might cause leaves to be shaped towards sizes and

structures that maximize the effectiveness of photosynthesis and the tolerance to increased shading by e.g.

maximizing displayed leaf area and reducing biomass costs of leaf support (Chazdon, 1991). Maximum stem

height represents a plant’s accessibility to light and exposure to heat load, humidity and wind speeds

(Westoby, 1998). For instance, in forests with a dense canopy layer, stem height of palms might be smaller

due to less light availability for growth whereas in disturbed forests with more canopy gaps erect solitary

palms can be more frequent (de Granville, 1992). Seed and fruit size further determine the establishment

success of animal-dispersed plants, because larger seed masses enable seedlings to better survive hazards

such as drought (Westoby, Leishman, Lord, 1996) and because dispersal distances rely on fruit and seed size

as large seeds can only be dispersed by animals large enough to swallow or transport them (de Almeida &

Galetti, 2007; Andreazzi et al., 2012; Galetti et al., 2013).

Several questions concerning functional trait distributions and their environmental correlates remain

unclear, and few studies have focused on functional traits across broad macroecologial scales (Ordonez et

al., 2009; Peppe et al., 2011; Moles et al., 2014). Previous studies have detected mean leaf size across

species to decrease towards dry (Giliberto & Estay, 1978; McDonald et al., 2003) and cold climatic (Peppe

et al., 2011) and acidic soil conditions (Liu et al., 2012) while high mean values were determined for warm,

moist areas with low annual seasonality (Murphy & Lugo, 1986; Dransfield et al., 2008; Balslev et al.,

2011), e.g. in low latitudinal moist rainforests such as the Amazon (Hulshof et al., 2013). In other studies it

was shown that tree height varied positively along a temperature and precipitation gradient (e.g. Kessler et

al., 2007). In other words, tree height was detected to peak under warm, moist and aseasonal climates

(Swenson & Weiser, 2010), while for palms in the Amazon community-level mean stem height was shown

to be low on poor soils (Balslev et al., 2011). Notably, several studies focused on fruit and seed size and their

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environmental drivers. Seed mass was found to be large in habitats with high mean annual temperature,

possibly due to increased metabolic costs and expenditure (Murray et al., 2004). Furthermore, precipitation

(Swenson & Weiser, 2010) as well as poor (Katabuchi et al., 2012) and acidic soils (Tautenhahn et al., 2008)

were shown to be positively related to mean and median seed mass and size, respectively. Soil variation is

often important for plant community compositions (Tuomisto & Ruokolainen, 2000) and trait distributions at

small geographical scales (Liu et al., 2012). But edaphic gradients also exist across broad spatial scales in the

Neotropics, notably the very nutrient-poor soils in large parts of the Cerrado and on the Guiana Shield

(Furley & Ratter, 1988; ter Steege et al., 2006) relative to the nutrient rich soils in the Chaco, northeastern

Brazil and the eastern Andes slopes (Ratter et al., 1978; Pennington et al., 2000). Poor soil conditions can

cause intermediate disturbance conditions, which may lead to less turnover of individuals trees, smaller

canopy gaps and more shading, thereby favoring larger seeds by reducing stress tolerance to seedlings (ter

Steege & Hammond, 2001; ter Steege et al., 2006). However, Quaternary climate change can also influence

large-scale means of seed distribution patterns across genera during glacial-interglacial time periods towards

large-seeded species in warm, dry areas due to survival benefits under such climatic conditions (Campbell,

1982).

To our knowledge, no study has so far focused on palm functional traits at a macroecological scale or

has linked palm functional trait patterns to long-term historical drivers (e.g. paleoclimate). Here, we test the

relationships of several environmental drivers (current climate, soil and paleoclimate) on the geographic

distributions of assemblage medians of three key functional palm traits (leaf size, stem height and fruit size).

We explore whether environmental predictor variables related to climate (Kissling et al., 2012a), soil

(Balslev et al., 2011; Eiserhardt et al., 2011) and paleoclimate (Kissling et al., 2012a,b; Blach-Overgaard et

al., 2013) are important for explaining functional trait distributions (Swenson & Weiser, 2010; Liu et al.,

2012). More specifically, we test the following hypotheses:

1) Median leaf size of species within grid cells is highest in currently warm, moist areas with low annual

seasonality and low soil acidity and sand content, but also in areas were these climatic conditions already

existed during the Quaternary.

2) Median stem height of palm species assemblages is low on poor soils, but high in tropical rainforests

because they have higher canopies than seasonally dry tropical and subtropical forests (Murphy & Lugo,

1986). Additionally, we expect high assemblage median stem height in areas with decreasing Quaternary

climate stability in warm, moist conditions (REF).

3) Median fruit sizes of palm species assemblages are large on poor soils and in habitats with high mean

annual temperature and precipitation. Furthermore, Quaternary climate change could have shaped

assemblage median fruit size patterns with high values in areas that have been exposed to strong

paleoclimatic oscillations.

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MATERIAL AND METHODS

Palm distribution and trait data

Distributional data for nearly all palm species (Arecaceae, n = 541) across the Americas were digitized in

ArcView 9.2 (ESRI Inc., Redlands, California, USA) based on the (partly estimated) range maps from

Henderson (1995). We excluded the coconut (Cocos nucifera L.) from our analysis due to its unexceptionally

large fruit size and its dispersal via floating seeds (Dransfield et al., 2008). Range maps were overlaid onto a

grid in cylindrical equal area projection with 110 × 110 km resolution (equivalent to c. 1° × 1° near the

equator) and the presence of each palm species was then recorded for each grid cell. We only included grid

cells with species richness >2 to calculate meaningful median values per grid cell (n = 1498). This represents

a total of 36,422 grid cell occurrences across all species.

For the palm traits, we focused on leaf size (maximum rachis length in m), stem height (maximum

height in m), and fruit size (volume in cm3, based on maximum fruit length, width, diameter, and shape

information). These traits are not identical, but in line, with the traits of the LHS plant ecology strategy

scheme (Westoby, 1998) and represent one trait for each category. We chose leaf size and fruit size rather

than specific leaf area and seed size because little information is available for palms for the latter two. Fruit

size can be seen as a proxy for seed size because many palm genera are mainly 1-seeded so that fruit and

seed size are often highly correlated (Tomlinson, 1990; Henderson, 1995). Trait data of palm species were

extracted from Henderson (2002) for the majority of species. Additional data were collated from other

sources, including monographs and species descriptions, the Aarhus University Herbarium and the palmweb

database from Royal Botanic Gardens Kew (http://palmweb.org/). A detailed overview of the trait data

sources for each species is provided in Table SX. For calculating fruit size, we derived a measure of fruit

volume based on information of fruit length, width and diameter in cm, respectively. Additional information

of three-dimensional fruit shapes (e.g. globose, ellipsoid, pyramidal, cylindrical) was then used together with

geometrical formulas to calculate the fruit size volume (in cm3) of each palm species. Globose fruit shapes

were calculated by the formula for spheres (V = 4/3π × radius³), ellipsoid shapes by the formula for

ellipsoids (V= 4/3π × height × length × width), pyramidal shapes by the formula for pyramids (V=1/3 ×

length2 × height), and cylindrical shapes with the formula for cylinders (V = π × radius² × height). In case

that trait values were missing for individual species, we used the mean of species in the same genus to

estimate the value of the missing species. This was done with leaf size for 87 species, stem height for 4

species, and fruit size for 18 species. A detailed overview of the mean and median trait values per genus as

well as the number of estimated species per genus is provided in Table S2. In a final step, we computed

median values for each of the three trait variables for all the species that were present in a given 110 × 110

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km-grid cell; these assemblage medians were then used as response variables in the statistical analyses (see

below). To assess whether acaulescent species (i.e. palms with no or only a very short stem concealed in the

ground) have a major influence on geographic trait variability in palms, we examined trait distributions of all

palms (Figure 1) versus those of non-aucaulescent palms only (see Figure S1).

Environmental determinants

To explain geographic variation in functional traits of New World palms, we focused on three categories of

predictor variables: present-day climate (three variables), soil (three variables), and paleoclimate (two

variables) (Table 1). These drivers have previously been shown to be important for explaining species

richness and assemblage composition in palms or geographic trait variation in other plant families (ter Steege

et al., 2006; Tautenhahn et al., 2008; Balslev et al., 2011; Kissling et al., 2012a,b). We used the same grid

(110 km × 110 km grid resolution) as for the palm distribution data to extract the environmental data. All

environmental variables were calculated in ArcGIS (version 10.1, ESRI, Redlands, CA, USA) and their

mean values were extracted for each grid cell.

Current climate

Current climatic factors have been shown to be important drivers of palm species distributions and diversity

patterns (Eiserhardt et al., 2011; Kissling et al., 2012a) as well as trait distributions of other plants (Giliberto

& Estay, 1978; Swenson & Weiser, 2010). To represent current climate, we used all 19 climate variables

from the WORLDCLIM database (version 1.4; http:// www.worldclim.org), a set of global climate layers

with a spatial resolution of c. 1 km2 (Hijmans et al., 2005). We performed a Principal Component Analysis

(PCA) based on the correlation matrix to reduce collinearity among the 19 climate variables. We retained the

first three PCA axes, which together explained 81.06% of the variability in the data (see Table S1). The PC1

axis was strongly positively related to mean annual precipitation, precipitation of the wettest quarter and

mean annual temperature (hereafter referred to as PC-ANNU, Table S1). The PC2 axis showed a positive

relation to temperature seasonality and precipitation seasonality (hereafter referred to as PC-SEAS, Table

S1). The PC3 axis showed a negative relation with precipitation of the driest month, hereafter referred to as

PC-DRYM (Table S1).

Soil

Soil variables play an important role for small-scale (Balslev et al., 2011) and large-scale species

distributions of palms (Eiserhardt et al., 2011), and for functional trait distributions in other plant families

(Tautenhahn et al., 2008). We focused on three topsoil variables, namely acidity of topsoil (pH), percentage

sand fracture in topsoil (sand%) and cation exchange capacity in topsoil (CEC) (see Table 1) in line with the

fact that palms mainly form short roots at ground level or slightly below (Dransfield et al., 2008). Soil data

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were derived from the Harmonized World Soil Database (FAO et al., 2012) and mean values within grid

cells were calculated for all three soil variables in ArcGIS. Correlations between the three soil variables were

low to moderate (Spearman rank: r < 0.53) and we therefore included all three soil variables in the analyses.

We initially also explored mean base saturation in the topsoil per grid cell as potential predictor variable, but

as it was highly correlated with soil pH (r = 0.86) we did not include this variable in analyses.

Paleoclimate

Paleoclimate has been shown to be an important predictor of regional, continental and global palm diversity

patterns (Kissling et al., 2012a; Blach-Overgaard et al., 2013; Rakotoarinivo et al., 2013), but to our

knowledge it has not been explored as a driver of trait distributions. Nevertheless, current climatic conditions

can be related to functional diversity patterns at macroecological scales (e.g. Swenson & Weiser, 2010). In

addition, palm species richness can be impacted by historical drivers such as Quaternary temperature

oscillations (Kissling et al., 2012a; Rakotoarinivo et al., 2013), so that palm functional trait patterns can also

be expected to be related to paleoclimate. To represent Quaternary climate change, we calculated the

anomalies (differences) between the climate during the Last Glacial Maximum (LGM; c. 21,000 years ago)

and the present-day climate. Using annual precipitation and annual mean temperature, we computed the

anomaly of temperature (LGM ANOM TEMP, in °C × 10) and the anomaly of precipitation (LGM ANOM

PREC, in mm year–1) as paleoclimatic predictor variables (see Table 1). The former can be seen as roughly

representative for the major climatic oscillations of the Quaternary (the last several 105 years) because these

temperature anomalies cover the full glacial-interglacial climate cycle with a geographic pattern that is

consistent with these orbitally-driven climatic oscillations over at least a large portion of the period (see

Jansson, 2003). We used two different climate simulations (the Community Climate System Model version

3, CCSM3, and the Model for Interdisciplinary Research on Climate version 3.2, MIROC3.2) of the

Paleoclimate Modeling Intercomparison Project (PMIP2; http://pmip2.lsce.ipsl.fr/) to quantify these

paleoclimatic changes (Braconnot et al., 2007). Both climate simulations provide temperature and

precipitation data for the LGM and data were resampled in ArcGIS with a bilinear interpolation from the

original 2.5″ resolution to the resolution of the contemporary climate data. We then calculated mean anomaly

values across these two climate simulations per 110 km × 110 km grid cell. Large positive anomaly values

indicate a higher precipitation and temperature in the present than in the past whereas small or negative

anomaly values indicate the opposite, i.e. higher precipitation and temperature in the past than in the present.

Statistical analysis

We analyzed geographic variation in three assemblage-level median palm traits (leaf size, stem height, fruit

size) and their relationships with environmental predictor variables related to climate (PC-ANNU, PC-SEAS,

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PC-DRYM), soil (pH, sand%, CEC) and Quaternary climate effects (LGM ANOM TEMP, LGM ANOM

PREC) using ordinary-least-squares (OLS) linear regression models as well as spatial autoregressive (SAR)

models. We excluded cells with 10% and less land area and those for which no environmental or any trait

variables (see below) were available. In a first step, we included all eight predictor variables in full OLS

models (separate models with all predictors for each trait) and then performed a model selection with the

Akaike Information Criterion (AIC) to identify the minimum adequate model (i.e. the one with the lowest

AIC value). All predictor and response variables were checked to approximate a normal distribution and

bivariate relationships were examined for non-linearity. As a consequence, the response variables leaf size

and fruit size and the explanatory variables CEC and LGM ANOM TEMP were log10-transformed. We

further tested for polynomial terms to account for non-linear relationships by examining the differences in

AIC between simple OLS models with and without polynomials. In all cases, AIC differences were < 4.55 %

and we therefore did not include any polynomial terms. Since spatial autocorrelation can affect significance

tests and coefficients estimates of statistical models (Legendre & Legendre, 1998; Kissling & Carl, 2008),

we used Moran’s I and residual maps based on the residuals of the selected minimum adequate OLS models

to quantify the presence of spatial autocorrelation (see Figure S2). Because Moran’s I values were significant

for OLS model residuals, we implemented SAR models of the error-type (Kissling & Carl, 2008). We used

the same variables as in the minimum adequate OLS regression models and included a spatial weight matrix

in the SARs to account for residual autocorrelation (Kissling & Carl, 2008). For defining the neighborhood,

we used the minimum distance needed to connect a grid cell to at least one nearest neighbor (132 km) and

row-standardization for the weighting (Kissling & Carl, 2008). We then used correlograms to quantify

spatial autocorrelation in the response variables (raw data), the residuals of the non-spatial OLS models, and

the residuals of the SAR models (see Figure S3). This allowed us to assess the amount of spatial

autocorrelation with increasing geographic distance by plotting distance classes (bins) of grid cells on the x-

axis and Moran’s I values on the y-axis (Kissling & Carl, 2008).

For the SAR models, we quantified how much of the explained variance could be attributed to the

predictor variables only, or to additional spatially-structured factors (e.g. unmeasured environmental

variables or dispersal limitation). We quantified the explained variance of the environmental predictors for

each selected SAR model (R2PRED) as well as the total explained variance (R2

FULL) of the full SAR models

(including environmental predictors and the spatial weights matrix) (Kissling & Carl, 2008). This was done

using pseudo-R2 values, which were calculated as the squared Pearson correlation between predicted and true

values (Kissling & Carl, 2008) (Table 2).

All statistical analyses were performed with R version 3.0.1 (R Core Team, 2013). Spatial analyses

were performed using the R package ‘spdep’ version 0.5-71 (2014, R. Bivand). Correlograms were

calculated with the function correlog() from the R package ‘ncf’ version 1.1-5 (Bjørnstad, 2005).

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RESULTS

Geographic variation of functional trait distributions

In contrast to the high species richness in the Andes and Amazon basin (Figure 1a), trait values mostly

peaked north of the Amazon and in southern Brazil (Figure 1b-d). Median leaf size peaked in areas north of

the Central Amazon basin (including eastern Colombia and western Venezuela), the Lesser Antilles within

the Caribbean, and in a broad belt from northeastern Bolivia to eastern Brazil. Low values for median leaf

size were located in Central Amazonia, southern Brazil and most parts of Central America (Figure 1a). The

palm species with the highest overall maximum leaf size in our dataset was Attalea funifera Mart. (12 m),

which is only found in eastern Brazil. The genus Attalea Kunth contains many species with large leaf sizes

and this increases assemblage median leaf size in many areas of the New World, like A. maripa Mart. along

the belt south of the Amazon and A. butyracea (Mutis ex L.f.) Wess.Boer in central Venezuela. One of the

main species-rich understory genera, which decreases assemblage median leaf sizes (e.g. in the Amazon) is

Bactris Jacq. (e.g. B. killipii Burret and elegans H.Wendl).

For maximum stem height, assemblage-level medians peaked along the Pacific coast, in the

Caribbean, northern Colombia and Venezuela and the Cerrado. In contrast, species-rich areas like the

Amazon basin, southeastern Brazil and Central America were dominated by relatively small-statured

understory palm species (Figure 1c). The palm species with the largest maximum stem height was Ceroxylon

quindiuense H.Wendl (50 m), which mainly occurs along the central and eastern Cordillera of the Columbian

Andes and this and other Ceroxylon species contribute essentially to high assemblage median stem heights in

this area. Along the belt of the Cerrado, the genus Attalea, while in the Caribbean Roystonea is strongly

represented. Low assemblage median stem heights were linked to high species richness in understory genera

such as Geonoma Wild. and Bactris in Central Amazonia, the understory genus Chamaedorea Wild. in

Central America, and the genus of Syagrus Mart. in south eastern Brazil, with several low-statured species

(e.g., S. vagan Bondar and S. werdermannii Burret) in this region.

In contrast to leaf size and stem height, median fruit size peaked in a broad band from the savannah

regions of the Brazilian Cerrado towards the Atlantic coast of eastern Brazil (Figure 1d). Several genera were

driving the distribution of high median fruit sizes within the Cerrado and the eastern Brazilian Atlantic coast,

including the genera Phytelephas (notably P. macrocarpa Ruiz & Pav.) and Attalea (e.g. A. funifera and A.

olifeira Barb.Rodr.). Furthermore, small values in assemblage median fruit size were found in Central and

southern Amazonia as well as along the eastern Andes slopes from Colombia to Peru where genera like

Geonoma, Bactris and Desmoncus Mart. occur with many small-fruited species (e.g. G. longipedunculata

Burret, B. simplicifrons Mart. and D. mitis Mart.).

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A total of 58 species (10.72%) in our dataset shows an acaulescent growth form which could

potentially influence geographic variability of assemblage-level traits such as leaf size, stem height and fruit

size. However, comparing median trait distributions of all palm species with those of non-acaulescent palm

species showed no qualitative differences in geographic patterns of those traits (Figure S1). Moreover,

median trait distributions of non-acaulescent palm species were highly correlated with those of all palm

species for all three traits (Spearman rank: rleaf = 0.84; rstem = 0.80; rfruit = 0.97). In the following, all results

refer to analyses including all palm species.

Trait and environmental correlates

For assemblage median leaf size, we expected and found trait values to increase with PC-ANNU, notably in

currently warm, moist areas for SAR (std. coef. = 0.251, Table 2; Figure 2a) and OLS (XX). Furthermore,

contrary to expectation, we also found a strong positive relation to temperature and precipitation seasonality

in SAR (PC-SEAS, std. coef. = 0.203) and OLS models (XX). However, we also expected, but did not find a

significant increase of median leaf size in areas with a high Quaternary climate stability of warm conditions.

Moreover, median leaf size was negatively correlated to topsoil sand fraction in the OLS, but less important

for SAR (sand%, std. coef. = -0.069), while CEC only showed a significant relation in OLS (XX). Other

included environmental variables related to soil (pH, CEC) and Quaternary climate change did not show any

statistically significant relationships with median leaf size (Table 2).

For median stem height, we expected an increase in assemblage medians in present and paleo-climatic

moist and warm areas, as well as in areas on fertile soils. Both parts of the hypothesis were not supported in

the SAR models, which only showed a strong environmental correlate was a positive relationship with PC-

SEAS (std. coef. = 0.113, Table 2; Figure 2b), indicating that assemblage stem height (like leaf size)

increases with increasing seasonality (Figure 2). Other included environmental variables did not show a

significant relation and only small standardized coefficients (std. coef. < 0.07, Table 2). On the other hand,

PC-SEAS (XX), PC-DRYM (XX), CEC (XX), LGM ANOM TEMP (XX) and to a lesser extent sand%

(XX) showed significant relations to assemblage median stem height, though SAR diminishes or removes all

these effects.

Geographic variation in assemblage median fruit size was hypothesized to show an increase in areas

with high annual temperature, unfertile soils and seasonal climates as well as in areas with warm, dry paleo-

climatic conditions. For the OLS models most of the variables showed a relation to assemblage mean fruit

size (Table 2), but most of the effects disappeared after using SAR models. While for SAR there was no

significant relations to annual temperature (as represented by PC-ANNU) and soil, there was a positive

relation to seasonality (PC-SEAS, std. coef. = 0.441, Table 2) and weaker to precipitation of the driest month

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(PC-DRYM, std. coef.= 0.070, Table 2). Furthermore, we detected a strong, positive relationship with LGM

ANOM TEMP (std. coef. = -0.117, Table 2), indicating that assemblage medians for fruit size increases in

areas with strong Quaternary temperature oscillations (Figure 2c).

The explanatory power of the environmental variables (R2PRED) in the SAR models varied among traits

(Table 2). A moderately large proportion of geographic variation in median leaf size was explained by the

included environmental factors (R2PRED = 0.43), and a similar amount reflected unknown spatially structured

variables (R2FULL- R2

PRED = 0.41). Geographic variation in median stem height was only related to the included

environment variables to a smaller extent (R2PRED = 0.21), with unknown spatially-structured variables

playing a bigger role (R2FULL- R2

PRED = 0.62). For median fruit size, the included environmental factors

explained moderate amount of its geographic variability (R2PRED = 0.37), with unknown spatially-structured

variables again providing more explanatory power (R2FULL- R2

PRED = 0.60).

DISCUSSION

We tested the relationships between geographic distributions of key functional traits (assemblage medians of

leaf size, stem height, and fruit size) and current climate, soil, and paleoclimatic changes, respectively, across

the New World for palms (Arecaceae), a major plant lineage of tropical and subtropical ecosystems. We

found that the geographic distributions of all trait variables were mainly related to current environment, with

seasonality (PC-SEAS) being an important driver for all three traits. In contrast, mean annual climate (PC-

ANNU) and soil were only important for leaf size. Additionally, for the first time we report evidence for

Quaternary climate change being linked to the current distribution of palm functional trait composition, with

larger average fruit sizes being found in areas with less pronounced climatic oscillations (Table 2, Figure 2c).

These findings show that current climate, notably seasonality, is the strongest determinant of geographic

variation in functional trait composition in palm assemblages across the New World, with paleoclimate and

soil playing smaller, but also important roles.

The geographic distribution of median leaf size was hypothesized to be high in aseasonal,

moist and warm climates with low soil acidity and low sand content. Indeed, we found strongly positive

relationships with mean annual temperature and precipitation (PC-ANNU, Table 2, Figure 2a) and a weak

negative relation to topsoil sand fraction (sand%), but surprisingly also a strong positive relation to

seasonality (PC-SEAS). The distribution for median leaf size was unrelated to glacial-interglacial climate

change (Table 2). These findings are in agreement with other studies showing that current climate is

important for palm species composition and richness (Eiserhardt et al., 2011; Kissling et al., 2012a) as well

as studies showing current climate-functional trait relations in plants more generally (Moles et al., 2014). For

example, a study of 690 eastern Australian plant species found leaf size to increase with increasing annual

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precipitation (McDonald et al., 2003). Similarly, another study of over 3000 globally distributed plant

species showed an increase in median leaf size with high values in annual temperature and precipitation

(Peppe et al., 2011). A decreasing leaf size with decreasing annual precipitation under hot climatic

conditions can be explained by less evaporation and water loss in small leaves (McDonald et al., 2003).

Therefore, species in humid areas are specifically able to optimize their photosynthetic activity by producing

large leaves (McDonald et al., 2003). Nevertheless, we did not find the highest assemblage median leaf size

in ever-wet tropical rainforests, but in areas with seasonal tropical climates, e.g., in northern and south-

eastern marginal parts of Amazonia. This could be due to the general morphology of palm leaves and their

adaptations to warm and hot climates (Dransfield et al., 2008), often including protection of the leaf surface

by strong cuticles and waxes that minimize water loss under dry conditions (Tomlinson, Horn & Fisher,

2011). Additionally, small-leaved understory palm species are mainly diverse in warm, moist aseasonal

climates, which have been argued to increase understory diversity due to less competition for water with the

canopy trees and less competitive exclusion within the understory, due to increased pest pressure and shade

limitation (Wright, 1992). On the other hand, in seasonal climates, low light levels in combination with

drought stress might not allow small-leaved palms to eke out and survive what suppresses understory

diversity in these areas. Allometric constraints will cause small palms to have small leaves, causing low

median leaf size in ever wet tropical areas with many understory species, as western and central Amazonia,

even if some tall, large-leaved species (e.g. Attalea butyracea, Attalea maripa) are also present.

Nevertheless, dense understory was also shown to maximize leaf area displayed on a whole-plant basis,

while minimizing the biomass costs of leaf support structures (Chazdon, 1991). Hence, the weak negative

relation to sand content could indicate small median leaf values on poor sandy soils, given that smaller

leaves could be an adaptation to low soil moisture and water availability due to greater water stress under

high temperatures (Giliberto & Estay, 1978). Clayey soils have higher water storage capacity and could

therefore provide increased water availability even during dry, seasonal conditions including longer drought

periods, while sandy soils show a very poor water storage capacity (Ritchie, 1981) and therefore might not

provide an optimal water availability for palms due to their shallow root systems (Dransfield et al., 2008).

Besides producing large leaves, an alternative strategy to increase photosynthetic capacity could be to

decrease in leaf width, while increasing leaf number (Malhado et al., 2009). Nevertheless, also other

environmental factors which were not tested here, such as topography (Tomer & Anderson, 1995), could

impact the water storage capacity of soils and therefore possibly indirectly influence median leaf size.

For assemblage median stem height we expected the highest values in moist, warm aseasonal

tropical rainforest areas, as these forests often have higher canopies than forests in more seasonal climates

(e.g. Murphy & Lugo, 1986), which are drier and show less net primary productivity (Moles et al., 2009).

This prediction was mostly not supported. Notably, assemblage median stem height was unrelated to mean

annual temperature and precipitation. However, there was a strong positive relation to PC-SEAS (Table 2;

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Figure 2b), indicating that assemblage median stem height increase with increasing seasonality (e.g. eastern

Brazil), similar to the findings for median leaf size. The positive relation to seasonality could reflect that

robust, large-stemmed species outcompete small, drought-sensitive species due to higher resistance to

environmental constraints of seasonal climates, notably drought (Balslev et al., 2011). For instance, the

genus Roystonea O.F.Cook, includes ten species within the Caribbean, which all show a solitary, robust stem

(15-40m) and are known to be dominant especially in disturbed landscapes under seasonal Caribbean climate

conditions (Henderson, Galeano-Garces & Bernal, 1997). Notably, Moles et al. (2009) found a similar

pattern in tree heights inside the tropics, involving a shift from the inner tropics with moist, warm climates

towards high community-level stem heights in the outer tropics with seasonal climates. This was explained

to reflect a switch in the plant ecological strategy towards the edge of the tropics, which might be driven by

environmental conditions or different life-history traits such as life span and time to first reproduction. This

assumption could also apply to palms, but needs further investigations. Notably, as already discussed for leaf

size, a low diversity of small-stemmed understory palm species and a low number of growth form types

could be the result of understory palm species being limited in competition with large canopy species in

seasonal climates due to sensitivity to both shade and drought stress (Wright, 1992), even if there are also

large-stemmed palm species (like Astrocaryum chambira Burret and Iriartea deltoidea Ruiz & Pav.) occur in

aseasonal and wet climates (Balslev et al., 2011). Furthermore, small stature is also an adaptation for shade

tolerance requires lower costs for biomass production (Chazdon, 1991). Additionally, Chazdon (1991) found

a correlation of decreasing leaf size with decreasing crown height, similar to the broad-scale patterns that we

have found here for palms across the Americas, suggesting that ….. the leaf size and median stem height

confirm these findings and suggest an adaptation of understory palms to dense forests and shading,

detectable not only by small assemblage median leaf sizes but also by low stem heights.

For assemblage median fruit size we expected large values on poor soils and in habitats with

high mean annual temperature and precipitation as well as Quaternary climate change having shaped

assemblage median fruit size pattern in the direction of high values in past warm and dry paleoclimates.

Geographically, we found increasing median fruit size from southern Amazonia towards eastern Brazil and

over the whole Cerrado. Large seed size is considered advantageous in situations where establishment

conditions are stressful due to e.g., low soil fertility (Liu et al., 2012) at the cost of lower seed dispersal

distances (Zona & Henderson, 1989; Beaune et al., 2013) and thus lower migration rates. Palm assemblage

median fruit size was not related to soil fertility and mean annual climate. In contrast, fruit size was indeed

positively related to seasonality and precipitation of the driest month, albeit letter only weakly so (Table 2).

Hereby, our findings for the New World palms are consistent with the idea that a large seed size is

advantageous in a seasonal climate with long dry periods, by increasing the likelihood of survival under

droughts (Westoby, 1998) and therefore reproduction success (Lloret, Casanovas & Penuelas, 1999). .

Interestingly, median fruit size was as the only trait investigated also linked to Quaternary glacial-

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interglacial-climate change, notably strongly positive to temperature change, i.e., with higher values of

median fruit size in areas with relatively high glacial cooling. In other words, the more temperature unstable

areas with stronger temperature oscillations over time showed larger median fruit size than climatically

stable areas (Table 2, Figure 2c). ,. Notably, larger seed mass and fruit size enable seedlings to better

survival hazards (e.g. droughts) while tropical understory palms are especially sensitive to drought (Wright,

1992; Westoby et al., 1996). Greater temperature oscillations might therefore favor palm species and clades

with larger fruit sizes which would result in a positive correlation between assemblage level fruit size and

LGM ANOM TEMP. For instance, African palm species were detected to be majoritarian large fruited, what

could be explained by Cenozoic drying having a strong effect on the trait composition of palm species

assemblages in this region (Kissling et al., 2012b). This finding is also consistent with the positive

correlation between phylogenetic clustering of palm assemblages and LGM ANOM TEMP that was found

for South America where a changing climate and habitat loss throughout the Cenozoic had strong impacts on

the phylogenetic structure of regional species assemblages in the tropics (Kissling et al., 2012b). In the

Neotropics, phylogenetic clustering increases with stronger effects of glacial-interglacial climate oscillations

and shows that specific clades perform better in climatically unstable regions, just as the Cerrado in our

study. Subsequently, many taxa are endemic to certain regions and local areas, such as the dominance of the

subfamily Cocoseae in the Neotropics (Kissling et al., 2012b) or the genus XX within the savanna area of the

Cerrado. Furthermore, phylogenetic clustering and functional trait distributions might also be related to

glacial environmental filtering and postglacial dispersal limitation. For New World palms, postglacial

migrational lag has been invoked to the current distribution and ongoing range dynamics in the rain-forest

understory palm Astrocaryum sciophilum Pulle (Charles-Dominique et al., 2003). These findings are similar

to those for other plants from higher latitudes which are more impacted by glaciations. Notably, postglacial

dispersal limitation has been shown to shape species ranges (Normand et al., 2011) and range filling in trees

across Europe (Nogués-Bravo et al., 2014). Our findings suggest that glacial survival and postglacial range

dynamics in New World palms are probably influenced by the size of their fruits and seeds, reflecting their

role in plant dispersal and stress tolerance. Importantly, large fruits may be subsequently conferred to

relatively low rates of range expansion due to dispersal limitations from these stable, glacial refugia

(Campbell, 1982; Nogués-Bravo et al., 2014). These findings are also consistent with the increasing

evidence that glacial and deeper-time paleoclimate still shape palms species richness of palms in the

Neotropics (Kissling et al., 2012a,b) and Africa (Blach-Overgaard et al., 2013; Rakotoarinivo et al., 2013).

In addition to the drivers discussed above, it has to be mentioned that geographic variation in

median trait values was also related to unknown spatially-structured variables, i.e. for leaf size 41% (R2FULL-

R2PRED), for stem height 62% and for fruit size 60% (Table 2). This unexplained variation could be attributed

to unmeasured environmental factors, dispersal limitation or diversification processes. Notably, we only

detected a relation of paleoclimate to assemblage median fruit size, but not to the other trait variables. Thus,

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the strong unexplained spatial components could also contain unexplained historical processes such as large-

scale dispersal limitation (Svenning & Skov, 2005). Another possible explanation could be that spatially

autocorrelated environmental variables are missing or that patterns of species are consistent with greater

ecological specialization, e.g. adaptations to high temperatures or soil sandiness (Svenning & Skov, 2005).

Furthermore, humans could impact median trait distributions by e.g. introductions and naturalizations of

species to enable them to grow beyond their native ranges (Svenning & Skov, 2005). Another aspect that

could impact functional trait distribution is fire, due to long dry and hot periods in seasonal climates (Grau &

Veblen, 2000; Furley, 2002), but also as caused by humans (e.g. Hoffmann, 1999; Michalski and Peres

2005). Human introduced fires are known to have influenced species richness and composition (Hoffmann,

1999), especially in southern Amazonia and the Cerrado (Ratter et al. 1997; Pennington et al., 2000;

Michalski and Peres, 2005). Not only species diversity might have been impacted by fire, but also

adaptations in ecology and functional traits, such as a thick, corky bark and scleromorphic leaves

(Pennington et al., 2000; Furley, 2002). Furthermore, fire frequency might determine whether a species will

decline towards extinction or become abundant under a particular fire regime, causing shifts in the plants’

size and have large effects on the physical structure of the vegetation as it was shwon for the Cerrado

(Hoffmann, 1999). Taller and thicker stems might be an advantage of robustness against (human induced)

fires while smaller species and individuals might not be able to handle fire disturbance and go extinct

(Williams et al., 1999). For instance, along the Xingu river in eastern Amazonian Brazil, an area with high

species diversity, human induced fires changed species composition towards extensive stands of tall species,

namely the babacu palm Attalea speciosa Mart. which shows several adaptations to fire, such as a tall stem

and large, thick endocarped fruits (Smith, 2015).

Overall, we used Westoby’s (1998) LHS plant strategy scheme to explain distribution of three

key functional traits by environmental correlates. We chose the trait variables leaf size, stem height and fruit

size as they were proposed in his study to capture the main trait axes of a plant species responses to different

factors such as e.g. competition and disturbance, and being supposed to be applied for every species.

Assemblage medians for palm functional traits in the New World were strongly related to current climate

and in particular to seasonality, with much weaker links to past climate change and soil. Theoretically, a

weak or missing soil effect on functional trait distributions in the New World could be due to the coarse

grain size of our analyses. Notably, Eiserhardt et al. (2011) reviewed that palm diversity can be influenced

by soil chemistry on a continental scale and palm community composition on a regional to local scale.

Notably, we only detected a relation of paleoclimate to assemblage median fruit size, but not to the other trait

variables. This suggests that distributions of assemblage trait medians like stem height and leaf size, are not

maintained over a million year time periods although unexplained historical components as well as co-

variation between paleo- and current climates could contribute, as explained above. Particularly, palms are

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known to be strong indicators for current temperature and precipitation (Dransfield et al., 2008). Therefore, a

fast shift in climatic conditions under future climate change could impact palm species distributions (Blach-

Overgaard et al., 2010) and along with it a shift in functional trait distributions (Diaz & Cabido, 1997). It is

likely that towards the end of the century the New World is undergoing an increase in mean annual

temperature and a decrease in mean annual precipitation over most areas, including a more seasonal climate

(especially in regard to precipitation), e.g. in the Andes and Eastern Amazonia (Magrin et al., 2014). This

could be problematic for many palm species because most are sensitive to drought and concentrated in the

wettest parts of the New World tropics (e.g., many understory palm species). Large-stemmed generalist palm

species that are widespread and common in tropical seasonal areas are expected to have an advantage under

drier and warmer conditions and may be able to migrate into new areas, if they can disperse fast enough to

track changing climates. In contrast, especially understory tree species can be highly drought-sensitive and

will suffer from increasing drought, as already documented in tropical moist forests in central Panama

(Condit, Hubbell & Foster, 1996). Furthermore, species with large fruits, which often have advantages in

seedling survival (Lloret et al., 1999), might be able to deal better with dry and more seasonal conditions.

Nevertheless, a considerable movement of large-fruited species with increasing seasonality seems to be

unlikely at a continental scale as our results show that present median fruit size distribution is still related to

Quaternary climate change, likely due to postglacial dispersal limitation, which will be exacerbated by

disperser loss due to current defaunation, especially of large mammal species (Galetti et al., 2006; Beaune et

al., 2013). Altogether, given the strong climate-trait relationships documented in this study we expect that

future climate change has a strong impact on the functional composition of palm communities and thus on

ecosystem dynamics in palm-inhabited parts of the New World, as palms are a keystone family with high

ecological importance here (Dransfield et al., 2008), notably as habitat and food resource for mammals and

birds (Zona & Henderson, 1989; Galetti et al., 2006).

ACKNOWLEDGEMENTS

Our research was supported by the European Research Council (grant ERC-2012-StG-310886-HISTFUNC

to J.-C.S.). W.D.K. acknowledges support from a University of Amsterdam (UvA) starting grant. We also

thank Aarhus University and several people for feedback and support to this study, notably Alejandro

Ordonez, Peder K. Bøcher, Wolf L. Eiserhardt and Henrik Balslev.

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Figure 1: Species richness (a) and community-level median values of (b) leaf size (in m), (c) maximum

stem height (in m) and (d) fruit size (in cm3) for palm assemblages across the New World. Quantile

classification is shown across a grid with 110×110 km cell size (equivalent to c. 1°×1° near the equator) and

a WGS 1984 projection.

Figure 2: Partial residual plots illustrating the relation of three community-level traits (a: leaf size, b: stem

height, and c: fruit size) with their most important environmental predictor variable (compare standardized

coefficients in Table 1). Partial residuals represent the relationship between a response and a predictor

variable when all other predictor variables in the model are statistically controlled for. Specifically, these

partial residual plots are plots of r + b × Environment versus Environment (x-axis), where r is the ordinary

residuals form the multiple-predictor model and b is the regression coefficient estimate for Environment

from the same multiple-predictor model. Abbreviations of predictor variables are explained in Table 1.

Figure 1:

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Figure 2:

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Table 1: Predictor variables to explain the geographic variation and environmental correlates of functional

trait distributions in palms across the New World

Abbreviation Predictor variables (units) Data source

Current climatePC-ANNU PCA axis mainly representing

annual precipitation (mm year-1), precipitation of the driest month (mm) and mean annual temperature (°C × 10)

Worldclim dataset (Hijmans et al., 2005)

PC-SEAS PCA axis mainly representing seasonality of temperature (standard deviation of monthly means, °C × 10) and precipitation (coefficient of variation of monthly, mm)

Worldclim dataset (Hijmans et al., 2005)

PC-DRYM PCA axis mainly representing precipitation of the driest month (mm)

Worldclim dataset (Hijmans et al., 2005)

SoilpH pH in topsoil (-log(H+)) Harmonized World Soil

Database (FAO et al., 2012)

sand% Sand fraction in topsoil (%) Harmonized World Soil Database (FAO et al., 2012)

CEC

Quaternary climate change

Cation exchange capacity in topsoil (cmol/kg)

Harmonized World Soil Database (FAO et al., 2012)

LGM ANOM TEMP Anomaly in TEMP between Last Glacial Maximum andpresent (K, originally in °C × 10)

Calculated in ArcGIS using the variables LGM TEMP andTEMP Worldclim dataset (Hijmans et al., 2005)

LGM ANOM PREC Anomaly in annual precipitation between Last GlacialMaximum and present (mm year1)

Calculated in ArcGIS using the variables LGM PREC andPREC from Worldclim dataset (Hijmans et al., 2005)

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Table 2: Multiple predictor models (ordinary least squares: OLS) and multiple predictor spatial autoregressive (SAR) models were used to explain the geographic variation of community-level functional traits (mean leaf size, mean stem height, and mean fruit size) in palm assemblages across the New World. Explanatory variables include current climate (PC-ANNU, PC-SEAS, PC-DRYM), soil (pH, sand%, CEC) and Quaternary climate change (LGM ANOM TEMP, LGM ANOM PREC). For each functional trait variable, a minimum adequate model was selected with the Akaike Information Criterion (AIC) based on a non-spatial model with all explanatory variables (OLS). This model was then fitted with a SAR model. The response variables leaf size and fruit size and the predictor variables CEC and LGM ANOM TEMP were log10-transformed. For the response variable leaf size, all included predictor variables were selected in the most parsimonious model whereas for stem height the predictors pH, sand % and LGM ANOM PREC and for fruit size the predictor pH were not selected (indicated by “--”). Sample sizes are 1498 grid cells of 110×110 km resolution in all analyses.

Explanatory variables Coefficients

Leaf size Stem height Fruit size

OLS SAR OLS SAR OLS SAR

Intercept 1.340 *** 1.215 *** 10.388 *** 11.110 *** 6.667 *** 7.557 ***

PC-ANNU 0.028 ***(0.026)

0.028 ***(0.251)

-0.004(-0.002) -- 0.108 *

(0.090)0.040(0.031)

PC-SEAS 0.022 ***(0.024)

0.017 ***(0.203)

0.304 ***(0.286)

0.185 ***(0.113)

0.501 ***(0.328)

0.441 ***(0.205)

PC-DRYM 0.002(0.001)

-0.005(-0.053)

0.110 *(0.101)

0.049(0.025)

0.194 **(0.177)

0.070 *(0.123)

pH -0.000(-0.000)

0.000(0.002)

0.104 *(0.097)

0.063(0.034)

-0.013(-0.009)

-0.023(-0.027)

sand% -0.019 **(-0.020)

-0.014 *(-0.069)

-0.000(-0.001) -- 0.005

(0.006) --

CEC 0.021 **(0.017)

-0.002(-0.005)

0.090 *(0.081)

0.046(0.031)

-0.026(-0.022)

-0.005(-0.004)

LGM ANOM TEMP 0.029(0.016)

0.001(0.021)

0.106 *(0.089)

0.081(0.069)

0.560 ***(0.411)

0.680 ***(0.561)

LGM ANOM PREC -0.000(-0.000)

-0.000(-0.030)

-0.000(-0.002) -- 0.003

(0.002) --

R2OLS 0.300 -- 0.287 -- 0.508 --

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R2PRED -- 0.428 -- 0.213 -- 0.365

R2FULL -- 0.842 -- 0.831 -- 0.969

AIC 5672 -2653 2578 4788 972 1821

Moran’s I 0.787 0.013 0.840 0.011 0.718 0.008

P (Moran’s I) *** n.s. *** n.s. *** n.s.

Abbreviations of predictor variables are explained in Table 1. For each model, the regression coefficients,

the explained variance of the OLS (R2OLS) and SAR models (R2

FULL,R2PRED), the AIC, Moran’s I, and the P-

value of Moran’s I are given. ***P < 0.001; **P < 0.01; *P < 0.05; n.s. not significant; ‘--’ not selected for

the Minimum Adequate Model.

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Appendix

Figure S1: Community-level median values for non-acaulescent palms of (a) leaf size (in m), (b) maximum

stem height (in m) and (c) fruit size (in cm3), and standard deviations of median leaf size (d), stem height (e)

and fruit size (f) for palm assemblages across the New World. Quantile classification is shown across a grid

with 110×110 km cell size (equivalent to c. 1°×1° near the equator) and a WGS 1984 projection.

Figure S2: Maps show the residuals of the OLS for our assemblage median trait distributions of leaf size (a),

stem height (b) and fruit size (c), and for the SAR models of leaf size (d), stem height (e) and fruit size (f).

The diameter of each dot indicates the amount of spatial autocorrelation at this particular area, while the

colors illustrate positive (black) and negative (grey) autocorrelation, respectively.

Figure S3: Moran’s I correlograms of the residuals of the model fit of the raw data (white circle), the OLS

model (grey circle) and the spatial autoregressive model (black circle) separately for the three trait variables

median leaf size (a), median stem height (b) and median fruit size (c).

Figure S4: Histograms for the three different trait variables illustrate the frequency and distribution of the

trait data values for leaf size (a), stem height (b) and fruit size (c) over the whole dataset. Data for traits leaf

size and fruit size are log10-transformed.

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Figure S2:

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Figure S3:

Figure S4:

Table S1: Principal component (PC) analysis for 19 current climate variables of the worldclim

database. Entries are eigenvalues, percentage of variance for each axis and cumulative across all,

and the correlation between the PC axes and the most important climate variables.

PC-ANNU PC-SEAS PC-DRYM

PCA result

Eigenvalue 3.001 1.895 1.640

Percent of variance 49.12 22.85 11.75

Cumulative percent of variance 46.49 69.34 81.09

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Correlation coefficient

Mean annual temperature 0.803 0.352 0.088

Temperature seasonality 0.109 0.795 -0.322

Mean annual precipitation 0.842 -0.412 -0.096

Precipitation of the wettest quarter 0.621 -0.169 0.229

Precipitation seasonality -0.205 0.693 -0.149

Precipitation of the driest month 0.379 -0.128 -0.542

Table S2: All genera including numbers of species per genus, number of estimated values per trait and genus and mean, median and standard deviation for all genera.

Genus Species no. Estimated Leaf size Stem height

Leaf size

Stemheight

Fruitsize

mean median SD mean median SD mean

Acoelorrhaphe 1 0.70 0.70 4.00 4.00 0.81

Acrocomia 3 1 2.41 3.00 1.66 9.67 11.00 9.07 82.83

Aiphanes 22 1 1.67 1.49 0.92 6.64 5.00 5.39 14.28

Allagoptera 5 1 0.85 0.83 0.30 2.00 0.00 3.46 53.55

Ammandra 1 4.00 4.00 1.50 1.50 5575.28

Aphandra 1 5.40 5.40 11.00 11.00 333.33

Asterogyne 5 3 1 0.75 0.75 0.21 4.60 3.00 3.13 5.84

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Astrocaryum 18 2 1 4.23 4.00 1.23 11.11 9.00 7.54 204.34

Attalea 29 2 6.72 6.48 2.78 9.78 10.00 10.50 957.32

Bactris 62 1.41 1.38 0.84 4.67 4.00 3.18 18.72

Barcella 1 2.00 2.00 0.00 0.00 103.08

Brahea 9 6 1 1.00 1.00 7.53 7.00 4.98 17.60

Butia 8 1.36 1.19 0.66 3.88 4.00 3.48 35.79

Calyptrogyne 8 5 2 0.95 0.95 0.25 2.73 2.50 1.81 4.58

Calyptronoma 3 2 4.60 4.60 12.33 12.00 2.52 4.15

Ceroxylon 11 1 3.11 2.95 0.94 20.00 15.00 11.78 12.57

Chamaedorea 76 7 2 0.64 0.50 0.56 2.84 2.00 2.87 2.26

Chelyocarpus 4 2 0.76 0.76 0.58 9.00 10.00 6.06 22.80

Coccothrinax 14 9 3 1.20 1.20 0.42 10.71 10.00 3.27 2.48

Colpothrinax 2 1.03 1.03 0.52 9.00 9.00 1.41 8.62

Copernicia 13 9 1.00 1.00 0.00 11.88 10.00 7.93 21.84

Cryosophila 9 7 2.00 2.00 0.00 8.11 7.00 3.48 24.66

Desmoncus 7 1.36 1.70 0.75 11.79 10.00 9.24 22.48

Dictyocaryum 3 3.50 3.00 1.32 22.33 22.00 2.52 44.94

Elaeis 1 5.80 5.80 6.00 6.00 47.19

Euterpe 7 1 2.98 3.30 0.88 16.86 20.00 4.10 5.27

Gaussia 5 4 3.10 3.10 12.80 14.00 5.36 4.25

Geonoma 49 4 0.80 0.65 0.53 3.85 3.00 2.54 1.77

Hemithrinax 3 1 1 1 1.30 1.30 0.10 9.00 8.00 5.57 3.80

Hyospathe 2 0.75 0.75 0.50 5.00 5.00 4.24 3.29

Iriartea 1 4.70 4.70 25.00 25.00 32.57

Iriartella 2 0.88 0.88 0.11 7.50 7.50 6.36 3.46

Itaya 1 2.10 2.10 4.00 4.00 15.46

Juania 1 5.00 5.00 15.00 15.00 10.58

Jubaea 1 5.00 5.00 15.00 15.00 186.84

Leopoldinia 3 1.80 1.30 1.32 8.33 8.00 1.53 90.89

Lepidocaryum 1 0.04 0.04 4.00 4.00 24.85

Leucothrix 1 1.10 1.10 11.00 11.00 0.29

Lytocaryum 2 1 0.99 0.99 5.00 5.00 0.00 44.12

Manicaria 1 8.00 8.00 10.00 10.00 10102.8

Mauritia 2 1.80 1.80 1.70 20.00 20.00 7.07 636.30

Mauritiella 3 1 0.11 0.11 0.04 12.67 10.00 6.43 1248.74

Neonicholsonia 1 1.20 1.20 0.50 0.50 1.54

Oenocarpus 9 4.71 4.00 2.69 12.33 10.00 7.14 29.67

Parajubaea 2 2.59 2.59 0.30 15.50 15.50 0.71 287.07

Pholidostachys 4 1.68 1.58 0.45 8.50 9.50 3.87 16.63

Phytelephas 6 1 1 6.28 6.25 1.07 6.60 5.00 5.03 2976.19

Prestoea 10 2.08 2.10 0.61 7.25 6.50 3.97 2.48

Pseudophoenix 4 2.08 2.01 0.54 14.75 14.00 9.22 16.65

Raphia 1 8.50 8.50 12.00 12.00 329.87

Reinhardtia 6 4 3 1.59 1.59 2.00 5.07 4.25 4.43 6.08

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Rhapidophylum 1 2.00 2.00 1.00 1.00 2.36

Roystonea 10 1 4.37 4.00 0.66 21.50 20.00 7.84 4.97

Sabal 15 10 2.53 2.70 1.26 11.93 15.00 6.99 6.72

Schippia 1 1.20 1.20 10.00 10.00 36.82

Serenoa 1 2.00 2.00 0.00 0.00 22.46

Socratea 5 2.98 2.80 0.62 20.80 20.00 3.42 86.02

Syagrus 30 1 2.01 1.90 1.06 7.95 7.00 7.81 98.90

Synechanthus 2 1 1.30 1.30 6.00 6.00 0.00 33.47

Thrinax 3 1.37 1.40 0.15 12.00 12.00 1.00 601.40

Trithrinax 3 1.07 1.00 0.12 9.00 6.00 5.20 3.58

Washingtonia 2 2.10 2.10 0.14 18.50 18.50 4.95 2.36

Welfia 1 5.00 5.00 20.00 20.00 31.56

Wendlandiella 1 0.30 0.30 1.50 1.50 0.81

Wettinia 21 2 3 2.95 3.00 0.73 12.38 12.00 4.55 72.80

Zombia 1 1.00 1.00 3.00 3.00 12.63

Table S3: All species within the dataset (n=541) and attendant references which had been used to fulfill the missing trait values of Henderson et al. (1995).

SpecName References

Acoelorrhaphe wrightii AAU Herbarium; Henderson et al. 1995

Acrocomia aculeata AAU Herbarium; Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Acrocomia crispa AAU Herbarium; Henderson et al. 1995

Acrocomia hassleri AAU Herbarium; Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Aiphanes acaulis AAU Herbarium; Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002; Jones 1995

831

832

833

834

835

836

837

838

839

840

841

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Aiphanes horrida AAU Herbarium; Borchsenius et al. 1998; Henderson et al. 1995

Aiphanes chiribogensis Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Aiphanes deltoidea Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Aiphanes duquei Henderson et al. 1995; Henderson 2002

Aiphanes eggersii Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002; Jones 1995

Aiphanes erinacea Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Aiphanes gelatinosa Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002; Jones 1995

Aiphanes grandis Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Aiphanes hirsuta AAU Herbarium; Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002; Jones 1995

Aiphanes leiostachys Henderson et al. 1995; Henderson 2002

Aiphanes lindeniana Henderson et al. 1995; Henderson 2002; Jones 1995

Aiphanes linearis Henderson et al. 1995; Henderson 2002; Jones 1995

Aiphanes macroloba Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002; Jones 1995

Aiphanes minima AAU Herbarium; Henderson et al. 1995; Henderson 2002; Jones 1995

Aiphanes parvifolia Henderson et al. 1995; Henderson 2002; Jones 1995

Aiphanes simplex AAU Herbarium; Henderson et al. 1995; Henderson 2002; Jones 1995

Aiphanes spicata Henderson et al. 1995; Henderson 2002

Aiphanes tricuspidata Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Aiphanes ulei AAU Herbarium; Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Aiphanes verrucosa Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002; Jones 1995

Aiphanes weberbaueri AAU Herbarium; Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002; Jones 1995

Allagoptera arenaria Henderson et al. 1995; Henderson 2002; Jones 1995; Lorenzi 2010

Allagoptera brevicalyx AAU Herbarium; Henderson et al. 1995; Henderson 2002; Jones 1995; Lorenzi 2010

Allagoptera campestris Henderson et al. 1995; Henderson 2002; Jones 1995; Lorenzi 2010

Allagoptera caudescens Henderson et al. 1995; Lorenzi 2010

Allagoptera leucocalyx AAU Herbarium; Henderson et al. 1995; Jones 1995; Lorenzi 2010

Ammandra decasperma AAU Herbarium; Borchsenius et al. 1998; Galeano and Bernal, 2010; Henderson 2002

Aphandra natalia Borchsenius et al. 1998; Henderson 2002; Lorenzi 2010

Asterogyne guianensis Henderson 2002

Asterogyne martiana Borchsenius et al. 1998; Henderson 2002; palmweb

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Asterogyne ramosa Henderson 2002

Asterogyne spicata Henderson 2002; palmweb

Asterogyne yaracuyense Henderson 2002

Astrocaryum acaule Henderson 2002; Lorenzi 2010

Astrocaryum aculeatissimum Henderson 2002; Lorenzi 2010

Astrocaryum aculeatum Henderson 2002; Jones 1995; Lorenzi 2010

Astrocaryum alatum Henderson 2002

Astrocaryum campestre Henderson 2002; Lorenzi 2010

Astrocaryum chambira Borchsenius et al. 1998; Henderson 2002; Lorenzi 2010

Astrocaryum confertum Henderson 2002

Astrocaryum gynacanthum Henderson 2002; Lorenzi 2010

Astrocaryum huaimi AAU Herbarium; Henderson 2002; Lorenzi 2010

Astrocaryum jauari AAU Herbarium; Borchsenius et al. 1998; Henderson 2002; Lorenzi 2010

Astrocaryum malybo Henderson 2002; palmpedia

Astrocaryum mexicanum AAU Herbarium; Henderson 2002; Jones 1995

Astrocaryum murumuru AAU Herbarium; Borchsenius et al. 1998; Henderson 2002; Lorenzi 2010

Astrocaryum paramaca Henderson 2002; Lorenzi 2010

Astrocaryum sciophilum Henderson 2002; Lorenzi 2010

Astrocaryum standleyanum Borchsenius et al. 1998; Galeano and Bernal, 2010; Henderson 2002

Astrocaryum triandrum Galeano and Bernal, 2010; Henderson 2002

Astrocaryum vulgare Henderson 2002; Lorenzi 2010

Attalea allenii Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002

Attalea amygdalina Henderson et al. 1995; Henderson 2002

Attalea attaleoides Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Attalea butyracea AAU Herbarium; Borchsenius et al. 1998; Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002

Attalea cohune Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002

Attalea colenda Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Attalea crassispatha Henderson et al. 1995; Henderson 2002

Attalea cuatrecasana Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002

Attalea dahlgreniana Henderson et al. 1995; Henderson 2002; Lorenzi 2010

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Attalea dubia Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Attalea eichleri AAU Herbarium; Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Attalea exigua Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Attalea funifera Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Attalea geraensis Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Attalea humilis Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Attalea iguadummat Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Attalea insignis AAU Herbarium; Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Attalea luetzelburgii Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002

Attalea maripa Borchsenius et al. 1998; Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002

Attalea microcarpa Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002

Attalea nucifera Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002

Attalea oleifera Henderson et al. 1995

Attalea phalerata AAU Herbarium; Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Attalea pindobassu Henderson 2002

Attalea racemosa Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Attalea septuagenata Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002

Attalea speciosa Henderson et al. 1995; Henderson 2002

Attalea spectabilis Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Attalea tessmannii Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Bactris acanthocarpa AAU Herbarium; Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Bactris acanthocarpoides Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Bactris aubletiana Henderson et al. 1995; Henderson 2002

Bactris bahiensis Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Bactris balanophora AAU Herbarium; Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Bactris barronis Henderson et al. 1995; Henderson 2002

Bactris bidentula Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Bactris bifida AAU Herbarium; Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Bactris brongniartii Henderson et al. 1995; Henderson 2002

Bactris campestris Henderson et al. 1995; Henderson 2002; Lorenzi 2010

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Bactris caryotifolia Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Bactris caudata Henderson et al. 1995; Henderson 2002

Bactris charnleyae Henderson et al. 1995; Henderson 2002

Bactris coloniata Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Bactris coloradonis Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Bactris concinna AAU Herbarium; Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Bactris constanciae Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Bactris corossilla Borchsenius et al. 1998; Henderson et al. 1995; Lorenzi 2010

Bactris cuspidata Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Bactris dianeura Henderson et al. 1995; Henderson 2002

Bactris elegans AAU Herbarium; Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Bactris ferruginea Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Bactris fissifrons Henderson et al. 1995; Henderson 2002

Bactris gasipaes AAU Herbarium; Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002; Jones 1995; Lorenzi 2010

Bactris gastoniana Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Bactris glandulosa Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002

Bactris glassmanii Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Bactris glaucescens Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Bactris gracilior Henderson et al. 1995; Henderson 2002

Bactris grayumii Henderson et al. 1995; Henderson 2002

Bactris guineensis Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002

Bactris hatschbachii Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Bactris hirta AAU Herbarium; Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Bactris hondurensis Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Bactris horridispatha Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Bactris killipii AAU Herbarium; Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Bactris kunorum Henderson et al. 1995; Henderson 2002

Bactris longiseta Henderson et al. 1995; Henderson 2002

Bactris macroacantha Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Bactris major AAU Herbarium; Henderson et al. 1995; Henderson 2002; Jones 1995

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Bactris maraja AAU Herbarium; Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Bactris mexicana Henderson et al. 1995; Henderson 2002

Bactris militaris Henderson et al. 1995; Henderson 2002

Bactris oligocarpa Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Bactris oligoclada Henderson et al. 1995; Henderson 2002

Bactris panamensis Henderson et al. 1995; Henderson 2002

Bactris pickelii Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Bactris pilosa Henderson et al. 1995; Henderson 2002

Bactris pliniana Henderson et al. 1995; Henderson 2002

Bactris plumeriana Henderson et al. 1995; Henderson 2002

Bactris ptariana Henderson et al. 1995; Henderson 2002

Bactris rhaphidacantha Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Bactris riparia Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Bactris setosa Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Bactris setulosa Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Bactris simplicifrons Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Bactris soeiroana Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Bactris syagroides AAU Herbarium; Henderson et al. 1995; Henderson 2002

Bactris tefensis Henderson et al. 1995; Henderson 2002

Bactris tomentosa Henderson et al. 1995; Henderson 2002

Bactris turbinocarpa Henderson et al. 1995; Henderson 2002

Bactris vulgaris Henderson et al. 1995; Henderson 2002

Barcella odora Henderson et al. 1995; Henderson 2002

Brahea aculeata Henderson et al. 1995; Henderson 2002; Jones 1995

Brahea armata Henderson et al. 1995; Henderson 2002

Brahea brandegeei Henderson et al. 1995; Henderson 2002; Jones 1995

Brahea calcarea Henderson et al. 1995; Henderson 2002

Brahea decumbens Henderson et al. 1995; Henderson 2002; Jones 1995

Brahea dulcis Henderson et al. 1995; Henderson 2002

Brahea edulis Henderson et al. 1995; Henderson 2002; Jones 1995

Brahea moorei Henderson et al. 1995; Henderson 2002; Jones 1995

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Brahea pimo Henderson et al. 1995; Henderson 2002; Jones 1995

Butia archeri Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Butia campicola Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Butia capitata Henderson et al. 1995; Henderson 2002; Jones 1995; Lorenzi 2010

Butia eriospatha Henderson et al. 1995; Henderson 2002; Jones 1995; Lorenzi 2010

Butia leptospatha Henderson et al. 1995; Henderson 2002

Butia microspadix Henderson et al. 1995; Henderson 2002; Jones 1995; Lorenzi 2010

Butia paraguayensis Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Butia purpurascens Henderson et al. 1995; Henderson 2002; Jones 1995; Lorenzi 2010

Butia yatay AAU Herbarium; Henderson et al. 1995; Henderson 2002; Jones 1995; Lorenzi 2010

Calyptrogyne allenii Henderson 2002

Calyptrogyne anomala Henderson 2002

Calyptrogyne condensata Henderson 2002

Calyptrogyne costatifrons Galeano and Bernal, 2010; Henderson 2002

Calyptrogyne ghiesbreghtiana Henderson 2002

Calyptrogyne kunorum Henderson 2005

Calyptrogyne pubescens Henderson 2002

Calyptrogyne trichostachys Henderson 2002

Calyptronoma occidentalis Henderson 2002; palmpedia

Calyptronoma plumeriana Henderson 2002; Jones 1995

Calyptronoma rivalis Henderson 2002; Jones 1995

Ceroxylon alpinum Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Ceroxylon amazonicum Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Ceroxylon ceriferum Henderson et al. 1995; Henderson 2002

Ceroxylon echinulatum Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Ceroxylon parvifrons Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Ceroxylon parvum AAU Herbarium; Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Ceroxylon quindiuense Henderson et al. 1995; Henderson 2002

Ceroxylon sasaimae Henderson et al. 1995; Henderson 2002

Ceroxylon ventricosum Henderson et al. 1995; Henderson 2002

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Ceroxylon vogelianum Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Ceroxylon weberbaueri Henderson et al. 1995; Henderson 2002

Chamaedorea adscendens Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea allenii Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Jones 1995

Chamaedorea amabilis Henderson et al. 1995; Henderson 2002; Jones 1995

Chamaedorea angustisecta AAU Herbarium; Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Jones 1995

Chamaedorea arenbergiana Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Jones 1995

Chamaedorea brachyclada Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Jones 1995

Chamaedorea brachypoda Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Jones 1995

Chamaedorea carchensis Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea castillo-montii Henderson et al. 1995; Hodel (1992a,b)

Chamaedorea cataractarum Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Jones 1995

Chamaedorea correae Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Jones 1995

Chamaedorea costaricana Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea dammeriana Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea deckeriana Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Jones 1995

Chamaedorea deneversiana Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea elatior Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Jones 1995

Chamaedorea elegans Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Jones 1995

Chamaedorea ernesti-augusti Henderson et al. 1995; Hodel (1992a,b); Jones 1995

Chamaedorea fractiflexa Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea fragrans Henderson et al. 1995; Hodel (1992a,b); Jones 1995

Chamaedorea geonomiformis Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea glaucifolia Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Jones 1995

Chamaedorea graminifolia Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Jones 1995

Chamaedorea guntheriana Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea hooperiana Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Jones 1995

Chamaedorea ibarrae Henderson et al. 1995; Henderson 2002; Hodel (1992b)

Chamaedorea keelerorum Henderson et al. 1995; Hodel (1992b)

Chamaedorea klotzschiana Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Jones 1995

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Chamaedorea lehmannii Henderson et al. 1995; Hodel (1992a,b)

Chamaedorea liebmannii Henderson et al. 1995; Hodel (1992a,b)

Chamaedorea linearis AAU Herbarium; Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea lucidifrons Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea macrospadix Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea metallica Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Jones 1995

Chamaedorea microphylla Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea microspadix Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Jones 1995

Chamaedorea murriensis Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea nationsiana Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea nubium Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Jones 1995

Chamaedorea oblongata Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Jones 1995

Chamaedorea oreophila Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Jones 1995

Chamaedorea pachecoana Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea palmeriana Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea parvifolia Henderson et al. 1995; Hodel (1992a,b)

Chamaedorea parvisecta Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea pauciflora AAU Herbarium; Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Borchsenius et al. 1998

Chamaedorea pinnatifrons AAU Herbarium; Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Borchsenius et al. 1998; Jones 1995

Chamaedorea pittieri Henderson 2002; Hodel (1992a,b)

Chamaedorea plumosa Henderson et al. 1995; Henderson 2002

Chamaedorea pochutlensis Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Jones 1995

Chamaedorea pumila Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea pygmaea Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea queroana Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea radicalis Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Jones 1995

Chamaedorea rhizomatosa Henderson et al. 1995; Hodel (1992a,b)

Chamaedorea rigida Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea robertii Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea rojasiana Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

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Chamaedorea sartorii Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Jones 1995

Chamaedorea scheryi Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea schiedeana Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Jones 1995

Chamaedorea seifrizii Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea selvae Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea simplex Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea stolonifera Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Jones 1995

Chamaedorea stricta Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea tenerrima Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea tepejilote Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Jones 1995

Chamaedorea tuerckheimii Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Jones 1995

Chamaedorea undulatifolia Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Jones 1995

Chamaedorea verecunda Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea volcanensis Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea vulgata Henderson et al. 1995; Henderson 2002; Hodel (1992a,b)

Chamaedorea warscewiczii Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Jones 1995

Chamaedorea whitelockiana Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Jones 1995

Chamaedorea woodsoniana Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002; Hodel (1992a,b); Jones 1995

Chelyocarpus chuco Henderson 2002; palmweb

Chelyocarpus dianeurus Henderson et al. 1995; Henderson 2002

Chelyocarpus repens Henderson et al. 1995; Henderson 2002; palmweb

Chelyocarpus ulei Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Coccothrinax argentata Henderson et al. 1995; Henderson 2002; Jones 1995

Coccothrinax argentea Henderson et al. 1995; Henderson 2002; Jones 1995

Coccothrinax barbadensis Henderson et al. 1995; Henderson 2002; Jones 1995

Coccothrinax borhidiana Henderson et al. 1995; Henderson 2002; Jones 1995

Coccothrinax crinita Henderson et al. 1995; Henderson 2002; Jones 1995

Coccothrinax ekmanii Henderson et al. 1995; Henderson 2002; Jones 1995

Coccothrinax gracilis Henderson et al. 1995; Henderson 2002

Coccothrinax gundlachii Henderson et al. 1995; Henderson 2002

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Coccothrinax hioramii Henderson et al. 1995; palmpedia

Coccothrinax inaguensis Henderson 2002; Jones 1995

Coccothrinax miraguama Henderson et al. 1995; Henderson 2002; Jones 1995

Coccothrinax pauciramosa Henderson et al. 1995; Henderson 2002

Coccothrinax salvatoris Henderson et al. 1995; Henderson 2002

Coccothrinax spissa Henderson et al. 1995; Henderson 2002; Jones 1995

Colpothrinax cookii Henderson et al. 1995; Henderson 2002

Colpothrinax wrightii Henderson et al. 1995; Henderson 2002

Copernicia alba AAU Herbarium; Henderson et al. 1995; Henderson 2002; Jones 1995

Copernicia baileyana Henderson et al. 1995; Henderson 2002; Jones 1995

Copernicia berteroana Henderson et al. 1995; Henderson 2002; Jones 1995

Copernicia brittonorum Henderson et al. 1995; Henderson 2002

Copernicia cowellii Henderson et al. 1995; Henderson 2002; Jones 1995

Copernicia ekmanii Henderson et al. 1995; Henderson 2002; Jones 1995

Copernicia gigas Henderson et al. 1995; Henderson 2002; Jones 1995

Copernicia glabrescens Henderson et al. 1995; Henderson 2002; Jones 1995

Copernicia hospita Henderson et al. 1995; Henderson 2002; Jones 1995

Copernicia macroglossa Henderson et al. 1995; Henderson 2002

Copernicia prunifera Henderson et al. 1995; Henderson 2002; Dransfield 1986

Copernicia rigida Henderson et al. 1995; Henderson 2002

Copernicia tectorum Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002

Cryosophila cookii Henderson et al. 1995; Henderson 2002

Cryosophila grayumii Henderson et al. 1995; Henderson 2002

Cryosophila guagara Henderson et al. 1995; Henderson 2002

Cryosophila kalbreyeri Henderson et al. 1995; Henderson 2002

Cryosophila macrocarpa Henderson et al. 1995; Henderson 2002

Cryosophila nana Henderson et al. 1995; Henderson 2002

Cryosophila stauracantha Henderson et al. 1995; Henderson 2002

Cryosophila warscewiczii Henderson et al. 1995; Henderson 2002

Cryosophila williamsii Henderson et al. 1995; Henderson 2002

Desmoncus cirrhiferus Borchsenius et al. 1998; Galeano and Bernal, 2010; Henderson 2002

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Desmoncus giganteus Borchsenius et al. 1998; Henderson 2002

Desmoncus mitis AAU Herbarium; Borchsenius et al. 1998; Henderson 2002

Desmoncus orthacanthos Borchsenius et al. 1998; Henderson 2002

Desmoncus phoenicocarpus Henderson 2002; Lorenzi 2010

Desmoncus polyacanthos AAU Herbarium; Borchsenius et al. 1998; Henderson 2002

Desmoncus stans Henderson 2002; palmweb

Dictyocaryum fuscum Henderson 1990; Henderson et al. 1995; Henderson 2002

Dictyocaryum lamarckianum AAU Herbarium; Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002; Henderson 1990

Dictyocaryum ptarianum Henderson et al. 1995; Henderson 2002; Henderson 1990

Elaeis oleifera Borchsenius et al. 1998; Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002

Euterpe broadwayi Henderson et al. 1995; Henderson 2002; palmweb

Euterpe catinga Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Euterpe edulis Henderson et al. 1995; Henderson 2002

Euterpe longibracteata Henderson et al. 1995; Lorenzi 2010

Euterpe luminosa Henderson et al. 1995; Henderson 2002

Euterpe oleracea Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Euterpe precatoria AAU Herbarium; Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Gaussia attenuata Henderson et al. 1995; Henderson 2002

Gaussia gomez-pompae Henderson et al. 1995; palmpedia

Gaussia maya AAU Herbarium; Henderson 2002

Gaussia princeps Henderson et al. 1995; Henderson 2002

Gaussia spirituana Henderson et al. 1995; Henderson 2002

Geonoma appuniana AAU Herbarium; Henderson 2002

Geonoma arundinacea Borchsenius et al. 1998; Henderson 2002

Geonoma aspidiifolia Henderson 2002; Lorenzi 2010

Geonoma baculifera AAU Herbarium; Henderson 2002; Lorenzi 2010

Geonoma brevispatha AAU Herbarium; Henderson 2002; Lorenzi 2010

Geonoma brongniartii AAU Herbarium; Borchsenius et al. 1998; Henderson 2002; Lorenzi 2010

Geonoma camana AAU Herbarium; Borchsenius et al. 1998; Henderson 2002; Lorenzi 2010

Geonoma chlamydostachys Galeano and Bernal, 2010; Henderson 2002

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Geonoma chococola Henderson 2002; palmweb

Geonoma concinna Galeano and Bernal, 2010; Henderson 2002

Geonoma congesta Borchsenius et al. 1998; Henderson 2002

Geonoma cuneata Borchsenius et al. 1998; Henderson 2002

Geonoma densa Borchsenius et al. 1998; Henderson 2002

Geonoma deversa AAU Herbarium; Borchsenius et al. 1998; Henderson 2002; Lorenzi 2010; Galeano and Bernal, 2010

Geonoma divisa Galeano and Bernal, 2010; Henderson 2002

Geonoma epetiolata Henderson 2002; palmweb

Geonoma ferruginea Henderson 2002; palmweb

Geonoma gamiova Henderson 2002

Geonoma interrupta AAU Herbarium; Borchsenius et al. 1998; Galeano and Bernal, 2010; Henderson 2002

Geonoma jussieuana Borchsenius et al. 1998; Galeano and Bernal, 2010; Henderson 2002

Geonoma laxiflora Borchsenius et al. 1998; Henderson 2002; Lorenzi 2010

Geonoma leptospadix AAU Herbarium; Borchsenius et al. 1998; Henderson 2002; Lorenzi 2010

Geonoma linearis Borchsenius et al. 1998; Henderson 2002

Geonoma longipedunculata Borchsenius et al. 1998; Galeano and Bernal, 2010

Geonoma longivaginata palmpedia; palmweb

Geonoma macrostachys Borchsenius et al. 1998; Henderson 2002; Lorenzi 2010; Galeano and Bernal, 2010

Geonoma maxima AAU Herbarium; Borchsenius et al. 1998; Henderson 2002; Lorenzi 2010; Galeano and Bernal, 2010

Geonoma myriantha Henderson 2002; Lorenzi 2010

Geonoma oldemanii AAU Herbarium; Henderson 2002

Geonoma orbignyana AAU Herbarium; Borchsenius et al. 1998; Henderson 2002

Geonoma paradoxa Borchsenius et al. 1998; Galeano and Bernal, 2010; Henderson 2002

Geonoma paraguanensis Henderson 2002

Geonoma pauciflora Henderson 2002; Lorenzi 2010

Geonoma poeppigiana Borchsenius et al. 1998; Henderson 2002

Geonoma pohliana Henderson 2002; Lorenzi 2010

Geonoma polyandra AAU Herbarium; Borchsenius et al. 1998; Henderson 2002

Geonoma rubescens Henderson 2002; Lorenzi 2010

Geonoma schottiana Borchsenius et al. 1998; Henderson 2002; Lorenzi 2010

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Geonoma scoparia Henderson 2002; palmweb

Geonoma simplicifrons AAU Herbarium; Henderson 2002; palmweb

Geonoma spinescens Henderson 2002; palmweb

Geonoma stricta AAU Herbarium; Borchsenius et al. 1998; Henderson 2002; Lorenzi 2010

Geonoma tenuissima Borchsenius et al. 1998; Henderson 2002

Geonoma triandra Galeano and Bernal, 2010; Henderson 2002

Geonoma triglochin AAU Herbarium; Borchsenius et al. 1998; Henderson 2002

Geonoma trigona Henderson 2002; palmweb

Geonoma umbraculiformis AAU Herbarium; Henderson 2002

Geonoma undata AAU Herbarium; Borchsenius et al. 1998; Henderson 2002

Geonoma weberbaueri AAU Herbarium; Borchsenius et al. 1998; Henderson 2002

Hemithrinax compacta Henderson et al. 1995

Hemithrinax ekmaniana palmpedia

Hemithrinax rivularis palmpedia

Hyospathe elegans AAU Herbarium; Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Hyospathe macrorhachis Borchsenius et al. 1998; Henderson et al. 1995

Iriartea deltoidea AAU Herbarium; Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002; Henderson 1990

Iriartella setigera AAU Herbarium; Henderson et al. 1995; Henderson 2002; Henderson 1990

Iriartella stenocarpa Henderson et al. 1995; Henderson 2002; Henderson 1990

Itaya amicorum Henderson et al. 1995; Henderson 2002

Juania australis Henderson et al. 1995; Henderson 2002

Jubaea chilensis Henderson et al. 1995; Henderson 2002

Leopoldinia major Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Leopoldinia piassaba Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Leopoldinia pulchra Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Lepidocaryum tenue AAU Herbarium; Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Leucothrinax morrisii palmpedia

Lytocaryum hoehnei Henderson et al. 1995; Henderson 2002

Lytocaryum weddellianum Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Manicaria saccifera Borchsenius et al. 1998; Henderson 2002; Lorenzi 2010

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Mauritia carana Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002

Mauritia flexuosa AAU Herbarium; Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Mauritiella aculeata Henderson et al. 1995; Henderson 2002

Mauritiella armata AAU Herbarium; Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Mauritiella macroclada Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Neonicholsonia watsonii Henderson et al. 1995; Henderson 2002

Oenocarpus bacaba AAU Herbarium; Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Oenocarpus balickii AAU Herbarium; Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Oenocarpus bataua AAU Herbarium; Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Oenocarpus circumtextus Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Oenocarpus distichus Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Oenocarpus makeru Henderson et al. 1995; Henderson 2002

Oenocarpus mapora AAU Herbarium; Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Oenocarpus minor AAU Herbarium; Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Oenocarpus simplex Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Parajubaea cocoides Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Parajubaea torallyi AAU Herbarium; Henderson et al. 1995; Henderson 2002

Pholidostachys dactyloides Borchsenius et al. 1998; Henderson 2002

Pholidostachys kalbreyeri Galeano and Bernal, 2010; Henderson 2002

Pholidostachys pulchra AAU Herbarium; Henderson 2002

Pholidostachys synanthera Borchsenius et al. 1998; Henderson 2002

Phytelephas aequatorialis Borchsenius et al. 1998; Henderson 2002

Phytelephas macrocarpa AAU Herbarium; Borchsenius et al. 1998; Galeano and Bernal, 2010; Henderson 2002

Phytelephas schottii AAU Herbarium

Phytelephas seemannii Henderson 2002; palmweb

Phytelephas tenuicaulis Borchsenius et al. 1998; Galeano and Bernal, 2010; Henderson 2002

Phytelephas tumacana Galeano and Bernal, 2010; Henderson 2002

Prestoea acuminata AAU Herbarium; Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Prestoea carderi Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Prestoea decurrens Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

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Prestoea ensiformis Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Prestoea longipetiolata Henderson et al. 1995; palmpedia

Prestoea pubens Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Prestoea pubigera Henderson et al. 1995; Henderson 2002

Prestoea schultzeana Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Prestoea simplicifolia Henderson et al. 1995; Henderson 2002

Prestoea tenuiramosa Henderson et al. 1995; Henderson 2002

Pseudophoenix ekmanii Henderson et al. 1995; Henderson 2002

Pseudophoenix lediniana Henderson et al. 1995; Henderson 2002

Pseudophoenix sargentii Henderson et al. 1995; Henderson 2002

Pseudophoenix vinifera Henderson et al. 1995; Henderson 2002

Raphia taedigera Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002; Lorenzi 2010; Tuley 1995

Reinhardtia elegans Henderson et al. 1995; Henderson 2002

Reinhardtia gracilis AAU Herbarium; Henderson et al. 1995; Henderson 2002

Reinhardtia koschnyana Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002

Reinhardtia latisecta AAU Herbarium; Henderson et al. 1995; Henderson 2002

Reinhardtia paiewonskiana Henderson et al. 1995; Henderson 2002

Reinhardtia simplex Henderson et al. 1995; Henderson 2002

Rhapidophyllum hystrix Henderson et al. 1995; Henderson 2002

Roystonea altissima Henderson et al. 1995; Henderson 2002

Roystonea borinquena AAU Herbarium; Henderson et al. 1995; Henderson 2002

Roystonea dunlapiana Henderson et al. 1995; Henderson 2002

Roystonea lenis Henderson et al. 1995; Henderson 2002

Roystonea maisiana Henderson et al. 1995; Henderson 2002

Roystonea oleracea Henderson et al. 1995; Henderson 2002

Roystonea princeps Henderson et al. 1995; Henderson 2002

Roystonea regia Henderson et al. 1995; Henderson 2002

Roystonea stellata Henderson et al. 1995; Henderson 2002

Roystonea violacea Henderson et al. 1995; Henderson 2002

Sabal bermudana Henderson et al. 1995; Henderson 2002; Zona 1990

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Sabal causiarum Henderson et al. 1995; Henderson 2002; Zona 1990

Sabal domingensis Henderson et al. 1995; Henderson 2002; Zona 1990

Sabal etonia Henderson et al. 1995; Henderson 2002; Zona 1990

Sabal gretherae Henderson et al. 1995; palmweb

Sabal maritima Henderson et al. 1995; Henderson 2002; Zona 1990

Sabal mauritiiformis Henderson et al. 1995; Henderson 2002; Zona 1990

Sabal mexicana Henderson et al. 1995; Henderson 2002; Zona 1990

Sabal miamiensis Henderson et al. 1995; Henderson 2002; Zona 1990

Sabal minor Henderson et al. 1995; Henderson 2002; Zona 1990

Sabal palmetto Henderson et al. 1995; Henderson 2002; Zona 1990

Sabal pumos Henderson et al. 1995; Henderson 2002; Zona 1990

Sabal rosei Henderson et al. 1995; Henderson 2002; Zona 1990

Sabal uresana Henderson et al. 1995; Henderson 2002; Zona 1990

Sabal yapa Henderson et al. 1995; Henderson 2002; Zona 1990

Schippia concolor AAU Herbarium; Henderson et al. 1995; Henderson 2002

Serenoa repens AAU Herbarium; Henderson et al. 1995; Henderson 2002

Socratea exorrhiza AAU Herbarium; Borchsenius et al. 1998; Henderson 1990; Henderson et al. 1995; Henderson 2002

Socratea hecatonandra Henderson et al. 1995; Henderson 2002; Henderson 1990

Socratea montana Henderson et al. 1995; Henderson 2002; Henderson 1990

Socratea rostrata Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002; Henderson 1990

Socratea salazarii Henderson et al. 1995; Henderson 2002; Henderson 1990

Syagrus amara Henderson et al. 1995; Henderson 2002

Syagrus botryophora Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Syagrus campylospatha Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Syagrus cardenasii AAU Herbarium; Henderson et al. 1995; Henderson 2002

Syagrus cocoides Henderson et al. 1995; Henderson 2002

Syagrus comosa Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Syagrus coronata Henderson et al. 1995; Henderson 2002

Syagrus duartei Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Syagrus flexuosa Henderson et al. 1995; Henderson 2002; Lorenzi 2010

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Syagrus glaucescens Henderson et al. 1995; Henderson 2002

Syagrus graminifolia Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Syagrus harleyi Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Syagrus inajai AAU Herbarium; Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Syagrus macrocarpa Henderson et al. 1995; Henderson 2002

Syagrus microphylla Henderson et al. 1995; Henderson 2002

Syagrus oleracea Henderson et al. 1995; Henderson 2002

Syagrus orinocensis Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Syagrus petraea AAU Herbarium; Henderson et al. 1995; Henderson 2002

Syagrus picrophylla Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Syagrus pleioclada Henderson et al. 1995; Henderson 2002

Syagrus pseudococos Henderson et al. 1995; Henderson 2002

Syagrus romanzoffiana Henderson et al. 1995; Henderson 2002

Syagrus ruschiana Henderson et al. 1995; Henderson 2002

Syagrus sancona AAU Herbarium; Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Syagrus schizophylla Henderson et al. 1995; Henderson 2002

Syagrus smithii AAU Herbarium; Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Syagrus stratincola Henderson et al. 1995; Henderson 2002

Syagrus vagans Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Syagrus werdermannii Henderson et al. 1995; Henderson 2002

Synechanthus fibrosus Henderson et al. 1995; Henderson 2002

Synechanthus warscewiczianus AAU Herbarium Database; Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Thrinax excelsa Henderson et al. 1995; Henderson 2002

Thrinax parviflora AAU Herbarium; Henderson et al. 1995

Thrinax radiata Henderson et al. 1995; Henderson 2002

Trithrinax brasiliensis Henderson et al. 1995; Henderson 2002

Trithrinax campestris Henderson et al. 1995; Henderson 2002

Trithrinax schizophylla Henderson et al. 1995; Henderson 2002

Washingtonia filifera Henderson et al. 1995; Henderson 2002

Washingtonia robusta Henderson et al. 1995; Henderson 2002

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Welfia regia Borchsenius et al. 1998; Galeano and Bernal, 2010; Henderson 2002

Wendlandiella gracilis Henderson et al. 1995; Henderson 2002; Lorenzi 2010

Wettinia aequalis Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Wettinia aequatorialis Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Wettinia anomala Borchsenius et al. 1998; Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002

Wettinia augusta AAU Herbarium; Henderson et al. 1995; Henderson 2002; AAU Herbarium Database; Lorenzi 2010; Galeano and Bernal, 2010

Wettinia castanea Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002

Wettinia disticha Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002

Wettinia drudei AAU Herbarium; Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Wettinia fascicularis Borchsenius et al. 1998; Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002

Wettinia hirsuta Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002

Wettinia kalbreyeri Borchsenius et al. 1998; Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002

Wettinia lanata Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002

Wettinia longipetala Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Wettinia maynensis Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Wettinia microcarpa Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002

Wettinia minima Borchsenius et al. 1998; Henderson et al. 1995; Henderson 2002

Wettinia oxycarpa Borchsenius et al. 1998; Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002

Wettinia panamensis Henderson et al. 1995; Henderson 2002

Wettinia praemorsa Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002

Wettinia quinaria Borchsenius et al. 1998; Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002

Wettinia radiata Borchsenius et al. 1998; Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002

Wettinia verruculosa Borchsenius et al. 1998; Galeano and Bernal, 2010; Henderson et al. 1995; Henderson 2002

Zombia antillarum AAU Herbarium; Henderson et al. 1995; Henderson 2002

References:

AAU Herbarium: The Aarhus University Herbarium (AAU), Ole Worms Allé 1, 8000 Aarhus C, Denmark

Borchsenius F, Pedersen HB, Balslev H. 1998. Manual to the palms of Ecuador. AAU Reports 37, Aarhus University Press, Aarhus.

Galeano G, Bernal R. 2010. Palmas de Colombia: guia de campo. Panamericana Formas e Impresos S.A., Bogota.

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Henderson A, 1990. Arecaceae Part I. Introduction and the Iriarteinae. Flora Neotropica 53: 1-101.

Henderson A, 1995: Field Guide to the Palms of the Americas.

Henderson A, 2002. Evolution and ecology of palms.

Hodel DR, 1992. Chamaedorea palms. The International Palm Society, Allen Press, Lawrence, Kansas.

Hodel DR. 1992. Additons to Chamaedorea palms: new species from Mexico and Guatemala and miscellaneous notes. Principes 36: 188-202.

Jones DL. 1995. Palms throughout the world. Smithsonian Institution Press, Washington, D.C.

Lorenzi H. 2010. Brazilian Flora Arecaceae (Palms). Nova Odessa, Instituto Plantarum.

Palmweb (http://palmweb.org/)

Palmpedia (http://www.palmpedia.net/)

Tuley P. 1995. The palms of Africa. The Trendrine Press, Zennor.

Zona S. 1990. A monograph of Sabal (Arecaceae: Coryphoideae). ALISO 12: 583-666.

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