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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 --
739740741742743744745746747748749750
751
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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(a) Residuals median leaf size
geographical x-coordinates
geog
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y-c
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-30
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(b) Residuals median stem height
geographical x-coordinates
geog
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y-c
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positive SACnegative SAC
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-30
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(c) Residuals median fruit size
geographical x-coordinates
geog
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(d) Residuals median leaf size
geographical x-coordinates
geog
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y-c
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-30
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(e) Residuals median stem height
geographical x-coordinates
geog
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(f) Residuals median fruit size
geographical x-coordinates
geog
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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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827
828
829830
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
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
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835
836
837
838
839
840
841
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
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
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
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
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
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
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
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)
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
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
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
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
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
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
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
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
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
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.
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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