17.

18.0

18.5

19.0

LINKING GEOGRAPHIC DISTRIBUTIONS AND FUNCTIONAL TRAITS:

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INSIGHTS FROM PUERTO RICAN FORESTS

19.0

Carite

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El Yunque

1.0

1.5

2.0

2.5

3.0

3.5

4.0

1500

1000

1500

2000

1000

Mean annual precipitation (mm yr!1 )

1500

2000

30
25
20
15

Frequency

10
5

Frequency

0.0

1.0

-1.0

-0.5

0.0

0.5

Slopes

0.8

0.4
0.1
0.0

30
20

10
8
6

Negative
slopes

1000

1500

0.0

xx

References:

Shipley (2010) From plant traits to vegeta0on structure: chance and selec0on in the assembly of
ecological communi0es. Cambridge University Press, New York.
Muscarella et al. (2014) ENMeval: An R package for conduc@ng spa@ally independent evalua@ons
and es@ma@ng op@mal model complexity for Maxent ecological niche models. Methods in Ecology
and Evolu0on. 5(11): 1198-1205.
Phillips et al. (2006) Maximum entropy modeling of species geographic distribu@ons. Ecological
Modelling. 190: 231-259.

We extend our gra@tude to the Departamento de Recursos Naturales y Ambiente for permission to
establish permanent plots in the forests of Puerto Rico under DRNA permit #2011-IC-046. This work

-66.0

SF DEB 1050957 to MU and NSF DEB 1311367 to MU and RM.

-65.5was supported by N-65.0

10

60

CWMpi

0.2

Merging func@onal community ecology with biogeography represents an important research direc@on
for improving our ability to predict the consequences of environmental change on biodiversity. Our
study represents a key step for linking local community structure with species geographic
distribu@ons.
-3
-2
-1
0
1
2
3

Acknowledgements:

0:10

-66.5

0.4

0.3
yy

0.3
yy

0.2
0.1

120

Relative
habitat
suitability
(ENMs)

80

abs(CWMp ti)
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=

Selec@on towards the CWM for dierent traits might promote local diversity, depending on how trait
axes are correlated. In ongoing work on this project, we are evalua@ng how trait axes are correlated at
dierent spa@al scales (among co-occurring species vs. between sites) as a way to further understand
the processes driving func@onal paYerns of community diversity. Addi@onally, we are incorpora@ng
demographic informa@on to confront poten@al limita@ons of correla@ve niche models. Finally, we are
con@nuing to expand our trait database to incorporate intraspecic trait varia@on as well as more
rened physiological traits.

0.0

50 km

CWM

100

0.9
0.8
0.7

R 2 = 0.93

0.6

17.5

17.5

30
20
5

10

H max (m)

200 350
100
50
25

2000

e.g.,
Species with
WD = 0.8

CWM

R 2 = 0.69

-1.0

OLS sSlopes
lope

For many more species than randomly expected, es@mated habitat suitability was lower for species
when they had trait values more distant from the local CWM trait value. This result supports for the
CWM-op@mality hypothesis and suggests that broad-scale environmental lters act to constrain local
func@onal diversity towards the CWM. However, a lesser number of species (although s@ll
signicantly more than randomly expected) had posi@ve slopes and thus appear to exploit successful
alterna@ve strategies.

0.4
1000

LMA (g m!2 )

1.0
0.8
0.6
0.4

Wood density (g cm!3 )

R 2 = 0.75

e.g., Wood density (g cm-3)

-66.0

-2.0

The tendency for func@onal traits to vary predictably along clima@c gradients suggests trait-mediated
tness dierences that act to constrain local func@onal diversity. However, the typically large
propor@on of within site trait varia@on (high func@onal varia@on within sites) implies the ac@on of
mechanisms that maintain local func@onal diversity. Reconciling these contras@ng paYerns represents
a major challenge for predic@on in community ecology.

-5suitability
0 at each
5 study
10 plot
Separately for each species and each trait, we compared es@mates o-10
f habitat
with the absolute dierence between the species-mean trait value and the CWM at each plot. Under the
xx
CWM-op@mality hypothesis, we predicted a nega@ve rela@onship between these values. We used OLS
regression to determine the sign and signicance of these rela@onships for each species-trait
combina@on and compared the results with a null model.

Relative basal area

of species i

-66.5

0.00020

i=1

0.00015

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Trait value of
species i

= pi trait i

Analyses:

High

Relative estimated
habitat suitability

0.00010

Community-weighted
mean trait value
(CWM)

Low

0.00005

For all 173 species, we measured three func@onal traits: WD = wood density [g cm-3], LMA = leaf mass per
area [g m-2], and Hmax= maximum height [m]. We calculated the community-weighted mean trait (CWM)
value in each plot based on species rela@ve basal area and species-mean trait values. CWM values for all
three traits varied signicantly with respect to mean annual precipita@on. In the gures below, red circles
( ) represent CWM values and grey points ( ) represent individual species .

1000

0.6

50 km

Func0onal traits:

77%

of signicant
slopes are
nega0ve

Conclusions, signicance, and future direc0ons:

p a c k a g e

Regularization Multiplier
Regularization
Multiplier

2000

18.0

18.0

Guanica

-2
Slopes

3000

Rio Abajo

81%

of signicant
slopes are
nega0ve

Maximum height
(m)
MAXHT

1.0

0.5

4000

LMA
(g
m-2)
LMA

Feature classes

-4

18.5

Mean
annual
precipitation
(mm yr!1)

18.5

ENM
e v a l

10 15 20 25 30 35

Frequency
Frequency

Soil parent material (categorical)

84%

of signicant
slopes are
nega0ve

80

CV of monthly
precipita0on (unitless)

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4000

3000

2000

1000

Cambalache Vega

L
LQ
H
LQH
LQHP
LQHPT

Minimum monthly
temperature (C)

19.0

Toro Negro

Mean daily temperature

range (C)

10 15 20 25 30

= p > 0.05 Wood density

-3
(g
WDcm )

Mean annual precipita@on (mm yr -1)

Guajataca

= p < 0.05

Mean annual precipita0on

(mm yr-1)

Cambalache / Vega

0.0

HERBARIUM COLLECTION LOCALITIES (N = 17,479)

60

yy

0.1

-66.5

Guilarte

-66.5

19.0

INPUT LAYERS

We established 12, 50m x 50m forest plots in 4 protected areas of Puerto Rico. These plots occur along a
precipita@on gradient that ranges from ca. 900 to 2,500 mm yr-1 on limestone soils. In each plot, we
counted, iden@ed, and measured all woody stems > 1 cm at 1.3 m above the ground. The plots
contained a total of 173 species.
1000 2000 3000 4000

Of the 173 total species, slightly more than half had signicant rela@onships (i.e., p < 0.05; both
posi@ve and nega@ve) between habitat suitability and CWM. These were many more than
predicted by random chance (null model not shown). Of the species with signicant rela@onships,
about 80% (for each trait) had nega@ve slopes, in line with the CWM-op@mality hypothesis.

We used the R package ENMeval (Muscarella et al. 2014) to conduct species-specic tuning of
environmental niche models (ENMs) in MAXENT (Phillips et al. 2006) for each of 173 species. We used
AICc to average models based on sefngs that balance model t and predic@ve ability. We used the
target group background approach to account for spa@al bias in collec@on locali@es.

Rio Abajo

0.2

Guanica

-67.0

Forest plots:

Contact: bob.muscarella@gmail.com

Results:

Environmental Niche Models:

40

0.3

Guajataca

Func@onal traits can provide physiological links between

species distribu@ons, abio@c gradients, and paYerns of
community diversity. One hypothesis that underpins
many models of plant community assembly based on
Relative
func@onal traits is that if community-weighted mean trait
fitness
values (i.e., plot-level trait values weighted by species
abundance; CWMs) vary predictably along an abio@c
gradient, then the local CWM reects the op@mal trait
value given the environmental condi@ons at that site (e.g.,
CWM
Shipley 2010). In other words, species with trait values
nearest to the CWM in a par@cular loca@on are expected
to have high tness because they occur at rela@vely high
-3
-2Species
-1
0
1
2
3
trait
value
abundance and contribute strongly to the local CWM. This
CWM-op@mality hypothesis is consistent with niche par@@oning along resource gradients as a driver of
xx
community diversity and it emphasizes processes that constrain local func@onal diversity (e.g.,
environmental ltering). Although dicult to evaluate in natural systems, ecological niche models
(ENMs) represent an unexploited opportunity to examine the core predic@on of the CWM-op@mality
hypothesis: for a given species, es0mated habitat suitability should be nega0vely

related to the dierence between its trait value and the local CWM.

ECOLOGY, EVOLUTION AND ENVIRONMENTAL BIOLOGY, COLUMBIA UNIVERSITY, NEW YORK, NY USA
2DEPARTMENT OF BIOSCIENCE, AARHUS UNIVERSITY, AARHUS, DENMARK

AICc
delta.AICc

18.5

0.4

Introduc0on:

18.0

OF

20

17.5

ROBERT MUSCARELLA1,2 AND MARA URIARTE1

0.2

1DEPARTMENT

2000

Mean Annual Precipitation

(mm yr-1)

1000

0 1500
0

4 20006

CWMpi
10:0

Increasing 1000
8

10

1500

2000