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Received: 4 December 2018    Revised: 27 March 2020    Accepted: 31 March 2020
DOI: 10.1111/jbi.13871
RE SE ARCH PAPER
The biogeography of ecoregions: Descriptive power across
regions and taxa
Jeffrey R. Smith1,2  | J. Nicholas Hendershot1,2
Gretchen C. Daily1,2,3,4
1
Department of Biology, Stanford University,
Stanford, CA, USA
2
Center for Conservation Biology, Stanford
University, Stanford, CA, USA
3
Woods Institute for the Environment,
Stanford University, Stanford, CA, USA
4
The Natural Capital Project, Stanford
University, Stanford, CA, USA
Correspondence
Jeffrey R. Smith, Department of Biology,
Stanford University, Stanford, CA, USA.
Email: jrsmith7@stanford.edu
Funding information
Gerhard Casper and John P. Morgridge
Fellowship; Bing Fellowship in Honor of
Paul Ehrlich; Stanford Graduate Fellowship;
National Science Foundation, Grant/Award
Number: GRFP DGE-1656518; Ward Wilson
Woods Jr. Environmental Studies Fund;
Stanford Department of Biology; Stanford
University
Handling Editor: Şerban Procheş
Journal of Biogeography. 2020;47:1413–1426. wileyonlinelibrary.com/journal/jbi
 | Nicole Nova1  |
Abstract
Aim: Biomes, biogeographical realms and ecoregions have become central concepts
of biotic organization and biodiversity research. Recent data-intensive analysis has
shown that, while ecoregions do delineate biotic communities, how distinct they are
from one another varies considerably across regions and taxa. Given their central
importance to global models of the earth system and to the development of conser-
vation plans, it is key to understand in what regions, and for which taxa, ecoregion clas-
sification schemes best describe the underlying variability in biological communities.
Location: Global.
Taxa: Plants, animals, fungi.
Methods: In this paper, we integrate ecoregion maps with data on spatially continu-
ous changes in environmental conditions, biodiversity and species traits to quantify
the descriptive power of ecoregions around the globe. Capitalizing on the troves of
newly available global biodiversity data, we model the abiotic and biotic factors that
describe how distinct ecoregions are from one another.
Results: From an abiotic perspective, we report compelling evidence that, first, ecore-
gions are more distinct in tropical zones with higher temperatures, less temperature
seasonality and greater rainfall seasonality. Second, we also find that ecoregions are
more distinct in regions with steeper slopes. From a biotic perspective, we find that
ecoregions tend to be more distinct for reptiles and amphibians than they are for
mammals or birds. Likewise, ecoregions tend to be more distinct for smaller-bodied
species and, to a lesser extent, species at lower trophic levels.
Main conclusions: While ecoregion-based conservation planning can provide a crucial tool
for developing holistic conservation interventions, we show here that the ability to capture
and describe communities is not uniform across regions or taxa. In particular, ecoregions
tend to best describe communities of small-bodied species, less vagile and tropical taxa
that are typically underrepresented in the scientific literature. While ecoregion classifica-
tion schemes will continue to provide invaluable conservation guidance, we must think
critically about when an ecoregional approach is best suited to informing management.
KEYWORDS
beta diversity, biodiversity, biogeographical realms, biomes, community composition,
conservation planning, environmental gradients, life history strategies
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1414       SMITH et al.
1 |  I NTRO D U C TI O N
Ecoregions have emerged as a powerful tool for both ecological
research and conservation (Olson et  al.,  2001). From a scientific
perspective, they have illuminated fundamental processes such
as speciation, community interactions and ecosystem function
(Crowther et  al.,  2015; Di Marco, Watson, Currie, Possingham,
& Venter,  2018; Holt et  al.,  2012; Murphy et  al.,  2019; Skeels &
Cardillo, 2019). From a management standpoint, they have enabled
development of science-based conservation plans designed to pro-
tect distinct regions, habitat types and taxa. Ecoregions also pro-
vide a framework for representing ecosystem-level diversity for
international conventions (Dinerstein et al., 2017, 2019; Lamoreux
et  al.,  2006; Olson & Dinerstein,  2002; Usubiaga-Liaño, Mace, &
Ekins, 2019). There remains uncertainty, however, around the ef-
fectiveness of ecoregion-based analysis in conserving biodiversity
across regions and taxa (McDonald et  al.,  2005), highlighting a
potential shortcoming of this approach. It is imperative to deepen
our understanding of the geographies, habitats and taxa for which
ecoregions provide a useful tool in defining biological communities.
Recent work shows that while ecoregion borders are generally
reflective of community-level diversity, there is a great amount of
variability that they fail to explain (Smith et al., 2018). This is likely
because ecoregions are not intrinsic properties of the biosphere
(MacArthur,  1972). Rather, they are simply classification schemes
designed to maximize both the amount of homogeneity within (i.e.
species makeup over an entire ecoregion is relatively consistent) and
the amount of heterogeneity across ecoregions (i.e. communities
found in different ecoregions are relatively dissimilar) (Bailey, 2004).
This aim of this classification system is similar to that used at other
levels of biological organization, such as populations, operational
taxonomic units or species. Such groupings may represent emergent
properties arising from a complex suite of environmental, ecologi-
cal and evolutionary factors (Bailey,  2004, 2014; Holdridge,  1947;
Ricketts et  al.,  1999;Whittaker,  1962). Thus, our goal here is to
identify factors that influence the distinctness of neighbouring
ecoregions.
We use two sets of hypotheses to address how abiotic (hypoth-
eses a1 and a2) and biotic (hypotheses b1 and b2) gradients struc-
ture ecological communities within and between ecoregions. Our
first abiotic hypothesis (a1) is that climatic factors are the stron-
gest predictors of ecoregion distinctness. This draws on a rich lit-
erature that demonstrates the overwhelming importance of climate
on macroecological patterns of biodiversity (Echeverría-Londoño
et  al.,  2018;Pinto-Ledezma, Larkin, & Cavender-Bares,  2018;Reu
et  al.,  2011;Wang, Brown, Tang, & Fang,  2009). We also expect,
however, that topography, soil conditions and human impacts
could be strongly correlated with ecoregion distinctness, particu-
larly for certain pairings of conditions and taxa (e.g. soil conditions
for plants). Furthermore, recent work suggests that evolutionary
history plays a key role in the relationship between abiotic con-
ditions and present-day patterns of biological diversity (Antonelli
et al., 2018).
Our second abiotic hypothesis (a2) is that environmental at-
tributes associated with the tropics (higher and less variable tem-
peratures) are also linked to more distinct ecoregions. This idea
originates from Janzen (1967), who posits that communities ex-
hibit higher spatial turnover in the tropics. Because species in the
tropics experience a lesser degree of temporal heterogeneity in
abiotic niche space, they should be constricted to more narrow en-
vironmental conditions (Ghalambor, Huey, Martin, Tewksbury, &
Wang, 2006;Huey, 1978;Sheldon, Huey, Kaspari, & Sanders, 2018).
By contrast, temperate species are adapted to great fluctuations in
temperature and therefore can occupy wider geographical ranges.
This hypothesis is further supported by the longer evolutionary
histories of tropics (ceteris paribus) that may yield more strongly co-
evolved communities (Condamine, Sperling, Wahlberg, Rasplus, &
Kergoat, 2012;Mittelbach et al., 2007).
We can similarly pose two hypotheses about the biotic fac-
tors that explain variability in ecoregion distinctness (Soininen,
McDonald, & Hillebrand,  2007). Our first hypothesis (b1) is that
ecoregions are more similar to one another, at a given distance of
separation, for species with strong associations with plant commu-
nities. For example, pollinators tied to a specific type of orchid, cat-
erpillars tied to a specific herbaceous plant or mycorrhizal fungi to a
single species of tree are more likely to be found with their partners
and thus be more strongly restricted to a particular ecoregion (Du,
Chen, Jiang, & Qiao, in press;Keinath et al., 2017;Purvis, Gittleman,
Cowlishaw, & Mace, 2000).
Our second biotic hypothesis (b2) is that ecoregions are a
more useful and robust classification for smaller-bodied species
(Keinath et  al.,  2017;Purvis et al., 2000). This derives from the
clear, positive, relationship between body size, vagility and range
size (Kelt & Vuren,  1999;Swihart, Slade, & Bergstrom,  1988).
Species with small bodies, home ranges and geographical ranges
are likely to have been exposed to a much smaller range of envi-
ronmental variability (Frishkoff et  al.,  2016), as well as to fewer
potential biotic partners (Staniczenko, Sivasubramaniam, Suttle,
& Pearson,  2017). Conversely, larger-bodied species likely expe-
rience larger variation in abiotic and biotic conditions, potentially
leading to less defined geographical boundaries around their
communities.
In this paper, we analyse data on environmental conditions
together with data on biodiversity (taxonomic and trait dimen-
sions) to explore the abiotic and biotic variables responsible for
predicting ecoregion distinctness. We use community similarity
both within and between ecoregions, to determine which regions
are most internally homogeneous and most heterogeneous with
nearby regions. We then explore the role of environmental con-
ditions and the groups of organisms underlying the distinctness
of ecoregions. From an abiotic standpoint, we hypothesize that
ecoregion distinctness will be (a1) explained principally by climatic
patterns and will (a2) higher in tropical areas. From a biotic stand-
point, we expect ecoregions to be more distinct for species that
(b1) are in more specialized functional guilds and (b2) have smaller
body sizes.
SMITH et al.
2 | M ATE R I A L S A N D M E TH O DS
2.1 | Ecoregion map
The map of ecoregions that we use for this analysis comes from
Dinerstein et  al.  (2017), which updates the map created by Olson
et  al.  (2001). The map derives principally from patterns of rainfall,
temperature, geologic history, broad vegetative patterns and expert
opinion on community distributions (Dinerstein et  al.,  2017;Olson
et al., 2001) and has been shown to be broadly reflective of large-
scale biogeographical patterns across all macroscopic, terrestrial
taxa (Smith et al., 2018). At its finest level, this map divides the world
into 846 ecoregions (Figure S1).
2.2 | Biodiversity data
We obtained biodiversity data from the Global Biodiversity
Information Facility (Amphibians: GBIF.org, 2018;Arthropods: GBIF.
org, 2018;Birds: GBIF.org, 2018;Fungi GBIF.org, 2018;Mammals
GBIF.org, 2018;Plants: GBIF.org, 2018;Reptiles: GBIF.org, 2018).
GBIF is the world's largest online repository of biodiversity informa-
tion, with data principally stored as point locations, or the geographi-
cal coordinates where a species was observed.
All data were downloaded from the GBIF online portal on 7 May
2018 using the following search terms: Plants – Kingdom Plantae;
Arthropods – Phylum Arthropoda; Mammals – Class Mammalia;
Birds – Class Aves; Reptiles– Class Reptilia, Amphibians Amphibia
and Fungi – Kingdom Fungi. While GBIF data are an immensely use-
ful resource, they require careful cleaning to ensure that observed
patterns reflect biological patterns and not anomalies in the way
in which data were collected (Zizka et  al.,  2019). Data were then
cleaned so that the resultant dataset contained only records that
(a) were collected since 1950, (b) were recorded with proper geo-
graphical coordinates accurate to within 10 km and (c) were entered
with a data type of ‘Observation’, ‘Human observation’, ‘Machine
Observation’, ‘Specimen’, ‘Preserved Specimen’, ‘Living Specimen’
or ‘Material Sample’. The two most significant groups that were ex-
cluded for the final reason were fossil records and observations with
TA B L E 1   To analyse which abiotic
Category
characteristics correlated with
ecoregion distinctness we picked Climatic
eight representative variables that
encompassed significant components
of Earth's climate, topography, soil,
vegetation and human impacts. We
calculated the mean value of each of
these variables within each ecoregion
and performed a principal component Topographic
analysis on the resultant values. We Soil conditions
ultimately included the first four principal
Vegetative
components in our analyses
Anthropocentric
|
      1415
an unknown basis. We also excluded data points which have been
clearly flagged by GBIF as having errors in the coordinates, such as
points falling in the middle of oceans, coinciding with the geometric
centre of a country, have exactly 0.0 latitude or longitude, or have
other coordinate issues as indicated in the GBIF portal. Summary
statistics for the GBIF dataset can be found in Table S1.
We complemented this point occurrence data using range map
data that is available globally for amphibians, birds, mammals and
reptiles from IUCN (2019). We converted each species polygon
range map to a 1  km by 1  km raster that we then used to place a
single point occurrence in the centre of each occupied pixel. Unlike
GBIF data, these data are less sensitive to taxonomic or geographical
sampling biases. However, there are some caveats to consider with
the range map data, principally the fact that both the range maps
and ecoregion delineations may rely on similar features (i.e. moun-
tain ranges, rivers, etc.). We therefore discuss and compare results
coming from both datasets throughout the Results and Discussion
sections. Summary statistics for the range map dataset can be found
in Table S2.
2.3 | Environmental data
We selected environmental variables representative of the mecha-
nisms we thought most likely would influence ecoregion strength
(Table  1). We used zonal statistics to calculate the mean value for
each ecoregion and variable. The slope layer was calculated using the
‘slope’ function in QGIS based on a 90-meter digital elevation model,
creating a 90-m resolution layer which was then resampled using
mean values to a 1-km resolution (USGS,  2010). The human foot-
print layer combines metrics of eight variables (built infrastructure,
population density, electric power, crop land, pasture land, roads,
railways and navigable waterways) into a single metric of human
impact on the landscape (Venter et  al.,  2016). Variables that were
not normally distributed were square root transformed to improve
model fit (annual precipitation, seasonality of temperature, slope
and human footprint). Summary histograms for the environmental
data are shown in Figure  S2. To make variable influence compara-
ble across all variables, we normalized each variable, setting mean
Layer name Reference
BIO01 = Annual Mean Temperature Hijmans et al. (2005)
BIO04 = Temperature Seasonality (standard Hijmans et al. (2005)
deviation*100)
BIO12 = Annual Precipitation Hijmans et al. (2005)
BIO15 = Precipitation Seasonality Hijmans et al. (2005)
(Coefficient of Variation)
Slope USGS (2010)
Soil pH×10 in H2O at 0.15 m Hengl et al. (2014)
MYD13Q1.006 Vegetation Indices 16-Day Didan (2015)
Global 250 m: NDVI
Human Footprint 2009 Venter et al. (2016)
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1416       SMITH et al.

conditions to zero, and representing values deviating from the mean
being converted to their corresponding z-score. Because environ-
mental variables often show large colinearity, especially at a global
scale, we proceeded to perform a principal component analysis. The
first four principal components explained ~90% of the variance and
were used in downstream analysis.
2.4 | Taxonomic and trait data
To examine biotic variability in ecoregion distinctness, we down-
loaded trait data for amphibians, birds, mammals and reptiles from
online published databases (Table 2). We chose to focus on feeding
guild and body size as they were globally available for all four taxa.
Trait data were then linked to GBIF and IUCN data based on pro-
vided Latin binomials, including synonymous scientific names when
provided. Species without matching trait information were dropped
from analyses. Summary statistics of the data broken down by trait
groups can be found in Table S3 (GBIF data) and Table S4 (range map
data).
2.5 | Within-ecoregion homogeneity analysis
To calculate the amount of taxonomic homogeneity within ecore-
gions, we further divided the Dinnerstein et  al.’s (2017) map by
splitting each ecoregion into two halves. This was done by dividing
the ecoregion with a line drawn through each ecoregion's centroid.
Initially, the ecoregion was split into eastern and western halves
(Figure S3). We then rotated the dividing line by 18 degrees, result-
ing in 10 different ways of dividing the ecoregion in half (Figure S3).
From each of these 10 divided sub-ecoregions, we calculated the
Jaccard index of dissimilarity and then averaged across these
10 values to derive a single, ecoregion-level, homogeneity score
(Figures S4–S14). This variable was then included as a predictor vari-
able in downstream analyses of between ecoregion heterogeneity.
2.6 | Across-ecoregion heterogeneity analysis
To test how abiotic factors affected dissimilarity across ecoregion
borders, we began by calculating Jaccard's index of dissimilarity
between all pairs of ecoregions from a presence–absence com-
munity matrix. Because our analyses here focus principally on the
differences between nearby ecoregions, we then subset these
comparisons to include only comparisons between ecoregions that
were fewer than 2,000  km apart (as calculated by the Haversine
formula).
From these data, we employed a Bayesian multiple linear re-
gression that used Jaccard's index of dissimilarity (log-transformed)
as the response variable, the mean homogeneity of the two ecore-
gions, the geographical distance between them and the mean of first
four principal components of the environmental conditions, along
with their squared terms (log[Jaccard]~distance+mean homogene-
ity+pc1+pc12+pc2+pc22+pc3+pc32+pc4+pc42). We included squared
terms here as we reasonably expected nonlinear impacts of certain
predictor variables. For example, pH levels may be strongly import-
ant at either very low or very high levels, but relatively unimport-
ant at intermediate levels. To present results in a more interpretable
format, we used the eigenvectors of the principal components to
determine the relative contribution of each variable to Jaccard's dis-
similarity index and present the graphs in Figures 1 and 2 using these
values.
(For our analyses of biotic contingency, we performed Bayesian
multiple linear regression between Jaccard's index of dissimilar-
ity (log-transformed) and the size of the ecoregion and either the
taxonomic grouping or trait grouping as the predictor variable and
Jaccard dissimilarity index as the response variable (log[Jaccard]
~distance+mean homogeneity+c(taxa) or log[Jaccard] ~distance+mean
homogeneity+c(guild) or Jaccard ~ distance+mean homogeneity+c(body
mass group)).
All models were constructed with uninformed priors and 10,000
iterations, with a burn-in of 2,000 iterations and a thinning rate of
0.5. Each model used four chains to ensure for model convergence.
Continuous variables were deemed to be significant if the 95% cred-
ible interval of the variable did not overlap with zero. Significant dif-
ferences among categorical groupings were defined as those with
non-overlapping 95% credible intervals. For quadratic features, the
values for which the combined effect of the linear and quadratic ef-
fect is different from zero (95% Bayesian credible interval) represent
the range of values for which this variable demonstrates a signifi-
cant effect on the model. Models were conducted using the ‘PyMC3’
package in Python using a No-U-Turn sampler (Salvatier, Wiecki, &
Fonnesbeck, 2016).

TA B L E 2   We obtained data on feeding guild and body size from published databases for terrestrial vertebrate taxa and used these
groupings to divide taxonomic groups into functional guilds. Taxa were split into large-bodied and small-bodied species by simply binning the
largest 50% as ‘large species’ and the smallest 50% as ‘small species’. Taxa were split into feeding guilds as described in the table

Taxa Body size Guild Source

Amphibians Total length Herbivore, predator (consumption of vertebrates = True) Oliveira et al. (2017)

Birds Body mass Fruit/nectar, invertebrate, plant/seed, vertebrates/fish/scavengers, omnivores Wilman et al. (2014)
Mammals Body mass Predator (consumption of vertebrates >10%), herbivore Wilman et al. (2014)
Reptiles (lizards) Snout–vent length Predator, herbivore, omnivore Meiri (2018)
SMITH et al. |
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F I G U R E 1   Ecoregion distinctness is explained principally by slope, climate, and human impact. Here we show the median magnitude
of the effect of a given variable on Jaccard dissimilarity between ecoregions, with higher values (lighter coloration) demonstrating that
variable is more important in explaining community turnover. Abiotic predictors are shown in order of importance from left to right
along the x-axis. Despite differences among taxa, climatic – together with slope and human impact – are the dominant forces structuring
community composition. The median values are reconstructed from the principal component eigenvectors of the regression log[Jaccard]
~distance+mean homogeneity+pc1+pc12+pc2+pc22+pc3+pc32+pc4+pc42 using GBIF data. The standard deviation for these values can be
found in Figure S20 [Colour figure can be viewed at wileyonlinelibrary.com]

3 | R E S U LT S 3.1.2 | Hypothesis a2: Ecoregions will be more

3.1 | Abiotic hypotheses
3.1.1 | Hypothesis a1: Ecoregion distinctness is
explained principally by macroclimate
We did not find any evidence that any of the eight environmental
variables considered significantly predicted the degree of internal
homogeneity – measured by Jaccard dissimilarity – within an ecore-
gion (Figures  S15–S18). There was also no clear spatial patterning
of internal ecoregion homogeneity (Figures  S4–S14). These results
were consistent irrespective of source of biodiversity data (from
GBIF or from IUCN range maps).
When we examined which variables contributed most to Jaccard
dissimilarity across ecoregion borders, we found that (for both GBIF
and IUCN range map data) the top five predictive variables were the
same, including three climatic variables (mean annual temperature,
temperature seasonality and rainfall seasonality) along with slope
and human impacts. The relative importance of these variables
differed depending on the source of the data, with slope being the
best predictor of GBIF-derived patterns and human impacts being
most important for the IUCN range-map-based data (Figure  1,
Figures S19–S21).
distinct in tropical zones
There was considerable variability in how taxa responded along
each individual abiotic gradient. Despite this, some general trends
emerged. First, we found that ecoregions more strongly differenti-
ated communities in regions with higher average temperatures, a
lower degree of temperature seasonality and a higher degree of
rainfall seasonality – all attributes associated with tropical areas
(Figure 2, Figure S23). We did not, however, find a strong signal of our
fourth climatic variable, total annual rainfall. We also found a strong
and uniform relationship between increasing slope and increasingly
distinct ecoregions. The most surprising relationship involved human
impacts, with ecoregions being the most similar to one another at
very low or very high levels of human impacts, and being most dis-
tinct at intermediate levels. We did not find any notable relationships
between either soil pH or NDVI (normalized differential vegetative
index) and the amount of dissimilarity explained by ecoregion bound-
aries (Figures S22 and S24).
When viewed spatially, we can see that our model predicts
ecoregion dissimilarity to be strongest in the tropics (Figure 3), with
relative consistency across data sources and taxa (Figures 4 and 5,
Figures  S25–S27). Furthermore, even while each taxon had differ-
ent responses to individual abiotic factors, all taxa were, with the
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1418       SMITH et al.

FIGURE 2 Here we show the relationship between taxon and the top four abiotic predictors of ecoregion distinctness (a) slope, (b) rainfall
variability, (c) human impact, and (d) mean annual temperature. Ecoregions are more distinct (more positive values) in warmer areas (top right in
(d)) and areas with a high degree or rainfall seasonality (top right in (b)). As slope increases, ecoregions become more distinct (to right in a). There
is a complex relationship with human impact, showing ecoregion distinctness being highest at intermediate levels of human impact and lower
and both extremes (c, for all taxa apart from birds). In this figure we show the median value (solid line) along with the 95% Bayesian credible
intervals (shading). The values are reconstructed from the principal component eigenvectors of the regression log[Jaccard]~distance+mean
homogeneity+pc1+pc12+pc2+pc22+pc3+pc32+pc4+pc42 using GBIF data [Colour figure can be viewed at wileyonlinelibrary.com]

exception of fungi, positively correlated with one another in terms
of the net model output (Figures S28 and S29). That is to say, ecore-
gions predicted to be dissimilar from one another for one taxon were
likely to be similarly dissimilar for all other taxa considered. Fungi
showed weakened, but still positively correlated, relationships in
terms of their similarity to other taxa.
3.2 | Biotic hypotheses
Even once we accounted for the tendency of each taxon to
have different homogeneity values, however, there were still de-
tectable dissimilarities in how well ecoregions differentiate biotic
communities. Once accounting for distance and the homogeneity
of each taxon, birds tended to be much more shared across ecore-
gion boundaries, followed by mammals, then jointly by reptiles and
amphibians. Plants were relatively dissimilar from neighbouring
ecoregions, but arthropods and fungi were very rarely shared across
ecoregions (Figure 6, Figure S30).

Surprisingly, we were able to discern only a few significant differ-

ences between the internal homogeneity of different taxa within
ecoregions (Tables S5 and S6). This was true regardless of whether
we considered data coming from GBIF or IUCN range maps. Most
notably birds were found to be much more homogeneous than were
the other taxa we considered.
3.2.1 | Hypothesis b1: Ecoregions will be more
distinct for species in lower trophic levels
We did not find any significant differences between any of our taxo-
nomic groupings in terms of the internal homogeneity of ecoregions.
SMITH et al. |
      1419

F I G U R E 3   Ecoregions are more distinct in the tropics. This map represents the average value of all seven taxonomic groups
considered (using GBIF data), after normalizing all values within each taxon. Lighter colors indicate ecoregions that are expected
to be more different from one another while darker areas should contain ecoregions more similar to one another. Values are
colored based on the expected Jaccard dissimilarity using the coefficients from the equation log[Jaccard] ~distance+mean
homogeneity+pc1+pc12+pc2+pc22+pc3+pc32+pc4+pc42 with a distance of 1,000 km and the mean value (between the two ecoregions) for
mean homogeneity and each principle component [Colour figure can be viewed at wileyonlinelibrary.com]

We were also surprised to not find any notable patterns with re-
spect to feeding guilds and between ecoregion heterogeneity. For
GBIF-based data, we found that only herbivorous reptiles showed
more distinct ecoregions than omnivorous or carnivorous reptiles
(Figure  S31). However, for range-map-based data, we found that
functional guild strongly predicted how well ecoregions can describe
biotic communities. We found that, for species of amphibains and
mamamls, ecoregions were more robust for herbivorous species than
predatory species (Figure 7a,c). Within birds, ecoregions performed
best for fruit-eating birds and worst for vultures, raptors, hawks and
allies (Figure 7b).
3.2.2 | Hypothesis b2: Ecoregions will be more
distinct for smaller-bodied species
We also did not find strong results based on GBIF data in the ability of
ecoregions to differentiate communities based on size (Figure S32).
However, when we used IUCN range map data, we uniformly found
that ecoregions better differentiated smaller-bodied species once
accounting for an ecoregion's internal homogeneity (Figure 8).
4 | D I S CU S S I O N
Ecoregions are a useful concept for both ecological research
and conservation management (Di Marco et  al.,  2018;Dinerstein
et al., 2017;Olson & Dinerstein, 2002). They are merely a classifica-
tion scheme, however, through which to maximize internal homoge-
neity and cross-boundary heterogeneity. Like classification schemes
at other levels of biological organization, they aim to describe the
outcomes of complex ecological and evolutionary relationships
between organisms and their environments. The processes that
influence how well they describe biological communities are by no
means straightforward, however, and involve complex interplay be-
tween abiotic and biotic processes over all time-scales (Bailey, 2004).
Our investigation here revealed strong support for our two abi-
otic hypotheses. First, we found that three of our major climatic
variables were consistently among the most important predictors
of ecoregion differentiation, though we note an unexpected and
outsized importance of slope and human impact (hypothesis a1).
Consistent with our second hypothesis, we found that tropical
ecoregion boundaries were likely stronger than ecoregion boundar-
ies in temperate regions (hypothesis a2).
Evidence for our biotic hypotheses was more mixed. We found
moderate evidence that ecoregions may better describe communities
of smaller-bodied and less-mobile animals, but only when we consid-
ered range map data, suggesting caution when interpreting the re-
sults with regards to our biotic hypotheses (hypothesis b1). Similarly,
we found weak support for our notion that herbivores should be
more strongly tied to ecoregions than predators (hypothesis b2).
Of all the environmental attributes we evaluated, climatic con-
ditions – in particular mean annual temperature, temperature
seasonality and rainfall seasonality – were among the most signif-
icant predictors of ecoregions distinctness. We found that warmer
areas with less temperature seasonality were more likely to have
more distinct ecoregions. This result is consistent with the expec-
tations of stronger zonation between communities in the tropics
(Buckley & Jetz, 2007;Qian, 2009). There are possible explanations
for this, however, beyond simply Janzen's (1969) mountain pass
hypothesis. It is also in line with the notion that the tropics have
experienced a much longer evolutionary history than the temper-
ate zones, considering especially the last glaciation (Condamine
et  al.,  2012;Mittelbach et  al.,  2007). This longer period would po-
tentially have allowed species to evolve into more interlinked food
webs, with greater degrees of specialization and dependency, than
 |
1420       SMITH et al.

F I G U R E 4   Ecoregion distinctness is largely consistent across taxonomic groups to a first order approximation along a
latitudinal gradient, but does show noticeable variability especially fungi (c). This map represents the GBIF-based model results
from (a) amphibians, (b) arthropods, (c) fungi, (d) mammals, (e) plants, and (f) reptiles. Lighter colors indicate ecoregions that are
expected to be more different from one another while darker areas should contain ecoregions more similar to one another. Values
are colored based on the expected Jaccard dissimilarity using the coefficients from the equation log[Jaccard] ~distance+mean
homogeneity+pc1+pc12+pc2+pc22+pc3+pc32+pc4+pc42 with a distance of 1,000 km and the mean value (between the two ecoregions) for
mean homogeneity and each principle component [Colour figure can be viewed at wileyonlinelibrary.com]

would be expected in ‘younger’ food webs in the temperate zone
(Novotny & Basset, 2005;Paine, 1966).
We were surprised, however, to find that slope and human
impacts were so related to the robustness of the Dinerstein et al.
(2017) ecoregion scheme. In regions with steeper slopes, we found
strong evidence of species clustering into distinct ecoregions. This
adds to a wealth of literature that suggests that elevational gradients
are among the strongest predictors of biodiversity patterns (Graham
et al., 2014;Peters et al., 2016;Quintero & Jetz, 2018).
When we pair these findings with those above on the outsize
strength of ecoregions in the tropics, we can return to a couple of
the earliest questions in modern biogeography. For example, are
mountain passes higher in the tropics? Our results suggest yes. Or,
as MacArthur succinctly contrasts the viewpoints of Whittaker
(mainly a temperate ecologist) and Holdridge (mainly a tropical ecol-
ogist), do more synchronous groupings of plants pervade the tropi-
cal zone than they do the temperate zone (MacArthur, 1972)? As he
points out, exploring the question was at the time data-limited, but
our analyses here suggest that, in fact, tropical biomes hold more ro-
bust, synchronous groupings. This is true not only for plants but also
for all taxa tested here, compared to their temperate counterparts.
The one pattern we find most difficult to explain with abiotic
mechanisms alone is the relationship between human impact on
the landscape and ecoregion distinctness. Ecoregion distinct-
ness is low in landscapes with high levels of human impact, fur-
ther supporting the evidence for homogenization of biodiversity
SMITH et al. |
      1421

F I G U R E 5   Expected dissimilarity
between communities within ecoregions
is largely consistent regardless of the data
source used to construct models. This
map represents the model results from
bird community dissimilarity using either
(a) GBIF data or (b) IUCN range map data.
Lighter colors indicate ecoregions that are
expected to be more different from one
another while darker areas should contain
ecoregions more similar to one another.
Values are colored based on the expected
Jaccard dissimilarity using the coefficients
from the equation log[Jaccard]
~distance+mean homogeneity+pc1+pc12+
pc2+pc22+pc3+pc32+pc4+pc42 with
a distance of 1,000 km and the mean
value (between the two ecoregions) for
mean homogeneity and each principle
component [Colour figure can be viewed
at wileyonlinelibrary.com]

F I G U R E 6   Ecoregions better
differentiate communities of arthropods,
plants, and fungi than they do vertebrates.
Further, they better describe reptiles
and amphibians than they do mammals
or birds. Here we show, once correcting
for geographic distance, how similar
ecoregions are expected to be from one
another, with higher values indicating
ecoregions are more similar in their
community makeup. Ecoregions are
most descriptive of communities of
plants, fungi, arthropods, amphibians,
and reptiles. The same borders do not
perform as well for mammal or bird
communities. Lines are colored by
taxa and show the median estimate
in a solid line and the 95% Bayesian
credible interval in shaded colors from
the equation Jaccard~distance+mean
homogeneity+c(taxa) using GIBF
data [Colour figure can be viewed at
wileyonlinelibrary.com]

in human-dominated systems (Capinha, Essl, Seebens, Moser, & distinct. One interpretation of this pattern is that many regions rel-
Pereira, 2015;Newbold et al., 2018). We also found, however, that atively devoid of human impact are expected to be relatively ho-
at the lowest levels of human impact, ecoregions were also not as mogenous (e.g. the boreal forests or tundra of Canada and Russia).
 |
1422       SMITH et al.
Alternatively, the data informing ecoregion delineations may be in-
adequately drawn in relatively unexplored and undocumented re-
gions (i.e. the Amazon rainforest or much of South-East Asia). We
find support for this explanation when we consider that these pat-
terns were more pronounced for range-map-based data than GBIF-
based data, potentially indicating confounding bias of expert maps in
underexplored regions (Beck, Böller, Erhardt, & Schwanghart, 2014).
If we assume these two mechanisms to both be at play, we do see
the expected result in areas with intermediate human impact ecore-
gions perform relatively well in segmenting distinct communities.
This potentially suggests a peak where human presence in a region
is sufficient to document its biodiversity, but no so overwhelming as
to alter it radically.
Consistent with our first biotic hypothesis (b1), we found evi-
dence that species in guilds more closely linked to plant communi-
ties (e.g. herbivores) were less likely to be shared across ecoregion
borders. We note that this pattern held only for range-map-based
data and not GBIF data, suggesting that more robust data may be
needed to elucidate this relationship. For mammals and amphibi-
ans, we found that predatory species were more ubiquitous across
ecoregion borders, but there were no significant differences among
different guilds of reptiles. The diet classifications for birds are more
F I G U R E 7   Functional guild strongly
predicts how well ecoregions can describe
biotic communities. We find that for
amphibians, and mammals ecoregions
are more distinct for herbivores than
predators (a, c). Within birds, ecoregions
perform best for fruit eating birds and
worst for vultures, raptors, hawks
and allies (b). The story is more mixed
for reptiles (d). Lines are colored by
trait grouping and show the median
estimate in a solid line and the 95%
Bayesian credible interval in shaded
colors. Data for these figures is based
off of IUCN range map data from the
equation Jaccard~distance+mean
homogeneity+c(guild) [Colour figure can
be viewed at wileyonlinelibrary.com]
complex, but again predatory and scavenging birds were the most
likely to be found across multiple ecoregions. One plausible expla-
nation for this finding is that predators, especially apex predators,
maintain large territories per individual or a group of few individuals,
such that predator population density is low compared to prey den-
sity (Arim, Abades, Laufer, Loureiro, & Marquet,  2010). Therefore,
it is not surprising that predators are more likely to span multiple
ecoregions.
Beyond the feeding guild of the organism considered, we found
that body size was also a useful predictor of how strongly ecoregions
divide communities. In general, we found that larger species were
more similar across ecoregion borders. This aligns well with our expec-
tations, given that body size correlates highly with range size (Kelt &
Vuren, 1999; Swihart et al., 1988) and species with larger ranges are by
definition more likely to span multiple ecoregions in their distribution.
This pattern was, however, evident only with IUCN range-map-based
data and not GBIF data, suggesting data deficiency may preclude
more nuanced distinctions between body mass classes. While these
individual patterns are robust, we cannot conclude definitively which
trait – body size or feeding guild – may underlie the driving biological
mechanism. Predatory species tend to also be larger, and therefore it
is possible that the large territories they maintain or landscapes over
SMITH et al.
F I G U R E 8   We found strong
relationships between body size and how
effectively ecoregions differentiated
assemblages of species. For amphibians
(a), birds (b), mammals (c), and reptiles (d)
we found that larger species were more
shared across ecoregion borders. Lines
are colored by trait grouping and show
the median estimate in a solid line and
the 95% Bayesian credible interval in
shaded colors. Data for these figures is
based off of IUCN range map data from
the equation Jaccard~distance+mean
homogeneity+c(bodySize) [Colour figure
can be viewed at wileyonlinelibrary.com]
which the roam are driven by their size, their dispersal abilities, their
feeding preferences or a combination thereof. Future analyses on spe-
cific clades of organisms wherein these different permutations of traits
(i.e. small predators, large predators, small herbivores and large herbi-
vores) are all present may help to resolve the relative importance of
feeding guild and body size.
Ecoregions have driven not only basic ecological research but
also land-use planning and conservation at large scales (European
Environment Agency,  2003; The Nature Conservancy,  2018; US
EPA, 2015; World Wildlife Fund,  2018). Here we show, however,
compelling evidence that there are certain regions and taxa for
which the concept is more or less useful. Fortuitously, the current
maps of ecoregions are best suited to address conservation goals
in tropical areas, for small organisms and for organisms lower in the
food chain. Given that these regions and taxa are the least likely
to be represented in global biodiversity datasets (Beck et al., 2014;
Troudet, Grandcolas, Blin, Vignes-Lebbe, & Legendre, 2017), their
usage in such decision contexts is well-justified. Ecoregion-based
conservation plans can fill a gap, informing conservation actions
where data are limited, and in so doing helping to preserve the
vital elements of nature that support the well-being of human-
ity in profound ways (Cardinale et  al.,  2012;Díaz, Fargione, Iii, &
Tilman, 2006).
|
      1423
AC K N OW L E D G E M E N T S
We thank Christopher Anderson and Glade Dlott for helpful com-
ments that improved the quality of this manuscript. J.R.S. was
supported by a National Science Foundation GRFP (GRFP DGE-
1656518), the Stanford Department of Biology, and the Center for
Conservation Biology at Stanford through its Ward Wilson Woods
Jr. Environmental Studies Fund. J.N.H. was supported by the
Gerhard Casper and John P. Morgridge Fellowship, the Stanford
Graduate Fellowship, and the Center for Conservation Biology
at Stanford through its Ward Wilson Woods Jr. Environmental
Studies Fund. N.N. was supported by the Bing Fellowship in
Honor of Paul Ehrlich. We also thank Stanford University and the
Stanford Research Computing Center for providing vital compu-
tational resources and support, including access to the Sherlock
Cluster.
DATA AVA I L A B I L I T Y S TAT E M E N T
The data that support the findings of this study are openly avail-
able in GBIF at the following DOIs: Amphibians: https://doi.
org/10.15468​/dl.as6uug, arthropods: https://doi.org/10.15468​/
dl.uhumps, birds: https://doi.org/10.15468​/dl.61rqcy, fungi: https://
doi.org/10.15468​/dl.akftay, mammals: https://doi.org/10.15468​/
dl.diqarx, plants https://doi.org/10.15468​/dl.uhenlf, and reptiles:
 |
1424       SMITH et al.
https://doi.org/10.15468​/dl.pn1aw9. The IUCN range map data
are openly available for academic use from the IUCN at https://
www.iucnr​edlist.org/resou​rces/spati​al-data-download and from
BirdLife at http://dataz​one.birdl​ife.org/speci​es/reque​stdis. The cli-
mate data used are available from https://world​clim.org/data/biocl​
im.html and are attributed to Hijmans, Cameron, Parra, Jones, and
Jarvis (2005). Elevation data are available from the USGS (2010)
at https://www.usgs.gov/cente​rs/eros/scien​ce/usgs-eros-archi​ve-
digit​al-eleva​tion-globa​l-multi​-resol​ution​-terra​in-eleva​tion?qt-scien​
ce_center_objec​t s=0#qt-scien​ce_center_objects. Soil conditions are
available from Hengl et al. (2014) at https://www.isric.org/explo​re/
soilg​rids. Vegetation characteristics are available from Didan (2015)
at https://lpdaac.usgs.gov/produ​c ts/myd13​q1v00​6/. Human foot-
print maps are available from Venter et al. (2016) at https://sedac.
ciesin.colum​b ia.edu/data/set/wilda​r eas-v3-2009-human​-footp​
rint. Amphibian trait data are available from Oliveira, São-Pedro,
Santos-Barrera, Penone, and Costa (2017) at https://www.nature.
com/artic​les/sdata​2017123. Bird and mammal trait data are avail-
able from Wilman et al. (2014) at http://www.esapu​bs.org/archi​ve/
ecol/E095/178/. Reptile trait data are available from Meiri (2018) at
https://onlin​elibr​ary.wiley.com/doi/abs/10.1111/geb.12773.
ORCID
Jeffrey R. Smith  https://orcid.org/0000-0001-5768-216X
J. Nicholas Hendershot  https://orcid.org/0000-0001-9570-8447
Nicole Nova  https://orcid.org/0000-0001-8585-1215
Gretchen C. Daily  https://orcid.org/0000-0003-1443-1111
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