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Received: 14 January 2022 Revised: 5 May 2022 Accepted: 16 May 2022

DOI: 10.1002/ecy.3827

ARTICLE

Fitness homeostasis across an experimental water gradient

predicts species’ geographic range and climatic breadth

Ian S. Pearse1 | Patrick McIntyre2 | N. Ivalú Cacho3 | Sharon Y. Strauss4

1
U.S. Geological Survey, Fort Collins
Science Center, Fort Collins,
Colorado, USA
2
Nature Serve, Western Regional Office,
Boulder, Colorado, USA
3
Instituto de Biología, 3er Circuito de CU
s/n, Universidad Nacional Aut
onoma de
México, Copilco Coyoac
an, Universidad
Nacional Aut
onoma de México, Mexico
City, Mexico
4 or a combination of these factors. We asked whether fitness expressed along a key
Center for Population Biology and
Department of Evolution and Ecology,
University of California, Davis,
California, USA
Correspondence
Ian S. Pearse
Email: ipearse@usgs.gov
Funding information
National Science Foundation,
Grant/Award Number: NSF-1545597
Handling Editor: James T. Cronin
Abstract
Species range sizes and realized niche breadths vary tremendously.
Understanding the source of this variation has been a long-term aim in evolution-
ary ecology and is a major tool in efforts to ameliorate the impacts of changing cli-
mates on species distributions. Species ranges that span a large climatic envelope
can be achieved by a collection of specialized genotypes locally adapted to a small
range of conditions, by genotypes with stable fitness across variable environments,
niche axis, water availability, could explain a species’ realized niche breadth, its
geographic range and climate breadth, in 11 species from a clade of jewelflowers
whose range sizes vary by two orders of magnitude. Specifically, we explored
whether the range size of a species was related to the ability of genotypes (mater-
nal families) to maintain fitness across a range of experimental water availabilities
based on 30-year historical field precipitation regimes. We operationally character-
ized fitness homeostasis through the coefficient of variation in fitness of a geno-
type (family) across the experimental water gradient. We found that species with
genotypes that had high fitness homeostasis, low variation in fitness over our
treatments, had larger climatic niche breadth and geographic range in their field
distributions. The result was robust to alternate measures of fitness homeostasis.
Our results show that the fitness homeostasis of genotypes can be a major factor
contributing to niche breadth and range size in this clade. Fitness homeostasis
can buffer species from loss of genetic diversity and under changing climates, pro-
vides time for adaptation to future conditions.

KEYWORDS
Brassicaceae, climate water deficit, comparative, drought, fundamental niche, mustards,
niche breadth, phenotypic plasticity, phylogeny, range limits, range size, realized niche,
Streptanthus

INTRODUCTION been central to understanding where organisms live

along environmental gradients (Hutchinson, 1959;
Species vary tremendously in their range sizes, the geo- Schrodt et al., 2019; von Humboldt, 1850), and how spe-
graphic areas over which they live and the range of envi- cies evolve to live in different habitats (Cacho &
ronments in which they persist. This observation has Strauss, 2014; Darwin, 1859). A better understanding of

© 2022 The Ecological Society of America. This article has been contributed to by U.S. Government employees and their work is in the public domain in the USA.

Ecology. 2022;e3827. https://onlinelibrary.wiley.com/r/ecy 1 of 11

https://doi.org/10.1002/ecy.3827
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2 of 11 PEARSE ET AL.

the basis of niche breadth has become increasingly
urgent because the mechanisms underlying niche
breadth and its evolution will determine how well and
how quickly species can respond to changing global envi-
ronments (Clavel et al., 2011; Colles et al., 2009). Niche
breadth must be limited in some capacity, as no species is
able to thrive in all environments (Reznick et al., 2000),
but what causes the tremendous variation in niche
breadth among species is still poorly understood
(Carscadden et al., 2020; Sexton et al., 2017).
Species-level niche breadth is a product of three main
processes or phenomena that act at different scales
(Figure 1; Bolnick et al., 2002; Sexton et al., 2017):
(1) plasticity or fitness homeostasis, the ability of a geno-
type to maintain similar fitness across a range of environ-
ments, allows genotypes to thrive across a wide range of
environments (Griffith & Sultan, 2012; Sides et al., 2014;
Sultan, 2001); (2) genetic variation within populations, in
which each genotype may persist in a subset of environ-
ments, but collections of genotypes can thrive across a
wider set of environmental conditions owing to variation
among genotypes (Sheth & Angert, 2014); and (3) local
adaptation at the population level, in which each popula-
tion is comprised of genotypes that thrive under a limited
number of environments (Angert et al., 2011;
Hereford, 2009). While more than one of these factors
likely contributes to the overall niche breadth of a spe-
cies, it is useful to differentiate these processes, both to
understand mechanisms of niche breadth evolution, and
particularly when considering conservation approaches
for species under rapidly changing climate (Williams
et al., 2008). For example, the consequences of loss of
genetic diversity within a population or species with
regards to its geographic occurrence (i.e., range shift, con-
traction, or expansion) will be determined by the degree
to which individual-level plasticity and fitness homeosta-
sis versus genetic differentiation among populations and
individuals shapes species-level niche breadth (Sheth &
Angert, 2014).
Niche breadth is closely related to the range size of
organisms. In general, species with larger range sizes persist
across a greater variety of environments (Sheth et al., 2014;
Slatyer et al., 2013). Most studies exploring niche breadth
use correlative approaches, taking biotic or abiotic
aspects of niche, as in species distribution models, and
using these to assign niche breadth. Others have added
an experimental approach to measuring niche breadth in
key environments (Chambers & Emery, 2016; Dixon &
Busch, 2017; Kelly et al., 2012; Sheth & Angert, 2014;
Tittes et al., 2019; Wei et al., 2017). For some species,
range size may be limited primarily by environmental
factors like water availability or temperature (Choat

(a) (b) (c)

b
Fitness

Range (maximum–minimum)
Environmental gradient

F I G U R E 1 Some hypothetical responses of genotypes within a population to an environmental gradient. (panel a) Solid lines reflect a
population comprised of genotypes or families that have broad niches and are stable (have high fitness homeostasis) across an
environmental gradient. Dashed lines reflect a population comprised of a collection of genotypes or families that each thrive under a
narrower set of conditions (the niche of each genotype is more specialized) and that are less stable. (panel b) Fitness homeostasis can
operationally be estimated as the opposite of the coefficient of variation (CV) in fitness of genotypes, or families, across the gradient,
schematically represented by arrows for three genotypes. Genotype a has less variation in fitness across the gradient and high fitness
homeostasis (low CV), a habitat generalist. Genotypes b and c both have lower fitness homeostasis and are specialized in some parts of the
gradient. Comparison of genotypes a and b suggest a trade-off between maximum fitness under optimal conditions and being a generalist
(panel b). Fitness homeostasis can also be measured as the range of environments over which some threshold fitness level is achieved by
each genotype (panel c); in our case, we assessed the range of water availabilities over which 70%, 80%, and 90% of peak fitness was
maintained. Genotypes with high fitness homeostasis (a) are able to maintain a threshold of fitness over a wide range of environments.
Genotype c has a narrower range of environments over which it can maintain the same threshold of fitness.

ECOLOGY
et al., 2012; Dilts et al., 2015; Engelbrecht et al., 2007;
Lutz et al., 2010). In these cases, performance curves
measured across range-limiting factors (Angert
et al., 2011; Huey & Stevenson, 1979) allow us to explore
experimentally whether the ability of genotypes to main-
tain fitness along that environmental gradient predicts
the range of environmental conditions in which a species
persists in nature. Interspecific comparisons of perfor-
mance curves can provide information on constraints in
the realized niche and help to explain why some species
have larger niches and range sizes than others (Tittes
et al., 2019).
One metric that captures performance of genotypes
across environments is fitness homeostasis, which is
the ability of a genotype to maintain similar fitness across
a range of environments (Hoffmann & Parsons, 1991;
Rejm
anek, 2000; Sultan & Bazzaz, 1993; Woods
et al., 2009). Across environments, fitness homeostasis
across environments is thought to be accompanied by
changes in gene expression underlying morphological or
physiological traits that influence fitness (Bradshaw, 1965;
Sultan, 1995; Turner et al., 2017), i.e., a reflection of plastic-
ity. The ability to maintain fitness homeostasis may also
reflect non-plastic phenotypes that are successful across a
range of environments, termed “phenotypic buffering”
(e.g., Reusch, 2014). Fitness homeostasis can be used as a
measure of niche breadth at the genotypic level (Rejm
anek
et al., 2005; Woods et al., 2009). The shape of the perfor-
mance curve relating fitness to an environmental gradient
can reveal how specialized a genotype is to its environ-
ment. A norm of reaction of fitness plotted against environ-
ment that is not steeply peaked indicates high fitness
homeostasis across a range of environments (for example,
Figure 1b, genotype a). Organisms that can maintain fit-
ness over a large range of environments have wide niche
breadth and are considered generalized genotypes.
One potential driver of variation in fitness homeosta-
sis is a generalist–specialist trade-off in which species or
individuals with high maximum fitness in some environ-
ments also have lower fitness homeostasis (i.e., a
narrower set of environments in which they can produce
high fitness (Angert et al., 2011; Sexton et al., 2017). The
concept of generalist–specialist trade-offs is central to
many theories of the evolution of niche breadth; how-
ever, empirical tests of such trade-offs are rare, and evi-
dence for generalist–specialist trade-offs is variable
among systems (Sexton et al., 2017).
Here, we use fitness homeostasis measured in an
experimental setting as a metric of niche breadth to ask
to what extent niche breadth of individual genotypes
across a key environmental axis can explain the species’
realized niche. More specifically, using a clade of mus-
tards that inhabit water-limited environments, we ask
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3 of 11
“Does the fitness homeostasis of genotypes within species
measured across a range of water availability in an exper-
imental setting predict that same species’ geographic
range size and climate occupancy across the landscape,
or its realized niche?” This experiment addresses the
extent to which the capacity of a species to persist over a
large range of environments is caused by individual-level
fitness homeostasis (Figure 1a, solid lines). We character-
ize range size and climatic niche breadth across the range
and measure fitness homeostasis across an experimental
water gradient for 11 species within a clade of mustards
native to mediterranean California. Because variation in
niche breadth is often considered in terms of
generalist–specialist trade-offs (Carscadden et al., 2020;
Sexton et al., 2017), we additionally ask whether there is
a tradeoff between maximum fitness of a species and its
fitness homeostasis over the same experimental water
gradient.
METHODS
Study system
We explore these questions using the Streptanthus (s.l.)
clade, which includes several closely related genera
(Caulanthus, Guillenia, Streptanthus [s.l.]) that are not
reciprocally monophyletic within the group (Cacho
et al., 2014). Members of this diverse clade occupy bare
harsh abiotic habitats, spanning southern sandy deserts
to northern, coarse-soiled south-facing scree slopes of
edaphically diverse substrates (Cacho & Strauss, 2014).
Along with other diverse clades of angiosperms, these
species have been classified as Madro-Tertiary, reflecting
their southern biogeographic origin and affinity for hot,
dry environments (Cacho & Strauss, 2014; Harrison
et al., 2010; Raven & Axelrod, 1978). Biogeographic ori-
gins are often overlooked in ecological studies, yet they
may characterize the responses of many species to envi-
ronmental variation. For example, in a diverse plant com-
munity, species with Madro-Tertiary, hot dry origins
increased in abundance over a 40-year census interval,
over which the climate got hotter and drier (Damschen
et al., 2010). By incorporating deep evolutionary history,
we can gain a richer understanding of niche evolution
and its constraints (see also Tittes et al., 2019). In addi-
tion, for among-species comparisons, understanding the
relationships between traits like range size and plasticity
requires understanding evolutionary constraints on those
relationships, something an explicit clade-wide perspec-
tive provides. For these reasons, we focus on species
within the Streptanthoid Complex subclade, which are
all located within the California Floristic Province and

4 of 11
for which we have detailed understanding of their evolu-
tionary relationships and biogeographic origin (Cacho
et al., 2014). We initially started with 15 species selected
to span the environments and subclades of the
Streptanthoid Complex major clade (Cacho et al., 2014).
Of the 15 taxa, four species had poor germination and
were dropped from the analyses; thus, we ultimately used
11 taxa: Caulanthus coulteri, C. crassicaulis, C. inflatus,
C. lasiophyllus, Streptanthus breweri, S. diversifolius,
S. farnsworthianus, S. glandulosus, S. hesperidis,
S. polygaloides, and S. tortuosus. The joint distributions of
these species span the Mojave desert to northern
California Sierra Nevada and cover a large range of
annual precipitation and temperature, with geographic
range sizes that vary more than two orders of magnitude.
Experimental measurement of fitness over
water levels
For each species, seeds were field-collected by maternal
plants from one population approximately centrally
located within the species’ range-wide climate water defi-
cit (CWD) space (Appendix S1: Figure S1, see
Measurement of realized niche: geographic range size and
climate water deficit niche breadth for more detail on how
CWD range was calculated). CWD is defined as the
annual evaporative demand that exceeds available water,
and is a combined metric based on soil parent material,
aspect, temperature, and precipitation and that integrates
drainage, changes in soil moisture, energy loading, and
climate (Flint et al., 2013).
Seeds from each of five maternal families per species
were germinated on an outdoor mist bench at the UC
Davis Orchard Park greenhouse facility starting on
25 November 2015 in 25  50 cm planting trays. When
germinants reached the cotyledon/first true leaf stage,
they were transplanted to 656-ml Conetainers (Stuewe &
Sons, St. Louis, MO), filled with 1:1:1 compost:peat moss:
sand to mimic high drainage rocky or sandy conditions
in the field, and grown in an outdoor lath house at
ambient fall/winter growing season temperatures from
28 December 2015–21 January 2016. Plants that did not
germinate or that died prior to 20 January 2016 were
replaced with new germinants. Plants were not fertilized
and were kept well watered during germination and
transplanting, as would be common with natural germi-
nation in heavy winter rainstorms typical of the mediter-
ranean climate in California. On 22 January, plants had
produced the first true leaves, and experimental watering
began. From each of the five maternal families, seven
plants were randomly assigned, one to each of seven water
treatments (8, 16, 60, 105, 149, 194, 239 mm/month),
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PEARSE ET AL.
resulting in a total of 35 plants per species. These
levels were chosen to represent a field-relevant range of
growing-season precipitation based on 30 years of precipi-
tation records experienced by this group of Streptanthus
(s.l.) species in the field. Our goal was to capture most or
all of the water availability niche for each species, yet also
impose the same treatments across the clade. For all taxa,
our lowest watering level was lower than the 30-year mini-
mum of the seed collection site, while our maximum treat-
ment was greater or equal (within 2%) to the 30-year
maximum monthly precipitation for all but two species
(Appendix S1: Table S1). The species S. glandulosus and
S. polygaloides experienced greater maximum rainfall at
least once over the 30-year period than our highest
watering level. Thus, our treatments largely capture and
exceed the historical water availability niche experienced
by these taxa. Water was applied to pots using a metered
drip irrigation system, and plants were grown outdoors
under the increasing heat of the growing season; thus,
water availability decreased over the course of the experi-
ment for all treatments under natural warming tempera-
tures, similar to spring–summer transition experienced by
plants in the field. Across the whole study, precise metered
irrigation allowed plants to be arranged in a completely
randomized design. Natural precipitation was excluded
from the experiment with a transparent roof.
Fitness was measured as total fruit production
in annuals or total biomass in biennials. To minimize
pollen limitation of annual species, all open flowers
were hand-pollinated with conspecific pollen using a
bee stick (excised Bombus thorax) twice per week (more
detail in Pearse et al., 2020). The species C. lasiophyllus
is highly autogamous, with fruits that set as the
small flowers open and without cross-pollination. We
collected and counted fruit prior to dehiscence. Fruit
counts were summed over the season yielding lifetime
fruit set (our estimate of fitness). Two taxa (S. tortuosus
and C. crassicaulis) were monocarpic biennials and
did not flower. In these species, roots and shoots of
plants were collected at the end of the experiment
(1 August 2016), dried at 45 C for 14 days, and weighed
to calculate total biomass. Total biomass of biennial
species is a measure of fitness, as biomass accrued in the
first year increases lifetime fecundity (e.g., Gross, 1981),
especially in monocarpic species.
A polynomial function was the best fit of the
fitness–water-availability relationship (see Pearse
et al., 2020 for model selection and alternate models con-
sidered). In a prior paper (Pearse et al., 2020), we found
in the same experiment that the water level at which
plants accrued fitness fastest (the inflection point of the
curves in Appendix S1: Figure S2) was highly correlated
with the 30-year mean precipitation at the seed collection

ECOLOGY
site, suggesting that our imposed watering regime was
field relevant.
Measurement of realized niche: geographic
range size and climate water deficit niche
breadth
We assembled a comprehensive database of our species’
collection locations from regional herbaria and aggre-
gated occurrence databases. Using the these collection
locations, we fit a convex hull around those points
(convHull function in the Dismo R package) to calculate
range size, estimated in square kilometers (Hijmans
et al., 2017). Range size included areas in California,
Nevada, and Oregon.
We extracted the climatic water deficit (CWD) at
each of these occurrence locations using a hydrologi-
cally based model of CWD for California (Appendix S1:
Figure S3; Flint et al., 2013). CWD is a combined metric
of temperature, precipitation, and soil properties that
together influence water availability and drought stress.
Most of our sampled populations were located near or
at the modal CWD experienced by that species across its
range (Appendix S1: Figure S1). For each species, we
estimated niche breadth across CWD using the ecospat
R package (Broennimann et al., 2018), which uses a
kernel-density function to relate the distribution of
locality records and environmental variables
(Broennimann et al., 2012). To estimate niche breadth
across these estimates, we used the Bayesian HPD
approach applied to the empirical values of the niche
kernel density to calculate the 95% probability interval
of the niche from the distributions (best fits were to a
single rather than multi-modal intervals) using the
emp.hpd function in the TeachingDemos R Package
(Snow, 2016; Appendix S1: Figure S4). The range of
most species (n = 8) were entirely within the state
of California, where CWD estimates were available.
For the remaining three species (S. tortuosus,
C. lasiophyllus, and C. crassicaulis), some collection
locations were outside of California, and excluded from
CWD breadth estimates.
Calculation of fitness homeostasis and
statistical analyses
Fitness variation was calculated as coefficient of variation
(CV) in fitness within each species, accounting for
family-level effects (Figure 1). CV is the standard devia-
tion of a set of measurements divided by its mean; a low
CV of fitness suggests that genotypes can maintain
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5 of 11
similar fitness across a range of watering levels, a mea-
sure of fitness homeostasis (see also Woods et al., 2009).
Species with individuals that express less variation in fit-
ness (low CV) have greater fitness homeostasis
(Schlichting & Levin, 1984; Wolfe & Mazer, 2005). To
focus on within-genotype effects and to reduce variation
caused by genetic diversity at the family level, we incor-
porated the effect of maternal family when calculating fit-
ness homeostasis. For each species, we calculated CV
from the residuals of fitness from a model in which fit-
ness was a function of maternal family
(fitness  maternal family). An independent way
to assess fitness homeostasis is to measure the range of
environmental variation over which some threshold of
maximum fitness is maintained (Sheth & Angert, 2014).
In our experiment, we assessed the range of water
availability over which fitness was maintained at three
thresholds—70%, 80%, and 90% of maximum fitness
(Figure 1C; sensu Sheth & Angert, 2014).
The latter method (range of water resulting in fitness
above a threshold) did not account for family-level varia-
tion. To assess whether family-level variation was likely
to contribute to this analysis, we tested whether the
water–fitness relationship varied among species and
among maternal genotypes within species. Using AICc,
we compared three models: (1) a full model in which fit-
ness was determined by water amount and its interaction
with maternal family within species; (2) a species model
in which fitness was determined by water amount and its
interaction with species (but not maternal family), and
(3) a model in which fitness was determined by water
amount and species, but the water–fitness relationship
was not allowed to differ among species (Appendix S1:
Table S2). By far the best model based on AICc had fit-
ness determined by water amount and its interaction
with species (model [2]; Appendix S1: Table S2), indicat-
ing that each species responds differently to variation in
water, but that, within species, there was minimal varia-
tion in the water–fitness relationship due to maternal
family.
To assess the relationships between fitness homeosta-
sis, geographic range size, and CWD niche breadth, we
used phylogenetic generalized least squares (PGLS)
models implemented in R packages ape and nlme
(Paradis et al., 2004; Pinheiro et al., 2012). We used a
Bayesian phylogeny of the Streptanthus (s.l.) clade
(Cacho et al., 2014) allowing the model to estimate phylo-
genetic signal as λ, using the function corPagel in the
R package ape (Paradis et al., 2004), and then rescaled
the phylogeny by λ (Revell, 2010). In two cases (presented
in Appendix S1: Figure S5, and noted with an asterisk),
estimation of λ caused pathological behavior in models,
and λ was set to one, an assumption of Brownian motion
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6 of 11 PEARSE ET AL.

evolution. Geographic range size and maximum fitness species in which we failed to detect a fitness peak, this
measured as total fruit production were log-transformed would indicate that our estimates of fitness homeostasis
for all analyses to meet assumptions of normality and were subject to bias due to incomplete estimation of fitness
homoscedasticity. Proportion variance explained was cal- curves. The estimate of fitness homeostasis did not differ
culated as predicted R 2 using the R package rr2 between curves in which a fitness peak was observed (six
(Ives, 2019). species) versus those with increasing or flat fitness curves
We also addressed other aspects of niche breadth evo- (five species) (pGLS, t9 = 0.8, p = 0.45).
lution. We asked if maximum fitness traded off with fit-
ness homeostasis, that is, if there was a cost to being a
generalist genotype under optimal conditions using the RESULTS
PGLS framework described above. In order to test the
alternate hypothesis that greater CWD niche breadth and Do species with greater fitness homeostasis across
geographic range size are achieved by species with higher watering levels in the lathhouse, have larger geographic
fecundity in amenable environments, we used the same ranges or environmental tolerances? Species with greater
PGLS models to assess whether CWD niche breadth and fitness homeostasis within maternal families (lower CV
geographic range size were positively related to maxi- fitness) over a range of water availability had larger geo-
mum fitness achieved. In this analysis, C. lasiophyllus, graphic ranges (Figure 2; pGLS t9 = 2.62, p = 0.028).
the small-flowered selfer, was a large outlier and was They also occurred over a greater range of water avail-
removed from the analysis (but included in the figure); it ability (CWD) in nature (Figure 3; pGLS t9 = 4.35,
was also an outlier in other fitness-related analyses in our p = 0.002). Range-wide CWD niche breadth was strongly
earlier study (Pearse et al., 2020). The two perennating positively correlated with range size (Appendix S1:
species (S. tortuosus and C. crassicaulis) whose fitness Figure S6). Experimentally measured fitness homeostasis
was measured in units of biomass were excluded from over the water gradient in our lathhouse was sufficient to
analyses of maximum fitness, as fruit number and bio- explain 59% of the variation in range-wide CWD niche
mass are different units. breadth and 43% of the variation in geographic range
Next, we explored, using pGLS models as above, size. Results were robust to an alternate measure of
whether an estimate of drought resistance, and not fit- fitness homeostasis, measured as the range of water
ness homeostasis, predicted CWD breadth and geo-
graphic range size. In a prior study (Pearse et al., 2020),
we used the inflection point of the curve of fitness versus
water availability as a measure of drought resistance. The
inflection point identifies the water level at which each
species is accruing fitness at its greatest rate relative to
water availability. In our earlier study, we found this met-
ric was strongly correlated with other drought adapta-
tions, such as timing and extent of flowering phenology
(Pearse et al., 2020).
Last, because we assessed performance curves over the
same range of water conditions for all species, there was
variation in how much these levels meshed with historical
field precipitation. For all species, our minimum water
was lower than historical minimum precipitation records
from collections sites over 30 years. For maximum water
levels, our experimental levels met (three species) or
exceeded (six species) maximum historical precipitation
levels at the field sites over 30 years for nine species, and
was less than that maximum for two species. To test
whether this introduced bias into our estimates of fitness F I G U R E 2 Streptanthus (s.l.) species comprised of maternal
homeostasis or other analyses, we categorized each perfor- families that better maintain fitness (have a lower coefficient of
mance curve as having or lacking a fitness “peak” and variation [CV] of fitness) when grown across seven experimental
asked, using a pGLS model as above, whether fitness water availabilities have larger geographic ranges (phylogenetic
homeostasis was greater for species with a fitness peak. If generalized least squares regression, t9 = 2.62, p = 0.028). Note
our estimates of fitness homeostasis were larger for those log scale of y-axis.

ECOLOGY
F I G U R E 3 Streptanthus (s.l.) species comprised of maternal
families that are more stable fitness (have lower coefficient of
variation [CV] in fitness) when grown experimentally across a
gradient of water availability grow across a greater range of climate
water deficit (CWD) across the whole species’ geographic range
(phylogenetic generalized least squares regression, t9 = 4.35,
p = 0.002).
availability required to maintain fitness above a certain
threshold (70%, 80%, or 90%) of maximum fitness.
The range of water required to maintain fitness above a
threshold was negatively related to CV of fitness
(70% maximum fitness: Pearson’s r = 0.74; 80% maxi-
mum fitness: Pearson’s r = 0.61; 90% maximum fitness:
Pearson’s r = 0.38) across the clade; thus, coefficient of
variation in fitness, as well as the range of water required
to maintain fitness, both capture the same relationship
between fitness homeostasis and the realized niche.
In our test of costs of being a generalist, we found a
marginal trade-off between maximum fitness (fruit pro-
duction) achieved across any water level and fitness
homeostasis across the gradient (CV fitness; t6 = 2.4,
p = 0.07, R 2 = 0.37; Figure 4), suggesting there may be
costs to being a habitat generalist. However, this relation-
ship was only apparent when excluding C. lasiophyllus,
whose fruit production strategy differs markedly from
other species in the study, thus the overall evidence for a
general trade-off is weak.
Neither of the alternative hypotheses we considered
explained niche breadth better than CV fitness.
Maximum fitness alone (total lifetime fruit production at
optimal water level) predicted neither CWD niche
breadth across the range (pGLS, t7 = 1.8, p = 0.12), nor
geographic range size (pGLS, t7 = 0.8, p = 0.43).
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7 of 11
F I G U R E 4 Streptanthus (s.l.) species with less stable fitness
(higher coefficient of variation [CV] fitness over watering levels)
had marginally greater maximum fitness (fruit production) at
optimal watering level (phylogenetic generalized least squares
regression, t6 = 2.4, p = 0.07). Note that Caulanthus lasiophyllus
(an outlier in maximum fitness due to mating systems) and the two
perennating species, whose fitness was measured in units of
biomass, were removed from this analysis.
Likewise, our measure of drought resistance did not pre-
dict range of CWD (pGLS, t9 = 1.7, p = 0.11).
We tested whether the incomplete estimation of fit-
ness curves for some species biased estimates of fitness
homeostasis.
DISCUSSION
Variation in the performance of genotypes, families, and
populations across environments contributes to shaping
species-level niche breadth, or the realized niche
(Hargreaves et al., 2014; Sexton et al., 2017; Slatyer
et al., 2013). Geographic range limits are often set by
individual niche limit: conditions beyond which
populations cannot maintain positive growth rates
(Hargreaves et al., 2014). For this clade of
mediterranean-climate mustards that typically grow in
dry, low-competition, rocky or sandy environments, we
find a direct link between the ability of plant genotypes
(families) within a species to maintain fitness along a
water availability gradient as measured experimentally,
and a species’ realized niche breadth, its geographic
range and climatic breadth. Individual genotypes (fami-
lies) within species with smaller ranges had lesser

8 of 11
ability to maintain fitness, i.e., had lower fitness homeo-
stasis, across a range of field-relevant water availability.
We also found some support for trade-offs between spe-
cialist and generalist niches: maximum fitness traded off
with fitness homeostasis. Species good at maintaining
fitness across a broader range of water availability had
marginally lower maximum fitness than species with
more variable performance across water availability
(Figures 1a and 4).
Other studies have linked similar measures of individ-
ual tolerances, measured from performance curves, to a spe-
cies’ geographic range. For example, Mimulus species with
broader thermal tolerances also had larger geographic
ranges (Angert et al., 2011). Widely distributed Acacia spe-
cies showed less variability in water- and drought-related
leaf traits than did more narrowly restricted species
(Pohlman et al., 2005). Thus, in several systems, it has been
shown that genotypic, or in our case, maternal family-level
plasticity is a good predictor of the range of conditions a
species occupies in the field. On the other hand, other
studies testing the relative importance of genetic variation
and fundamental niche of genotypes in determining niche
breadth show that both of these factors can contribute to
the species-level realized niche and that their relative
importance may be species specific (Angert et al., 2011;
Sheth & Angert, 2014). As a whole, individual niche
breadth as determined by individual-level or maternal
family-level performance curves appears to be an impor-
tant component of the species-level realized niche.
Nevertheless, there are still relatively few instances in
which the niche has been quantified in this way, even if
performance curve studies may provide some of the most
valuable information about how an organism responds to
a changing climate.
In contrast to our results, there appears to be little rela-
tionship between measurements of individual fitness
homeostasis measured experimentally and the realized
niche in some taxa and different mechanisms might con-
tribute to niche limits above and beyond physiological tol-
erance (Sheth et al., 2020). For example, across the clade
Lasthenia, all species studied exhibited optimal perfor-
mance under the same experimental water inundation con-
ditions, despite occupying quite different hydrological field
niches (Tittes et al., 2019). Thus, in Lasthenia, the funda-
mental niche as measured by individual performance was
not closely linked to the realized field hydrological niche
and was a trait that seemed to be evolutionarily
constrained. As more studies take these approaches, a
meta-analysis incorporating habitat type and productivity
as well as the relative roles of genetic diversity, local adap-
tation, and plasticity would be a next step to a more
nuanced understanding of the drivers of niche breadth and
when knowledge of individual plasticity allows predictive
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PEARSE ET AL.
power to our understanding of how species occupy variable
environments (Hargreaves et al., 2014).
Our study suggests that performance curves, how fit-
ness of an organism responds to the environment, predict
aspects of the realized niche of organisms, the range of
environments in which they live and their geographic
range size. In using this approach more broadly, it will be
important to consider the range of environments sampled
across the performance curve. For example, in our study,
we see a substantial decrease in fitness of all Streptanthus
species at low water. However, with high amounts of
water, we see a decline in fitness in only roughly half of
Streptanthus species. This contrasts with results from per-
formance curve studies of various organisms that exhibit
a unimodal relationship between some abiotic variable,
typically temperature or water, and fitness (Angert
et al., 2011; Kingsolver & Buckley, 2017; Sheth &
Angert, 2014). Our experiment used field-realistic
watering levels that spanned very dry to very wet condi-
tions for species across this clade. Our maximum
watering levels met or exceeded the maximum rainfall
rates experienced by most of the species over a 30-year
interval. For two species, we might not have captured
high water levels that caused physiological stress
(Appendix S1: Table S1). However, we did not see
declines in fitness in some species in which our maxi-
mum water treatment greatly exceeded historical maxi-
mum precipitation (e.g., C. inflatus and C. coulteri). It
could be that biotic interactions like competition
(e.g., Cacho et al., 2014) or pathogens are more important
in limiting the realized niche in wetter parts of the water
resource gradient in nature. In our study, we considered
a gradient of water availability that was greater than
what occupants of the field site would typically experi-
ence, and we show that the range of water over which a
Streptanthus genotype maintains high fitness still predicts
aspects of its realized niche and range size.
A large caveat for our results is that we only sampled
one population per species; populations were selected to
be central within the climate distribution of each species
(Appendix S1: Figure S1). Our study design, as a trade-off
to replicating at the species level, has no replication
among populations within species. Thus, the question
arises how well one can infer species-level properties
from single populations. A rejoinder to this criticism is
that, if our study populations were not representative of
the species as a whole, it would be unlikely that we
would have found the results that we did: 59% of the vari-
ation in range-wide CWD niche breadth and 43% of the
variation in geographic range size could be explained by
fitness homeostasis as measured in the lathhouse on the
single population per species. Moreover, a qualitative
impression of performance curves of other studies reveals

ECOLOGY
that variation in performance curves among populations
within species is generally less than that among species
(e.g., Figure 4; Sheth & Angert, 2014). In addition,
because we used field-collected seed, we also cannot rule
out maternal effects or epigenetic contributions to the
patterns that we observed. Repeating this study with
more populations per species and more families would
increase the power to separate local adaptation and
individual-level effects, other factors that likely contrib-
ute to predicting species-level properties from samples
within species.
Biologists are increasingly asked to assess the adap-
tive capacity of species in terms of their ability to with-
stand changing climate, novel species interactions, and
loss of habitat (Williams et al., 2008). We suggest that
species that maintain fitness optima over only a narrow
range of conditions may be particularly vulnerable to
predicted hotter, drier future climates as well as loss of
genetic diversity, and they may be good candidates for
assisted translocation. Conversely, for plastic or generalist
species that can maintain fitness over a range of environ-
ments, plasticity may buy time for in situ adaptation to
future climates.
A C K N O WL E D G M E N T S
We thank Jessica Aguilar for help in conducting the
greenhouse experiment, and feedback from anonymous
reviewers who improved this manuscript. Any use of
trade, firm, or product names is for descriptive purposes
only and does not imply endorsement by the
U.S. Government.
CONFLICT OF INTEREST
The authors declare no conflict of interest.
DATA AVAILABILITY STATEMENT
Data (Strauss et al., 2022) are available in Dryad at
https://doi.org/10.5061/dryad.4b8gtht7s.
ORCID
Ian S. Pearse https://orcid.org/0000-0001-7098-0495
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SU PP O R TI N G I N F O RMA TI O N
Additional supporting information can be found online
in the Supporting Information section at the end of this
article.
How to cite this article: Pearse, Ian S.,
Patrick McIntyre, N. Ivalú Cacho, and Sharon
Y. Strauss. 2022. “Fitness Homeostasis across an
Experimental Water Gradient Predicts Species’
Geographic Range and Climatic Breadth.” Ecology
e3827. https://doi.org/10.1002/ecy.3827