Thermal performance of larval longfin dace (Agosia

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Environ Biol Fish
DOI 10.1007/s10641-014-0270-7
Thermal performance of larval longfin dace (Agosia
chrysogaster), with implications for climate change
Matthew J. Troia & James E. Whitney & Keith B. Gido
Received: 25 November 2013 / Accepted: 28 April 2014
# Springer Science+Business Media Dordrecht 2014
Abstract Temperature is an important factor affecting
the distribution of freshwater fishes. The longfin dace
(Agosia chrysogaster) is endemic to the Gila River basin
of the southwestern USA and northern Mexico and
occupies a range of thermal environments from cool
mountain tributaries to warm desert rivers but information about its thermal biology is limited, particularly for
larvae. We quantified the effect of rearing temperature
on survival, growth capacity, and critical thermal maximum (CTM) of larval longfin dace. Broodstocks of
longfin dace were collected from two sites in the upper
Gila River in New Mexico from which larvae were
hatched and reared for 22 days in indoor aquaria at
constant temperatures ranging from 18.0 to 31.0 °C.
Growth capacity peaked at 27.0 °C and was 21 % greater for larvae hatched from the upstream compared to the
downstream broodstock, indicating intraspecific variability in growth capacity. CTM increased with rearing
temperature and ranged from 33.9 to 39.9 °C, indicating
that thermal acclimation influences maximum thermal
tolerance. CTM and acclimation response ratio of larvae
are lower than those of adult longfin dace measured in a
previous study, suggesting that larvae are more sensitive
and less responsive to thermal stress than adults. Water
temperatures in 2012 from six sites in the upper Gila
River basin did not exceed 27.0 °C and larval growth
capacities in May of 2012 ranged from 5 to 28 % of the
maximum growth capacity. We assert that rising
M. J. Troia (*) : J. E. Whitney : K. B. Gido
Division of Biology, Kansas State University,
116 Ackert Hall, Manhattan, KS 66506, USA
e-mail: troiamj@ksu.edu
temperatures may increase larval growth rates, although
this will depend on resource limitation and shifts in
community interactions.
Keywords Growth capacity . Critical thermal
maximum . Thermal acclimation . Larval fish . Gila River
Introduction
Temperature exerts a strong influence on the physiology, life history, population dynamics, and distribution of
freshwater fishes (Brett 1956; Myrick and Cech 2000).
Maximum temperature tolerance, measured as critical
thermal maximum (CTM), is probably the most studied
aspect of thermal biology because its effect on survival
is direct and acute (Lutterschmidt and Hutchison 1997;
Beitinger et al. 2000). In temperate streams, species
exhibit innate differences in maximum thermal tolerance, which constrain their distribution along the river
continuum (Rahel and Hubert 1991). Individuals of the
same species can also differ in maximum thermal tolerance stemming from reversible (Bennett and Beitinger
1997; MacNutt et al. 2004) or non-reversible (Kinne
1962; Schaefer and Ryan 2006) acclimation as well as
genetically-based local adaptation (Otto 1973; Fangue
et al. 2006). Maximum thermal tolerance can also increase with age (Hokanson et al. 1973), but this aspect
of thermal tolerance has been assessed in a limited
number of species (Rombough 1997).
Thermal performance of a species can be characterized as a thermal reaction norm that describes a change
Environ Biol Fish
in a physiological rate or behavior along a gradient of
temperatures (Angilletta 2009). Because physiological
rates and behavior vary over a range of non-lethal temperatures and these performance differences can influence ecological processes (e.g., Taniguchi and Nakano
2000), characterization of thermal optima and
breadths of ecologically-relevant physiological and
behavioral traits is crucial for understanding the
distribution of species across thermally-variable
landscapes. For example, feeding rate is a commonly measured thermal reaction norm for freshwater fishes because it is a practical surrogate for
estimating energetic requirements (Alvarez et al.
2006). Metabolic rate and growth capacity are
informative thermal reaction norms for testing evolutionary hypotheses such as countergradient variation (Schaefer and Walters 2010; Baumann and
Conover 2011) and macroecological theories such
as the metabolic theory of ecology (Brown et al.
2004; Schaefer 2012). Thermal reaction norms in
growth rate of larvae also are ecologically meaningful because larval survival can drive population
dynamics of many freshwater fishes (Velez-Espino
et al. 2006; Piffady et al. 2010). Larval fish that
grow faster are more resistant to starvation (Einum
and Fleming 1999), able to exploit a broader array
of food resources (Wankowski 1979), and avoid
size-limited predators (Werner and Gilliam 1984).
Characterization of thermal tolerance and reaction
norms of fishes is necessary to predict effects of rising
water temperatures due to anthropogenic climate
(Morgan et al. 2001; Caissie 2006) and land cover
(LeBlanc et al. 1997) change. This is particularly important in arid regions where, in addition to rising air
temperatures, reduced flows will further increase thermal maxima and variability in streams. Much of the
arid southwestern United States is drained by the
Colorado River and its major tributaries, including
the Gila River. This region contains many endemic
fishes that are imperiled due to fragmentation, flow
alteration, and the presence of non-native species
(Minckley and Deacon 1968; Pool and Olden 2011).
Several studies that characterized the acute maximum
thermal tolerances of Colorado River fishes provide a
basis for predicting how species will respond to altered thermal regimes (Otto 1973; Deacon et al.
1987). Carveth et al. (2006) measured maximum thermal tolerance and acclimation response ratio (a measure of thermal acclimation) of ten native and four
non-native species and found that these thermal performance metrics were variable among species but
were not significantly greater for native compared to
non-native species, indicating that native desert fishes
are not necessarily more resistant to rising
temperatures. Other studies have compared growth
and survival rates of native Colorado River fishes
exposed to chronic temperature differences. For
example, Widmer et al. (2006) showed that survival
and growth of loach minnow (Tiaroga cobitis) over
30 days was suppressed at temperatures above
28.0 °C, but mortalities occurred when temperatures
were above 30.0 °C. Few studies have evaluated the
thermal performance of larval desert fishes (but see
Bestgen 2008), despite the influence of this life stage
on population dynamics (Velez-Espino et al. 2006;
Piffady et al. 2010).
We studied lethal and non-lethal aspects of the thermal biology of the longfin dace (Agosia chrysogaster,
Girard 1856), a cyprinid endemic to the lower Colorado
River basin. This species occupies a variety of habitats
along the river continuum, from high-gradient mountain
tributaries to low-gradient desert river mainstems
(Minckley and Barber 1971). As a stream-size generalist, they experience a range of thermal environments,
making the study of its thermal biology informative for
predicting distributional responses to rising temperatures. Longfin dace also influence ecosystem properties, excreting up to 10 % of the nitrogen that is
taken up by algae (Grimm 1988), which makes the
study of their physiology and distribution relevant to
the understanding of desert stream ecosystems.
Previous investigators reported maximum thermal
tolerance and plasticity (due to thermal acclimation)
in thermal tolerance of adult longfin dace from a
tributary of the Gila River in Arizona (Carveth et al.
2006); however, the thermal performance of larvae
and temperature dependence of other performance
metrics of longfin dace remain unknown. We had
three objectives for this study: (1) measure thermal
reaction norms in larval survival and growth capacity to characterize the thermal optimum and breadth,
(2) measure CTM of larvae reared at a range of
temperatures to test for plasticity in maximum thermal tolerance due to acclimation, and (3) compare
thermal optimum to stream temperature regimes
throughout the upper Gila River basin to predict
potential changes in larval performance in response
to rising stream temperatures in the future.
Environ Biol Fish
Methods
Study area, collection of broodstock, stream temperature
data
We collected broodstocks of adult (>50 mm total
length) longfin dace from two sites on the mainstem
of the upper Gila River in southwestern, New
Mexico. The upstream broodstock collection site
(1,329 m above sea level) is located 20.1 river km
from the downstream broodstock collection site
(1,239 m above sea level) (Fig. 1). We used a seine
to collect 18 to 25 individuals from each site between
18 and 22 March 2013. Adults were transported to
the experimental stream facility at Konza Prairie
Biological Station in Kansas, USA and housed in
experimental stream channels (see Matthews et al.
2006 for description of experimental stream
Fig. 1 Upstream (open circle)
and downstream (filled circle)
broodstock collection sites of
longfin dace (Agosia
chrysogaster) and locations of
three tributary (open squares) and
three mainstem (closed squares)
temperature recording sites from
the upper Gila River in
southwestern New Mexico. Gray
shading indicates extent of the
Gila River Basin and dashed box
indicates the upper Gila River and
extent of the inset map
channels). Spawning occurred between 26 and 30
April 2013 and hatched larvae were removed from
experimental stream channels on 1 May 2013 and
transported to an indoor laboratory at Kansas State
University in Manhattan, Kansas where growth capacity and thermal tolerance experiments were carried out.
Stream water temperature was recorded every hour
from January 2012 to January 2013 at six sites within
the distributional limits of longfin dace that ranged in
elevation from 1,360 to 1,735 m above sea level (Paroz
et al. 2006; Whitney 2010). These temperature recording sites are located upstream of the two broodstock
collection sites, with three located on the Gila River
mainstem (hereafter ‘Mainstem-Up’, ‘Mainstem-Mid’,
and ‘Mainstem-Down’) and three located on tributaries
(hereafter ‘West Fork’, ‘Middle Fork’, and ‘East Fork’)
of the Gila River (Fig. 1).
Environ Biol Fish
Experimental procedures
Growth capacity and survival We measured growth
capacity as the temperature-specific growth rate at unlimited feeding conditions (Baumann and Conover
2011). At the start of the experiment, a subset of 12 to
15 larvae from each broodstock were euthanized with a
lethal dose of MS-222 (tricaine methanosulfonate) and
preserved in 5 % buffered formalin to estimate starting
body size. From each broodstock, ten larvae were reared
in aerated 2 L aquaria that were incubated in 75 L water
baths maintained at 18.0, 20.4, 21.5, 23.1, 23.7, 25.4,
26.4, 28.8, 29.4, or 30.1 °C. Larvae were fed an excess
of live brine shrimp nauplii (Ocean Star International,
Inc., Snowville, UT) twice per day (08:00 and 20:00 h)
for 22 days. Unconsumed food and waste were siphoned
and a 50 % water change was conducted once per day
(08:00 h). We measured the proportion of individuals
surviving in each replicate aquarium after 22 days.
Following the 22-day growth capacity experiment, a
subset of five to seven individuals were retained for
measurement of CTM and remaining individuals were
euthanized and preserved in formalin. Preserved specimens were eviscerated, padded dry with a paper towel
and weighed to the nearest 0.1 mg. Daily growth rate of
each individual was calculated as the eviscerated wet
mass on day 22 minus the mean eviscerated wet mass of
individuals euthanized at the start of the growth capacity
experiment divided by 22 days.
Acute thermal tolerance We measured CTM using the
loss of righting response (Lutterschmidt and Hutchison
1997). Following the 22 day growth capacity experiment, temperatures in all aquaria were equilibrated to
24.0 °C for 36 h to minimize the effect of acute thermal
and handling stress on CTM measurements. Fish were
also fasted for 36 h prior to trials. Five to seven individuals from each temperature treatment were selected to
represent the range of body sizes present in each aquarium, allowing us to statistically control for the potential
effect of body size on CTM. Each test individual was
placed in a 70 ml cup that submerged in an 18 L water
bath and heated at a rate of 0.7 °C·min−1 starting at
24.0 °C. Oxygen concentrations were measured at all
temperatures with a dissolved oxygen probe (Yellow
Springs Instruments, Yellow Springs, Ohio, USA) and
remained >98 % saturated throughout the duration of
each trial. The temperature at which individuals lost
righting response was recorded and test fish were
immediately removed, euthanized, preserved, and later
eviscerated and weighed for body size.
Data analysis
We used growth capacity of the largest individual from
each aquarium as the maximum growth capacity for
each temperature treatment. To characterize the temperature dependence of maximum growth capacity,
we used nonlinear least squares regression with a 3
parameter Gaussian function (Eq. 1).
2
growth capacity ¼ B e−ðtemperature−AÞ =2C
2
ð1Þ
This method allows for the estimate of the optimal
temperature (A), maximum performance at the optimum
(B), and the breadth of performance (C), and is useful
for characterizing thermal reaction norms in physiological processes (Angilletta 2009; Schaefer 2012).
Parameter estimates and non-overlapping standard errors were used to infer statistically significant differences in optimum growth capacity and maximum
growth capacity between larvae from the upstream and
downstream broodstocks.
We combined data from the two broodstocks and
used linear regression to test for a relationship between
rearing temperature and two response variables: survival
and CTM. If significant linear relationships were detected, we used analysis of covariance (ANCOVA) to test
for differences in slopes and y-intercepts between
broodstocks. Lastly, we calculated the acclimation response ratio (ARR), which is the slope of a linear
regression equation describing the relationship between
acclimation temperature and CTM. ARR is an index of
the capacity for thermal acclimation with higher values
indicating greater ability to acclimate to changing temperatures (Claussen 1977).
We used the experimentally-derived thermal reaction
norms in growth capacity and stream water temperatures
from the six temperature recording sites to estimate the
potential larval growth capacity throughout the Gila
River basin and compare these estimates to the
experimentally-derived optimum. The spawning season
of longfin dace lasts from December to July and peaks in
April (Lewis 1978), so we calculated mean daily water
temperatures from 1 to 31 May 2012 at each of the six
temperature recording sites as an estimate of typical
rearing temperatures for larvae. Daily increase in larval
body mass was calculated from the mean temperature of
Environ Biol Fish
Fig. 2 Relationship between rearing temperature and growth
capacity of larval longfin dace. Open and closed circles represent
maximum growth rates from the upstream and downstream
broodstock collection sites, respectively. Dashed and solid lines
are best fit lines for the upstream and downstream populations,
respectively, using non-linear regression with 3-parameter Gaussian functions
each day in May 2012 using the 3-parameter Gaussian
functions fit to the thermal reaction norm in growth
capacity. These daily growth rates were summed for
May 2012 at each temperature recording site (hereafter
‘Total May Growth’). We estimated Total May Growth
using the Gaussian functions from the upstream and
downstream broodstocks to explore the range of growth
rates stemming from variability in growth capacity between broodstocks.
significantly between broodstocks (t-test; t19 =−0.73;
P=0.48). Fitted 3-parameter Gaussian functions indicated statistically significant relationships between
rearing temperature and maximum growth capacity
for the upstream (non-linear least squares regression,
R2a = 0.89, P < 0.001) and downstream (non-linear
least squares regression, R 2 a = 0.90, P < 0.001)
broodstocks. Maximum growth capacity peaked at
26.6 °C and was 6.3 mg ·day−1 for the upstream
broodstock and at 27.0 °C and was 5.2 mg·day−1
for the downstream broodstock. Ninety five percent
confidence intervals of the upstream and downstream broodstocks overlapped for optimum temperature but not maximum growth capacity, indicating
that maximum growth capacity was significantly and
17 % greater for the upstream broodstock compared
to the downstream broodstock. Optimum temperatures did not differ significantly between these two
broodstocks (Fig. 2). Mean 22-day survival rate was
82 % and was not significantly correlated with
rearing temperature (linear regression; R2adj =0.01;
P=0.30).
Results
Growth capacity and survival
Mean larval body size at the start of the experiment for
the upstream (1.32 mg) and downstream (1.26 mg)
broodstocks were not significantly different (t-test;
t21 =0.58; P=0.57). Mean larval body size after 22 days
ranged from 26.0 to 141.1 mg for the upstream
broodstock collection site and 24.3 to 124.2 mg for the
downstream broodstock collection and did not differ
Environ Biol Fish
Fig. 3 Mean daily water temperature for 2012 at (A) three tributary and (B) three mainstem temperature recording sites in
the upper Gila River basin. The dashed line indicates
Fig. 4 Total growth capacity of larvae estimated for May 2012 at
six temperature recording sites in the upper Gila River basin. Total
growth is the summed estimate of daily growth which was calculated from mean daily water temperature at each temperature
recording site and the experimentally-derived 3-parameter Gaussian functions fit to the thermal reaction norms in growth capacity
from the upstream (open bars) and downstream (filled bars)
broodstock collection sites. Sites are ordered from highest (top)
to lowest (bottom) elevation. See Fig. 1 for locations of temperature recording sites
experimentally-derived optimum for larval growth capacity.
See Fig. 1 for locations of temperature recording sites
Fig. 5 Relationship between rearing temperature and CTM (measured as loss of righting response) of 22 day old longfin dace
acclimated at 24.0 °C for 36 h. Open and closed circles represent
progeny from individuals collected from the upstream and downstream broodstock collection sites, respectively. The solid line
represents a best fit line for both populations and the slope represents the acclimation response ratio (ARR)
Environ Biol Fish
Mean daily water temperature in 2012 reached a
maximum of 26.1 °C in the Middle Fork of the Gila
River, whereas mean daily water temperature in 2012 at
the West Fork reached a maximum of only 22.8 °C
(Fig. 3). Mean daily water temperature in May of 2012
ranged from 15.3 °C in the West Fork to 19.1 °C in the
Middle Fork. Total May Growth ranged from 15.3 mg
and 9.2 mg (based on equations derived from upstream
and downstream broodstocks, respectively) in the West
Fork to 46.3 mg and 39.9 mg in the Middle Fork. By
comparison, Total May Growth at the experimentallyderived optimum (27.0 °C) would be 193.4 mg and
162.7 mg based on equations from the upstream and
downstream broodstocks, respectively (Fig. 4).
Acute thermal tolerance
CTM increased with rearing temperature and ranged
from 33.9 to 39.8 °C (Fig. 5). Because CTM was significantly correlated with body size (linear regression;
R2adj =0.27; P<0.001), effects of broodstock and rearing
temperature on CTM were assessed using the residuals
of the linear relationship between CTM and body size.
Residualized CTM increased linearly with rearing temperature (ANCOVA; F3,91 =6.70; P <0.001), indicating
that maximum thermal tolerance depends on acclimation and is independent of body size. Thermal tolerance
did not differ between broodstock collection sites
(ANCOVA; F3,91 =1.15; P=0.25), nor was there a significant interaction between rearing temperature and
broodstock (ANCOVA; F 3,91 = −0.92; P = 0.36).
Acclimation response ratio was 0.37, meaning that
CTM increases by 0.37 °C for every 1.0 °C increase in
acclimation temperature.
Discussion
Growth capacity
Growth capacity of larval longfin dace exhibited a
Gaussian-shaped response to rearing temperature.
Reduced growth capacity above-optimal temperatures,
independent of food availability, is caused by slowing of
enzyme activity and loss of structural integrity of cell
membranes, whereas growth rate at below-optimal temperatures is limited by metabolic rate (Angilletta 2009).
Mean daily temperatures during May 2012 were variable among sites, which transmitted to variable
estimates of Total May Growth among sites in the upper
Gila River basin. In particular, temperatures and Total
May Growth in the West Fork were lower than in the
Middle Fork and mainstem sites. Although peak
spawning has been documented in April (Lewis 1978),
the spawning season is protracted (December through
July) and populations likely vary in peak spawning date
corresponding to the variable temperature cues in different streams (Heggberget 1988). Nevertheless, daily
water temperatures were lower in the West Fork
throughout the entirety of the spawning season so, regardless of peak spawning date, larvae hatched at the
same time and presented with equal food resources
should exhibit reduced growth in the West Fork compared to the Middle Fork and mainstem sites.
Maximum growth capacity was higher for the upstream compared to the downstream broodstocks. This
finding supports the hypothesis of metabolic cold adaptation (also called countergradient variation), which
posits that populations exposed to colder environments
(generally associated with higher elevation or latitude)
will evolve elevated metabolic rates and growth capacities at the same optimal temperature compared to populations exposed to warmer environments. Elevated
metabolic and growth rates at higher elevations can be
adaptive because growing seasons are shorter
(Baumann and Conover 2011). Three conditions must
be satisfied for metabolic cold adaptation to evolve: (1)
gene flow is restricted between populations, (2) temperature regimes differ between populations, and (3) selection for faster growth at higher elevations must be strong
enough to outweigh the costs (Arnott et al. 2006).
Although our thermal reaction norms support the metabolic cold adaptation hypothesis, more information is
necessary to confirm this hypothesis. Gene flow between the two populations in this study that were separated by only ~ 20 km is likely; however, population
genetic data to estimate the level of genetic isolation
between populations as well as thermal reaction norms
from other populations, particularly those that are located farther upstream and are exposed to lower temperatures, would elucidate whether metabolic cold adaptation occurs among populations of longfin dace in the
upper Gila River basin.
Acute thermal tolerance
Previous estimates of CTM for adult longfin dace collected in tributaries of the Gila River in Arizona and
Environ Biol Fish
acclimated at 25.0 °C and 30.0 °C were 38.2 °C and
40.5 °C, respectively (Carveth et al. 2006). Lower
CTMs in our study could stem from the earlier developmental stage evaluated in our study compared to that
of Carveth et al. (2006). In other temperate freshwater
species, larvae exhibit reduced thermal tolerance compared to juveniles and adults due to a limited ability to
acclimate metabolic rates (Hokanson et al. 1973;
Rombough 1997). Acclimation response ratio was also
lower for our New Mexico population (0.37) compared
to the Arizona population (0.44) (Carveth et al. 2006),
which supports the prediction that larvae are less responsive to temperature changes than adults (Hokanson
1973; Rombough 1997). Alternatively, lower CTMs in
our study could be due to local adaptation to lower
temperatures in the higher elevation populations of
New Mexico compared to Arizona (Fangue et al.
2006) or differences in rearing temperatures experienced by test fish between these two studies (Schaefer
and Ryan 2006). The contribution of these alternative
explanations cannot be tested independent of age, using
the currently available information.
Longfin dace in our study were variable in thermal
tolerance (CTM), which was related to the different
temperatures experienced during the 22-day rearing period. Do these findings suggest that longfin dace at
colder sites (e.g., West Fork) are more susceptible to
short term (i.e., several days to weeks) temperature
fluctuations compared to those at warmer sites (e.g.,
Middle Fork)? This depends on the relative contribution
of reversible acclimation versus non-reversible (i.e.,
developmental) acclimation to the variation in CTM that
we observed (Schaefer and Ryan 2006; Angilletta
2009). If non-reversible acclimation is the overriding
cause for the positive relationship between rearing temperature and thermal tolerance, then fish reared at low
temperatures would be susceptible to short term temperature fluctuations that surpass the CTM because their
ability to acclimate to rising temperatures over the
course of several days would be limited. By contrast,
if acclimation is reversible, then fish reared at low
temperatures would acclimate to short-term
temperature increases and would not be as vulnerable
to thermal stress. Schaefer and Ryan (2006) measured
the independent effects of reversible and non-reversible
acclimation in zebrafish (Danio rerio) and demonstrated
that reversible acclimation has a stronger effect on CTM
than does non-reversible acclimation. Therefore, if this
phenomenon is general among cyprinids, it is likely that
the majority of variation in thermal tolerance of larval
longfin dace in this study was due to reversible acclimation (regardless of the thermal environment in which
they developed), which would buffer longfin dace from
short-term temperature fluctuations.
Implications for environmental change
Anthropogenic environmental change including changes in riparian and catchment land cover, surface water
diversions and impoundments, and rising air temperatures—has and will continue to increase the temperatures of freshwater habitats worldwide (Poole and
Berman 2001). With regard to stream fish responses to
climate change, much focus has been placed on the
potential negative effects on cold-water species such as
salmonids that occupy high elevation streams in western
North America (MacNutt et al. 2004; Wenger et al.
2011). Comparatively less is known about the potential
responses of cool- and warm-water stream fishes to
warming (but see Buisson et al. 2008). Lyons et al.
(2010) used species distribution models to forecast the
distributional changes of stream fishes in Wisconsin
under several warming scenarios and predicted that all
cool-water species will decline in distribution whereas
warm-water species may increase or decline. Our results
predict that warming will increase larval growth capacity of longfin dace because current water temperatures
throughout the upper Gila River basin never exceeded
their optimum of 27.0 °C during 2012. Thus, humaninduced warming would increase growth capacity of
larval longfin dace. If food is not limiting, faster growth
should transmit to increased population-level performance because larval growth capacity often is positively
associated with age-0 recruitment (Wankowski 1979;
Werner and Gilliam 1984; Einum and Fleming 1999)
and intrinsic rate of population increase (Velez-Espino
et al. 2006; Piffady et al. 2010). From the perspective of
acute thermal tolerance, it appears unlikely that
warming will negatively impact longfin dace in the
upper Gila River basin because water temperatures during 2012 did not approach CTM, regardless of the
rearing temperature. We also show that longfin dace
have greater acclimation potential than other desert fishes (Carveth et al. 2006), which suggests that plasticity
may act as a stronger buffer against the negative impact
of rising temperatures for this species compared to other
native and non-native fishes of the desert southwest
(Culumber and Monks 2014). Increasing stream
Environ Biol Fish
temperatures are likely to result in complex changes in
community level processes such as resource abundance,
competition and predation (Davis et al. 1998).
Nevertheless, with all else equal, warming should generally favor longfin dace in the upper Gila River basin.
Acknowledgments We thank Josh Perkin for assistance with field
collections, Michael Denk and Rebecca Zheng for assistance with
experimental procedures and data collection, and Jake Schaefer for
assistance with experimental design. The Konza Prairie Biological
Station provided use of the experimental stream facility. This research
was funded by the National Science Foundation (DEB#1311183), the
Southwestern Association of Naturalists, Prairie Biotic Research Inc.,
the Kansas Academy of Science, and the Bureau of Reclamation
Water Smart program. Longfin dace were collected and housed under
the permission of the New Mexico Game and Fish Department
(permit #3351), Konza Prairie Biological Station (permit ID#221)
and the Institutional Animal Care and Use Committee (permit #2996)
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