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Projected impacts of climate change on stream salmonids with implications for resilience-based management

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The sustainability of freshwater fisheries is increasingly affected by climate warming, habitat alteration, invasive species, and other drivers of global change. The State of Michigan, USA, contains ecologically, socioeconomically valuable coldwater stream salmonid fisheries that are highly susceptible to these ecological alterations. Thus, there is a need for future management approaches that promote resilient stream ecosystems that absorb change amidst disturbances. Fisheries professionals in Michigan are responding to this need by designing a comprehensive management plan for stream brook charr (Salvelinus fontinalis), brown trout (Salmo trutta), and rainbow trout (Oncorhynchus mykiss) populations. To assist in developing such a plan, we used stream-specific regression models to forecast thermal habitat suitability in streams throughout Michigan from 2006–2056 under different predicted climate change scenarios. As baseflow index (i.e., relative groundwater input) increased, stream thermal sensitivity (i.e., relative susceptibility to temperature change) decreased. Thus, the magnitude of temperature warming and frequency of thermal habitat degradation were lowest in streams with the highest baseflow indices. Thermal habitats were most suitable in rainbow trout streams as this species has a wider temperature range for growth (12.0–22.5°C) compared to brook charr (11.0–20.5°C) and brown trout (12.0–20.0°C). Our study promotes resilience-based salmonid management by providing a methodology for stream temperature and thermal habitat suitability prediction. Fisheries professionals can use this approach to protect coldwater habitats and drivers of stream cooling and ultimately conserve resilient salmonid populations amidst global change.
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Projected impacts of climate change on stream
salmonids with implications for resilience-based
management
Andrew K. Carlson
1
, William W. Taylor
1
, Kelsey M. Schlee
1
, Troy G. Zorn
2
, Dana M. Infante
1
1
Department of Fisheries and Wildlife, Center for Systems Integration and Sustainability, Michigan State University, East Lansing, MI 48824, USA
2
Michigan Department of Natural Resources, Marquette Fisheries Research Station, Marquette, MI 49855, USA
Accepted for publication November 16, 2015
Abstract The sustainability of freshwater fisheries is increasingly affected by climate warming, habitat alteration,
invasive species and other drivers of global change. The State of Michigan, USA, contains ecologically,
socioeconomically valuable coldwater stream salmonid fisheries that are highly susceptible to these ecological
alterations. Thus, there is a need for future management approaches that promote resilient stream ecosystems that
absorb change amidst disturbances. Fisheries professionals in Michigan are responding to this need by designing a
comprehensive management plan for stream brook charr (Salvelinus fontinalis), brown trout (Salmo trutta) and
rainbow trout (Oncorhynchus mykiss) populations. To assist in developing such a plan, we used stream-specific
regression models to forecast thermal habitat suitability in streams throughout Michigan from 2006 to 2056 under
different predicted climate change scenarios. As baseflow index (i.e., relative groundwater input) increased, stream
thermal sensitivity (i.e., relative susceptibility to temperature change) decreased. Thus, the magnitude of temperature
warming and frequency of thermal habitat degradation were lowest in streams with the highest baseflow indices.
Thermal habitats were most suitable in rainbow trout streams as this species has a wider temperature range for
growth (12.022.5 °C) compared to brook charr (11.020.5 °C) and brown trout (12.020.0 °C). Our study
promotes resilience-based salmonid management by providing a methodology for stream temperature and thermal
habitat suitability prediction. Fisheries professionals can use this approach to protect coldwater habitats and drivers
of stream cooling and ultimately conserve resilient salmonid populations amidst global change.
Key words: brook charr; brown trout; rainbow trout; Michigan; temperature; resilience
Introduction
As climate warming, habitat fragmentation, invasive
species and other drivers of global change alter aqua-
tic ecosystems throughout the world, managing fish-
eries for resilience has become an important
conservation framework (Hansen et al. 2015). Resili-
ence is the capacity of a system to absorb distur-
bances and retain its structure and function (Holling
1973). Managing aquatic ecosystems for resilience is
particularly important when they contain species that
are sensitive to ecological stressors. For instance,
with relatively low thermal tolerance thresholds
(Raleigh 1982a,b; Raleigh et al. 1986), salmonid
fishes may experience reductions in growth and sur-
vival due to temperature elevation caused by climate
change. Thus, managing these species and their
ecosystems for thermal resilience is an important
task. Collaboration among scientists, managers, poli-
cymakers and public stakeholders will be important
for developing management approaches commensu-
rate with the wide geographical distribution and high
socioeconomic value of salmonid populations (Isaak
et al. 2015; Snyder et al. 2015).
Temperature exerts a fundamental physiological
influence on fish metabolism, which regulates
growth, survival and reproduction of individuals
(Dodds & Whiles 2010) and ultimately the dynamics
Correspondence: A. K. Carlson, Department of Fisheries and Wildlife, Center for Systems Integration and Sustainability, Michigan State University, East Lansing,
MI, 48824 USA. E-mail: carls422@msu.edu
190 doi: 10.1111/eff.12267
Ecology of Freshwater Fish 2017: 26: 190204 Ó2015 John Wiley & Sons A/S. Published by John Wiley & Sons Ltd
ECOLOGY OF
FRESHWATER FISH
of populations and communities. For example, water
temperature is an important factor determining fish
distribution and assemblage composition (Magnuson
et al. 1979). Temperatures above species-specific
thermal maxima cause mortality; temperatures below
maxima alter fish growth, reproduction, abundance
and population size structure (Magnuson et al. 1997).
In addition, thermal warming can indirectly decrease
fish growth and survival by degrading water quality
(e.g., reduced dissolved oxygen; Ficklin et al. 2013).
Climate change is predicted to increase stream tem-
peratures and thereby alter thermal habitat suitability
(Lyons et al. 2010) and fish community composition
(Isaak et al. 2012). Moreover, dams, culverts, and
other anthropogenic barriers increase water tempera-
tures and decrease population and habitat connectiv-
ity (Lessard 2000; Hayes et al. 2006). Regardless of
mechanism, stream salmonids are sensitive to thermal
warming because they are adapted to cold and cool
water environments (Raleigh 1982a,b; Raleigh et al.
1986; Wehrly et al. 2007). Climate-driven increases
in water temperature may reduce salmonid growth,
reproduction, and survival in streams currently near
thermal optima. Thus, it is imperative that fisheries
professionals develop management strategies to pro-
mote thermal resilience and thereby conserve salmo-
nid populations in a warming climate.
The State of Michigan, USA, has ecologically and
socioeconomically valuable populations of brook charr
(Salvelinus fontinalis), brown trout (Salmo trutta), and
rainbow trout (Oncorhynchus mykiss) distributed
throughout 31,000 km of streams (Godby et al. 2007;
Tyler & Rutherford 2007). Projected air temperature
warming resulting from climate change is predicted to
increase stream temperatures in the Midwestern Uni-
ted States by 0.84.0 °C (Pilgrim et al. 1998; Lyons
et al. 2010). If temperatures exceed thermal optima for
these species, growth, reproduction, and/or survival
will decline, particularly during the warmest month of
the year (i.e., July; Zorn et al. 2011). Thus, Michigan
is an ideal study area for investigating how thermal
warming will impact coldwater streams and salmonid
population dynamics. This information will enable
fisheries professionals to develop resilience-based
management programmes: collaborative efforts among
scientists, managers, policymakers and public stake-
holders to maintain stream ecosystem structure and
function amidst global change.
The goal of this study was to evaluate the effects of
projected air temperature warming on salmonid ther-
mal habitat suitability in coldwater streams in Michi-
gan to facilitate the development of a resilience-based
management programme. Our first objective was to
measure the accuracy of stream-specific airwater tem-
perature regression models by backcasting stream tem-
peratures in 2006 and 2012, years with pre-existing air
and stream temperature metrics. Our second objective
was to forecast stream temperatures in 2036 and 2056
and project thermal habitat suitability for brook charr,
brown trout and rainbow trout growth and survival.
We predicted that water temperatures would increase
overall from 2006 to 2056 but unevenly among
streams and time periods (e.g., 20122036, 2036
2056) due to system-specific patterns of thermal warm-
ing and temporal variability in projected carbon diox-
ide (CO
2
) emissions (Arblaster et al. 2014). We
expected that thermal habitat impairment would occur
more frequently in streams with extensive surface run-
off than in groundwater-dominated, thermally buffered
systems (Sear et al. 1999; Krider et al. 2013).
Methods
Study area
Our study encompassed 30 coldwater streams
throughout the State of Michigan, USA (Fig. 1).
Streams were chosen in three Michigan Department
of Natural Resources (MDNR) management regions
[i.e., Upper Peninsula (UP), northern Lower Penin-
sula (NLP) and southern Lower Peninsula (SLP);
Table 1] to conduct the study at a statewide scale rel-
evant for salmonid management that also spanned a
latitudinal thermal gradient in which temperatures
increased from north (47.03°N) to south (42.64°N).
In addition, streams were selected to encompass a
hydrological gradient from surface runoff to ground-
water dominance by evaluating their relative base
flow, the component of streamflow attributable to
groundwater. We used a United States Geological
Survey report (Neff et al. 2005) to obtain each
streams base flow index (BFI), the mean rate of base
flow divided by the corresponding mean rate of total
streamflow. Baseflow index ranges from zero to one
with increasing groundwater input (Wahl & Wahl
1988); streams in our study ranged from 0.44 to 0.73
(Table 1). All BFI calculations were made using a
digital filter hydrograph separation technique (Arnold
& Allen 1999; Kelleher et al. 2012) whereby daily
streamflow records were partitioned into groundwater
and surface runoff components. Moreover, all streams
were important from a management standpoint as
they supported recreational fisheries for brook charr,
brown trout, rainbow trout or a combination of these
coldwater species. Only streams that had necessary
historical data for development of stream-specific
temperature regression models (i.e., field-measured
air and water temperatures) were selected. We devel-
oped a list of streams meeting these criteria (i.e., lati-
tudinal gradient, hydrological gradient, recreational
importance and historical data) using information
from the MDNR Better Fishing Waterswebpage
191
Resilience-based salmonid management
(MDNR 2015) and through personal communication
with MDNR Fisheries Division personnel (Tracy
Kolb, Todd Wills) and employees of the Michigan
Council of Trout Unlimited (Trout Unlimited 2015).
A minimum of five streams per region were selected
for each species to ensure regional replication. One
exception was the SLP, where brook charr and rain-
bow trout are not widely distributed and only four
streams contained one or both of these species. In
total, 16, 18 and 11 streams contained brook charr,
brown trout and rainbow trout, respectively. Eleven
streams supported more than one species, and four
supported all species (Table 1).
Stream-specific regression models
Historical air and water temperatures for each stream
were used to create stream-specific regression mod-
els. Daily air temperatures collected in summer
months (i.e., June, July and August) from 1990 to
2010 were compiled using the United States Histori-
cal Climate Network online interface (http://
cdiac.ornl.gov/). Air temperatures were measured at
the gauging station closest to each streams headwa-
ters, where MDNR water temperature gauges
recorded daily temperatures in summer months from
19902010. The most upstream gauge on each stream
was selected because temperatures are typically cool-
est and most optimal for salmonids in headwater
reaches. As such, we focused on these areas because
if their thermal habitat is degraded, downstream habi-
tat and salmonid populations will also be impaired.
Hydrological unit codes (HUCs) for each streams
sub-basin (HUC8) and subwatershed (HUC 12) were
identified using the National Hydrography Dataset
Plus Version 1 (NHDPlusV1) and the Watershed
Boundary Dataset (USEPA 2005). The North Ameri-
can Anthropogenic Barrier Dataset was used to locate
and omit gauges directly below dams, which elevate
temperatures compared to upstream reaches (Lessard
2000). For streams without MDNR gauges, a substi-
tute gauge on the nearest stream within the same sub-
watershed was used. Stream-specific regression
models were developed by pairing mean summer air
and water temperatures from recent years (i.e., 2002
2010) in Microsoft Excel (Table 2). Air temperature
coefficients represented indices of stream thermal
sensitivity (i.e., relative susceptibility to temperature
change; Kelleher et al. 2012) and were coupled with
air temperature projections (see below) to predict
future water temperatures. In addition, historical
warming in each stream was evaluated by comparing
air temperatures in 1976 and 2006 and multiplying
this temperature change by each streams air tempera-
ture coefficient to determine the magnitude of stream
warming.
Fig. 1. Map of brook charr, brown trout
and rainbow trout streams used for air
stream temperature modelling in Michigan,
USA. Streams and corresponding
identification numbers are listed in
Table 1. Triangles denote locations of
MDNR stream temperature gauges.
192
Carlson et al.
Air temperature projections
Three coupled climate models (CCMs) were used to
backcast mean June, July and August air tempera-
tures in 2006 and 2012 and forecast mean tempera-
tures in the same months in 2036 and 2056 for each
sub-basin: the Third Generation Coupled Global Cli-
mate Model (CGCM3, Canadian Centre for Climate
Modelling and Analysis), the CM2 Global Coupled
Climate Model (CM2, Geophysical Fluid Dynamics
Laboratory at the National Oceanic and Atmospheric
Administration) and the Hadley Centre Coupled
Model version 3 (HadCM3, Met Office, United King-
doms National Weather Service). All CCMs were
based on the World Climate Research Programme
(WCRP) Coupled Model Intercomparison Project
phase 3 (CMIP3) multimodel data set. Spatial down-
scaling was performed using the Bias-correction spa-
tial disaggregation (BCSD) approach to adjust the
resolution of the climate model (~200 9200 km) to
a scale germane for Michigan streams (12 912 km;
Maurer et al. 2007). The United States Forest
Services (USFS) Eastern Forest Environmental
Threat Assessment Center (EFETAC) in North
Carolina supplied mean June, July and August air
temperatures for Michigan sub-basins containing
coldwater salmonid streams. Projections were made
based on the CGCM3, CM2, and HadCM3 models
using area-weighted means for all years assuming the
Special Report of Emission Scenarios A2 and B1 cli-
mate forcing scenarios. The A2 scenario describes a
future world with rapid economic growth and effi-
cient energy technologies and predicts atmospheric
CO
2
concentrations to be 820 ppm in 2100. In con-
trast, the B1 scenario projects a convergent world
with a service and information economy and reduced
material consumption and predicts atmospheric CO
2
concentrations 550 ppm in 2100. Combining scenar-
ios was informative as they represent upper and
lower emission thresholds for stream temperature
prediction.
Stream temperature and thermal habitat suitability
projections
Stream temperatures were backcasted in 2006 and
2012 and forecasted in 2036 and 2056 by inputting
CCM air temperature predictions into stream-specific
Table 1. Descriptive information about 30 salmonid streams used for temperature modelling in Michigan, USA. Region refers to Michigan location [i.e., Upper
Peninsula (UP), northern Lower Peninsula (NLP) and southern Lower Peninsula (SLP)]. Sub-basin denotes the National Hydrography Dataset sub-basin (i.e.,
8-digit hydrological unit code). Map number refers to stream identifiers in Fig. 1. Species refers to salmonids present in each stream [i.e., brook charr (BKC),
brown trout (BNT) and rainbow trout (RBT)]. BFI denotes baseflow index, the mean rate of base flow divided by the corresponding mean rate of total
streamflow (Neff et al. 2005).
Stream Region Subbasin Map number Species BFI
Bark River UP Cedar-Ford 18 BNT 0.60
Bear Creek SLP Kalamazoo 14 BNT 0.58
Black River NLP Black 1 BKT 0.63
Boardman River NLP Boardman-Charlevoix 2 BKT, BNT 0.58
Bryan Creek UP Escanaba 19 BKT 0.62
Canada Creek NLP Black 3 BKT 0.63
Carp River UP Carp-Pine 20 BKT, BNT, RBT 0.64
Cedar Creek SLP Lower Grand 10 BKT, BNT 0.50
Cedar River SLP Cedar-Ford 11 BNT 0.60
Chocolay River UP Betsy-Chocolay 25 RBT 0.62
Davenport Creek UP Brevoort-Millecoquins 26 RBT 0.65
Duke Creek SLP Lower Grand 15 BKT, BNT 0.50
East Branch Fox River UP Manistique 27 BKT, BNT 0.73
Elm River UP Keweenaw Peninsula 21 RBT 0.45
Escanaba River UP Escanaba 22 BKT, BNT 0.44
Iron River UP Brule 28 BKT 0.59
Little Indian River UP Manistique 29 BKT 0.73
Little Muskegon River NLP Muskegon 4 RBT 0.62
Manistee River NLP Manistee 5 BKT, BNT, RBT 0.65
Martin Creek SLP Pere Marquette-White 16 BKT 0.61
Pere Marquette River NLP Pere Marquette-White 6 BNT 0.61
Pigeon River NLP Cheboygan 7 BNT, RBT 0.65
Pine River SLP Pine 12 BNT 0.65
Pine River NLP Manistee 8 BKT, BNT, RBT 0.49
Prairie Creek SLP Lower Grand 13 BNT 0.50
Rogue River SLP Lower Grand 17 BNT, RBT 0.50
Salmon Trout River UP Keweenaw Peninsula 23 BKT 0.45
Tahquamenon River UP Tahquamenon 24 BNT 0.55
West Branch Sturgeon River NLP Cheboygan 9 BKT, BNT, RBT 0.65
Yellow Dog River UP Dead-Kelsey 30 RBT 0.52
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Resilience-based salmonid management
Table 2. Stream-specific temperature regression models, with summary statistics, historical warming and projected future warming for each system. F(i.e., F
1,7
) and Pvalues refer to the air temperature (A) parameter
used to predict stream temperature (S). DAand DSdenote changes in air and stream temperatures over corresponding time periods. For 20062012, 20122036 and 20362056, predicted changes in Aand Sare
provided for the A2 and B1 climate forcing scenarios (B1 in parentheses).
Stream name Regression SE FPR
2
ΔA1976
2006
ΔS1976
2006 ΔA20062012 ΔS20062012 ΔA20122036 ΔS20122036 ΔA20362056 ΔS20362056
Bark River S=0.32A +12.98 0.05 49.41 <0.01 0.86 +1.72 +0.55 +1.09 (+1.23) +0.35 (+0.39) +2.74 (+3.16) +0.88 (+1.01) 0.69 (0.65) 0.22 (0.21)
Bear Creek S=0.23A +11.61 0.03 46.74 <0.01 0.85 +0.94 +0.22 +0.58 (+0.82) +0.13 (+0.19) +2.40 (+3.10) +0.55 (+0.71) +0.72 (0.29) +0.17 (0.07)
Black River S=0.29A +8.74 0.04 57.04 <0.01 0.88 +3.17 +0.92 +0.60 (+0.59) +0.17 (+0.17) +2.58 (+3.79) +0.75 (+1.10) 0.77 (1.15) 0.22 (0.33)
Boardman River S=0.14A +11.99 0.02 72.85 <0.01 0.90 +3.17 +0.44 0.56 (0.77) 0.08 (0.11) +2.49 (+3.77) +0.35 (+0.53) 0.87 (1.04) 0.12 (0.15)
Bryan Creek S=0.42A +7.69 0.08 28.86 <0.01 0.78 +2.33 +0.98 +1.01 (+1.48) +0.43 (+0.62) +2.83 (+3.11) +1.19 (+1.31) 0.62 (0.83) 0.26 (0.35)
Canada Creek S=0.60A +6.91 0.08 61.72 <0.01 0.88 +3.17 +1.90 0.79 (0.59) 0.47 (0.36) +2.58 (+3.79) +1.55 (+2.27) 0.77 (1.15) 0.46 (0.69)
Carp River S=0.28A +12.06 0.04 41.63 <0.01 0.84 +2.00 +0.56 +0.65 (+1.14) +0.18 (+0.32) +2.78 (+3.62) +0.78 (+1.01) 0.49 (1.29) 0.14 (0.36)
Cedar Creek S=0.56A +6.32 0.09 39.10 <0.01 0.83 +0.72 +0.40 +0.58 (+0.65) +0.33 (+0.37) +2.48 (+3.14) +1.39 (+1.76) +0.48 (0.49) +0.27 (0.28)
Cedar River (SLP) S=0.25A +11.42 0.03 97.82 <0.01 0.92 +2.61 +0.65 0.68 (0.23) 0.17 (0.06) +2.74 (+3.16) +0.69 (+0.79) 0.69 (0.65) 0.17 (0.16)
Chocolay River S=0.23A +10.29 0.03 77.99 <0.01 0.91 +2.72 +0.63 0.64 (+0.43) 0.15 (+0.10) +2.71 (+3.36) +0.62 (+0.77) 0.42 (1.31) 0.10 (0.30)
Davenport Creek S=0.13A +8.97 0.01 99.55 <0.01 0.92 +2.00 +0.26 +0.55 (+0.79) +0.07 (+0.10) +2.63 (+3.53) +0.34 (+0.46) 0.54 (1.26) 0.07 (0.16)
Duke Creek S=0.48A +4.45 0.08 37.27 <0.01 0.82 +0.72 +0.35 +0.65 (+0.74) +0.31 (+0.36) +2.48 (+3.14) +1.19 (+1.51) +0.48 (0.49) +0.23 (0.24)
East Branch Fox River S=0.33A +7.73 0.04 86.45 <0.01 0.91 +2.00 +0.66 +0.56 (+0.87) +0.19 (+0.29) +2.61 (+3.44) +0.86 (+1.14) 0.50 (1.27) 0.17 (0.42)
Elm River S=0.82A +2.11 0.06 193.20 <0.01 0.96 +2.72 +2.23 +0.99 (+1.79) +0.82 (+1.47) +3.19 (+2.65) +2.62 (+2.17) 0.75 (0.39) 0.62 (0.32)
Escanaba River S=0.88A +3.03 0.14 39.26 <0.01 0.83 +2.72 +2.40 +1.01 (+1.48) +0.89 (+1.30) +2.83 (+3.11) +2.49 (+2.73) 0.62 (0.83) 0.54 (0.73)
Iron River S=0.30A +12.76 0.04 52.58 <0.01 0.87 +1.89 +0.57 0.77 (0.03) 0.23 (0.01) +3.13 (+2.82) +0.94 (+0.85) 0.74 (0.35) 0.22 (0.11)
Little Indian River S=0.06A +14.86 0.01 83.65 <0.01 0.91 +2.72 +0.16 0.61 (+0.21) 0.04 (+0.01) +2.61 (+3.44) +0.16 (+0.21) 0.50 (1.27) 0.03 (0.08)
Little
Muskegon River
S=0.34A +12.09 0.04 69.13 <0.01 0.89 +2.56 +0.87 0.79 (0.80) 0.27 (0.27) +2.48 (+3.41) +0.84 (+1.16) 0.12 (0.72) 0.04 (0.24
Manistee River S=0.13A +10.67 0.02 57.70 <0.01 0.88 +3.17 +0.41 +0.76 (+0.49) +0.10 (+0.06) +2.47 (+3.56) +0.32 (+0.46) 0.54 (0.81) 0.07 (0.11)
Martin Creek S=0.38A +9.61 0.05 62.93 <0.01 0.89 +2.11 +0.80 0.82 (0.83) 0.31 (0.31) +2.48 (+3.32) +0.94 (+1.26) +0.05 (0.55) +0.02 (0.21)
Pere
Marquette River
S=0.18A +12.50 0.02 51.89 <0.01 0.86 +2.11 +0.38 0.82 (0.83) 0.15 (0.15) +2.48 (+3.32) +0.45 (+0.60) +0.05 (0.55) +0.01 (0.10)
Pigeon River S=0.11A +11.93 0.01 60.49 <0.01 0.88 +3.17 +0.35 +0.67 (+0.50) +0.07 (+0.06) +2.52 (+3.78) +0.28 (+0.42) 0.83 (1.11) 0.09 (0.12)
Pine River (NLP) S=0.22A +10.89 0.03 73.41 <0.01 0.90 +2.11 +0.46 +0.76 (+0.49) +0.17 (+0.11) +2.47 (+3.56) +0.54 (+0.78) 0.54 (0.81) 0.12 (0.18)
Pine River (SLP) S=0.40A +9.23 0.04 119.15 <0.01 0.94 +2.56 +1.02 +0.68 (+0.72) +0.27 (+0.29) +2.49 (+3.25) +1.00 (+1.30) +0.23 (0.69) +0.09 (0.27)
Prairie Creek S=0.74A +0.95 0.07 106.03 <0.01 0.93 +1.94 +1.44 +0.58 (+0.65) +0.43 (+0.48) +2.48 (+3.14) +1.83 (+2.32) +0.48 (0.49) +0.35 (0.37)
Rogue River S=0.23A +13.17 0.04 27.46 <0.01 0.77 +0.72 +0.17 +0.65 (+0.74) +0.15 (+0.17) +2.48 (+3.14) +0.57 (+0.72) +0.48 (0.49) +0.11 (0.11)
Salmon Trout River S=0.29A +8.23 0.05 33.60 <0.01 0.80 +2.33 +0.68 +0.99 (+1.79) +0.29 (+0.52) +3.19 (+2.65) +0.92 (+0.77) 0.75 (0.39) 0.22 (0.11)
Tahquamenon River S=0.50A +12.29 0.05 115.88 <0.01 0.93 +2.00 +1.00 +0.75 (+1.32) +0.38 (+0.66) +2.72 (+3.51) +1.36 (+1.75) 0.41 (1.36) 0.21 (0.68)
West Branch
Sturgeon River
S=0.06A +11.93 0.01 67.49 <0.01 0.89 +3.17 +0.19 +0.67 (+0.50) +0.04 (+0.03) +2.52 (+3.78) +0.15 (+0.23) 0.83 (1.11) 0.05 (0.07)
Yellow Dog River S=0.38A +9.90 0.06 45.58 <0.01 0.85 +2.33 +0.89 0.63 (+0.39) 0.24 (+0.15) +2.98 (+2.96) +1.13 (+1.13) 0.63 (0.81) 0.24 (0.31)
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Carlson et al.
regression models. Mean July stream temperatures
were projected because this month is typically the
warmest and most thermally stressful for salmonids
in Michigan (Zorn et al. 2011). To incorporate the
range of air temperatures projected by each CCM and
the intrinsic uncertainty and unique characteristics
(i.e., atmospheric pressure, sea ice rheology, forest
canopy density and soil layering) of each model, pre-
dictions were averaged across the three CCMs. Spe-
cies-specific thermal habitat suitability status was
assigned for each stream based on conditions for
growth and survival associated with projected July
temperatures. Status 1, 2, 3, and 4 corresponded with
optimal growth, suboptimal (i.e., reduced) growth, no
growth and extirpation, respectively (Table 3). Uni-
ted States Fish and Wildlife Service (USFWS) Bio-
logical Reports (e.g., Raleigh 1982a,b; Raleigh et al.
1986) contained thermal habitat status temperature
ranges for juveniles and adults of each species. Other
sources (i.e., Fry et al. 1946; Baldwin 1957; Wurts-
baugh & Davis 1977; Elliott & Hurley 2000; Hay
et al. 2006) contained temperature ranges for juve-
niles and/or adults and were used to confirm tempera-
tures reported in the USFWS reports. When threshold
temperatures (e.g., thermal minima, maxima) differed
between juveniles and adults, we reported juvenile
temperatures under the premise that resilient salmo-
nid fisheries can only be conserved if young fish
survive to adulthood.
Analyses
The accuracy of stream-specific regression models
was evaluated by comparing each streams projected
temperature and thermal habitat suitability status in
2006 to its actual (i.e., field measured) temperature
and associated habitat status obtained from the
MDNR database. The association between BFI and
air temperature regression coefficients, which are
indices of stream thermal sensitivity, was assessed
using simple linear regression (Kelleher et al. 2012).
Results
Model accuracy
Stream-specific models accurately projected tempera-
ture and thermal habitat suitability status in brook
charr, brown trout, and rainbow trout streams. Under
the A2 scenario, the mean deviation between pre-
dicted and actual temperatures was 0.46 °C
(SD =0.56; Table 4). Under the B1 scenario, the
mean deviation between predicted and actual temper-
atures was 0.58 °C (SD =0.59; Table 4). Under
the A2 and B1 scenarios, stream-specific models pre-
dicted thermal habitat status with 93.0 per cent over-
all accuracy in streams with brook charr (94.0 per
cent accuracy, N=15), brown trout (89.0 per cent
accuracy, N=16) and rainbow trout (100.0 per cent
accuracy, N=11; Table 4).
Thermal habitat suitability: brook charr
Stream-specific regression models projected that cli-
mate-induced air temperature elevation will have sub-
stantial effects on stream temperature and thermal
habitat suitability, with impacts varying by BFI (i.e.,
streams with lower BFI were more thermally sensitive;
Fig. 2) as well as species, region, time period and cli-
mate forcing scenario. From 1976 to 2006, the mean
temperature of brook charr streams increased by
0.74 °C as mean air temperature increased by 2.34 °C
(Table 2). In the UP under the A2 and B1 scenarios,
thermal habitat suitability was predicted to be optimal
in the East Branch Fox and Little Indian rivers and
suboptimal in the Iron River from 20062056 (Fig. 3a,
b). From 2006 to 2012, thermal habitat was projected
to be optimal in Bryan Creek and the Salmon Trout
River and suboptimal in the Carp and Escanaba rivers
under both scenarios. From 2012 to 2036, thermal
habitat was projected to become suboptimal in Bryan
Creek with predicted warming by 1.19 °C (A2 scenar-
io) and 1.31 °C (B1 scenario; Table 2). Similarly, ther-
mal habitat was forecasted to become unsuitable for
growth in the Escanaba River with projected warming
by 2.49 °C (A2 scenario) and 2.73 °C (B1 scenario;
Table 2). Habitat suitability was predicted to remain
the same from 2036 to 2056 under both scenarios in all
brook charr streams evaluated.
In the NLP from 2006 to 2056, thermal habitat
was projected to be optimal in most streams (Black,
Boardman, Manistee, Pine, West Branch Sturgeon
Table 3. Thermal habitat suitability status (habitat status) designations and
corresponding temperature ranges (temperature) and growth conditions
(growth) for juvenile and adult brook charr (BKC; Fry et al. 1946; Baldwin
1957; Raleigh 1982a), brown trout (BNT; Raleigh et al. 1986; Elliott &
Hurley 2000; Hay et al. 2006) and rainbow trout (RBT; Wurtsbaugh &
Davis 1977; Raleigh 1982b).
Species Habitat status Temperature Growth
BKC 1 11.0 °C<16.5 Optimal
2 16.5 °C<20.5 Suboptimal
3 20.5 °C<25.3 None
4°C25.3 Extirpation
BNT 1 12.0 °C<17.0 Optimal
2 17.0 °C<20.0 Suboptimal
3 20.0 °C<26.2 None
4°C26.2 Extirpation
RBT 1 12.0 °C<16.4 Optimal
2 16.4 °C<22.5 Suboptimal
3 22.5 °C<25.0 None
4°C25.0 Extirpation
195
Resilience-based salmonid management
rivers) under the A2 and B1 scenarios (Fig. 3c,d).
From 2012 to 2036, thermal habitat was forecasted to
become unsuitable for growth in Canada Creek under
the B1 scenario with predicted warming by 2.27 °C
(Fig. 3d; Table 2). In the SLP under the A2 and B1
scenarios, thermal habitat was predicted to be subop-
timal from 2006 to 2056 in Martin Creek and Cedar
Creek (Fig. 3e,f). From 2012 to 2036 under the B1
scenario, thermal habitat was forecasted to become
suboptimal in Duke Creek with predicted warming
by 1.51 °C (Fig. 3f; Table 2).
Thermal habitat suitability: brown trout
From 1976 to 2006, the mean temperature of brown
trout streams increased by 0.65 °C as mean air tem-
perature increased by 2.09 °C (Table 2). In the UP
under the A2 and B1 scenarios, thermal habitat suit-
ability was predicted to be optimal in the East Branch
Fox River from 2006 to 2056 (Fig. 4a,b). From 2006
to 2012, thermal habitat suitability was projected to
be optimal in the Bark River, suboptimal in the
Escanaba River and unsuitable in the Tahquamenon
Table 4. Actual versus projected stream temperatures and thermal habitat suitability (THS) status for brook charr (BKC), brown trout (BNT) and rainbow trout
(RBT) as predicted by stream-specific airwater temperature regression models. Temperatures are predicted in 2006 under the A2 (820 ppm atmospheric CO
2
by 2100) and B1 (550 ppm atmospheric CO
2
by 2100) scenarios. Dsymbols denote differences between projected and actual stream temperatures obtained
from the Northeast Climate Science Center. The first and second THS numbers represent statuses associated with actual and projected temperatures,
respectively.
Stream name Actual A2 ΔTHS B1 ΔTHS
Bark River 19.91 18.95 0.96 BNT: 2,2 18.88 1.03 BNT: 2,2
Bear Creek 15.75 16.54 0.79 BNT: 1,1 16.54 0.79 BNT: 1,1
Black River 15.05 14.46 0.59 BKC: 1,1 14.36 0.69 BKC: 1,1
Boardman River 15.02 14.88 0.14 BKC: 1,1 14.85 0.17 BKC: 1,1
BNT: 1,1 BNT: 1,1
Bryan Creek 16.66 15.27 1.39 BKC: 2,1 15.06 1.60 BKC: 2,1
Canada Creek 19.95 19.41 0.54 BKC: 2,2 19.09 0.86 BKC: 2,2
Carp River 17.61 17.05 0.56 BKC: 2,2 17.00 0.72 BKC: 2,2
BNT: 2,2 BNT: 2,2
RBT: 2,2 RBT: 2,2
Cedar Creek 18.30 18.27 0.04 BKC: 2,2 18.32 0.02 BKC: 2,2
BNT: 2,2 BNT: 2,2
Cedar River (SLP) 17.15 16.59 0.56 BNT: 2,1 16.46 0.69 BNT: 2,1
Chocolay River 15.23 14.77 0.46 RBT: 1,1 14.52 0.71 RBT: 1,1
Davenport Creek 11.50 11.39 0.11 RBT: 1,1 11.34 0.16 RBT: 1,1
Duke Creek 15.32 14.69 0.63 BKC: 1,1 14.73 0.59 BKC: 1,1
BNT: 1,1 BNT: 1,1
East Branch Fox River 14.23 13.76 0.47 BKC: 1,1 13.62 0.61 BKC: 1,1
BNT: 1,1 BNT: 1,1
Elm River 17.48 16.96 0.52 RBT: 2,2 16.40 1.10 RBT: 2,2
Escanaba River 19.95 18.88 1.07 BKC: 2,2 18.45 1.50 BKC: 2,2
BNT: 2,2 BNT: 2,2
Iron River 18.61 18.53 0.08 BKC: 2,2 18.34 0.27 BKC: 2,2
Little Indian River 16.16 16.09 0.07 BKC: 1,1 16.03 0.13 BKC: 1,1
Little Muskegon River 19.89 19.24 0.65 RBT: 2,2 19.22 0.67 RBT: 2,2
Manistee River 13.55 13.29 0.26 BKC: 1,1 13.29 0.26 BKC: 1,1
BNT: 1,1 BNT: 1,1
RBT: 1,1 RBT: 1,1
Martin Creek 18.33 17.86 0.47 BKC: 2,2 17.87 0.46 BKC: 2,2
Pere Marquette River 16.62 16.41 0.21 BNT: 2,2 16.41 0.21 BNT: 2,2
Pigeon River 14.45 14.12 0.33 BNT: 1,1 14.10 0.35 BNT: 1,1
RBT: 1,1 RBT: 1,1
Pine River (NLP) 15.97 15.26 0.71 BKC: 1,1 15.27 0.70 BKC: 1,1
BNT: 1,1 BNT: 1,1
RBT: 1,1 RBT: 1,1
Pine River (SLP) 18.42 17.55 0.87 BNT: 2,2 17.56 0.86 BNT: 2,2
Prairie Creek 18.35 16.57 1.78 BNT: 2,1 16.64 1.71 BNT: 2,1
Rogue River 18.36 17.95 0.41 BNT: 2,2 17.97 0.39 BNT: 2,2
RBT: 2,2 RBT: 2,2
Salmon Trout River 14.62 13.51 1.11 BKC: 1,1 13.31 1.31 BKC: 1,1
Tahquamenon River 20.02 21.16 1.14 BNT: 3,3 20.86 0.84 BNT: 3,3
West Branch Sturgeon River 13.30 13.12 0.18 BKC: 1,1 13.11 0.19 BKC: 1,1
BNT: 1,1 BNT: 1,1
RBT: 1,1 RBT: 1,1
Yellow Dog River 18.00 17.33 0.67 RBT: 2,2 16.96 1.04 RBT: 2,2
196
Carlson et al.
River under both scenarios (Fig. 4a,b). During the
same time period, thermal habitat was forecasted to
become suboptimal in the Carp River under the B1
scenario with predicted warming by 0.32 °C
(Table 2). From 2012 to 2036, thermal habitat was
projected to become unsuitable for growth in the
Escanaba River with predicted warming by 2.49 °C
(A2 scenario) and 2.73 °C (B1 scenario; Table 2).
From 2036 to 2056, habitat suitability was predicted
to remain the same under both scenarios in all brown
trout streams evaluated.
In the NLP from 2006 to 2056, thermal habitat was
forecasted to be optimal under the A2 and B1 scenar-
ios in all brown trout streams evaluated, including the
Boardman, Manistee, Pere Marquette, Pigeon, Pine
and West Branch Sturgeon rivers (Fig. 4c,d). In the
SLP under the A2 and B1 scenarios, thermal habitat
suitability was predicted to be optimal in Duke Creek
and suboptimal in the Rogue River from 2006 to 2056
(Fig. 4e,f). From 2006 to 2012, thermal habitat was
projected to be optimal in the Cedar River and subop-
timal in Cedar Creek and the Pine River under both
scenarios. During the same time period, thermal habi-
tat was predicted to become suboptimal in Prairie
Creek under the B1 scenario with projected warming
by 0.48 °C (Table 2). From 2012 to 2036, thermal
habitat was forecasted to become suboptimal in the
Cedar River and Bear Creek and unsuitable for growth
in Cedar Creek under both scenarios. From 2036
2056 under the A2 scenario, thermal habitat was pro-
jected to become optimal in the Cedar River and
unsuitable for growth in Cedar Creek with predicted
cooling and warming by 0.17 °C and 0.27 °C respec-
tively (Fig. 4e; Table 2). Thermal habitat suitability
was forecasted to remain the same from 2036 to 2056
under the B1 scenario in all brown trout streams eval-
uated (Fig. 4f).
Thermal habitat suitability: rainbow trout
From 1976 to 2006, the mean temperature of brown
trout streams increased by 0.64 °C as mean air tem-
perature increased by 2.42 °C (Table 2). In the UP
under the A2 and B1 scenarios, thermal habitat suit-
ability was predicted to be optimal in the Chocolay
River and Davenport Creek and suboptimal in the
Carp, Elm and Yellow Dog Rivers from 20062056
(Fig. 5a,b). Similarly, thermal habitat was projected
to be optimal (Manistee, Pigeon, Pine and West
Branch Sturgeon rivers) and suboptimal (Little
Muskegon River) in the NLP from 2006 to 2056
under the A2 and B1 scenarios (Fig. 5c,d). Thermal
habitat in the Rogue River, the only rainbow trout
stream in the SLP, was predicted to be suboptimal
under both emissions scenarios from 20062056
(Fig. 5e,f).
Discussion
Our study is the first broad-scale investigation of cli-
mate-driven stream temperature warming in Michigan
with implications for salmonid management. Water
temperatures were projected to increase by 0.19
5.94 °C in 30 coldwater salmonid streams over the
next 40 years due to predicted air temperature warm-
ing. This finding supports a previous study in the
State of Minnesota, USA (Pilgrim et al. 1998), in
which temperatures in 39 streams increased by 0.30
6.90 °C under a doubling of atmospheric CO
2
con-
centrations comparable to the A2 climate forcing sce-
nario. In addition, our study supports a previous
investigation in the State of Wisconsin, USA (Lyons
et al. 2010), in which thermal habitat degradation
was projected for brook charr and brown trout in 282
streams forecasted to warm by 0.804.00 °C over the
next 45 years. In our research, the magnitude of
warming and thermal habitat impairment varied
among streams and time periods, with the greatest
warming and habitat degradation occurring from
1976 to 2006 (2.72 °C) and projected from 2012 to
2036 (2.492.62 °C) in the Elm and Escanaba rivers,
systems with the lowest BFI (<0.45). In contrast, the
magnitude of warming from 1976 to 2006 (0.16
0.66 °C) and projected warming from 2012 to 2036
(0.160.86 °C) was considerably lower in the East
Branch Fox and Little Indian rivers, systems with the
highest BFI (0.73). This finding supported our
hypothesis that temperature and thermal habitat alter-
ations would vary spatially and temporally due to
system-specific patterns of warming. It also rein-
forces previous research indicating fish assemblages
in Michigan streams with stable discharge from
groundwater inputs are more resilient to thermal
warming than those in streams with less stable dis-
Fig. 2. Relative influence of baseflow index on thermal sensitiv-
ity (i.e., air temperature regression coefficients) of streams in the
State of Michigan, USA. The regression equation is: Thermal sen-
sitivity =1.78 9Baseflow index +1.38 (P<0.01, R
2
=0.44).
197
Resilience-based salmonid management
charge (Zorn et al. 2012). Thermal habitat degrada-
tion was projected to occur least frequently in rain-
bow trout streams as this species has a wider
temperature range (12.022.5 °C) for growth and sur-
vival compared to brook charr (11.020.5 °C) and
brown trout (12.020.0 °C) (Wurtsbaugh & Davis
1977).
Warmer air temperatures are projected to increase
stream temperatures both directly (i.e., advection) and
indirectly through effects on precipitation, evapora-
Fig. 3. Projected temperatures in individual Michigan brook charr streams in 2006, 2012, 2036 and 2056 under A2 (820 ppm atmospheric
CO
2
by 2100) and B1 (550 ppm by 2100) scenarios. Plots are organised by region and emissions scenario: (a) Upper Peninsula, A2 scenar-
io; (b) Upper Peninsula, B1 scenario; (c) northern Lower Peninsula, A2 scenario; (d) northern Lower Peninsula, B1 scenario; (e) southern
Lower Peninsula, A2 scenario; (f) southern Lower Peninsula, B1 scenario. Dotted lines represent transitions between thermal habitat suit-
ability statuses. Stream abbreviations are as follows: BRC =Bryan Creek, BLR =Black River, BOR =Boardman River, CAC =Canada
Creek, CAR =Carp River, CEC =Cedar Creek, DUC =Duke Creek, EFR =East Branch Fox River, ESR =Escanaba River, IRR =Iron
River, LIR =Little Indian River, MAR =Manistee River, MAC =Martin Creek, PIR =Pine River (northern Lower Peninsula),
STR =Salmon Trout River, WSR =West Branch Sturgeon River.
198
Carlson et al.
tion and transpiration. Predicted effects of climate
change include more variable (and overall lower) pre-
cipitation (Parry et al. 2007; Stoner et al. 2013),
increased evaporation (Compagnucci et al. 2001) and
decreased discharge. These impacts would increase
stream temperature by reducing the volume of water
exposed to solar radiation. However, in Michigan
streams near the Great Lakes, precipitation will likely
Fig. 4. Projected temperatures in individual Michigan brown trout streams in 2006, 2012, 2036 and 2056 under A2 (820 ppm atmospheric
CO
2
by 2100) and B1 (550 ppm by 2100) scenarios. Plots are organised by region and emissions scenario: (a) Upper Peninsula, A2 scenar-
io; (b) Upper Peninsula, B1 scenario; (c) northern Lower Peninsula, A2 scenario; (d) northern Lower Peninsula, B1 scenario; (e) southern
Lower Peninsula, A2 scenario; (f) southern Lower Peninsula, B1 scenario. Dotted lines represent transitions between thermal habitat suit-
ability statuses. Stream abbreviations are as follows: BEC =Bear Creek, BOR =Boardman River, BRC =Bryan Creek, CAR =Carp
River, CEC =Cedar Creek, CER =Cedar River, DUC =Duke Creek, EFR =East Branch Fox River, ESR =Escanaba River,
MAR =Manistee River, PMR =Pere Marquette River, PGR =Pigeon River, PNR =Pine River (southern Lower Peninsula), PIR =Pine
River (northern Lower Peninsula), PRC =Prairie Creek, ROR =Rogue River, TAR =Tahquamenon River, WSR =West Branch Stur-
geon River.
199
Resilience-based salmonid management
intensify in fall and winter due to predicted increases
in snowfall (Norton & Bolsenga 1993; Primack
2000), increasing surface runoff and water volume
and decreasing temperature compared to systems
without this source of precipitation (Kurylyk et al.
2013). Increased canopy and soil surface evaporation
is projected to decrease groundwater recharge and
increase water temperature (Ge et al. 2013), although
streams high BFI are expected to warm at a slower
rate than those with low BFI due to groundwater-dri-
ven thermal buffering (Menberg et al. 2014). Tran-
spiration of water from plants has been reported to
contribute to stream warming by reducing streamflow
and groundwater recharge (Federer & Lash 1978;
Bond et al. 2002). Effects of climate warming on
transpiration and stream temperature will depend lar-
Fig. 5. Projected temperatures in individual Michigan rainbow trout streams in 2006, 2012, 2036 and 2056 under A2 (820 ppm atmo-
spheric CO
2
by 2100) and B1 (550 ppm by 2100) scenarios. Plots are organised by region and emissions scenario: (a) Upper Peninsula,
A2 scenario; (b) Upper Peninsula, B1 scenario; (c) northern Lower Peninsula, A2 scenario; (d) northern Lower Peninsula, B1 scenario; (e)
southern Lower Peninsula, A2 scenario; (f) southern Lower Peninsula, B1 scenario. Dotted lines represent transitions between thermal habi-
tat suitability statuses. Stream abbreviations are as follows: CAR =Carp River, CHR =Chocolay River, DAC =Davenport Creek,
ELR =Elm River, LMR =Little Muskegon River, MAR =Manistee River, PGR =Pigeon River, PIR =Pine River, ROR =Rogue
River, WSR =West Branch Sturgeon River, YDR =Yellow Dog River.
200
Carlson et al.
gely on changes in the species composition and leaf
characteristics (e.g., stomatal conductance, water-use
efficiency) of riparian plants and trees (Compagnucci
et al. 2001; Kirschbaum 2004). We encourage
researchers to incorporate the thermal effects of tran-
spiration, riparian shading and watershed land cover
(Wehrly et al. 1997; Wiley et al. 2010) into future
stream temperature modelling to improve predictive
accuracy.
Projected increases in Michigan stream tempera-
tures will result in thermal habitat conditions less
conducive for salmonid growth and survival during
the warmest period of the year (Raleigh 1982a,b;
Raleigh et al. 1986), particularly in thermally sensi-
tive streams with low BFI values. This will likely be
manifested by declines in availability and changes in
spatial arrangement of optimal thermal habitat, with
potential effects on salmonid distribution and sur-
vival. For instance, if stream warming restricts cold-
water habitats to specific areas (e.g., headwater
reaches), salmonid populations will become isolated,
which could increase interspecific competition and
decrease survival (Tsuboi et al. 2013; Dugdale et al.
2015). Previous research in Michigans Muskegon
River suggested warmer water temperatures resulting
from climate change will restrict the distribution and
decrease the survival of brook charr, brown trout,
and rainbow trout (Steen et al. 2010). Researchers in
the State of Wisconsin, USA, predicted that by 2060
the total length of streams suitable for brown trout
would decline by 8, 33, and 88 per cent under limited
(summer air temperatures increase 1.0 °C and water
0.8 °C), moderate (air 3.0 °C and water 2.4 °C), and
major (air 5.0 °C and water 4.0 °C) climatic warming
(Lyons et al. 2010). Results were more extreme for
brook charr, with distributional declines of 44 and 94
per cent under limited and moderate warming, and
complete extirpation under major warming. Although
our models did not project extirpation of any species
in the Michigan streams evaluated, summer growth
limitation may be coupled with decreased reproduc-
tion as lower oxygen levels resulting from stream
temperature warming and/or groundwater withdrawal
may reduce egg survival or prevent spawning
(Raleigh 1982a,b; Raleigh et al. 1986). Although
annual salmonid growth may remain stable or
increase due to longer growing season length and
increased prey availability, projected reductions in
growth and survival during the warmest period of the
year have important management implications.
Management implications
We encourage scientists, biologists, policymakers and
public stakeholders to collaboratively develop resili-
ence-based management programmes to conserve sal-
monid populations amidst global change. Our
research represents a step in this direction as it pro-
jects consequences of stream thermal warming for
salmonid growth and survival in Michigan. Our
results indicate it is important for fisheries profession-
als to manage streams for thermal resilience by form-
ing publicprivate partnerships to protect watershed
land cover types that facilitate high groundwater
recharge (e.g., grasslands; Waco & Taylor 2010; Sii-
tari et al. 2011), preserve riparian vegetation and
associated shading (Blann et al. 2002), and maintain
longitudinal connectivity to promote salmonid move-
ment to cold headwater reaches (Drake & Taylor
1996; Hayes et al. 1998). We recommend these
strategies play prominent roles in resilience-based
management programmes for coldwater streams and
their important salmonid fisheries. As documented in
our study, streams with low thermal sensitivity (i.e.,
high BFI) will likely maintain thermal conditions
(e.g., cool summer temperatures, seasonal flow stabil-
ity; Wiley et al. 1997; Baker et al. 2003) that are
more conducive for salmonid growth and survival
than systems with high thermal sensitivity in the next
40 years. Thus, we encourage managers to allocate
resources (i.e., time, money and personnel) to priori-
tise protection of streams with low thermal sensitivity
and inform public stakeholders about realistic expec-
tations for stream fish communities amidst global
change (e.g., salmonid decline, centrarchid expan-
sion; Pease & Paukert 2014). Moreover, because
streams with high BFI were less susceptible to tem-
perature change, we suggest that managers use spatial
BFI maps as tools for understanding stream thermal
sensitivity across large geographical areas. Moreover,
we encourage managers to increase the spatial and
temporal coverage of air and stream temperature
monitoring networks and thereby expand their utility
for salmonid management.
Fisheries professionals and public stakeholders can
also collaboratively implement additional strategies
for resilience-based salmonid management. They can
promote thermally resilient salmonid populations by
removing dams and installing fish ladders at roadside
crossings and culverts to restore stream habitat con-
nectivity, particularly in cold headwater reaches,
which function as thermal refugia during warm sum-
mer months. Fisheries professionals can also foster
salmonid population resilience by implementing regu-
lations (e.g., protected slot limits, reduced creel lim-
its) that reduce harvest and increase survival during
thermally stressful periods. In addition, we recom-
mend fisheries professionals protect a diversity of sal-
monid size classes, genetic stocks and prey species
that tolerate a wide range of temperatures predicted
from climate change models (Hansen et al. 2015).
We encourage fisheries managers and policymakers
201
Resilience-based salmonid management
to provide incentives (e.g., financial assistance,
open space tax deduction, fast-track permitting;
Knight 2009) for land developers and property
owners to protect coldwater habitat and thermal
buffering mechanisms on their lands. Moreover,
because it is infeasible to protect salmonid popula-
tions and thermal habitat in all streams, it will be
necessary for fisheries professionals to implement a
triage approach with specific criteria for stream pro-
tection (e.g., species composition, habitat quality
and recreational importance). In summary, resili-
ence-based salmonid management programmes will
require effective collaboration among scientists,
biologists, policymakers and public stakeholders.
Our research promotes resilience-based management
by providing a methodology to project stream tem-
perature and thermal habitat suitability. Fisheries
professionals can use this approach to protect cold-
water habitats and drivers of stream cooling and
ultimately conserve resilient salmonid populations
amidst global change.
Acknowledgements
We thank Jennifer Moore Myers (USFS EFETAC) for assis-
tance with air temperature projection models and data acqui-
sition. We thank Kyle Herreman and Wesley Daniel
(Michigan State University), Tracy Kolb and Todd Wills
(Michigan Department of Natural Resources) and Henry
Quinlan (United States Fish and Wildlife Service) for provid-
ing initial environmental and trout population data critical for
development of this study. We acknowledge the Programme
for Climate Model Diagnosis and Intercomparison (PCMDI)
and the WCRPs Working Group on Coupled Modelling for
assisting in procurement of the WCRP CMIP3 multimodel
data set.
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... Stream temperature affects fishes directly by influencing biological processes including metabolism, feeding, and growth (Elliot 1981), as well as indirectly by influencing prey availability and competitive interactions (Taniguchi et al. 1998). Changes in stream temperature regimes can alter fish abundance and distribution (Jackson and Mandrak 2002;Lyons et al. 2010;Stewart et al. 2015Stewart et al. , 2016a, sometimes resulting in declines of stream salmonids near the upper end of their thermal tolerances Carlson et al. 2017aCarlson et al. , 2017b. Therefore, forecasting stream temperature alterations is imperative for predicting how stream salmonid populations may respond to climate change and prioritizing management actions from local to regional scales (Snyder et al. 2015;Carlson et al. 2019Carlson et al. , 2020. ...
... After daily stream temperatures were forecasted and correction factors applied for each time step within each AOGCM, another R script was used to calculate summary statistics characterizing the predicted temperature regime of each stream reach over time (e.g., temperature magnitude, variability, and rate of change, Tsang et al. 2016). Here, July mean stream temperature is reported because it is an important driver of fish population dynamics in general, and stream salmonid reproduction, growth, and survival in particular (Lyons et al. 2009;Carlson et al. 2017aCarlson et al. , 2017bCarlson et al. , 2019Carlson et al. , 2020. July mean stream temperature was used to categorize streams into thermal classes (i.e., coldwater, cold-transitional, warm-transitional, and warmwater) that were developed based on Michigan and Wisconsin fish assemblages (Lyons et al. 2009) and were relevant for fisheries management and environmental regulation. ...
... For instance, coldwater streams that become warm-transitional or warmwater streams will likely become less thermally suitable for brook charr Salvelinus fontinalis, although thermal refugia arising from groundwater inputs may provide locally suitable conditions for growth and survival (Wilbur et al. 2020;O'Sullivan et al. 2021). Even if coldwater streams become cold-transitional and still support coldwater fishes such as brook charr, brown trout Salmo trutta, and rainbow trout Oncorhynchus mykiss, growing conditions may become suboptimal, contributing to changes in fish community composition (Carlson et al. 2017a(Carlson et al. , 2017b(Carlson et al. , 2020. Clearly, populations of stream salmonids and other coldwater fishes will decline if climate change causes water temperatures to exceed species-specific thresholds for survival or reproduction and thermal refugia become limited. ...
Chapter
Despite abundant research exploring how climate change may affect population demographics of stream fishes (e.g., survival, growth, and relative abundance), tools are needed to provide accurate, fine-scale stream temperature predictions over large spatial extents encompassing multiple fisheries management agencies and jurisdictions. As such, a fine-scale modeling approach based on artificial neural networks was developed to forecast reach-specific thermal conditions under different climate-change scenarios across the Upper Midwest, USA (i.e., Michigan, Minnesota, and Wisconsin). Specifically, models collated landscape and climatic data to predict individual stream temperatures across watersheds and states with relatively high accuracy (r2 = 0.71–0.76). Collectively, models predicted widespread increases in stream temperature across the study region averaging 1.5 °C through 2087, with substantial warming in Wisconsin and Minnesota but moderate warming in Michigan streams. For instance, the extent of warm-transitional (July mean water temperature 19.5–21.0 °C) and warmwater (>21.0 °C) habitats in Wisconsin—currently 40% of stream kilometers—was projected to reach up to 63% in 2042 and 71% in 2087, with concomitant reductions in growth and survival of stream salmonids (e.g., brook charr Salvelinus fontinalis, brown trout Salmo trutta, and rainbow trout Oncorhynchus mykiss). Overall, our modeling approach provides managers with a stream-specific yet spatiotemporally broad tool for predicting the magnitude and distribution of stream warming amid climate change that can be used to prioritize efforts for stream salmonid management and conservation. Such information is critical for developing state and regional climate adaptation strategies for Upper Midwest streams and coordinating thermal habitat management activities (e.g., groundwater conservation and riparian/watershed rehabilitation) among fisheries and aquatic resource agencies and across jurisdictions.
... The current and future impacts of these climate changes are severe. They include increased surface water body temperature resulting in decreased fish survival (Carlson et al., 2017), increased white-tailed deer populations leading to disease outbreak (Weiskopf et al., 2019), wetland vulnerability due to volatile water levels and flooding (Kling et al., 2003), algal blooms and toxic dead zones (Kling et al., 2003) (Michalak et al., 2013), drier overall land surfaces due to increased temperatures and evaporation rate (Pryor, 2009) and natural barriers to dispersal, which could prevent heat-sensitive mobile species from migrating northward (Lee et al., 2011). More specifically, climate change vulnerability work done by the Michigan Natural Features Inventory and the Michigan Department of Natural Resources indicates that 17% of terrestrial game species and 61% of the Species of Greatest Conservation Need (both terrestrial and aquatic) are vulnerable to climate change and will therefore experience range or population reductions as warming continues (Hoving et al., 2013). ...
... Thus, resilience in aquatic systems relies on water quality levels remaining high enough for sustained habitat diversity both within and between sites, and for a diverse assemblage of aquatic species. Of particular concern for our model is the potential for warming temperatures to devastate coldwater fisheries (Carlson et al., 2017). Many of the denizens of these streams, including the imperiled unionid freshwater mussels, are unable to tolerate decreased water quality, often borne out through increased turbidity and pollution, increased water temperatures, and subsequent lower dissolved oxygen levels (Daniel et al., 2018). ...
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... Among the most sensitive species of mountain rivers to ecological changes is the trout (family Salmonidae), which is highly represented in Slovakian streams, and often used as a bioindicator [23][24][25][26] . Based on ichthyological measurements, the habitat preference of both Brown trout and Minnow was evaluated. ...
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The article focuses on adapting generalized depth preference and flow velocity characteristics from Brown trout (Salmo trutta morpha fario) to the Minnow (Phoxinus phoxinus). The results obtained were used to model habitat suitability with 1D and 2D models. Since 1995, research on assessing aquatic habitat quality has been ongoing on 77 mountain streams in Slovakia. This assessment employs a System of Environmental Flow Analysis (SEFA), which is based on the Instream Flow Incremental Methodology (IFIM). In most sections, Brown trout occurred in representative numbers. Brown trout habitat preferences were derived and generalized, and represented by suitability curves. Minnows were present in sufficient numbers for the derivation of suitability curves only on some streams, making direct generalization from the measured data unrepresentative. The measurement results showed that Minnow has similar depth and flow rate preferences to Brown trout. Therefore, it can be assumed that it is possible to adapt the generalised suitability curves from Brown trout to Minnow. This expansion enabled us to broaden the assessment of habitat quality using the SEFA model to mountain streams that are dominated by Minnows and where there is insufficient Brown trout presence. Verification of parameter adaptation from Brown trout to Minnow was conducted in 11 sections of mountain streams. We discuss the performance of habitat quality modeling, concerning the fundamental hydraulic characteristics of streams, using both 1D and 2D models. The results of the 2D modeling are presented for a sub-mountain stream.
... Land-use changes (Stranko et al. 2008), nonnative species (Kirk et al. 2018;Zorn et al. 2020), acidic deposition (Baker et al. 1996), and other factors have led to widespread declines in the distribution of Brook Trout Salvelinus fontinalis across the southern and central Appalachian mountains in North America (Hudy et al. 2008). Ongoing land-use conversion of forested habitats continues to threaten Brook Trout populations, and many studies have predicted further declines as anthropogenic climate change progresses (Flebbe et al. 2006;Snyder et al. 2015;Bassar et al. 2016;Carlson et al. 2017). Although protected areas largely prevent land-use conversion with their boundaries, climate change, acidic deposition, and other processes that operate at regional or global scales may pervade protected area boundaries (Hansen and DeFries 2007). ...
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Objective Landscape context structures fish abundance and dynamics, and understanding trends in fish abundance across the landscape is often prerequisite for effective conservation. In this study, we evaluated the status and trends of Brook Trout Salvelinus fontinalis in Shenandoah National Park to understand how these are structured across bedrock geology, elevation, and stream size. Methods We used long‐term monitoring data from 94 sites in Shenandoah National Park to evaluate trends in Brook Trout abundance over a 27‐year period (1996–2022) and assess the importance of local environmental covariates using a hierarchical Bayesian N‐mixture model based on depletion sampling. Focal covariates were chosen for their demonstrated importance in structuring fish populations in Shenandoah National Park and elsewhere. Bedrock geology controls sensitivity to acid deposition, watershed area is related to stream habitat features such as complexity and flow variability, and elevation creates gradients in temperature. Result Models revealed significant decreases in adult Brook Trout abundance over time (95% credible intervals < 0) for 31 of 94 sites (33%), and at least three sites exhibited apparent extirpations over the study period. Estimated Brook Trout abundance declined by 50% or more in approximately 70% of streams across the park over the study period. Sites with the warmest water temperatures exhibited the fastest declines in abundance. However, large watersheds on poorly buffered bedrock exhibited significant gains in abundance over time, suggesting some recovery from acid deposition due to improvements in air quality. Conclusion Our analysis revealed large and divergent changes in Brook Trout abundance over recent decades and suggests the importance of local water temperature and acid sensitivity as probable causal mechanisms. These results highlight the importance of considering local factors when evaluating long‐term trends in stream fish populations. Results of this study can assist the development of targeted conservation actions within Shenandoah National Park and elsewhere.
... Climate change represents a significant threat to Pacific salmon (Oncorhynchus spp.) due to rising water temperatures and changes in river flows (Carlson et al., 2017;Kundzewicz et al., 2008). For example, dramatic decreases in commercial, recreational, and Indigenous subsistence catches over the past 15 years have been attributed to the direct and indirect effects of climate change on salmon survival (Grant et al., 2021). ...
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Understanding the adaptive potential of populations and species is pivotal for minimizing the loss of biodiversity in this era of rapid climate change. Adaptive potential has been estimated in various ways, including based on levels of standing genetic variation, presence of potentially beneficial alleles, and/or the severity of environmental change. Kokanee salmon, the non‐migratory ecotype of sockeye salmon ( Oncorhynchus nerka ), is culturally and economically important and has already been impacted by the effects of climate change. To assess its climate vulnerability moving forward, we integrated analyses of standing genetic variation, genotype‐environment associations, and climate modeling based on sequence and structural genomic variation from 224 whole genomes sampled from 22 lakes in British Columbia and Yukon (Canada). We found that variables for extreme temperatures, particularly warmer temperatures, had the most pervasive signature of selection in the genome and were the strongest predictors of levels of standing variation and of putatively adaptive genomic variation, both sequence and structural. Genomic offset estimates, a measure of climate vulnerability, were significantly correlated with higher increases in extreme warm temperatures, further highlighting the risk of summer heat waves that are predicted to increase in frequency in the future. Levels of standing genetic variation, an important metric for population viability and resilience, were not correlated with genomic offset. Nonetheless, our combined approach highlights the importance of integrating different sources of information and genomic data to formulate more comprehensive and accurate predictions on the vulnerability of populations and species to future climate change.
... Historical declines in the abundance of brook trout in the Lake Superior basin are primarily attributed to overharvest due to a lack of adequate fishery regulations from the late 1800s through mid-1900s, combined with heavy landscape modification associated with early logging and mining activities (Huckins et al., 2008). In comparison, contemporary declines are likely driven by habitat fragmentation resulting from man-made barriers such as dams and culverts (Newman et al., 2003; but see Zorn et al., 2020), competition with introduced non-native salmonids (Miller et al., 2019;Zorn et al., 2020), and anthropogenic climate change causing habitat loss due to large flood events and warmer water temperatures (Carlson et al., 2017). ...
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... Freshwater environments are expected to experience alterations in the flow and thermal regimes of streams and rivers owing to climate change [1,2]. Given that freshwater ecosystems are dependent on the flow and thermal regimes [3][4][5], changes in these are expected to affect the growth, migration, spawning, survival, and distribution of freshwater fish [6][7][8]. Specifically, environmental variables such as water depth, velocity, and temperature are important to freshwater fish habitats [9,10]. Therefore, the effects of climate change on the habitat of freshwater fish should be evaluated for future risk assessment. ...
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Climate change, which has the potential to alter water flow and temperature in aquatic environments, can influence the freshwater fish habitat. This study used an ecological habitat suitability model (EHSM), which integrates hydraulic (water depth and velocity) and physiologic (water temperature) suitability, to investigate the impact of climate change on two native freshwater fish species (Zacco platypus and Nipponocypris koreanus) in South Korea. The model predicted that in 2080 (2076–2085), the decrease in average ecological habitat suitability (EHS) will be higher for N. koreanus (19.2%) than for Z. platypus (9.87%) under the representative concentration pathway (RCP) 8.5 scenario. Under the same condition, EHS for Z. platypus and N. koreanus at 36.5% and 44.4% of 115 sites, respectively, were expected to degrade significantly (p < 0.05). However, the habitat degradation for Z. platypus and N. koreanus was much lower (7.8% and 10.4%, respectively) under the RCP 4.5 scenario, suggesting a preventive measure for carbon dioxide emission. Partial correlation analysis indicated that the number of hot days (i.e., days on which the temperature exceeds the heat stress threshold) is the variable most significantly (p < 0.05) related to EHS changes for both species. This study suggests that the EHSM incorporating the effect of water temperature on the growth and heat stress of fish can be a promising model for the assessment of climate change impacts on habitat suitability for freshwater fish.
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As climate change alters the thermal environment of the planet, interest has grown in how animals may mitigate the impact of a changing environment on physiological function. Thermal acclimation to a warm environment may, for instance, blunt the impact of a warming environment on metabolism by allowing a fish to shift to slower isoforms of functionally significant proteins such as myosin heavy chain. The thermal acclimation of brook trout (Salvelinus fontinalis) was examined by comparing swimming performance, myotomal muscle contraction kinetics and muscle histology in groups of fish acclimated to 4, 10 and 20 °C. Brook trout show a significant acclimation response in their maximum aerobic swimming performance (Ucrit), with acclimation to warm water leading to lower Ucrit values. Maximum muscle shortening velocity (Vmax) decreased significantly with warm acclimation for both red or slow-twitch and white or fast-twitch muscle. Immunohistochemical analysis of myotomal muscle suggests changes in myosin expression underly the thermal acclimation of swimming performance and contraction kinetics. Physiological and histological data suggest a robust acclimation response to a warming environment, one that would reduce the added metabolic costs incurred by an ectotherm when environmental temperature rises for sustained periods of time.
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Freshwaters are being transformed by multiple environmental drivers, creating uncertainty about future conditions. One way of coping with uncertainty is to manage for resilience to unanticipated events while facilitating learning through adaptive management. We outline the application of these strategies to freshwater recreational fisheries management using a case study in Wisconsin, USA, where black bass (Micropterus spp.) populations are increasing, while Walleye (Sander vitreus) populations are decreasing. Managing for heterogeneity in functional groups (e.g., age classes and prey species of sport fishes), fishery objectives, and regulations can increase resilience, although heterogeneity must be balanced with replication to facilitate learning. Monitoring designed to evaluate management objectives and inform about critical uncertainties, when combined with heterogeneity, creates opportunities for adaptive management, another critical resilience strategy. Although barriers exist to implementing resilience strategies, management designed to accommodate uncertainty and illuminate its consequences is needed to maintain critical fisheries in a rapidly changing world.
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