Comparison of triploid and diploid rainbow trout (Oncorhynchus mykiss) fine-scale movement, migration and catchability in lowland lakes of western Washington

Abstract

Fisheries managers stock triploid (i.e., infertile, artificially produced) rainbow trout Oncorhynchus mykiss in North American lakes to support sport fisheries while minimizing the risk of genetic introgression between hatchery and wild trout. In Washington State, the Washington Department of Fish and Wildlife (WDFW) allocates approximately US $3 million annually to stock hatchery-origin rainbow trout in > 600 lakes, yet only about 10% of them are triploids. Many lakes in Washington State drain into waters that support wild anadromous steelhead O. mykiss that are listed as threatened under the U.S. Endangered Species Act. As a result, there is a strong interest in understanding the costs and benefits associated with stocking sterile, triploid rainbow trout as an alternative to traditional diploids. The objectives of this study were to compare triploid and diploid rainbow trout in terms of: (1) contribution to the sport fishery catch, (2) fine-scale movements within the study lakes, (3) rate of emigration from the lake, and (4) natural mortality. Our results demonstrated that triploid and diploid trout had similar day-night distribution patterns, but triploid trout exhibited a lower emigration rate from the lake and lower catch rates in some lakes. Overall, triploid rainbow trout represent a viable alternative to stocking of diploids, especially in lakes draining to rivers, because they are sterile, have comparable home ranges, and less often migrate.

Full text

Peaseetal. Movement Ecology (2023) 11:57
https://doi.org/10.1186/s40462-023-00418-w
RESEARCH
Comparison oftriploid anddiploid
rainbow trout (Oncorhynchus mykiss) ne-scale
movement, migration andcatchability
inlowland lakes ofwestern Washington
Jessica E. Pease1, James P. Losee1,2*, Stephen Caromile1, Gabriel Madel1, Michael Lucero1, Anna Kagley3,
Michael G. Bertram2,5,6, Jake M. Martin2, Thomas P. Quinn4, Daniel Palm2 and Gustav Hellström2
Abstract
Fisheries managers stock triploid (i.e., infertile, artificially produced) rainbow trout Oncorhynchus mykiss in North
American lakes to support sport fisheries while minimizing the risk of genetic introgression between hatchery
and wild trout. In Washington State, the Washington Department of Fish and Wildlife (WDFW) allocates approximately
US $3 million annually to stock hatchery-origin rainbow trout in > 600 lakes, yet only about 10% of them are triploids.
Many lakes in Washington State drain into waters that support wild anadromous steelhead O. mykiss that are listed
as threatened under the U.S. Endangered Species Act. As a result, there is a strong interest in understanding the costs
and benefits associated with stocking sterile, triploid rainbow trout as an alternative to traditional diploids. The objec-
tives of this study were to compare triploid and diploid rainbow trout in terms of: (1) contribution to the sport fishery
catch, (2) fine-scale movements within the study lakes, (3) rate of emigration from the lake, and (4) natural mortality.
Our results demonstrated that triploid and diploid trout had similar day-night distribution patterns, but triploid trout
exhibited a lower emigration rate from the lake and lower catch rates in some lakes. Overall, triploid rainbow trout
represent a viable alternative to stocking of diploids, especially in lakes draining to rivers, because they are sterile, have
comparable home ranges, and less often migrate.
Keywords Introgression, Creel, Angler catch rates, Telemetry, Stocking
Introduction
Fisheries managers have stocked rainbow trout Onco-
rhynchus mykiss in rivers and lakes to support conserva-
tion and recreational objectives for over a century [34].
e native range of rainbow trout is restricted to west-
ern North America and eastern Russia, but rainbow
trout currently inhabit much of the world and persist as
self-sustaining populations outside the native range as a
result of these stocking programs [8, 31]. However, there
is also extensive stocking within their native range. For
example, over 2 million rainbow trout are stocked annu-
ally in Washington State, USA [41].
Open Access
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Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation,
distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source,
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zero/1.0/) applies to the data made available in this article, unless otherwise stated in a credit line to the data.
Movement Ecology
*Correspondence:
James P. Losee
james.losee@dfw.wa.gov
1 Washington Department of Fish and Wildlife, OlympiaWashington, WA,
USA
2 Department of Wildlife, Fish and Environmental Studies, Swedish
University of Agricultural Sciences, Umeå, Sweden
3 NOAA Fisheries, Northwest Fisheries Science Center, Seattle, WA, USA
4 School of Aquatic and Fishery Sciences, University of Washington,
Seattle, WA, USA
5 Department of Zoology, Stockholm University, Stockholm, Sweden
6 School of Biological Sciences, Monash University, Melbourne, Australia
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Peaseetal. Movement Ecology (2023) 11:57
Rainbow trout stocking has been linked to important
conservation gains [1, 10], and significant economic ben-
efits [17]. For example, in Washington State, rainbow
trout stocking is responsible for over US$1.1 billion of
revenue [11]. However, in many parts of the world, there
has been growing concern that stocked rainbow trout
pose potential risks to natural ecosystems through com-
petition, predation, and spawning with native species
[8, 23, 26]. In the United States, introgression between
stocked rainbow trout and with natively threatened ana-
dromous rainbow trout (steelhead) and coastal cutthroat
trout, O. clarkii clarkii, is a major issue for maintaining
genetic integrity and overall fitness [14, 30, 38, 43]. How-
ever, given funding limitations and public satisfaction
with rainbow trout stocking programs, formal evaluation
of the costs and benefits of these popular programs are
lacking [4, 39].
One strategy that managers use to reduce hybridiza-
tion between native and hatchery-origin fish is to stock
sterile, triploid rainbow trout rather than traditional dip-
loids, particularly in lakes draining into waters accessible
to anadromous conspecifics (i.e., wild steelhead listed
as reatened under the Endangered Species Act in the
Puget Sound region of Washington, and elsewhere). For
instance, the state of Idaho adopted a policy in 2001
stocking only sterile, not diploid, rainbow trout in flow-
ing waters. In Washington State, where most steelhead
populations are listed as reatened, the WDFW allo-
cates approximately US$3 million annually to stock
hatchery-origin rainbow trout in > 600 lakes but less than
10% of the fish stocked are triploids [11]. Increasing the
use of triploid trout in popular trout fisheries may help
conserve the genetic integrity of native populations but
the effect on catch rates is unclear. For instance, Dillon
etal., [9] found no significant differences in catch rate
and fishery duration between the two trout ploidy strains
in Idaho streams. On the other hand, Koenig etal. [21]
and Koenig and Meyer [22] documented differences in
survival across habitat conditions and higher catch rates
of diploid than triploid trout in lake systems. Differ-
ences between triploid and diploid rainbow trout catch-
ability are poorly understood and difficult to assess but
could include different rates of survival, migration from
the lake, and feeding, and in-lake movement patterns. To
ensure conservation objectives while maintaining suc-
cessful fisheries when switching from diploid to triploid
rainbow trout, post-stocking mortality, migration rate,
and recruitment to the fishery of triploids and diploids
need to be compared.
Many tools have been developed to assess individual
fish movements, growth, and survival, including a vari-
ety of tags, transmitters, and marking techniques [5,
27]. Acoustic telemetry has accelerated research on fish
behavior as it can reveal patterns of fish behavior, habitat
use, predation, and migration [5, 6, 19, 24]. For example,
acoustic telemetry has revealed precise survival rates of
stocked rainbow trout in rivers and lakes, interactions
with natural populations, and diel movement patterns
[16, 20, 40]. e uncertainty around the catchability and
movement patterns of triploid trout in popular sport
fisheries and the potential for these sterile fish as an alter-
native to traditional stocking of diploid trout objectives
make acoustic telemetry a suitable assessment technique,
especially if paired with studies on the catchability and
movement patterns of triploid and diploid trout. Accord-
ingly, the objectives of this study were to compare diploid
and triploid rainbow trout with respect to their (1) con-
tribution to lake sport fisheries, (2) fine-scale movements
in the lake, (3) rate of migration from the lake, and (4)
natural mortality. Movements patterns of stocked diploid
and triploid trout revealed in this study will improve the
ability of inland fisheries managers to maximize catch
rates or rainbow trout while meeting management objec-
tives associated with conservation.
Methods
Creel sampling
Goldendale, fall spawning strain, triploid (mixed sex,
thermally heat shocked) and diploid rainbow trout were
reared to similar size at Eels Springs Hatchery in Shelton,
Washington on spring water. Equal numbers of triploid
and diploid trout (36,372 of each) were stocked into 15
western Washington lakes (Table1) to achieve a ratio of
50:50 triploid to diploid, targeting a total stocking den-
sity of 22.26 fish/hectare (Table1). Triploid trout were
marked for field identification by removing the adipose
fin 6months prior to stocking. Stocked trout fell within
the “catchable” size with a stocking rate of 1.04 fish per
kilogram ± 0.03 SD (mean ± SD; triploid = 1.05 ± 0.03 and
diploid = 1.04 ± 0.03). All fish were stocked 1week prior
to the opening day of trout season (24 April 2021).
We conducted creel surveys on 15 western Washington
lowland lakes in Pierce, Kitsap, urston, Jefferson, and
Mason counties (Table1), ranging in area from 4.45 ha
(Aldrich Lake) to 95.51ha (Ohop Lake). ese lakes sup-
port popular fisheries on the opening day of trout fish-
ing (4th Saturday in April). Species composition varies
between lakes but includes centrarchids, cyprinids, cot-
tids and wild, native anadromous species such as coastal
cutthroat trout and coho salmon O. kisutch.
Angler interviews were conducted from 08:00 to
12:00h on opening day (24 April 2021) at all study lakes
to estimate the catches of triploid and diploid trout. As
reported by Losee and Phillips [25], this sampling period
coincides with the peak of inland trout harvest in west-
ern Washington and thus the best index of the fishing
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Peaseetal. Movement Ecology (2023) 11:57
season. Samplers interviewed anglers and recorded both
boat and shore angler trip time, lure type, and numbers
of fish caught and released, and retained. All retained fish
were checked for clipped (triploid) and non-clipped (dip-
loid) adipose fins. Informative flyers notified anglers of
the presence and identification of acoustically tagged fish,
and how to report and return tags that were recovered.
is information was shared in a WDFW blog (https://
wdfw. medium. com/ the- secret- lives- of- rainb ow- trout-
36a2d 00fd9 bf) to encourage anglers to report caught
trout.
Acoustic tracking
e acoustic tracking component of this study took
place in two of the 15 lakes, Ward (N 47.008767°,
W-122.875442°) and Ohop (N 46.905224°,
W-122.273341°) lakes (Fig.1). Triploid (n = 40) and dip-
loid (n = 40) trout were acoustically tagged (V9-6L, sig-
nal delay of 220–340s, battery life 912days, Innovasea,
Canada, Halifax) at the hatchery. Specifically, trout were
anesthetized with MS-222 (0.07 g/L) and supported
upside down by a closed cell foam block during surgery,
during which they were given anesthetic by gravity feed
over the gills (0.02g/L). After an incision was made in
the abdomen forward of the pelvic girdle muscle, a trans-
mitter was inserted, antibiotic injected (25 mg/kg oxy-
tetracycline), and the incision sutured with 2–3 stitches
(4-0 RB-1 Taper antibacterial Ethicon Vicryl Plus vio-
let braided, Johnson & Johnson, United States, New
Brunswick, New Jersey). e incision was treated with
antibacterial ointment (Bacitracin®), and weight and
length were recorded. Following tagging, fish were held
with aerated water until swimming upright and respon-
sive. All tags and surgery tools were disinfected with
Nolvasan® (chlorhexidine diacetate) and rinsed in saline
solution before use and between fish. Tagged triploid
fish ranged from 122 to 377 g (mean ± SD: 207 ± 45.5)
and length (mm) 222–292 (mean ± SD: 250.23 ± 14.37).
Diploid fish weight ranged from 128–376g (mean ± SD,
260.0 ± 20.3) and length 227–300 mm (mean ± SD,
225.5 ± 62.3). Individuals were only tagged if they weighed
more than 120g to ensure that the internal tag did not
exceed 3% of the dry body weight of the fish [35]. Prior to
stocking, individuals were placed in a recovery tank and
monitored for 30 d before being transported and stocked
in the study lakes. Twenty triploids and twenty diploids
were stocked each in Ohop Lake and Ward Lake on 20
April, on the same day as untagged individuals (Table1),
4days prior to opening day of fishing.
Ward Lake in urston County, Washington (27.11ha,
20.4m maximum depth) is a mixed species fishery man-
aged for kokanee O. nerka and as a put-and-take fishery
for rainbow trout. In Ward Lake, stocking of adult rain-
bow trout as a put-and-take fishery has occurred annu-
ally since 1935. Additional species found in the lake
include rock bass Ambloplites rupestris, largemouth bass
Micropterus salmoides, bluegill Lepomis macrochirus
and coastal cutthroat trout (WDFW, unpublished data).
Table 1 Surface Hectare of studied lakes and stocking density of triploid and diploid trout in western Washington prior to opening
day of trout fishing (April 24th) in 2021
Lake name County Size Number of sh stocked Stocking density
Surface hectare Triploids Diploids Total Fish/Hectare
Clear Lake Thurston 70 4760 4760 9520 136.0
Hicks Lake Thurston 65 4400 4400 8800 135.9
Ward Lake Thurston 27 1835 1835 3670 135.4
Crescent Lake Pierce 19 1293 1293 2586 136.0
Ohop Lake Pierce 96 6490 6490 12,980 135.9
Tarboo Lake Jefferson 8 558 558 1116 135.8
Buck Lake Kitsap 8 512 512 1024 136.0
Panther Lake Kitsap 41 2775 2775 5550 135.9
Wildcat Lake Kitsap 44 3285 3285 6570 149.1
Aldrich Lake Mason 4 292 292 584 136.1
Benson Lake Mason 32 2195 2195 4390 135.9
Devereaux Lake Mason 40 2693 2693 5386 135.7
Haven Lake Mason 28 1898 1898 3796 134.0
Robbins Lake Mason 7 454 454 908 136.0
Tiser Lake Mason 43 2932 2932 5864 135.9
Total 36,372 36,372 72,744
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Fig. 1 Study area map A indicating the area in Washington where both study lakes were located. Panel B shows Ward Lake (a) located in Thurston
County, Washington, andOhop Lake (d) located in Pierce County, Washington. Also, shown in panel B are the two additional receivers located
downstream of Ohop Lake (b, c): one receiver is located at the confluence of Ohop Creek with the mainstem as the Nisqually River (c) and a second
at river km 19 of the mainstem Nisqually River (b). All other receivers in Ohop Lake are shown as black dots in panel C. The five Ward Lake receivers
are shown as black dots in Panel D
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Ohop Lake in Pierce County, Washington (area: 95.51ha,
maximum depth: 7.6m) is managed as a mixed species
fishery with a rainbow trout emphasis. Rainbow trout
have been stocked in Ohop Lake since 1995 to provide
put-and-take fishing opportunity. Additional species
found in the lake include brown bullhead Ameiurus neb-
ulosus, largemouth bass, largescale sucker Catostomus
macrocheilus, sculpins Cottus spp., yellow perch Perca
flavescens, pumpkinseed sunfish Lepomis gibbosus, black
crappie Pomoxis nigromaculatus, coho salmon and cut-
throat trout (WDFW, unpublished data). e southern
end of Ohop Lake flows through Ohop Creek into the
Nisqually River (Fig.1).
Five acoustic receivers (VRTx, Innovasea, Canada,
Halifax) were deployed in Ward Lake and 22 in Ohop
Lake on April 19, 2021 (Fig.1). Internal synchronization
tags were used to synchronize receiver internal clocks.
Prior to deployment of the receiver arrays, range test-
ing was conducted using the same acoustic transmit-
ters being implanted into study fish (V9-6L, signal delay
of 220–340 s). In Ward Lake range testing suggested
targeting 200 m to achieve a detection range greater
than 90%. To achieve a detection range of 80% we tar-
geted 150 m in Ohop Lake. Receivers were deployed
approximately 230 m apart in WardLake and 200 m
apart in Ward Lake. To detect fish leaving Ohop Lake we
deployed one receiver at the confluence of Ohop Creek
and the Nisqually River (Fig.1) and one receiver in the
lower mainstem of the Nisqually River (46.98, − 122.64).
Detection probabilities for receivers varied between the
lakes. In Ward Lake detection probabilities were > 70%,
up to 220m from a tag (Ward Lake; mean ± SE, 85 ± 15%)
and > 50% when 130m away in Ohop Lake (mean ± SE,
85 ± 15%).
Data analysis
We evaluated the contribution of each ploidy strain to
the catch by summing the total number of triploid (adi-
pose fin clipped) and diploid (unclipped) rainbow trout
reported to be caught during creel surveys at study lakes
on opening day. A chi-square test was used to assess the
probability of capture for triploids relative to diploids
with the odds ratio, ɸ = Ѡ1/Ѡ2, where Ѡ1 represents
the relative contribution of stocked fish from each group
(triploid versus diploid) to the total stocked and Ѡ2
represents the relative contribution of fish caught in the
test fishery from each group to the total number of fish
caught.
Acoustic telemetry was used to detect tagged trout in
Ohop and Ward lakes and estimate the rates of mortality
and emigration. Angler reporting of tagged fish caught,
and detection history allowed for an assignment of
“fate” for individuals removed from the lake. Individuals
that were last detected at the receiver in the outlet and
then never detected in the lake again were classified as
migrants. Tagged fish returned by anglers were classified
as having been caught. Sedentary fish, based on acous-
tic detections, were classified as natural mortalities. All
other tags that went undetected during the study period
were classified as “unknown removal”.
Fine‑scale positioning
Raw acoustic telemetry detection data were downloaded
and sent for processing to Innovasea for VEMCO Posi-
tioning System (VPS). VPS utilizes hyperbolic position-
ing to get a weighted-average position for a fish based on
the time difference of arrival at multiple receivers for a
single ping of a transmitter. VPS provides an estimate of
the horizontal position error (HPE) associated with each
of the positions [36]. Differences in space use between
triploid and diploid trout were determined using fine-
scale positions and kernel utilization distribution (KUD),
which describes the probability of a rainbow trout in a
location of the lake based on a utilization distribution
[42]. Areas of high importance, known as core areas,
were represented by 50% of the KUDs. Home ranges
were signified by 95% KUDs. e “ks” package in R was
used to calculate both home ranges and core areas for
both ploidy strains and for day and nighttime periods.
Day and night were defined using the “suncalc” pack-
age in R, defined by local sunrise and sunset. Individu-
als with fewer than 50 detections were excluded from the
analysis because the data were insufficient to accurately
determine a KUD. ArcGIS 10.8.2 was used to create ker-
nel density maps for both ploidy strains and time periods
(day and night) at both study lakes to qualitatively visu-
alize the spatial distribution of the fine-scale positional
data. Kernel density rasters had an output cell size of
0.1m and show the least to most dense areas of use by
each rainbow trout and between the two time periods.
e home range distributions were not normally dis-
tributed, so we compared triploids and diploids in each
lake and between day and night periods with a series of
Mann–Whitney-Wilcoxon tests.
Results
A total of 891 anglers were interviewed across the 15
study lakes where similar densities of triploid and diploid
rainbow trout were stocked (Table1). On opening day
of fishing (April 24th) creel samplers reported 742 trout
total, of which fewer were triploid (316, 42.59%) than
diploid (426, 57.41%; Chi-square = 15.8, p < 0.001). O dds
ratio revealed that across the 15 lakes stocked, triploid
trout were caught at a rate 15.3% lower than would have
been expected based on stocking. Lake specific patterns
of trout contribution (triploids versus diploids) varied;
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diploids contributed more than triploids in 6 of 15 lakes
(p < 0.05, Chi-square test; Fig. 2), and triploids contrib-
uted significantly more only in Crescent Lake (73.5% of
observed catch, Chi-square = 15.8, p < 0.05; Fig. 2). In
8 of 15 study lakes, triploid and diploid trout contribu-
tion rates did not differ from expected based on stocking
(p > 0.05, Chi-square test).
In the two study lakes, acoustic receivers recorded
more than 300,000 individual detections from the 80
tagged trout, with an average of 3,850 detections per
trout (± 1802). All tagged fish were detected the first
day after stocking, and 19 tagged fish were still present
55 days later, on 15 June (9 in Ward and 10 in Ohop;
Fig. 3). Overall apparent survivorship was similar for
triploids and diploids but different between lakes with
50% of tagged trout in Ohop Lake no longer available to
the fishery 21 d after stocking because of capture, appar-
ent natural mortality (i.e., tag became motionless in the
lake), migration, or unknown removal (Fig.4). In Ward
Lake, fish survived longer; 50% were still available 36 d
after stocking (Fig.4).
Anglers reported recoveries of tagged trout in both
Ward (4 diploid and 4 triploid) and Ohop Lake (2 dip-
loid), all within two months of stocking (Fig.3). e last
detection locations indicated that 25% (10/40) of stocked
trout migrated from Ohop Lake, mostly (8/10) within 21
d of stocking (Fig.3) and mostly (7/10) diploid trout. Two
diploid rainbow trout were reported as taken by anglers
at Ohop Lake. Nearly half (44%: 35/80) of the tagged
trout were removed from lakes by unknown causes and
24% (19/80, 10 triploids and 9 diploids) survived until
the end of the study (Ward Lake: 118days, Ohop Lake:
55days).
e HPE values for synchronization tag position data
collected in the two weeks prior to the start of the study
were compared to twice the distance root mean square of
measured error (HPEm) [3, 28].VPS calculated positions
for study fish were filtered by HPE less than 10 to signifi-
cantly reduce positioning errorwhich resulted in 55.4%
(58,743) of positions in Ward Lake having a HPE less than
10. In Ohop Lake 75.4% (27,264 positions) had an HPE
less than 10. Qualitative spatial analysis of the data indi-
cated more variability in lake usage areas for diploid trout
in comparison to triploid trout. However, the time of
day did not greatly impact the patterns of usage (Fig.5).
Areas of high use were focused on the central portions of
both study lakes with fish moderately using some littoral
regions of the lake (e.g., southern shore of Ward Lake and
eastern shore of Ohop. Overall, home range did not dif-
fer significantly between diploids and triploids (Mann–
Whitney–Wilcoxon test, W = 1623, p = 0.30 or between
day and night periods (Mann–Whitney–Wilcoxon test,
Fig. 2 Relative proportion of triploid (black) versus diploid (grey) caught in selected Western Washington Lakes. Horizontal red line represents a 1:1
ratio between expected and realized for triploid catch
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Fig. 3 Tagged triploid and diploid rainbow trout across the study period from 24 April–23 August 2021 in A Ohop Lake, Pierce County and B Ward
Lake, Thurston County, Washington. Each horizontal line represents an individual fish for the period that they remained in the study area, and fish
are grouped by their fate in the study
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W = 2095, p = 0.15; Fig. 6). Home ranges were signifi-
cantly greater for both diploids and triploids in Ohop
Lake than Ward Lake (Mann–Whitney-Wilcoxon
test, W = 3522, p < 0.005, Fig. 6), likely because Ohop
Lake is larger (95.51ha vs. 27.11 ha for Ward Lake). In
neither lake was there a significant difference in ploidy
strain (Mann–Whitney–Wilcoxon test, Ward: W = 350,
p = 0.05; Ohop: W = 349, p = 0.27) or time period (Mann–
Whitney–Wilcoxon test, Ward: W = 604, p = 0.14; Ohop:
W = 426, p = 0.78).
Fig. 4 Trout survivorship for Ohop Lake (A) and Ward Lake (B) diploid (black-dashed) and triploid (grey) rainbow trout. With a dotted line indicating
the time at which 50% of individuals were no longer in the study
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Discussion
e results of this multi-faceted study, combining stand-
ard creel monitoring with fine scale tracking, indicated
that triploid trout were a viable alternative to traditional
diploids for maintaining angler opportunity while reduc-
ing the conservation concern associated with genetic
introgression. Standard creel monitoring in 15 lakes
showed that catch rate of diploids was greater than 50%
in most lakes but triploid trout still contributed greatly
to fisheries. Secondly, fine scale tracking showed that
triploids had a decreased rate of emigration out of the
lake, similar survivorship, and similar diel movements in
comparison to diploid trout. However, the small number
of trout leaving reduced our ability to demonstrate a dif-
ference in migration (3 triploids vs. 7 diploids), and this
might be fruitful area of future work.
Our study represents the first use of acoustic telemetry
to document the movement patterns of triploid rainbow
trout and provides important insights into the catch-
ability of stocked triploid trout relative to traditional
diploids. Consistent with previous studies comparing
catch rates of triploid versus diploid trout, our results
indicated that triploid trout stocked in lakes can return
to the creel at a somewhat reduced or similar rate than
that of diploids [9, 21, 22]. By combining standard creel
monitoring and fine scale acoustic telemetry, our results
help to understand why rates of catchability between
triploids and diploids often differ. Specifically, fewer trip-
loid trout left the lake, and remaining trout had similar
home ranges between the two ploidy strains; both these
qualities may be perceived as desirable for fisheries man-
agement objectives associated with the need to balance
conservation and fishing objectives. ese findings have
important implications for managers weighing the cost
and benefits of differing stocking plans.
In Ohop Lake, where migrating rainbow trout have
access to waters used by anadromous conspecifics, 25%
(10/40) of the tracked trout were last detected in the out-
let of the lake. Extrapolating the observed rate of migra-
tion to the total number of trout stocked in Ohop Lake
(12,980), as many as 3245 hatchery trout might have left
Ohop lake in 2021. e present study had a relatively
Fig. 5 Kernel density of triploid (left) and diploid (right) rainbow trout during the day (A) and night (B) in both study lakes. Kernel density rasters
had an output cell size of 0.1 m and show the least to most dense areas of use by each rainbow trout ploidy strains and between the two time
periods
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Page 10 of 13
Peaseetal. Movement Ecology (2023) 11:57
small sample of tracked fish, therefore, we recommend
caution in such an extrapolation, and regard these
results as tentative. None of the tracked rainbow trout
were detected at the confluence of Ohop Creek and Nis-
qually River or in the lower Nisqually River, so it is likely
that stocked trout the left Ohop Lake remained in the
creek or experienced low survival in the fluvial environ-
ment of Ohop creek, consistent with other studies [2,
18, 37]. Regardless of the exact number of trout that left
Ohop Lake in the current study, rates of emigration are
Fig. 6 Home ranges (m2) for both triploid and diploid rainbow trout in both the day and night periods at Ohop Lake (A) and Ward Lake (B). Note
differing Y-axis between panels. Home range did not differ significantly between diploids and triploids (Mann–Whitney–Wilcoxon test, W = 1623,
p = 0.30) or between the two time periods (Mann–Whitney–Wilcoxon test, W = 2095, p = 0.15)
Content courtesy of Springer Nature, terms of use apply. Rights reserved.

Page 11 of 13
Peaseetal. Movement Ecology (2023) 11:57
significant in Ohop Lake. Risks associated with these
findings (e.g., competition, genetic introgression) may
be partially mitigated by stocking sterile triploid trout.
Trout stocking plans are designed to achieve goals based
on angler opportunity and satisfaction. erefore, stock-
ing strategies that limit emigration and reduce gene flow
from domesticated hatchery stocks to wild trout while
achieving these angler-related goals are preferable.
Triploid trout stocked in Ohop and Ward lakes demon-
strated comparable home ranges relative to diploid trout.
However, density maps (Fig.4) showed less variability in
the distribution of triploid home ranges, perhaps further
limiting the potential to leave the waterbody they were
stocked in, relative to diploids. is reduced migration
rate for triploids may provide a benefit for fisheries man-
agers. Additionally, the reduced variability in home range
may have contributed to the slightly overall lower catch
rates for triploids observed in the current study, if dip-
loids distributed in a way that enhanced their potential
to be caught. Given the conservation concern associated
with wild steelhead and cutthroat trout in waterbod-
ies connected to important put-and-take rainbow trout
fisheries [39], managers may benefit from prioritizing
the available triploid rainbow trout for stocking in lakes
where both the conservation risks and likelihood of emi-
gration are the greatest. In addition, consideration should
be given to the potential for mitigating for reduced catch
of triploids by considering other factors that influence
catch rates, such as stocking density [29], stocking sea-
son [44], prey availability [13], fish size [7, 25] and stock-
ing location [15] to fine-tune triploid stocking plans.
Together these results suggest raising fish to a larger
size, stocking near fishing access points and stocking just
prior to the opening of the fishery are likely to support a
reduction in the total fish that need to be released, thus
mitigating increased cost or reduced catch rate associ-
ated with stocking triploids. In doing so, managers could
maximize chances of achieving management objectives
associated with both conservation and opportunity.
Our study was not designed to identify causes of
variability in catch rates between triploid and diploid
trout. However, others have explored this topic and the
results have important management implications to
consider before applying these results to other systems.
Previous studies suggested that triploids may have a
reduced aerobic capacity and decreased tolerance to
chronic stress [12, 33], therefore catch rates and move-
ment patterns could be affected by variability in habitat
conditions (e.g. temperature, pH). In the current study,
the catch rate of diploids was greater than 50% in most
study lakes over a broad range of environmental con-
ditions. For example, diploids made up > 75% of trout
sampled in one of the smallest lakes in this current
study, Buck Lake (7.69ha) and the largest lake, Ohop
Lake (131.93ha) suggesting lake size alone is not a good
predictor of triploid trout catchability. Koenig et al.
[21] found stocking density to be the most important
factor explaining variability of triploid trout catchabil-
ity, but we observed differences in movement patterns
and catch for tagged triploids relative to diploids across
two different sized lakes stocked at similar density and
variable catch rates across the broader set of lakes. is
information highlights the need to better understand
factors affecting catch rates of triploid rainbow trout to
increase precision around stocking programs. While it
is beyond of the scope of this study, future work should
further investigate factors affecting both catch rate and
home range of triploids and diploids to clarify potential
causes for the patterns reported here.
Triploid rainbow trout represent an important tool for
fisheries managers faced with increasing threats to wild
populations of salmonids and growing pressure for fish-
eries managers to design sustainable fishing opportu-
nity. Pairing acoustic telemetry with a traditional stock
assessment tool (i.e., creel survey), we demonstrated
that triploid trout were a viable alternative when stock-
ing rainbow trout in western Washington lakes. Com-
pared to diploid trout, triploids were caught at a reduced
rate overall but exceeded or met expectations in many
waterbodies (Fig. 2). With a comparable home range
and reduced rate of emigration, our results provide sup-
port for a modification of trout stocking where concerns
over genetic introgression with wild stocks exist [32, 39].
A strategic approach by managers to integrate triploids
into current stocking plans while prioritizing values (e.g.
conservation vs. opportunity) has potential for main-
taining or improving catch rates of these popular sport
fisheries while providing increased protection for native
populations.
Acknowledgements
This project would not have been possible without the assistance of the team
at WDFW’s Eels Springs hatchery, especially Michael Lucero and Steve Smo-
therman. Additionally, we would like to thank Kinsey Frick for assisting with
tagging effort. We are also grateful for support and fruitful conversations with
WDFW colleagues Kenny Behen, Riley Freeman, John Pahutski, Tara Livingood-
Schott, Jeremiah Shrovnal, Megan Moore (NOAA), and the Swedish University
of Agricultural Sciences team in Tomas Brodin’s lab. The WDFW Fish Program
leadership team and Toby Harbison provided thoughtful comments on earlier
versions of the manuscript.
Author contributions
JP and JL wrote the main manuscript text and prepared figures. AK, JP, JL,
GM, MB, JM, TQ, DP, GH, ML, SC, designed the study. AK, JP, GM, JL, ML, DP,
GH, assisted in performing surgeries and deploying tracking equipment. ML
facilitated and oversaw the raising and production of both triploid and diploid
rainbow trout used in the study. DP, GH, TQ, provided equipment, training, and
funding for the project. All authors reviewed the manuscript.
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Page 12 of 13
Peaseetal. Movement Ecology (2023) 11:57
Funding
Open access funding provided by Swedish University of Agricultural Sciences.
Funding and/or equipment was provided by WDFW, University of Washington
and the Coastal Cutthroat Coalition.
Availability of data and materials
The data that support the findings of this study will be available upon request.
Declarations
Ethical approval and consent to participate
Ethical and legal approval was obtained from Washington Department of Fish
and Wildlife prior to the start of the study.
Competing interests
The authors declare no competing interests.
Received: 29 March 2023 Accepted: 2 September 2023
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