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Prey and non-native fish predict the distribution of Colorado pikeminnow

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Article in Ecology of Freshwater Fish · July 2014

DOI: 10.1111/eff.12093

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Ecology of Freshwater Fish 2013 Ó 2013 John Wiley & Sons A/S. Published by John Wiley & Sons Ltd

ECOLOGY OF
FRESHWATER FISH

Prey and non-native fish predict the distribution

of Colorado pikeminnow (Ptychocheilus lucius)
in a south-western river in North America
Nathan R. Franssen1, Scott L. Durst2
1
Department of Biology and Museum of Southwestern Biology, University of New Mexico, 167 Castetter Hall, Albuquerque, NM 87131, USA
2
U.S. Fish and Wildlife Service, New Mexico Ecological Services Field Office, San Juan River Basin Recovery Implementation Program, 2105 Osuna Road NE,
Albuquerque, NM 87113, USA

Accepted for publication July 27, 2013

Abstract – Colorado pikeminnow (Ptychocheilus lucius) has been extirpated from a large portion of its historical
range in the Colorado River basin, USA. A repatriation effort via stocking of juvenile P. lucius in the San Juan
River, NM, CO and UT has resulted in limited recruitment of individuals into an adult population. Understanding
biotic and abiotic factors that limit their persistence in the Colorado River basin will be a critical step in providing
for their recovery. To elucidate potential recruitment barriers in the San Juan River, we assessed relationships
between the numbers of two age classes of P. lucius and prey, competitors and predators collected at a 1.6 km
reach scale between 2003 and 2012. We used an information theoretical approach to rank candidate models testing
the relative importance of these biotic conditions in predicting the spatial distribution of P. lucius. We found
positive relationships between the numbers of P. lucius ≤200 mm total length (TL) collected and catch per unit
effort (CPUE) of native prey among reaches. For P. lucius >200 mm TL (individuals that are likely completely
piscivorous), we found positive associations between the numbers of P. lucius collected and CPUE of total prey
and CPUE of potential non-native competitors in each reach. Our data suggest size-specific affinities of P. lucius
for native and non-native prey as well as the potential for negative interactions between P. lucius and non-native
competitors may contribute to limited recruitment of juvenile P. lucius into an adult population in the San Juan
River.

Key words: non-native fishes; negative binomial regression; AIC; desert streams

The federally protected Colorado pikeminnow

Introduction
Habitat alteration, introduction and establishment of
non-native fishes and changes to natural flow regimes
have contributed to the decline of native fishes
worldwide (Dudgeon et al. 2006; Fullerton et al.
2010). Approximately 40% of North American fishes
are currently at risk (i.e., endangered, threatened,
vulnerable; Jelks et al. 2008), and fishes in the south-
western USA are exceptionally imperilled because of
their high degree of endemism (Minckley & Deacon
1968; Holden & Stalnaker 1975; Fagan et al. 2002).
Identifying and ameliorating factors that limit the per-
sistence and recovery of endangered fishes will be a
major challenge in the coming decades.
(Ptychocheilus lucius) is a large piscivorous cyprinid
that is endemic to the Colorado River basin in the
south-western USA (United States Fish & Wildlife
Service 1967; Tyus 1991). Currently, wild popula-
tions are limited to areas upstream of Lake Powell,
UT and AZ, in the Colorado and Green River sub-
basins, UT and CO. Remnant individuals of the
historical population in the San Juan River NM, CO
and UT were considered extremely rare (Tyus et al.
1982; Platania et al. 1991), and population restora-
tion efforts via stocking mostly age 0 (i.e., young-
of-year) hatchery-reared fish began in 1996 and
continue today. While augmentation has increased
the numbers of P. lucius in the river, stocked

Correspondence: N. R. Franssen, Department of Biology and Museum of Southwestern Biology, University of New Mexico, 167 Castetter Hall, Albuquerque,
NM 87131, USA. E-mail: nrfranssen@gmail.com

doi: 10.1111/eff.12093 1
Franssen & Durst
individuals are rarely encountered more than three
years after stocking, but female P. lucius do not
become reproductive until at least six years of age
(Vanicek & Kramer 1969). The biotic or abiotic fac-
tors limiting the persistence of stocked individuals
and evidently creating a recruitment bottleneck are
unknown.
Age 0 P. lucius often use low-velocity habitats in
lower portions of large rivers (Haynes et al. 1984;
Osmundson et al. 1998), whereas adults can be
highly mobile and often use mainstem river habitats
(Osmundson et al. 1998). Information on subadult
habitat use is limited, but they also appear to be
highly mobile and can use mainstem as well as small
tributary habitats (Fresques et al. 2013). Small
P. lucius are mostly insectivorous until they become
large enough to consume fish prey (~ 100 mm TL;
Vanicek & Kramer 1969; Muth & Snyder 1995), and
individuals >200 mm TL can be entirely piscivorous
(Vanicek & Kramer 1969). It is currently assumed
the San Juan River has adequate habitat and
resources to support all life stages of P. lucius, but
understanding what factors limit population persis-
tence, and particularly juvenile recruitment into
adults, will be requisite to protect and recover
P. lucius.
Changes in the composition of the San Juan
River’s fish community have occurred with non-
native fish introductions. The historical fish commu-
nity of the San Juan River was relatively depauperate
(i.e., up to eight species; Sublette et al. 1990), but
more recent investigations have documented the pres-
ence of 19 non-native fishes (Ryden 2000). While
some small-bodied non-native fishes may be potential
prey for P. lucius, smaller P. lucius individuals may
be gape limited due to the body morphology of non-
natives compared with native fishes (e.g., non-native
red shiner Cyprinella lutrensis versus native bluehead
sucker Catostomus discobolus and native flannel-
mouth sucker C. latipinnis; Franssen et al. 2007).
Thus, P. lucius individuals may rely on prey they
shared an evolutionary history with rather than
recently introduced prey. While the relative abun-
dance of small-bodied native and non-native fishes
may dictate prey options to P. lucius, other large-
bodied fishes may pose a predation risk to or
compete for resources with P. lucius. Non-native
channel catfish (Ictalurus punctatus) are the second
most frequently encountered large-bodied species in
the system, comprising an average of 22% of all
large-bodied fishes collected annually (Ryden, D.W.
unpublished). Similar to young P. lucius, I. punctatus
in all life stages feed on invertebrates (Sublette et al.
1990) and therefore have the potential to compete for
prey resources if these species occupy the same habi-
2
tats. Adult I. punctatus, although highly omnivorous,
may also have deleterious impacts on P. lucius
through direct predation. Indeed, the fish component
of I. punctatus diet tends to increase when
individuals reach 300 mm TL (Bailey & Harrison
1945). Additionally, Gido & Propst (2012) identified
a weak negative trend in the annual catch per unit
effort (CPUE) of I. punctatus >300 mm TL and
native and non-native small-bodied fishes in the San
Juan River.
Concurrent with alterations to the native fish com-
munity, the San Juan River’s natural flow regime has
been altered with its impoundment by Navajo Dam
in 1962. However, reservoir releases since 1998 are
currently managed to often follow recommendations
to mimic a natural flow regime characterised by high
spring flows and low summer base flows (Propst &
Gido 2004; Gido & Propst 2012). Yet, spring peak
flows are still attenuated, and summer flows are
higher compared with historical levels (Franssen
et al. 2007). High spring flows facilitate increased
CPUE of native prey fishes, while low summer flows
tend to increase the CPUE of non-native small-bod-
ied fishes (Propst & Gido 2004; Gido & Propst
2012). Changes to the natural flow regime and intro-
duced fishes have potentially altered the annual quan-
tity and availability of native and non-native prey for
juvenile P. lucius.
The numerous changes to the biotic and abiotic
conditions in the San Juan River make identifying
factors that limit recruitment and thus population per-
sistence of P. lucius difficult. Our goal was to gain
insight into factors that may limit population
persistence of stocked P. lucius by assessing environ-
mental conditions that drive spatial variation in the
numbers of P. lucius collected. Because P. lucius
can be highly mobile (moving both within and
among river reaches; Tyus 1991), we used a broad-
scale (1.6 km) assessment of CPUE of other fishes to
predict the numbers of P. lucius in each reach. We
used riverwide data on the CPUE of potential fish
prey (i.e., native and non-native small-bodied fishes),
potential competitors (age 0 and juvenile I. punctatus)
and predators (adult I. punctatus) to assess what
factors, if any, are associated with the spatial distribu-
tion of age 1 (≤200 mm TL) and age 2+ (>200 mm
TL) P. lucius in the San Juan River.
We predicted age 1 P. lucius would be positively
associated with native prey (due to gape limitations
for non-native prey) while negatively associated with
age 0 and adult I. punctatus. Conversely, we pre-
dicted that age 2+ individuals would be positively
associated with total prey (i.e., native and non-native
prey), but negatively associated with juvenile and
adult I. punctatus.

Methods
Study site
The San Juan River originates in south-west Colorado
and is a major tributary to the Colorado River draining
99,200 km2 in Colorado, Utah, Arizona and New
Mexico (Carlson & Carlson 1982). The closing of
Navajo Dam in 1962 inundated 56 km of the river,
which currently flows for 365 km from Navajo Dam
to Lake Powell (Fig. 1). We used river km (Rkm) to
designate locations along the San Juan River and con-
sidered Rkm 0.0 the inflow near to Lake Powell. The
relative numbers of fishes vary longitudinally along
the river course (Gido & Propst 2012; Ryden 2012),
likely due to differences in substrate, habitat availabil-
ity, gradient and water temperature (Bliesner & Lama-
rra 2000). Due to hypolimnetic releases from Navajo
Dam, water temperature increases moving down-
stream during the summer months. The river exhibits
some braiding upstream of river Rkm 85.0, but is
mostly canyon bound between Rkms 0.0 and 85.0.
Fish sampling primarily occurred between Rkm 289.7
and 0.0, but analyses were restricted to Rkms below
261.7 (see below for rational).
P. lucius catch numbers
We used data from riverwide large-bodied fish sur-
veys conducted each fall (September or October) via
Fig. 1. Study map of the San Juan River where catch per unit effort (CPUE) of prey, competitors and predators were investigated to predict
the catch numbers of age 1 and age 2+ Ptychocheilus lucius. The river flows from east to west, and river km 85.0 is the beginning of
canyon-bound reaches before the river enters Lake Powell.
Predicting the spatial distribution of P. lucius
raft-mounted electrofishing units between 2003 and
2012 to assess the spatial distribution of P. lucius
(Fig. 2). We had two obvious size classes of P. lucius
during the study period, and because P. lucius likely
have size-specific ecologies, we grouped individuals
≤200 mm TL as age 1 (mean = 168 mm,
range = 112–200) and individuals >200 mm TL as
age 2+ (mean = 275, range = 201–658). To allow for
undisturbed small-bodied fish sampling during the
same sampling periods, only three of every four 1.6
Rkm sections (a reach henceforth) were sampled by
raft electrofishing. In all reaches, two electrofishing
rafts sampled each shoreline of the river and the num-
ber of fishes and effort (seconds) from both rafts were
summed at the ends of each reach.
Large-bodied predators and competitors
To assess potential effects of non-native I. punctatus
on the spatial distribution of P. lucius, we used data
from the same riverwide large-bodied fish surveys
conducted between 2003 and 2012 (Fig. 2). For our
study, juvenile I. punctatus were individuals
≤300 mm TL and adults >300 mm TL (De Roth
1965; Elrod 1974). These data allowed us to simulta-
neously assess the catch numbers of P. lucius and
CPUE of potential predators of age 1 and age 2+
P. lucius (i.e., adult I. punctatus) and competitors
(i.e., juvenile I. punctatus) of age 2+ P. lucius in
each reach.
3
Franssen & Durst

Fig. 2. Spatial variation in catch numbers of age 1 and age 2+ Ptychocheilus lucius, and catch per unit effort (CPUE) of native and non-
native prey, age 0, juvenile and adult I. punctatus from the San Juan River between 2003 and 2012. Values of 0.001 (seining data) or 1.0
(electrofishing data) were added to independent variables to facilitate plotting on a log scale.

channels encountered were sampled. All fishes were

Small-bodied prey and competitors
To quantify the spatial CPUE of potential prey of age
1 and age 2+ P. lucius and competitors of age 1
P. lucius, small-bodied fishes were collected with
seine (4.6 9 1.8 m, mesh = 3.2 mm) every fall annu-
ally between 2003 and 2012 (Fig. 2). One of every
four reaches (i.e., the reach not sampled by electro-
fishing each year) was sampled with 8–16 seine hauls
along about 150 m of shoreline. In addition to these
designated sample sites, all backwaters and secondary
counted and measured, and the area of each seine haul
was recorded. The CPUE of small-bodied fishes was
quantified in each reach by dividing the total numbers
of individuals of each species by the total area
sampled. We quantified the CPUE of native prey (i.e.,
C. latipinnis, C. discobolus and speckled dace
Rhinichthys osculus), non-native prey (C. lutrensis,
fathead minnow Pimephales promelas, western mos-
quitofish Gambusia affinis) and potential competitors
of age 1 P. lucius (age 0 I. punctatus). Cyprinella

4

lutrensis was the most common non-native prey
(mean CPUE = 0.27 fish per m2), while R. osculus
and Catostomus sp. were the most common native
fishes (0.12 and 0.05 fish per m2, respectively).
Because not all of the factors we considered to
impact the spatial distribution of P. lucius were quan-
tified in every reach each year, we only included
reaches where all variables were collected within a
single year. We also assumed that responses to prey,
competitors and predators of P. lucius would not
vary over time and therefore did not evaluate tempo-
ral variation. Annual sampling occurred primarily
between Rkm 289.7 and Rkm 0.0, but a selective
barrier (a weir and native fish passage located at
Rkm 268.1) limits upstream movement of non-native
fishes. To avoid potential influences of the weir, we
only included reaches below Rkm 261.7 in analyses
(i.e., five river miles below the weir). Inclusion of all
reaches below the weir did not qualitatively alter
results (data not presented).
Data analyses
Electrofishing data were Log10(x + 1)-transformed,
and seining data were Log10(x + 0.001)-transformed
prior to analyses. To quantify relationships between
CPUE of fishes used as independent variables and
longitudinal position of reaches, we conducted a prin-
cipal component analysis (PCA) using a correlation
matrix. Only axes with eigenvalues >1.0 were
retained for interpretation. We also tested for multi-
collinearity among independent variables used to pre-
dict the catch numbers of both age classes of
P. lucius using Pearson’s correlations and considered
Predicting the spatial distribution of P. lucius
correlations with r < 0.70 as not being highly corre-
lated (the strongest correlation was r < 0.50).
We used negative binomial regression to predict the
catch numbers of age 1 and age 2+ P. lucius in every
reach in all years using the CPUE of potential prey,
competitors and predators. We used catch numbers
per reach rather than CPUE of P. lucius (i.e., individu-
als per hour of electrofishing), because the CPUE dis-
tribution was not amenable to statistical analyses (zero
inflated and right skewed). Initial goodness-of-fit
assessments using a Pearson’s chi-square test revealed
that both the age 1 and age 2+ catch numbers fit a neg-
ative binomial distribution (both P > 0.50; models
including all independent variables were fit).
To assess which factors contributed to the catch
numbers of P. lucius among reaches, we used an
information theoretical approach (Burnham & Ander-
son 2002). We first built nine models for age 1
P. lucius we thought would influence their catch
numbers in each reach (Table 1). These models were
built with effects from CPUE of prey (native and
non-native prey), competitors (age 0 I. punctatus
from seining surveys) and potential predators (adult
I. punctatus from electrofishing surveys). We con-
structed four models to predict the catch numbers of
age 2+ P. lucius using total prey CPUE (sum of
native and non-native prey fishes from seining sur-
veys), competitors (juvenile I. punctatus from elec-
trofishing surveys) and predators (adult I. punctatus
from electrofishing surveys). Although all P. lucius
individuals were present in the river for at least one
year, P. lucius are usually only stocked in the upper
portions of the river. Therefore, we included reach
(i.e., Rkm) as a variable in every model to control for

Table 1. Candidate models and variables included in each (X) developed to predict the spatial catch numbers of age 1 and age 2+ Ptychocheilus lucius from
catch per unit effort (CPUE) of prey, competitors and predators in each reach in the San Juan River, NM and UT. Ictalurus punctatus are abbreviated ‘Ictpun’.

Prey Competitors Predators

Age class Model River km Native prey Non-native prey Total prey Ictpun age 0 Ictpun juvenile Ictpun adult

Age 1 1 (Global) X X X X X
2 (Null) X
3 X X
4 X X
5 X X
6 X X
7 X X X
8 X X X
9 X X X
10 X X X
11 X X X
Age 2+ 1 (Global) X X X X
2 (Null) X
3 X X X
4 X X X
5 X X
6 X X

5
Franssen & Durst
longitudinal variation that may have been associated
with stocking. We also included a null model (reach
only) and a global model (all variables) to bring the
total to 11 candidate models for age 1 and six candi-
date models for age 2+ P. lucius (Table 1). We used
Akaike’s information criterion (AIC) to select the
best candidate models by comparing all of the candi-
date models simultaneously for each age class of
P. lucius. AIC scores were adjusted for small sample
sizes (AICc), and Akaike weights (wi) were quanti-
fied. Models with the lowest AICc and highest wi
were considered having the best support. Only mod-
els with DAICc values <2.0 and wi values >0.10 were
retained for interpretation. Following model selection,
we used model averaging to assess the direction and
magnitude of variable effects from the top-ranked
models. Model averaging was conducted by quantify-
ing the mean and 95% CI of coefficients from all
models of each variable of interest (coefficients were
set to zero in models that did not contain the variable
of interest and were included in the calculations;
Burnham & Anderson 2002). We conducted all sta-
tistical analyses in R (R Development Core Team
2011).
Results
When assessing relationships between independent
variables using PCA, three axes were retained
explaining 76.9% of the variance (PC I = 39.1%, PC
II = 21.8% and PC III = 16.0%). The first PC axis
was positively associated with total prey and reach
(i.e., Rkm) while negatively associated with age 0
I. punctatus (Table 2). The second axis was posi-
tively associated with reach and native prey while
negatively associated with age 0 and juvenile
I. punctatus. The last axis was negatively related to
non-native and total prey while positively associated
with adult I. punctatus and reach. Generally, native
prey tended to increase linearly upstream and peaked
in the highest reaches we investigated. Conversely,
non-native prey and all size classes of I. punctatus
tended to show higher CPUE in the middle reaches
Table 2. Results from principal component analysis (PCA) of independent
variables used to predict the spatial catch numbers of age 1 and age 2+
Ptychocheilus lucius.
Variable PC I (39.1%) PC II (21.8%)
Total prey 0.889 0.052
River km 0.706 0.422
Non-native prey 0.700 0.253
Native prey 0.688 0.201
Adult Ictalurus punctatus 0.572 0.384
Juvenile I. punctatus 0.206 0.881
Age 0 I. punctatus 0.338 0.562
6
with lower CPUE at both lower and higher reaches
of the river (Fig. 2). All fishes and size classes
tended to have low CPUE in canyon-bound reaches
below Rkm 85.0.
A total of 526 age 1 and 398 age 2+ P. lucius was
collected during the study period, and up to 24 and
10 individuals of each age class were collected in a
single reach, respectively (Fig. 2). The catch numbers
of age 1 P. lucius tended to be highest in the upper
reaches of the river, while age 2+ catches were fairly
uniform in the upper half of the river.
Age 1 P. lucius
We identified one model with support that only
included native prey to predict the catch numbers of
age 1 P. lucius (Table 3). Model averaging revealed
that native prey had a positive association with age 1
P. lucius catches in each reach (Table 4; Fig. 3). Our
estimate suggested that for every 1-unit increase in
native prey CPUE, the log catch number of age 1
P. lucius increased by 0.223.
Age 2+ P. lucius
We also retained only one model with support to pre-
dict spatial catch numbers of age 2+ P. lucius
(Table 3). This model contained both total prey and
juvenile I. punctatus, and model averaging indicated
that both had positive effects on the catch numbers of
age 2+ P. lucius (Table 4; Fig. 4). Coefficient esti-
mates indicated that for every 1-unit increase in the
CPUE of total prey and juvenile I. punctatus, the log
catch numbers of age 2+ P. lucius increased by
0.214 and 0.222, respectively.
Discussion
We assessed the spatial distribution of age 1 and age
2+ P. lucius in the San Juan River using data on
native and non-native prey, and potential non-native
competitors and predators of P. lucius. We found
positive associations with age 1 P. lucius and native
prey while catch numbers of age 2+ individuals were
positively associated with total prey and juvenile
I. punctatus.
We found no relationships between age 0 and adult
I. punctatus and catch numbers of age 1 P. lucius,
PC II (16.0%) suggesting that the spatial distribution of age 1
P. lucius was not shaped by interactions with these
0.407
0.371
size classes of I. punctatus at the reach scale. How-
0.479 ever, the positive relationship between age 1
0.130 P. lucius and native prey suggests that this size class
0.646 of P. lucius is tracking native versus non-native food
0.232
0.319
resources. Gape-limited age 1 P. lucius individuals
may occur in reaches with high numbers of native
 Predicting the spatial distribution of P. lucius
Table 3. Negative binomial regression models ranked by AICC that predict
the catch numbers of age 1 and age 2+ Ptychocheilus lucius at the reach
scale in the San Juan River. Only models with DAICC <2.0 and Akaike
weights (wi) >0.10 were retained for interpretation. The values of DAICC
reflect the change in AICC compared with the highest ranked model, and K
is the number of parameters in each model (including the intercept).

Age class Model K AICC DAICC wi

Age 1 7) Native prey + Rkm 3 929.85 0.00 0.75

Age 2+ 9) Total prey + 4 851.37 0.00 0.93
Juvenile Ictalurus
punctatus + Rkm

Table 4. Estimates and 95% CI for negative binomial parameter

coefficients from the variables that were in models that received the most
support using AIC model selection for each age class of Ptychocheilus
lucius.

Age class Parameter Estimate 95% CI

Age 1 Native prey 0.223 0.188

Age 2+ Total prey 0.214 0.158
Juvenile Ictalurus punctatus 0.222 0.166

Fig. 4. Relationships between catch numbers of age 2+ Ptychoc-

heilus lucius per reach and catch per unit effort (CPUE) of total
prey and juvenile I. punctatus. Negative binomial regression lines
are drawn from models using only the independent variable to
predict the dependent variable.

river, suggesting that this region may contain more

Fig. 3. Relationships between catch per unit effort (CPUE) of
native prey and age 1 Ptychocheilus lucius catch numbers per
reach. The negative binomial regression line was drawn from a
model using only the independent variable to predict the depen-
dent variable.
small-bodied fishes because these prey may be more
susceptible to predation due to their more fusiform
body shapes compared with the more laterally com-
pressed non-native C. lutrensis (Franssen et al.
2007). However, this positive association could also
be the result of similar food resources between small-
bodied native fishes and age 1 P. lucius. Indeed, all
three native small-bodied fishes and age 1 P. lucius
feed on invertebrates when available (Sublette et al.
1990). Both native prey CPUE and catch numbers of
age 1 P. lucius were highest in upper reaches of the
suitable habitat or food resources compared with
lower and canyon-bound reaches. If age 1 P. lucius
are reliant on native fishes as prey at this early life
stage, mimicking the natural high spring flow may
help increase numbers of native forage fish available
for age 1 P. lucius. While the amount of discharge
Navajo Reservoir can release each spring cannot
match historical peak flows, high spring discharge
positively correlates with the annual CPUE of native
fishes in the San Juan River (Gido & Propst 2012),
potentially linking increased prey availability for age
1 P. lucius to spring releases. If lack of native prey
availability is contributing to low recruitment of
P. lucius, flow management may be able to facilitate
increased prey availability for age 1 P. lucius.
Similar to age 1 P. lucius, age 2+ P. lucius indi-
viduals showed positive associations with total prey
at the reach scale. This suggests that the different size
classes of P. lucius are potentially tracking prey
resources, a pattern similar to P. lucius in the Upper

7
Franssen & Durst
Colorado River, UT and CO, where adults often
reside in reaches with higher prey numbers (Osmund-
son et al. 1998). Larger P. lucius individuals are
likely not as gape limited for non-native prey com-
pared with age 1 individuals (Franssen et al. 2007),
which may allow both non-native and native prey to
be a vital food source for non-gape-limited age 2+
individuals. Similar to high spring flows increasing
native fish numbers, extended summer low flows
often increase the numbers of non-native small-bod-
ied fishes (Gido & Propst 2012), but summer flows
are maintained at higher than historical levels for irri-
gation. While management of Navajo Reservoir to
reduce summer flows would more closely mimic
pre-dam conditions, this action would be inconsistent
with management efforts seeking to reduce popula-
tions of non-native fishes in the San Juan River.
Quantifying the potential contribution of non-native
prey to piscivorous P. lucius versus their detriment to
other life stages of P. lucius or other native fishes
may lend insight into this potential management
action.
Age 2+ P. lucius individuals were also positively
associated with juvenile I. punctatus. Because both
of these fishes tend to co-occur in the same reaches,
there is the potential for negative interactions
between the two species. Predation on P. lucius by
juvenile I. punctatus is likely rare due to the limited
fish prey they ingest (Tyus & Nikirk 1990), but they
may have other deleterious impacts through competi-
tion for invertebrate prey or simply space, especially
when resources are limiting (Tyus & Nikirk 1990;
Tyus & Saunders 2000). Moreover, P. lucius have
been observed in the field with I. punctatus lodged in
their throats by the pectoral and dorsal spines of
I. punctatus (Ryden & Smith 2002), indicating con-
sumption of juvenile I. punctatus by P. lucius may
be a source of mortality. However, controlled experi-
ments have suggested that this phenomenon may not
be common (Pimental et al. 1985). We suggest that
the positive association between age 2+ P. lucius and
juvenile I. punctatus is likely linked to similar habitat
or prey preferences between the species. Although
documented effects of interactions between juvenile
I. punctatus and age 2+ P. lucius are limited, they
may be partitioning habitats within reaches, which
would reduce any potential negative interactions.
Assessment of habitat use by both I. punctatus and
P. lucius at a finer spatial scale would likely lend
insight into their habitat preferences and elucidate
potential interactions between the species.
Contrary to our prediction, the catch numbers of
age 1 and age 2+ P. lucius were not associated
with the CPUE of predators (i.e., adult I. punctatus
>300 mm TL). Ictalurus punctatus individuals
become increasingly piscivorous when they reach
8
300 mm TL, which suggests that they could poten-
tially prey on age 1 and age 2 P. lucius. While
we found no negative effects of adult I. punctatus
on the catch numbers of P. lucius, we cannot rule
out negative interactions between the two species.
We only investigated patterns of distribution of
both species in the fall during one snapshot in
time; therefore, other factors could potentially drive
distributions during other times of the year.
Indeed, the mobile nature of P. lucius (Osmundson
et al. 1998; Fresques et al. 2013) and I. punctatus
(Wendel & Kelsch 1999) likely provides the
potential for more spatial and temporal interactions
between these species. In attempts to reduce poten-
tial negative interactions between I. punctatus and
native fishes, intensive mechanical non-native fish
removal has been implemented in the San Juan
River since 2001. While it is difficult, direct
assessment of predatory or competitive interactions
between I. punctatus and native fishes will be nec-
essary before the impacts of I. punctatus on the
population persistence of P. lucius can be evalu-
ated. However, our analysis does not indicate that
the CPUE of adult I. punctatus limits the numbers
of age 1 and age 2 P. lucius at the scale we
examined.
As predicted, our results suggested the spatial
distribution of P. lucius was linked to potential
prey, but they did not support our prediction of
negative relationships between the catch numbers
of P. lucius and CPUE of juvenile and adult
I. punctatus. Although correlative, the association
between age 2+ P. lucius and juvenile I. punctatus
was positive, indicating that I. punctatus were not
reducing the numbers of P. lucius collected at the
reach scale we investigated. Current efforts to
mechanically remove I. punctatus from the San
Juan River assume negative interactions between
I. punctatus and P. lucius (and other native fishes),
but specific interactions between these species are
unknown. Identifying and understanding the biotic
and abiotic mechanisms that are limiting survival
of all life stages of P. lucius in the San Juan
River and the Colorado River basin as a whole is
essential for their recovery. Still, complexity of
interactions between native and non-native species
and their environment in large, open systems such
as the San Juan River make this task difficult.
Acknowledgements
We thank the staff from the U.S. Fish and Wildlife Service,
New Mexico Fish and Wildlife Conservation Office. This pro-
ject was only possible with the assistance and dedication of
numerous individuals from U.S. Fish and Wildlife Service,
Colorado River Project office; New Mexico Department of

Game and Fish, Conservation Services Division. Funding for
this work was provided through authorising legislation for the
San Juan River Basin Recovery Implementation Program and
administered by U.S. Bureau of Reclamation, Salt Lake City,
Utah. This article reflects the views of the authors and do not
necessarily reflect the view of the U.S. Fish and Wildlife
Service. We thank E. Gilbert and D. Ryden for graciously
providing data, and thoughtful comments by D. Propst and K.
Gido greatly improved the manuscript.
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