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Western Kentucky UniversityTopSCHOLAR®
Masters Theses & Specialist Projects Graduate School
5-1-2008
Longitudinal Variation in Fish Community Alongthe Upper Green River Below the Green RiverLakeJoshua WilseyWestern Kentucky University
Follow this and additional works at: http://digitalcommons.wku.edu/theses
Part of the Medical Sciences Commons
This Thesis is brought to you for free and open access by TopSCHOLAR®. It has been accepted for inclusion in Masters Theses & Specialist Projects byan authorized administrator of TopSCHOLAR®. For more information, please contact [email protected].
Recommended CitationWilsey, Joshua, "Longitudinal Variation in Fish Community Along the Upper Green River Below the Green River Lake" (2008).Masters Theses & Specialist Projects. Paper 387.http://digitalcommons.wku.edu/theses/387
LONGITUDINAL VARIATION IN FISH COMMUNITY ALONG THE UPPERGREEN RIVER BELOW THE GREEN RIVER LAKE.
A Thesis Presented toThe Faculty of the Department of Biology
Western Kentucky UniversityBowling Green, Kentucky
In Partial FulfillmentOf the Requirements for the Degree
Master of Biology
ByJoshua Delmar Wilsey
May 2008
LONGITUDINAL VARIATION IN FISH COMMUNITY ALONG THE UPPERGREEN RIVER BELOW THE GREEN RIVER LAKE.
Date Recommended
Director of Thesis
)ean, Graduate Studies and Research Date
Table of Contents
1) Abstract2) Introduction3) Study Area4) Habitat Characterization5) Temperature6) Fish Sampling7) Analytical Methods8) Results9) Discussion10) Appendix11) Literature Cited
pg. ii-iiipg. 3-5pg. 5-6pg. 6-7Pg-7Pg-7pg. 7-9pg. 9-13pg. 13-18pg. 19-39pg. 40-43
LONGITUDINAL VARIATION IN FISH COMMUNITY ALONG THEUPPER GREEN RIVER BELOW THE GREEN RIVER LAKE.
Joshua D. Wilsey May 2008 Pages 43
Directed By: Scott Grubbs, Philip Lienesch and Albert MeierDepartment of Biology Western KentuckyUniversity
The impoundment of flowing waters can cause decreased variation in flow and
temperature, reduce habitat availability, and decreased mean temperature below a
reservoir. These types of alterations can influence fish communities. Twelve sites were
sampled along the Upper Green river in order to address the potential influence of
impoundment on fish communities below the Green River Lake. The habitat features:
mean depth, flow, dominant substrate, embeddedness, percent riffle, run and pool were
measured. Temperature probes were placed at each site in order to measure temperature
over an extended period of time. Backpack electrofishing and seining was used to obtain
fishes. The lowest richness and Shannon diversity were from the two most upstream
sites. Principal components analysis of the wadeable fish data suggests possible
differences in species composition at the two most upstream sites when compared to the
other downstream sites. Mean daily temperature was significantly different between
sites (p = 0.05). Significant differences in seasonal daily means, minimums and
maximum temperatures suggests that there are three thermal sections along the Green
River within the study area: (1) a cooler section influenced by release patterns from the
Green River Dam, (2) a warmer section downstream of the Green River Dam receiving
surface inputs from the non-karst tributaries, and (3) the cooler section flowing through
the well-developed karst region. Linear regressions only showed significant relationships
n
between Etheostoma helium abundances and mean daily temperature (p = 0.002),
between Campostoma oligolepis abundances and mean daily temperature (p = 0.043) and
between Etheostoma zonale and % gravel. Finding only four significant linear
relationship between species abundances and environmental variable suggest other
environmental variable not measured may be important for determining species
composition in the Green River.
in
Introduction
The damming of streams is an anthropogenic disturbance is considered one of
major causes for the imperilment of native species in the southern United States because
of frequency of its occurrence across a wide range of stream sizes (Warren et. al. 2000).
Not only are impoundments widespread geographically but they influence a wide range
of communities including: macroinvertebrate communities (Vinson 2001; Lehmkuhl
1972), riparian communities within flood plains (Richter and Richter 2000) and fish
communities (Quist et. al. 2005; Eberle et. al. 2002). Dams have the potential to impact
streams by altering natural flow regime, habitat availability, and in-stream thermal
patterns (Allan 1995). Flow alteration is one of the most common and obvious changes
caused by impoundment. With the exception of hydroelectric dams, typically
impoundment causes decreases in the annual variation of water levels downstream
(Baxter 1977). The presence of a dam can lessen the impact of seasonal variation in water
level from increased water flow due to heavy rain events and decreased water levels that
occur during periods of drought. This buffering of hydrologic variability then leads to
decreased frequency of extreme high and low water events (Richter et al. 1996). These
kinds of changes to flow regime often negatively impact fish communities (Powers et al.
1988; Osmundson et al. 2002; Franssen et al. 2006; Freeman & Marcinek 2006).
Hydrologic changes below an impoundment can alter a stream's morphology. The
natural variation between high and low water events are important to the maintenance of
habitat features within a stream, including riffles, sand bars, and floodplain riparian zones
(Richter and Richter 2000). As water level increases so does its ability to pick up
sediments, organic material and other particles. Periodic high water events are often
necessary to prevent coarser substrates from being covered over by finer substrates. High
flow events causes a stream to transport these materials from further upstream and
deposit them in sand bars, stream banks and slack water areas once water levels recede
(Osmundson et al. 2002). Impoundments routinely disrupt this process by altering natural
flow regimes and by acting as a sink trapping finer sediments being transported from
upstream. This deposition is often necessary for the maintenance of those features
composed of finer substrates. Without a source of finer substrates from upstream these
features can erode away, leaving behind only courser substrates. When habitat structures
like sand bars, and riffles are lost it causes a loss of in-stream habitat heterogeneity. This
loss of habitat can influence fish communities (Schlosser 1982; Pearsons et al. 1992;
Bunn & Arthington 2002; Koel 2004), especially for those species that specialize in
utilizing these types of habitat.
Dams may alter downstream temperature regimes by decreasing thermal variation
both daily and seasonally (Jaske & Goebel 1967; Baxter 1977). Thermal alteration is
largely due to the decreased surface area to volume of reservoirs and thermal
stratification that typically decrease the influence of fluctuations in air temperature on
water temperature. In addition, if water is being released from the hypolimnion it is often
much cooler than the surface water of streams. These kinds of alterations in thermal
pattern can influence the distribution and diversity of fishes by causing reduced
abundances and absences of those species sensitive to this type of alteration (Baxter
1977; Paller& Saul 1996).
Many studies have previously studied from a temporal perspective the
downstream effects of impoundment on fish communities (Tieman & Gillette 2004;
Taylor et. al. 2001; Martinez et. al. 1994). Few studies have addressed the influence that a
disturbance gradient created by damming can have on the longitudinal distribution fish
communities. The purpose of this research was to address the influence of impoundment
on the longitudinal distribution of a riverine fish community in central Kentucky, U.S.A.
The key environmental variables that were the focus of this study were thermal
variability and habitat size. They were addressed in detail at several positions
downstream from the reservoir, by determining if the furthest upstream sites possessed
the lowest mean daily water temperatures, the least variation in water temperature, and
the least percent composition of fine substrates than sites further downstream, in order to
assess the possible disturbance gradient created by the reservoir. These key
environmental variables were compared to the fish communities at each site in order to
find if the changes in the environmental variables being caused by dam releases resulted
in decreases or absences of certain fish species or even changes to entire guilds of fishes.
These types of changes in the fish community would indicate that the presence of the
Green River Lake was impacting fish communities below it.
Study Area
All field work was conducted along the mainstem of the Green River in central
Kentucky, U.S.A. between the outflow of the Green River Lake and Mammoth Cave
National Park (Fig. 1). The Green River is a 6th-7lh order stream within the study area.
The Green River originates in Casey and Lincoln Counties and flows nearly 600 km west
to its confluence with the Ohio River in Henderson County.
Within the study area the Green River flows through three Level-IV Interior
Plateau Ecoregions (Woods et al. 2002). In longitudinal order, the Green River flows
through the Eastern Highland Rim, Western Pennroyal Karst Plain, and Crawford-
Mammoth Cave Uplands Ecoregions. The Eastern Highland Rim is underlain by
Mississippian-age sandstones. In contrast, the Western Pennyroyal Karst Plain is
characterized by Mississippian-age limestone with extensive networks of underground
fluvial systems. Surface stream density is very low. The Crawford-Mammoth Cave
Uplands is geologically characterized by Mississippian-age sandstone in upland
environments and Mississippian limestone in karst valleys.
Habitat Characterization
Twelve sites were sampled along the Green River within the study area (Fig. 1).
Habitat variables were quantified during summer baseflow conditions. Sampling reaches
at each site were defined as 15 times the wetted width of the stream or 1000 m,
whichever was smaller, and included both wadable and non-wadable portions. The
percent composition of riffle, run, and pool habitat features of the entire sampling reach
were visually estimated. Within the wadable section five transects were placed
perpendicular to flow and spaced every 30-40 m. Along each transect the dominant
substrate was estimated visually. Depth was measured to the nearest 0.5 cm and velocity
was quantified with a Marsh-McBirney Flo-Mate Model 2000 Portable Flowmeter.
Within the non-wadable portions of the sampling reach depth was measured at an
additional five transects spaced every 30-40 m. Large woody debris, root wads, undercut
banks and other habitat features along both sets of transects were noted. Percent
composition of each substrate class sampled, mean depth, mean velocity, and mean
discharge were calculated from the transect data collected. The percent composition of
riffle, run, and pool habitat features within each sampling reach were also visually
estimated.
Temperature
HOBO® Water Temp Pro v2 Logger temperature probes were used to measure
temperature over an extended period of time at each sampling site. The probes were
placed in-stream in May 2005 and removed in August 2006. Probes were redeployed in
August 2006 and retrieved in September 2007, yet because data could not be obtained
from a subset of three probes all thermal information for this time period was excluded
from analyses. Temperature was recorded every hour. Mean daily temperature, minimum,
maximum, standard deviation, and coefficient of variation were calculated per site for
each month, season, and the entire 15-mo period.
Fish Sampling
Fish were collected at each site during May - August 2007. Within wadable
sections of each sampling reach fish were sampled using 30-60 minutes of seining and 15
minutes of backpack electrofishing. Common and easily identifiable fish were identified
in the field. All remaining fish collected were fixed in 10% buffered formalin and taken
back to the lab for identification.
Analytical Methods
Habitat and temperature data were combined in one environmental matrix. The
mean daily, minimum, and maximum temperatures were compared between sites using
the analysis of variance (ANOVA). The mean daily, minimum and maximum
temperatures for each site were further partitioned by season and ANOVAs were
8
performed to explore seasonal trends in the data. Tukey's honestly significant difference
(Tukey's HSD) test was performed on all ANOVAs to detect statistical pair-wise site
differences. Bonferroni corrections were applied to minimize type I error. Principle
components analysis (PCA) was performed on the environmental data to explore
longitudinal relationships along the Green River.
Species richness, evenness and Shannon-Weaver diversity indices were calculated
from the fish data set. Rank abundance curves were plotted to explore changes in
abundance that occurs longitudinally along the Green River. In order to assess fish
community structure each species was assigned to guilds within four categories (trophic
guild, substrate preference, stream flow velocity preference, and reproductive guild;
Table 1). Life history information from multiple sources was used to assign species to
guilds (Frose & Pauly 2008; Natureserve 2008; Pflieger 1997; Etnier & Starnes 1993).
The fish data were ratified by removing those species that occurred only at one
site and were logio (x+1) transformed prior to analyses, were x = species abundance per
site. Principle components analysis was performed to explore longitudinal relationships
in species composition. Simple linear regressions were conducted to assess relationships
between species abundance data and environmental variables. Only those species and
variables that had strong PCA loadings were used. An objective mechanism was not
determined apriori when interpreting which variables possessed strong loadings. The
same loading values were not used between PCA axes, instead the first natural block of
the most extreme values positive or negative were selected. The number of those
variables selected was entirely based on the relative strength of their loading. Simple
linear regressions were also performed between species richness, Shannon-Weaver
diversity and fish guild data and environmental variable data. Bonferroni corrections
were applied to minimize Type I error. Indicator species analysis was performed on the
fish data sets in order to identify those taxa that were indicative of distinct groups among
the sites sampled. Ordination, calculation of indices and the indicator species analysis
were performed with PC-ORD Version 5 for Windows (MJM Software Design 1999)
while SPSS 12.0 (SPSS Inc. 2003) was used for the ANOVA's and linear regressions.
Results
Although wetted width varied among the upstream sites due to difference in
channel morphology, the downstream sites were markedly larger (Table 2). Mean depth
(50-101 cm) and discharge (1.0-5.8 m3/s) varied predictably, increasing in a downstream
direction (Table 2). Gravel was the primary substrate at most sites. With one exception
the two most upstream sites, GR 8.14 and GR 8.15, possessed the lowest percentage of
fine substrates.
Mean daily temperatures varied less between sites (16.3 - 17.6°C) than both
minimum (0.2 - 5.4°C) and maximum (28.4 - 32.4°C) temperatures. The lowest mean
temperatures occurred at the two most upstream sites, GR 8.14 and GR 8.15. Conversely,
these two sites also possessed the highest minimum daily temperatures and except for the
two most downstream sites (GR 8.0D and GR 8.1) exhibited the least variability (Figure
2; Table 2).
Over the whole 15-mo period, mean daily (Fdf=n = 1.79, p = 0.05) and daily
maximum (Fdf n = 2.09, p = 0.02) temperatures were significantly different between
sites, but daily minimums were not significant (Fdf=n = 1.40, p = 0.17). There were a
consistent series of within-season differences. Mean daily (Fdf=n = 6.99, p < 0.0005),
10
daily minimum (Fdf=n = 8.45, p < 0.0005) and maximum temperatures (Fdf=n = 5.43, p <
0.0005) were significantly different between sites for spring. Within summer, mean daily
(FdfHi = 44.89, p < 0.0005), daily minimum (Fdf=n = 40.105, p < 0.0005) and maximum
temperature (Fdf=i i = 43.92, p < 0.0005) were significantly different between sites.
Similarly, autumn mean daily (Fdf=n = 2.77, p = 0.001), daily minimums (Fdf=n =3.19,
p < 0.0005), and daily maximum temperatures were significantly different between sites
(Fdf=ii = 2.54, p = 0.004). Means for daily temperature (Fdf=n = 4.79, p < 0.0005), daily
minimum (F = 5.05, p < 0.0005) and daily maximum temperatures (Fdf=n = 4.72, p <
0.0005) were also significant during winter days (Table 3).
The first three PC A axes explained 74.3% of the variation of the environmental
data (Table 4). Strong positive scores were exhibited for sites GR 8.0D, GR 8.1, and GR
8.2 along PCA Axis 1. Strong negative scores were observed for GR 8.9 and GR 8.10.
GR 8.3 and GR 8.15 possessed strong positive scores along PCA Axis 2. Along PCA
Axis 3 there were strong positive scores for GR 8.14 and GR 8.15 and a strong negative
score for GR 8.3 (Table 4; Figure 3). Mean wetted width, percent pool, and both standard
deviation and coefficient of variation of the temperature data were strong contributors to
PCA Axis 1. Percent cobble, percent small boulder, and mean daily temperature
produced high loadings along PCA Axis 2 while percent gravel was the sole variable
important to PCA Axis 3.
There were 61 species total collected across the 12 sites. The four most common
species sampled were Etheostoma zonale, Pimephales notatus, Lepomis megalotis, and
Campostoma oligolepis, comprising 46.6 % of the total number of individuals collected.
Several species were obtained from all 12 sites, including C. oligolepis, E. blenniodes, E.
11
caeruleum, E. zonale, L. megalotis and P. notalus. Cottus carolinae was collected
mainly from the downstream sites. C.oligolepis, E. helium, E. zonale, L. megalotis and P.
notatus were most common in the middle sites. Pomoxis annularis was obtained only
from the two most upstream sites (GR 8.14 and GR 8.15) (Figure 4; Table 5).
Species richness ranged from 14 to 27 species per site. Shannon-Weaver diversity
ranged from 2.19 to 2.69, while evenness showed no obvious pattern between sites and
was between 0.95 and 0.96 (Table 6). The lowest richness and Shannon diversity were
from the two most upstream sites, GR 8.14 and GR 8.15. Benthic insectivores comprised
the largest trophic guild at most sites. Prominent species within the benthic insectivore
guild ordered from most abundant to least include: Etheostoma zonale, Cottus carolinea,
E. blennoides, and E. helium. Generalist insectivores composed the second largest trophic
guild (Figure 5). Lepomis megalotis and Notropis photogenis were two important species
within this group. Those species that had a preference for gravel substrates made up the
highest percentage of the fish community at all sites except GR 8.15. C. carolinae, N.
atherinoides, E. rafinesquei, and E. helium composed most of the individuals collected
that possessed a preference for gravel substrates. The next largest group was those
species preferring cobble substrates, followed closely by substrate generalists (Figure 6).
E. zonale, C. oligolepis, N. photogenis, and E. blenniodes were important cobble species
and Pimephales notatus and L. megalotis were the most abundant substrate generalists.
GR 8.15 possessed a greater proportion of silt and substrate generalist guilds than all
other sites. The presence of Pomoxis annularis and Lepomis macrochirus were primarily
responsible for the greater percentage of silt species. Percent composition within velocity
guilds revealed an even distribution between fast, moderate, slow, and no flow
12
preferences. Only a small percentage of species were flow velocity generalists (Figure 7).
GR 8.15 possessed greater percent composition of slow and generalist stream flow guilds
than other sites. Lepomis megalotis, Fundulus caienatus, and Gambusia affinis were
prominent species within the slow/no flow guild and Pimephales notatus the most
abundant species within the flow generalist guild. The two upstream sites, GR 8.14 and
GR 8.15, possessed higher percentages of nest spawners and substrate choosers than were
present at most other sites and fewer open water/substrate spawners (Figure 8). Lepomis
megalotis, and P. notatus were the most common nest spawner species collected. Within
the substrate chooser guild, E. zonale and E. blenniodes were important species.
Important species within the open water/substrate spawner group included: C. oligolepis,
Cottus carolinea, and N. photogenis.
The PC A of the fish data showed 60.9% of the variation was explained along the
first three axes (Table 4). Strong negative loadings were exhibited for sites GR 8.3, GR
8.14 and GR 8.15 along PCA Axis 1. Along PCA Axis 2, Sites GR 8.14 and GR 8.15
possessed strong positive loadings and GR 8.3 possessed a strong negative loading. A
plot of axes 1 and 2 isolated GR 8.3, GR 8.14 and GR 8.15 for the other sites sampled
(Figure 9). Campostoma oligolepis, Notropis atherinoides, N. photogenis, Phenacobius
uranops, Fundulus notatus, E. bellum, E. rafinesquei were strong contributors to PCA
axis 1. Notropis volucellus, Pylodictis olivaris, Gambusia affinis, Labdidesthes sicculus,
E. caeruleum, E. zonale produced strong loadings along PCA Axis 2. Increased
abundances of E. rafinesquei, G. affinis, and L.sicculus were important in determining
where GR 8.3 was located along PCA axes 1 and 2. Decreased abundances of E. bellum,
E. zonale and C. oligolepis were important in determining where sites GR 8.14 and
GR8.15 were located along PCA axes 1 and 2.
Linear regressions revealed three significant relationships between the abundance
of individual fish species and environmental variables. Both Etheostoma helium (b =
0.02, r2 = 0.62, p = 0.002) and Campostoma oligolepis (b = 0.01, r2 = 0.35, p = 0.04)
abundances were linearly related to mean daily temperature. The only remaining
significant linear relationship was between C. oligolepis and stream velocity (b = 4.11, r2
= 0.53, p = 0.007). Mean daily temperature was able to significantly explain values of
both species richness (b = 6.57, r2 = 0.54, p = 0.007) and Shannon-Weaver diversity (b =
0.35, r2 = 0.57, p = 0.005). Pertaining to guild data, there were significant positive
relationships between substrate choosers and percent riffle (b = 0.41, r2 = 0.43, p = 0.02)
and between nest spawners and percent cobble (b = 0.30, r = 0.38, p = 0.03). The
indicator species analysis revealed that the two upstream sites, GR 8.14 and GR 8.15,
were isolated from the remaining 10 sites (Figure 10). Etheostoma helium (p = 0.015) and
E. zonale (p = 0.015) were significant indicators of the ten more downstream sites while
P. annularis was the sole significant indicator of the two upstream sites (p = 0.015).
Discussion
This decrease in fine substrates and increased dominance of coarser substrates
frequently occurs directly below a dam outflow. The two most upstream sites, GR 8.14
and GR 8.15, possessed lower percentages of fine substrates than the other sites sampled
and as a result possessed greater percentages of coarser substrates. This results because
dams interrupt the transport of these fine substrates from further upstream and the
13
14
sediment-free waters leaving the dam outflow tend to pick up fine substrates mainly in a
downstream manner (Allan 1995).
Results from the 2005-2006 temperature data analyses at first glance appear to be
somewhat mixed. The Tukey's HSD revealed patterns in seasonal daily means,
minimums and maximums at GR 8.14 and GR 8.15 that were significantly different from
most of the other sites. The mean temperatures and maximums were significantly lower,
and minimums were significantly higher, indicating that the Green River Lake outflow
altered the thermal regime at these sites. This type of thermal pattern is common in the
tailwaters of streams, especially those with hypolimnetic release patterns (Jaske &
Goebel 1967; Baxter 1977). Thermal similarities between these two upstream sites and
the two most downstream sites (GR 8.0D and GR 8.1), however, appear to somewhat
contradict this conclusion unless the base geology of this study region is considered.
These two downstream sites are located in Crawford-Mammoth Cave Uplands Level-IV
Ecoregion, a well-developed karst region of Mississippian-age limestone. Surface stream
density is very low is this region. The thermal regime of the Green River flowing through
this region is heavily influenced by groundwater inputs. O'Driscoll and DeWalle (2006)
similarly found that stream sites that received significant groundwater inputs were less
responsive to changes in air temperature, often resulting in cooler water temperatures in
the summer and warmer temperatures in the winter. This type of a pattern is similar to
that of streams being influenced thermally by hypolimnetic release from reservoirs. This
suggests that there are three thermal sections along the Green River within the study area:
(1) a cooler section influenced by release patterns from the Green River Dam, (2) a
warmer section downstream of the Green River Dam receiving surface inputs from the
15
non-karst tributaries, and (3) the cooler section flowing through the well-developed karst
region.
The differences in species richness, Shannon-Weaver diversity, guild
composition, the PC A ordination of species abundances, and indicator species analysis all
indicate that the two upstream sites, GR 8.14 and GR 8.15, possess fish communities
distinct from each of the remaining study sites. Mean daily temperature possessed a
significant positive relationship to both richness and Shannon-Weaver diversity. Richness
and Shannon-Weaver diversity were lowest at GR 8.14 and GR 8.15. Of the species that
loaded strongly on PC A Axis 1, both Campostoma oligolepis and Etheostoma helium
abundances were significantly related in a positive direction to mean daily temperatures.
This makes it very possible that at least part of the variation explained along PC A Axis 1
was related to mean daily temperature. Decreases in diversity and changes to community
composition often occur within the tailwaters of a reservoir after the establishment of the
impoundment (Edwards 1978; Quinn & Kwak 2003).
Campostoma. oligolepis, one of the species possessing significant linear
relationships with environmental variables, is a stoneroller species found in drainages
within the Midwestern and southeastern United States (NatureServe 2008). They occur in
swift flowing rocky riffles within creeks, and small to medium streams (Fowler & Taber
1985). They graze gravel and rock substrates comsuming periphyton present on these
substrates (Fowler & Taber 1985). Lower mean daily temperatures could be negatively
influencing C. oligolepis abundances indirectly by limiting periphyton growth.
Rosemond (1994) found temperature to be a factor that positively influenced periphyton
biomass and abundance.
16
Etheostoma helium the other species possessing a significant linear relationship to
mean daily temperature, is endemic to the Green and Barren River Basins and inhabits
and spawns in fast-flowing reaches with coarse substrates (Fisher 1990; Page and Burr
1992). This species was absent only from the two upstream reaches, GR 8.14 and GR
8.15. Etheostoma bellum spawns between April and June when water temperatures are
between 20-25 °C. Although the upstream reaches possess the highest proportions of
coarse substrates, thermal release patterns from the Green River Lake produced stream
temperatures that are arguably too cool for effective spawning of this species. During
2006 the daily mean April - June temperature of GR 8.14 and GR 8.15 possessed only 37
and 25 days, respectively where mean daily temperature was between 20-25 °C, while the
remaining sites possessed between 51 and 63 days.
The significantly positive relationship between both E. bellum and C. oligolepis
and mean daily temperature both indicate that temperature is one factor influencing fish
communities in the Green River. This influence becomes clearer when the changes in
these two species are considered along with the results from the ANOVAs of the
temperature data that suggest three thermal regions. Abundances in both E. bellum and
C. oligolepis are highest in the middle sites where mean temperatures are warmer and
their abundances are lowest in both the most upstream two sites (GR 8.14, GR 8.15) and
the two most downstream sites (GR 8.0D, GR 8.1) where mean water temperatures are
the coolest. Why these two species possess lower abundances in sites with cooler mean
temperatures is unclear. The decreased abundances of these two species could be the
result of temperature influencing the larval development of these species. Darters have
been shown to possess an inverse-log relationship between temperature and incubation
17
time in days (Page 1983). Temperature has been shown to influence the larval growth
rates of other fish species (Claramunt & Wahl 2000; Lasker 1964). The influence of
temperature on growth rates is well known in marine and estuarine fishes (Klimogianni
et. al. 2004; Morehead & Hart 2003; McGurk 1984; Laurence & Rogers 1976). The
growth rate of larval fishes often determines the time of first feeding in these fish, which
is an important factor in determining their ability to avoid starvation (Rogers & Westin
1981; Laurence 1978; Houde 1974).
Other environmental parameters (e.g., substrate heterogeneity and hydrologic
variability) not specifically measured in this study may also be important factors
influencing fish community composition along the Green River. Several studies have
shown that hydrologic patterns can influence fish community structure (Poff & Allan
1997; Bain et al. 1988). Poff & Allan (1995) found that those sites that are hydrologically
variable have a greater composition of fishes that possess generalized feeding strategies,
are silt-tolerant, are slow stream velocity species and are associated with silt or are
substrate generalists. The greater percent composition of both species preferring silt and
substrate generalists and the greater percent composition of slow velocity preference
guilds at the most upstream site (GR 8.15) could suggest fishes at this site are
experiencing greater flow variability instead of more stable hydrologic conditions as
would be expected. However, direct measurement of hydrologic variability is necessary
to quantify such a relationship.
In conclusion this study was able to detect differences in key environmental
variables between upstream sites and downstream sites. Lower percent composition of
fine substrates, decreased mean daily temperatures, and minimum temperatures both
18
throughout the entire sampling period and within particular seasons within the upstream
sites were important changes found to be occurring as a result of the presence of the
Green River Lake. Being able to explain species richness, Shannon-Weaver diversity and
abundances of E. bellum and C. oligolepis, with mean daily temperature indicates that
releases from the Green River Lake are influencing the fish communities found below it
by causing decreased mean daily temperatures in the upstream sites.
\
Big Brush CreekLynn Camp Creek f Big Pitman Cieek
Mammoth CaveNational Park
Legend
O Sampling_SitesGreen River and Tributaries
Mammoth Cave National Park
Green River Lake
Upper Green River Subbasin
DDCD3Q.
5'
0 25 5 10 15 20
Figure 1. Map of study area along the upper Green River within central Kentucky, U.S.A.
20
Table 1. The four categories andthe guilds fish species wereassigned to within those categories.
ID Description
Trophic Guild123
45678
Hebivore-detritivoreOmnivoreGeneral InsectivoreSurface/water columninsectivoreBenthic InvertivorePiscivorePlanktivoreParasite
Substrate12345
CobbleGravelSandSiltGeneral
Current Velocity Preference1234
FastModerateSlow-NoneGeneral.
Reproductive Guild
123456
Open water/ substratumspawnerBrood HiderSubstratum ChooserNest SpawnerExternal BrooderInternal Bearer
Table 2. Environmental parameters sampled in 2007 from twelve sampling reaches along the Upper Green River. Samplingreaches arranged in order from downstream to upstream.
Par am
HabitatWidth
DepthVeloc
Disch
%Riff%Run%Pool%Fine
%Gr
%Cb
%SmB
Dist
Description
mean wetted width(m)mean depth (cm)mean velocity(m/s)mean totaldischarge (m3/s)% riffle habitat% run habitat% pool habitat% substrate as silt,sand and clay% substrate asgravel% substrate ascobble% substrate assmall boulderdistance from dam(km)
GR8.0D
47.8
930.20
5.8
35452038
62
0
0
155
GR8.1
47.5
760.35
5.0
20552520
80
0
0
130
GR8.2
45.3
1010.23
3.5
15701518
78
4
0
124
GR8.3
33.4
870.21
2.7
503020
6
28
30
36
96
GR8.7
34.5
690.16
1.8
10603018
64
6
12
64
GR8.9
29.9
670.33
1.7
204040
0
66
28
6
47
GR8.10
23.1
680.26
1.6
25403526
74
0
0
47
GR8.11
26.8
540.27
1.9
206020
8
72
20
0
43
GR8.12
36.3
500.25
2.0
30403034
66
0
0
39
GR8.13
34.9
600.19
1.7
30403034
62
0
4
24
GR8.14
32.3
500.20
1.0
354025
2
92
6
0
4
GR8.15
28.8
690.17
1.1
354025
2
74
22
2
2
to
Table 2. Continued
Par am Description GR GR GR GR GR GR GR GR GR GR GR GR8.0D 8.1 8.2 8.3 8.7 8.9 8.10 8.11 8.12 8.13 8.14 8.15
Temp.Min
Max
Mean
StDev
CV
minimum meandaily temperaturemaximum meandaily temperaturemean dailytemperaturestandard deviationof dailytemperaturescoefficient ofvariation of dailytemperatures
2.0
28.4
16.9
6.9
0.41
2.6 1.7 1.2 1.3 1.3 1.6 1.6 1.8 3.0 5.3 5.8
30.2 28.9 30.3 31.1 31.7 31.7 31.1 32.1 31.8 31.6 32.4
16.8 16.9 17.0 17.0 17.6 17.6 17.3 17.7 17.5 16.5 16.3
6.8 7.1 7.5 7.2 8.1 8.1 7.7 8.2 8.0 6.9 6.7
0.40 0.42 0.44 0.42 0.46 0.46 0.45 0.46 0.46 0.42 0.41
K)
23
OO
o
00
O
oc
O
Qo
o
o
oo
o
00
o
00
O
in
oo
a
00
a
en• i
00
O
00
o
Temperature °C
i/"i O ^"i O */"J O */"» Or*"t r*"( (N CNI •—« - H
Temperature °C
>/"i O V") O ^ J O «O Or^ f^i rs r-J -—i i—'
Temperature °C
V%
I P
\
V\
\.
Dat
e
1)
Q
•utaQ
ju13Q
Oo
c/33Of
1
oo
1co
o
0>
-pC3O
"3u
ratu
r
1
ean
d
•
ex
24
GR8.9A CV-
StDev
GR8.10A
•"""" GRS
°oPool y
A MeanGR8.12
A GR8.3
%SmB
%CB \ i o R l f f
13 /iir / \
^ / ^Veloc
A GR8.15
A G R 8 J > ^
( \
°o Fuie
^ °oGR
s Depth
GR8
S^Disch" Width
%Ruii
.OD •
AGR8.2
A
GR8.1
Axis 1
Figure 3. Plot of PCA Axes 1 and 2 of environmental data set
Table 3. Tukey's HSD results for mean daily, mean daily minimum, and mean daily maximum temperatures within each season.
SpringMeanTukey HSDDaily MinimumTukey HSDDaily MaximumTukey HSD
SummerMeanTukey HSDDaily MinimumTukey HSDDaily MaximumTukey HSD
GR8.0D
17.94b
17.70b
17.82abc
24.12ab
23.40bed
24.99a
GR8.1
18.07b
17.31b
18.86be
23.78ab
22.87abc
24.72a
GR8.2
17.87b
17.00b
18.80be
24.37be
23.49cd
25.28ab
GR8.3
17.68b
16.69b
18.73be
24.97cd
24.12d
25.83b
GR8.7
17.69b
16.84b
18.56be
24.47be
23.60cd
25.45ab
GR8.9
18.10b
17.07b
19.21c
26.33e
25.29e
27.44cd
GR8.10
17.93b
16.85b
19.09be
26.26e
25.23e
27.34cd
GR8.11
17.82b
16.71b
19.04be
25.27d
23.93d
26.72c
GR8.12
18.05b
16.80b
19.38c
26.57e
25.25e
27.91d
GR8.13
17.50b
16.17b
18.92be
26.16e
24.88e
27.52cd
GR8.14
15.61a
14.40a
17.29ab
23.94ab
22.50a
25.87b
GR8.15
15.25a
14.45a
16.33a
23.52a
22.72ab
24.78a
Table 3. Continued.
FallMeanTukey HSDDaily MinimumTukey HSDDaily MaximumTukey HSD
WinterMeanTukey HSDDaily MinimumTukey HSDDaily MaximumTukey HSD
GR8.0D
12.48a
12.04a
13.00ab
8.06c
7.68b
8.48b
GR8.1
12.47a
11.74a
13.35abc
8.01c
7.56be
8.49b
GR8.2
12.34a
11.69a
13.07abc
7.82be
7.29abc
8.40b
GR8.3
12.21a
11.64a
12.79a
7.63abc
7.02abc
8.31b
GR8.7
13.49ab
12.93ab
13.97abc
7.36abc
6.86abc
7.90be
GR8.9
13.03ab
12.28a
13.77abc
7.29abc
6.70a
7.89be
GR8.10
13.29ab
12.72ab
13.91abc
7.22abc
6.65a
7.80be
GR8.11
13.18ab
12.37ab
14.09abc
7.31abc
6.77ab
7.87be
GR8.12
13.25ab
12.45ab
14.13abc
7.18abc
6.59a
7.77be
GR8.13
13.65ab
12.93ab
14.39abc
7.06ab
6.43a
7.77be
GR8.14
14.97ab
14.26ab
15.74be
6.82a
6.45a
7.35a
GR8.15
15.39b
15.06b
15.90c
6.82a
6.63a
7.10a
O\
27
Table 4. Diversity indicies describing fish sampling data.
Site
GR8.0DGR8.1GR8.2GR8.3GR8.7GR8.9GR8.10GR8.11GR8.12GR8.13GR8.14GR8.15
Richness
182222241924222723221314
Evenness
0.9590.9380.9580.9630.9470.9510.9540.9590.9560.9570.9620.951
Shannon-WeaverDiversity ('H)
2.7732.9
2.963.0622.7893.0212.9473.1622.9992.9582.4692.509
Table 5. Abundances of individual fish species collected in 2007 from the twelve sampling sites along the Upper Green River.
Species
Ambloplites rupestrisCampostoma oligolepisCarassius auratusCottus carolinaeCyprinella spilopteraCyprinella venustaCyprinella whippleiErimystax dissimilisEtheostoma heliumEtheostoma blenniodesEtheostoma caeruleumEtheostoma flabellareEtheostoma maculatumEtheostoma raflnesqueiEtheostoma squamicepsEtheostoma stigmaeumEtheostoma zonaleFundulus catenatusFundulus notatusGambusia affinisHypentelium nigricansLabedesthes sicculusLepomis cyanellusLepomis macrochirus
Abbrev
AmbrupeCam_oligCar_auraCot_caroCypspilCyp_venuCyp_whipEri dissEth_bellEth_blenEth caerEth_flabEth macuEth_rafiEth_squaEthstigEth_zonaFun_cateFun notaGamaffiHyp_nigrLab_siccLepcyanLep macr
GR8.0D
060
17000333100200
2423
100000
GR8.1
543
1103
001414003102
17000
11027
GR8.2
240
17000422402102
331000600
GR8.3
4003007017908
3801
5568
371
2940
GR8.7
110
100000
1222
105201
560001010
GR8.9
06900000
1679101000
17100
208602
GR8.10
25400
13200
1412110015
2327
076000
GR8.11
257031001
13662070
1847
1082600
GR8.12
11009701544311100
211003600
GR8.13
8800001279320008
447001
1201
GR8.14
7300002003200600
100001000
GR8.15
210000000
1910070540200011
oo
Table 5. Continued
Species
Lepomis megalotisLepomis microlophusLuxilus chrysocephahisLythrurus faciolarisLythrurus fumeusMicropterus dolomieuMicropterus punctulatusMicropterus salmiodesMoxostoma sp.Notropis atherinoidesNotropis hoopsNotropis photogenisNotropis rubellusNotropis telescopusNotropis volucellusNoturus elegansNoturus eleutherusPercina caprodesPercina evidesPhenocobius mirabilisPhenocobius uranopsPimephales notatusPomoxis annularisPylodictus olivarus
Abbrev
Lep_megaLepmicrLux_chryLyt_faciLyt_fumeMicdoloMicjpuncMic_salmMox_sp.Not_atheNot_boopNot_photNot_rubeNotjeleNotjvoluNtu_elegNtu_eleuPer_caprPer evidPhe_miraPheuranPim_notaPomannuPyloliv
GR8.0D
80500210
321500000000000900
GR8.1
180012530000
42400000110100
GR8.2
1300101500
1704000000401100
GR8.3
2505000809000009000100
1001
GR8.7
500000009607006020100
2500
GR8.9
2500001318
380
19000001152
9000
GR8.10
350000020069
10000000101
5801
GR8.11
13000020009160
144100041
1700
GR8.12
90000000000
22004110002
2200
GR8.13
4000000200703067001000
8500
GR8.14
501200000000000000000
1510
GR8.15
1310000010000000000000420
too
30
Table 6. Scores of PCA axes 1,2 and 3 of environmental and biologicaldata sets for sites sampled along the Upper Green River
Site
GR 8.0DGR8.1GR8.2GR8.3GR8.7GR8.9GR8.10GR8.11GR8.12GR8.13GR8.14GR8.15
% var.explained
EnvironmentalAxis 1
3.2603.0803.350
-1.0800.530
-3.190-2.420-0.810-2.110-1.5600.3500.610
32.8
Axis 2
0.170-1.440-0.8505.000
-0.270-0.450-1.660-0.600-1.700-0.8100.5002.100
21.0
Axis 3
-2.250-0.220-0.170-1.9200.6000.280
-0.3301.280
-1.380-1.2202.7902.530
16.4
Axis 1
1.3800.4000.3002.5200.670
-4.450-3.430-4.060-1.530-2.1805.2805.100
20.9
BiologicalAxis 2
0.4003.6800.370
-7.8000.5701.920
-0.160-1.1700.310
-1.3201.3902.180
15.6
Axis 3
-0.800-4.860-2.220-2.6700.880
-2.4301.7602.4701.4402.1901.9902.260
12.8
31
Table 7. Loadings for PCA axes 1, 2 and3 of environmental data set for sitessampled along the Upper Green River.
WidthDepthVelocDisch%Riff%Run%Pool%Fine%Gr%Cb%SmBMeanStDevCV
Axis 1
0.7760.622
-0.1480.673
-0.1950.584
-0.7380.3880.198
-0.164-0.346-0.621-0.828-0.895
Axis 2
-0.1850.155
-0.568-0.3510.390
-0.404-0.207-0.523-0.4390.7440.629
-0.648-0.416-0.268
Axis 3
0.3250.4840.0000.5490.447
-0.446-0.1600.379
-0.819-0.2840.3450.3300.2620.204
32
Table 8. Loadings for PCA axes 1, 2 and 3 of biological data set for sites sampledalong the Upper Green River.
Amb_rupeCam_oligCot_caroCyp_spilCyp_whipEri_dissEth_bellEth_blenEth caerEth_flabEth_macuEth_rafiEth_stigEthe zonFun cateFun_notaGam_affiHyp_nigrLab_siccLep_cyanLep macr
Axis 1
-0.3600.880
-0.1080.473
-0.5630.5200.8330.114
-0.1440.601
-0.323-0.7370.2760.2560.398
-0.6660.1990.5650.122
-0.6000.126
Axis 2
-0.0830.2410.276
-0.169-0.4770.303
-0.348-0.201-0.723-0.341-0.282-0.389-0.200-0.673-0.502-0.551-0.6650.115
-0.673-0.2690.479
Axis 3
-0.2610.1250.525
-0.3100.1520.552
-0.114-0.324-0.190-0.5520.547
-0.103-0.4220.1460.2970.2010.4010.3370.1750.3530.457
Lep_megaLux_chryLyt_faciMic_doloMic_puncMic_salmMox_sp.Not_atheNot boopNot_elegNot_eleuNot_photNot_teleNotvoluPer_caprPer evidPhemiraPhe_uranPirn notaPomannuPyloliv
Axis 1
0.397-0.559-0.3780.212
-0.0320.0660.0330.6470.4620.3600.0470.7020.413
-0.0090.4930.0380.5810.6990.582
-0.548-0.101
Axis 2
-0.432-0.4440.5430.401
-0.3830.278
-0.364-0.075-0.236-0.0960.0160.264
-0.282-0.703-0.105-0.0210.090
-0.003-0.4100.408
-0.700
Axis 3
0.2790.3070.1330.5430.7940.0920.4940.253
-0.202-0.422-0.2640.258
-0.438-0.2830.1600.6220.2600.081
-0.277-0.4590.163
m Relative Abundance Relative Abundance Relative Abundance^3* C *t 0 s ! •""• ^3" f^l C**) r~^ ^f C^i C^~i r~~i
o o o o o o o o o o o o o o o
ai'D
rn
ao00Pi
oo
o06
o
oo
Pi
o
ooOiO
LT;
00
O
ooa:a
CN
00
O
om
oo
.Qi
oo
0CN -ig
0
0
0
en
3O
c-OC3X)03
oCN , as
—
Lynn Camp Creek B l * ? I I K h C r " k Big Pitman deek
8.2 ^ J H.9
8.01)
Cave
National Park
Legeuil
Herbivore/Detntivore
Omnivore
| Generalist Insectivore
9 SurfaceMfcter Column Insectivore
U BenthicInsectivore
| Rsovore
| Parasite
Green River and Tnbutanes
Mammoth Cave National Park
Green River Lake
Upper Green River Subbasm0 45 9
IWIometers18
Figure 5. Map depicting pie charts showing percent composition of trophic guilds communities at each site.
-u
Lynn Camp Ci eek BigBrush C reek B j g Famm Cnfk
8.7 8-9
801) GieeuRnei
Muiiiinntli CNational Pmk
iver8.12
Cobbled'avdSandSilr
| Substrate Genei'alistGieen River and TiibntaiiesISLuoinoth Cave National ParkGreen River L aleeUpper Green Ri\rei' Sub basin
Figure 6. Map depicting pie charts showing percent composition of substrate preference guilds within fishcommunities at each site.
Mammoth CaNational Piu k
Lyim Csunp Cr*dt B i r B l n s h C r ' f k B i 8 Pitman Creek
831 «T 8."
Legend
Velocity Guild
| Fast FlowModerate Flow
QB Slow/No FlowQFlow Generallst
— Green River and TributariesMammoth Cave National Park
g Green River Lake_ Upper Green River Subbasin 4.5 9
I Kilometers18
Figure 7. Map depicting pie charts showing percent composition of velocity preference guilds within fishcommunities at each site.
Mammoth CaveNational Park
Lynn Camp Creek f Big Pitman Creek
8-3A • J ' 8.7
8.0Di
Legend
Reproductive Guild
Open Water/Substrate Spawnerg Brood Hider3 Substrate Chooser~] Nest Spawner
0 Internal Bearer• Green River and TributariesMammoth Cave National Park
| Green River LakeUpper Green River Subbasin
I Kilometers18
Figure 8. Map depicting pie charts showing percent composition of reproductive guilds within fishcommunities at each site. U)
38
Cam olig
A GR8.9s^5:Eri_diss
Hyp_nigrl^^S.
A GR8.1
Lep macri
Mic d
\Phe m i r a \ ^ \ ^ v \
Phe uran—^^/^*GR8.12Notjrthe""^55^
A G R 8 _ 1 0 _ _ —Eth belT" ^^**
Eth flab—-^""^A-, ~, *GR8OR8.11 A /
Fun cate J//
Ethe_zon yLabsicc
^ ^ ' i ^
< <
13/0/
/ /
'/II 1im afT6
/Not_
V
\
\\lAWmm7/'1volu
Eth
Mic salm
/ " //
^GR8.7^^^**^ A
Lyt_faci^ Pom
^ ^
gR8.2 A GR80D
VV^Arat
Eth macuS
' \ \Cypwhip
caer
rupe
Lep cyan\ ~
^Eth ran\
Fun_nota
*GR8.15A
annu ^GR8.14
Axis 1
Figure 9. Plot of PCA axes 1 and 2 offish data.
4.3E-02
100
3.3E-01
75
Distance (Objective Function)
6.1E-01 9E-01
Information Remaining (%)
50
1.2E+00I
GP 8 ODGP 8 3 1GR8."7
GR8.11 ,GR8.13 1
OK&.li 'GR8.9 ,GR8.10 1GR8.1 ,GP 8 "• 'GR8.14
E. bellum
Group
1 •>
E. zonale
. P. annularisGR8.15 1
Figure 10. Dendrogram depicting results of indicator species analysis of the 2007 fish data.
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