Canadian Journal of Zoology
Large-Scale Changes in the Littoral Fish Communities of Lakes in Southeastern Ontario
Journal: Canadian Journal of Zoology
Manuscript ID cjz-2017-0080.R1
Manuscript Type: Article
Date Submitted by the Author: 27-Oct-2017
Complete List of Authors: Finigan, Paul; Queen's University, Freshwater Fisheries Conservation Lab, Department of Biology Mandrak, Nicholas; University of Toronto at Scarborough, Ecology and EvolutionaryDraft Biology Tufts, Bruce; Queen's University, Freshwater Fisheries Conservation Lab, Department of Biology
FISH < Taxon, BIODIVERSITY < Discipline, FRESHWATER < Habitat, Keyword: Invasive Species, Climate Change, Land Use
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Large-Scale Changes in the Littoral Fish Communities of Lakes in Southeastern Ontario
Paul A. Finigan1, Nicholas E. Mandrak2, Bruce L. Tufts1 1Freshwater Fisheries Conservation Lab, Department of Biology, Queen's University, Kingston, ON, Canada 2 Department of Biological Sciences, University of Toronto Scarborough, Toronto, ON, Canada
SUMMARY
Biodiversity loss is a serious issue for freshwater fishes in temperate
climates and there is a need for more information in this area. A study was
conducted to assess fish community changes in the littoral zone of 22 lakes over a
45-year period (1969-1979 vs. Draft 2014). To compare fish communities, historical
seining records were compiled for 22 inland lakes and compared to contemporary
data sampled using the same protocol. Fish abundance data analyzed using a
multivariate approach identified a shift from cyprinid-dominated communities to
centrarchid-dominated communities between time periods. There was no evidence
to support a strong influence of invasive species on these communities, but there
have been significant changes in temperature and land use around these lakes since
the historical data sets were collected. This is an important contribution to our
understanding of biodiversity change in North American freshwater fish
communities and may influence fisheries management approaches in the future.
Keywords: Fish, Biodiversity, Freshwater, Cyprinidae, Centrarchidae, Invasive
Species, Climate Change, Land Use
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INTRODUCTION
Freshwater ecosystems are among the most ecologically threatened on the planet (Abell et al. 2008). There is evidence that North America’s freshwater species are disappearing at a faster rate than species in tropical rain forests (Ricciardi and
Rasmussen 1999). Increasing human populations contribute to species decline in many ways including habitat loss/degradation, alien species introductions, and climate change (Jelks et al. 2008; Alofs et al. 2014). Understanding the underlying mechanisms involved in biodiversity loss in aquatic ecosystems is crucial for future management efforts, but the lack ofDraft long-term monitoring of fish communities makes it difficult to determine these mechanisms. Evidence provided by Jeppesen et al. (2012) indicated that fish assemblages in European lakes have been shifting in recent decades and that an important contributing factor is probably increasing temperature. In North America, studies examining fish community changes in lakes are limited and have mainly focused on introduced predatory species and their impact on biodiversity under current and future climate scenarios (e.g. Whittier et al. 1997; Jackson and Mandrak 2002; Trumpickas et al. 2011; Alofs and Jackson
2014).
Identifying changes in fish communities over time and the causes of those changes is challenging due to the general lack of detailed historical data. Often only historical occurrence data are available as abundance and sampling effort were typically not reported (Patton et al. 1998; Tingley and Beissinger 2009). Limitations
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of using only occurrence data, for which sampling effort is not known, include false
absences and the inability to describe fish community changes resulting from shifts
in relative abundance rather than introductions or extirpations. In the absence of
abundance data, occurrence data are still useful to gain valuable insight and are
considered relatively robust when comparing data collected with varying effort
(Jackson and Harvey 1997; Heino et al. 2010). Some studies have used abundance
data to describe fish community changes in Europe (Jeppesen et al. 2012). However,
in North America, the only studies using abundance data to examine fish community
changes over time have been focused on larger gamefishes (Robillard and Fox
2006) or on smaller fishes in a single waterbody (Trumpickas et al 2012). Thus,
there is a need for larger-scale studiesDraft that examine changes in small fish
communities over time in North America using abundance data.
The aim of the present study is to determine if there have been changes in
the littoral fish communities in a group of 22 Ontario lakes where historical species
abundance data exist for small fishes. Historical fish community data with known
sampling gear type, effort, date, and location were compiled and the surveys
repeated using the historical methods. We hypothesize that there may have been
changes in these fish communities and that these changes may not be a result of
invasive species.
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METHODS
Fish Community Data
A dataset was compiled that consisted of littoral fish community data for 22 lakes in southeastern Ontario that have been sampled at least once historically
(1969-1979) (Fig. 1). Repeating the historical methods used in the Ontario Ministry of Natural Resources Lake Inventory (Dodge et al. 1985), a contemporary data set was compiled based on fish sampling in summer 2014. All contemporary sampling was conducted within the same time period and similar water temperatures as the
Lake Inventory; June through August and with water temperatures above 20°C. To make direct comparisons betweenDraft time periods, only seining data were used from the historical sampling. The number of sites per lake varied (1-11) based on historical effort. Depending on the site (n=55), either a 9.14 m x 1.83 m, 6 mm bag seine or an 18.29 m x 1.83 m straight seine was used as indicated in the historical records. Species sampled were identified and counted for each site and then pooled by lake. Vouchers were taken for each species at each site and verified in the lab.
Environmental Data
Previous research has shown that surface air temperatures can be used as a proxy for near-surface lake water temperatures (Livingstone and Lotter 1998;
Sharma et al. 2008). To examine the potential influence of climate change on fish communities, near-surface air temperatures from each lake were compared between time periods. Temperature from climate stations across southeastern
Ontario were averaged over 10-year historical (1969-1979) and contemporary
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(2004-2014) time periods. Inverse Distance Weighting (IDW) was used to create an
interpolated temperature surface in ArcGIS (ERSI 2011), then a spatial join was
used to assign temperatures to individual lakes. All temperature data were obtained
from Environment Canada (http://climate.weather.gc.ca/). Climate variables
included mean annual, mean seasonal, and mean July temperatures. Mean annual
and seasonal temperatures are a good measure of ice duration (Benson et al. 2012),
and growing season and mean July temperature have been related to winter survival
of Smallmouth Bass (Micropterus dolomieu (Lacepéde, 1803)) (Shuter et al 1980).
Other environmental variables included were Zebra Mussel (Dreissena polymorpha (Pallas, 1771)) occurrenceDraft and shoreline alterations. Zebra Mussel occurrence was recorded at each sample site and absences were validated with
current distribution maps (EDDMapS Ontario 2015). Zebra Mussel was not present
during the Lake Inventory as it was not found in Ontario until the 1980s (Neary and
Leach 1992). There were no known fish introductions into the lakes. Land use and
the number of cottages on each lake for the two time periods were used to
characterize shoreline alterations. Land use around each lake was identified using a
30m buffer in ArcGIS (ERSI 2011), as this area is known to influence aquatic biota
communities (Lee et al. 2004). Land-use data from the 1966 Canadian land
inventory (DUC and OMNR 2011) was used for the historical period and from the
2000 Southern Ontario land resource information system (OMNR 2000) for the
contemporary period. The number of cottages for each lake was counted as part of
the historical sampling. The current numbers of cottages were counted using 2007
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aerial photography (Google Earth 2015) and 2014 GIS parcel fabric layers showing property boundaries in South Frontenac County (http://www.frontenacmaps.ca/).
Statistical Analyses
To test for significant abundance changes in individual species, species abundance was expressed as Catch-per-unit-effort (CPUE), where ‘unit effort’ is the site, calculated for each fish species in both sampling periods. The mean CPUE per lake for each species was compared between the historical and contemporary sample periods using a paired nonparametric Wilcoxon sign-rank test (α=0.05;
Wilcoxon 1992). Tests for CPUE were conducted multiple times, once for each species, increasing Type-I error dueDraft to multiple hypothesis testing. This error was controlled by applying a false discovery rate (FDR) (Benjamini and Hochberg 1995).
To examine shifts in fish communities among lakes and time periods, ordination methods based on similarity matrices were used to visually identify patterns in the data. A Bray-Curtis similarity index (Bray and Curtis 1957) was used on root-transformed relative fish abundance to down-weight the importance of highly abundant species. Non-metric Dimensional Scaling (NMDS) was used to examine the relationship between fish community and time in two-dimensional space (Kruskal and Wish 1978). To test for significant changes in the fish community, Adonis (p<0.05) was used to compare historical and contemporary distance matrices used in the nMDS analysis (Anderson 2001). Adonis was also used to assess the potential effect of Zebra Mussel on fish community change by comparing contemporary fish communities in lakes with and without Zebra Mussel.
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Matrices were further analyzed using percentage analysis to identify which species
contributed the most to the dissimilarity (Clarke 1993).
A paired student t-test was used to examine the influence of the explanatory
variables temperature, Secchi depth, and shoreline use. Alternatively, a paired
Wilcoxon sign-rank test was used where data did not follow a normal distribution.
All statistical analyses were performed with Program R (R Core Team 2013). The
Community Ecology Package {vegan} was used for multivariate analyses, including
Adonis and similarity percentage (Oksanen et al. 2013).
RESULTS
Species Abundance Draft
A total of 6088 fish records was used in this study: 2458 individual fishes
representing 29 species in the historical data set; and, 3630 fishes representing 24
species in the contemporary data set (Table 1). Several species were not caught
during the re-sampling: Alewife, Bridle Shiner, Creek Chub, Northern Pearl Dace,
Northern Redbelly Dace, Rainbow Smelt, and Iowa Darter (common names
according to Page et al. (2013); scientific names provided in Table 1). Conversely,
the contemporary sampling detected the presence of Bowfin and Northern Pike,
which were absent in the historical data set, but are considered native to the region
(Mandrak and Crossman 1992).
Changes in species abundance were observed. The top three fish species that
increased in CPUE between the time periods belong to the family Centrarchidae, and
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the top three fish species that decreased belong to the family Cyprinidae (Fig. 2).
CPUE increases in Bluegill and CPUE decreases in Bluntnose Minnow were found to be significant (Wilcoxon rank-sign; P < 0.05).
Fish Community
There was a significant difference between historical and contemporary fish communities, where time period accounts for 16% of the variation (Adonis; P <
0.001). Bluegill, Pumpkinseed, Bluntnose Minnow, and Golden Shiner contribute over 50 % of the difference observed between time periods. The NMDS results showed a strong shift from cyprinid-dominated communities to centrarchid- dominated communities (Fig 3), consistentDraft with species CPUE results (Fig. 2). All lake communities (n=22) had a large reduction in cyprinid CPUE and an increase in centrarchid CPUE, with the exception of one outlier, Tetsmine Lake. Centrarchids have not been recorded in Tetsmine Lake. As a result, this lake’s fish community exhibited little change over the two time periods (Fig 3).
Temperature
Temperatures increased significantly across the study region between historical and contemporary sampling periods. Annual temperatures increased 1.2 °C (SE +/-
0.07) (Table 2), with spring, summer, fall, and winter temperatures increasing by
0.7°C (SE +/- 0.07), 0.4 °C (SE +/- 0.02), 1.0°C (SE +/- 0.07), and 3.0 °C (SE +/- 0.24), respectively. In addition, mean July temperature increased on average 0.6 °C (SE
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+/- 0.06). All reported temperature increases were significant between the two time
periods (paired t-test; P < 0.05).
Shoreline Use
Land use around the perimeter of the study lakes has changed since the
historical sampling period. There has been a 5 % increase in pastures, 4 % increase
in wetlands, and 18% decrease in forested areas within a 30 m buffer from the
water. Cottages have significantly increased by 65 % in the study region (Table 2),
which is equivalent to one more cottage for every 1.75 km of shoreline (paired
Wilcoxon rank-sign; P = 0.001). Draft
Invasive Species
Two invasive fish species, common in Lake Ontario (Mills et al. 2005), were
detected in the historical sampling. Alewife was seined in Big Rideau Lake and both
Alewife and Rainbow Smelt were seined in Gananoque Lake. Neither species was
detected in the 2014 sampling. Zebra Mussel, a more recent invasive species, not in
the historical records, is now established in 13 of the 22 study lakes (Table 2). The
introduction of Zebra Mussel to lakes has led to reports of increased water clarity
(Zhu et al. 2006). In the present study however, there were no significant
differences in Secchi depths between the two sampling periods (paired t-test; P >
0.05). Furthermore, there were no significant differences in Secchi depths between
the two time periods in lakes with and without Zebra Mussel (paired t-test; P >
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0.05). There were also no significant differences in contemporary fish communities between lakes with and without Zebra Mussel (Adonis; P > 0.05).
DISCUSSION
The historical data used in this study provides a new perspective on how freshwater fish communities are changing. By compiling detailed fish assemblage records for 22 lakes in southeastern Ontario, 1969-1979, and repeating the sampling effort in 2014, this study shows that there has been a profound shift from cyprinid-dominated to centrarchid-dominated communities in this region.
To date, studies describing large shifts in fish communities in North America have been largely focused on the impactsDraft of introduced species. Several of these studies have described the impact of introduced centrarchids on naïve native cyprinid communities under current climate (e.g. Trumpickas et al. 2011) or climate change (e.g. Jackson and Mandrak 2002; Alofs et al. 2014) scenarios. In our results, there is no evidence that invasive species contributed significantly to the observed fish community change. Two invasive species, Rainbow Smelt and Alewife, established in the 1960s, were not detected in 2014. Both fishes prefer large, open cool waters (Scott and Crossman 1998) that may be threatened by warming climates in inland areas. Only one new invasive species was detected in 2014, Zebra
Mussel. Zebra Mussel was first detected in Lake Ontario and the St. Lawrence River in the late 1980s and has been progressively invading inland lakes (Griffiths et al.
1991). Although this filter feeder is known to increase water clarity, we found no evidence of changes in clarity levels in our study lakes. Furthermore, there was no
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difference in the contemporary fish communities in lakes with and without Zebra
Mussel.
The present study provides evidence that there have been significant
changes in water temperature and shoreline use since the historical time period.
Since the changes we observed in fish communities were consistent across many
lakes and not attributed to invasive species, it is possible that these factors have
contributed to the pronounced shifts in fish communities that we observed.
Increases in the abundance of centrarchids without introduction could be
explained by small, but significant, temperature increases in our study lakes. Winter survival of young-of-year (YOY) centrarchidsDraft is largely dependent on their size and duration of winter (Shuter et al. 1980). Younger fishes deplete their energy reserve
more rapidly than older fishes, so the general strategy for YOY centrarchids over
winter is to build large lipid stores in the summer and fall (Schultz and Conover
1999; Suski and Ridgway 2009). Lipid stores can be estimated using July
temperatures, which are correlated with production and growth of YOY centrarchid
species such as Smallmouth Bass (Shuter et al. 1980). The significant increase in
mean July temperature in southeastern Ontario has likely resulted in YOY
centrarchids with greater lipid stores better able to survive winter. Changes in
winter duration over time may be another important factor leading to improved
survival of YOY centrarchids (Shoup and Wahl 2011). Assel and Robertson (1995)
concluded that ice duration on inland lakes has been decreasing in the Great Lakes
region. They found that every 1oC increase in fall/winter temperatures resulted in
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11-15 more ice-free days, meaning the lakes in this study could have experienced
22-30 more ice-free days compared to the historical period. This increase in ice-free days, combined with larger YOY centrarchids going into winter, may result in higher overwinter survival and, ultimately, larger populations of centrarchids.
There are several reasons why the observed reductions in cyprinids in our study lakes could be a direct result of the increases in centrarchids. Young-of-year
Bluegill can outcompete Golden Shiner for food resources (Paszkowski 1986).
Cyprinids are also a common prey species for the majority of adult centrarchids such as Black Crappie, Largemouth Bass, Rock Bass, and Smallmouth Bass (Scott and Crossman 1998). Bluegill also preysDraft on YOY cyprinids (Scott and Crossman 1998). Thus, the increased survival of YOY centrarchids, as a result of increasing temperatures, could initially lead to competition with cyprinids for food, then increased predation on cyprinids as the centrarchids grow. The observed decline in cyprinids over time may therefore be a result of increasing temperatures favouring increased centrarchid abundance, but this potential explanation will require further investigation.
Shoreline alterations influence the littoral habitat for freshwater fishes by affecting water temperature, vegetation cover and the breeding and nursery habitat for many fishes, including cyprinids. This study found shoreline alterations changed between time periods; the number of cottages increased and riparian forest cover decreased. Altered littoral fish habitat generally favors larger-bodied centrarchid communities, which is reflected in our 2014 samples (Poe et al. 1986 and
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Smokorowski and Pratt 2007). These alterations may also explain why Bluntnose
Minnow has experienced the greatest declines. The Bluntnose Minnow lays its eggs
on the underside of structures in shallow water; these structures are flat rocks or
coarse woody debris (CWD) in many lakes (Scott and Crossman 1998). There is a
positive correlation between CWD and riparian tree density and a negative
correlation between CWD and cottage density (Christensen et al. 1996). The 18%
decrease in riparian forested area and 65% increase in cottages should therefore
result in less CWD for the Bluntnose Minnow. The observed changes in land use
may also be contributing to increasing water temperatures. Thus, it is possible that
both the increase in cottages and decrease in forested areas in southeastern Ontario
over the last 40 years have also contributedDraft to the shift from cyprinid- to
centrarchid-dominated communities.
Historical data are widely used in fisheries research, but few studies have
explored the influence of historical data structure on the interpretation of fish
community change over time. The historical data used in this study were rare;
detailed data on sampling gear and effort, and site location were available. Further
research should be conducted to determine the influence of limited data availability
on identifying historical changes in fish communities and to identify methods to
overcome potential shortcomings in historical datasets. In this study, detailed
historical data using gear types other than a seine were not available for the study
lakes; therefore, offshore sites were not sampled and an examination of changes in
the entire fish community was not possible. Although there have been significant
changes in temperature and land use since the historical data sets were collected, it
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is important to point out that our results do not provide sufficient evidence to conclude that there is a direct linkage between these factors and the observed changes in fish communities and that there could be other contributing factors. We suggest that future studies should examine these relationships in more depth using different proxies for climate change and land use and larger data sets.
Unfortunately, these studies may also be limited by the availability and quality of historical data.
Fisheries management decisions regarding biodiversity loss should be based on an understanding of how fish communities are changing and the factors that are the drivers of these changes. This Draftstudy provides further insights about the underlying causes of biodiversity loss in freshwater fishes in lentic systems. A consistent shift from cyprinid-dominated communities to centrarchid-dominated communities has occurred in these lakes that is not a direct result of invasive species as found in previous studies on North American lakes (Jackson 2002;
Dextrase and Mandrak 2006; Jelks et al. 2008). It is possible that warming temperatures and changes in land use have contributed to these fish community shifts, but additional research is warranted to determine the relative importance of these environmental factors on fish communities in North America. Improving our understanding of these linkages should be a priority as it will provide important direction for future strategies to manage and conserve fish populations.
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ACKNOWLEDGEMENTS
We thank B. Cumming, S. Arnott, and S. Bridgeman for their support throughout the
development of this project and reviews of earlier drafts. We also thank C. DeMille of
Ontario Ministry of Natural Resources for compiling historical fisheries records; A. Drake
for processing the climate data; E. Taylor for his technical support with GIS software; J. Colm
and the rest of the Freshwater Fisheries Conservation Lab at Queen’s University for their
field support; and the staff in the Species-at-Risk Program of Fisheries and Oceans Canada
for their technical support and field training. This project was funded by the Species-at-Risk
Program of Fisheries and Oceans Canada and the Greenberg Family Fund for the
Conservation of Freshwater FisheriesDraft at Queen’s University.
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Draft 24 https://mc06.manuscriptcentral.com/cjz-pubs Page 25 of 31 Canadian Journal of Zoology Figure 1. Southeastern Ontario lakes (n=22) sampled for fish communities, 1969-1979, and re- sampled in 2014. Figure 2. Change in species catch-per-unit-effort (CPUE) in 22 lakes in southeastern Ontario between two sampling periods (1969-1979 and 2014). Controlling for false detection rate, changes in CPUE in Bluegill and Bluntnose Minnow are significant (* Wilcoxon rank-sign; P < 0.05). See Table 1 for scientific names. Figure 3. Ordination (nMDS) using a root transformed Bray-Curtis matrix showing spatial and temporal patterns in fish community compositions in 22 study lakes in southeastern Ontario (Stress = 0.03). ● Historical communitiesDraft (1969-1979); Contemporary communities (2014). 25 https://mc06.manuscriptcentral.com/cjz-pubs Canadian Journal of Zoology Page 26 of 31 Table 1. The number of individuals of each fish species caught in historical (1969-1979) and contemporary (2014) sampling periods in 22 southeastern Ontario lakes. Scientific and common names according to Page et al. (2013). Family Scientific Name Common Name Historical Contemporary Amiidae Amia calva L., 1766 Bowfin 0 2 Atherinopsidae Labidesthes sicculus (Cope, 1870) Brook Silverside 33 3 Catostomidae Catostomus commersonii White Sucker 2 13 (Lacepéde, 1803) Centrarchidae Pomoxis nigromaculatus (Lesueur, 1829) Black Crappie 1 91 Lepomis macrochirus Rafinesque, 1819 Bluegill 93 1398 Lepomis gibbosus L., 1758 Pumpkinseed 242 584 Micropterus salmoides (Lacepéde, 1802) Largemouth Bass 61 275 Micropterus dolomieu Lacepéde,Draft 1802 Smallmouth Bass 20 32 Ambloplites rupestris (Rafinesque, 1817) Rock Bass 14 105 Clupeidae Alosa pseudoharengus (Wilson, 1811) Alewife 41 0 Cyprinidae Notropis heterodon (Cope, 1865) Blackchin Shiner 210 37 Notropis heterolepis Blacknose Shiner 42 21 Eigenman and Eigenman, 1893 Pimephales notatus (Rafinesque, 1820) Bluntnose Minnow 732 173 Notropis bifrenatus (Cope, 1867) Bridle Shiner 25 0 Semotilus atromaculatus (Mitchill, 1818) Creek Chub 9 0 Pimephales promelas Rafinesque, 1820 Fathead Minnow 3 3 Chrosomus neogaeus (Cope, 1867) Finescale Dace 9 20 Notemigonus crysoleucas (Mitchill, 1814) Golden Shiner 193 161 Margariscus nachtriebi (Cox, 1896) Northern Pearl Dace 1 0 Chrosomus eos Cope, 1861 Northern Redbelly Dace 42 0 Esocidae Esox americanus vermiculatus Grass Pickerel 3 2 Lesueur, 1846 Esox lucius L., 1758 Northern Pike 0 5 Fundulidae Fundulus diaphanous (Lesueur, 1817) Banded Killifish 252 277 Ictalurididae Ameiurus nebulosus (Lesueur, 1819) Brown Bullhead 30 64 Ameiurus natalis (Lesueur, 1819) Yellow Bullhead 3 1 Osmeridae Osmerus mordax (Mitchill, 1814) Rainbow Smelt 10 0 Percidae Etheostoma exile (Girard, 1859) Iowa Darter 5 0 Etheostoma nigrum Rafinesque, 1820 Johnny Darter 11 3 Percina caprodes (Rafinesque, 1818) Logperch 42 9 Percina flavescens (Mitchill, 1814) Yellow Perch 327 339 Umbridae Umbra limi (Kirtland, 1840) Central Mudminnow 2 1 26 https://mc06.manuscriptcentral.com/cjz-pubs C) C) ° Annual Temperature ( Temperature Annual 27 Number of Cottages of Number Draft Canadian Journal of Zoology ) presence. Hist: Historical; Cont: Contemporary. Hist: Historical; Cont: Contemporary. ) presence. https://mc06.manuscriptcentral.com/cjz-pubs (m) Hist Hist Cont Hist Cont Change Hist Cont Change Depth Secchi 22 southeastern Ontario lakes in historical (1969-1979) and contemporary (2014) time periods. periods. time (2014) contemporary and (1969-1979) in historical lakes Ontario southeastern 22 Dreissena polymorpha Dreissena 81 + 50 5 3.5 32 5 44 47 5.5 37 60 84 4.9 17 + 2.7 % 28 + 3 26 2.5 5 - - 6.17 77 1 14 5.3 7.14 6 - + 4.1 37 6 12 3.5 4.5 0.96 - % 1100 % 164 6.05 9 4.5 7 6.58 6.52 7.05 - 6 % 50 8.05 8.23 7 1.01 6.40 13 - 1.47 1.70 % 0 7.61 % 117 6.42 6.35 6.58 1.22 7.97 7.26 7.89 1.55 0.91 1.31 5.5 5.5 3.2 4.5 0 0 % 0 6.30 7.25 0.95 646 646 227 + 964 3.5 617 191 + 1.6 7 + 2.7 + * 4.7 175 166 4.4 2 3.3 229 0 2.5 190 222 88 % 31 787 5.3 51 342 0 179 6.5 + 6.04 58 % 80 4.3 % 103 % 0 3.2 20 220 7.07 4.8 119 % 14 6.50 3.5 6.42 6.10 38 12 551 1.03 + 120 7.52 6.46 7.97 954 2.1 7.07 2.6 % 90 206 + 12 7.89 2.8 1.02 + 0.5 1.55 0.97 6.30 - % 72 4.3 12 % 0 1.42 38 3.5 7.25 6.5 6.29 18 6.36 - 105 45 0.95 7.47 % 50 7.38 200 62 1.18 - 6.38 % 1.02 90 % 38 7.86 6.17 6.26 6.35 1.48 7.16 8.37 7.75 0.99 2.11 1.40 6497 + 3.5 8 - - - 6.13 7.33 1.21 Lake Lake (ha) Area ZM ZM: 2014 Zebra Mussel Mussel ( 2014 Zebra ZM: Table 2. Characteristics of of Characteristics 2. Table Burridge Christie Cranberry Davern Dog Gananoque Grippen Higley Killenbeck Kingsford Leo Long RockLower Opinicon Pearkes BigRideau Singleton South Temperance Tetsmine Beverley Upper Wolfe Some historicalNote: contemporary and data missingisto limited due dataaccess Secchi depth* > depthmax Page 27 of 31 Page 28 of 31 28 Draft Canadian Journal of Zoology https://mc06.manuscriptcentral.com/cjz-pubs Page 29 of 31 Canadian Journal of Zoology SMITH FALLS 7 N Christie Cranberry Long Davern Big Rideau Wolfe 29 Burridge Upper Beverley BROCKVILLE 15 Draft Temperance Kingsford Opinicon Tetsmine Singleton Grippen Higley Pearkes Lower Rock Killenbeck St. Lawrence Leo South Gananoque Dog 401 KINGSTON 0 5 10 20 Km Lake Ontario https://mc06.manuscriptcentral.com/cjz-pubs Canadian Journal of Zoology Page 30 of 31 (23.7) 10 10 7.5 7.5 5 5 2.5 2.5 0 Draft 0 Change in CPUE ‐2.5 ‐2.5 ‐5 ‐5 ‐7.5 ‐7.5 ‐10 ‐10 Darter Bowfin Bluegill Alewife Logperch Rock Bass Creek Chub Iowa Yellow Perch * Bridle Shiner White Sucker Black Crappie Pumpkinseed Grass Pickerel Northern Pike Johnny Darter Golden Shiner Finescale Dace Rainbow Smelt Banded Killifish Yellow Bullhead Brown Bullhead Brook Silverside Blackchin Shiner Fathead Minnow Blacknose Shiner Largemouth Bass Smallmouth Bass https://mc06.manuscriptcentral.com/cjz-pubs Bluntnose Minnow Northern Pearl Dace Central Mudminnow Northern Redbelly Dace * Page 31 of 31 Canadian Journal of Zoology Draft 221x191mm (96 x 96 DPI) https://mc06.manuscriptcentral.com/cjz-pubs