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Theses and Dissertations
2016 Fish Ladders Designed for Drop Culverts and Central Stoneroller Ecology Across a Latitudinal Gradient John Arthur Lorenzen South Dakota State University
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Recommended Citation Lorenzen, John Arthur, "Fish Ladders Designed for Drop Culverts and Central Stoneroller Ecology Across a Latitudinal Gradient" (2016). Theses and Dissertations. Paper 984.
This Thesis - Open Access is brought to you for free and open access by Open PRAIRIE: Open Public Research Access Institutional Repository and Information Exchange. It has been accepted for inclusion in Theses and Dissertations by an authorized administrator of Open PRAIRIE: Open Public Research Access Institutional Repository and Information Exchange. For more information, please contact [email protected]. FISH LADDERS DESIGNED FOR DROP CULVERTS AND CENTRAL
STONEROLLER ECOLOGY ACROSS A LATITUDINAL GRADIENT
BY
JOHN ARTHUR LORENZEN
A thesis submitted in partial fulfillment of the requirements for the
Master of Science
Major in Wildlife and Fisheries Sciences
Specialization in Fisheries Science
South Dakota State University
2016
iii
To Karissa, my wife, for the constant love and support that got me through this and to my parents, Raymond and Sue for guiding me through life and making me who I am today
iv
ACKNOWLEDGEMENTS
I would like to especially thank my wife Karissa. She has been a part of my life ever since I stepped foot at SDSU and I cannot imagine having finished my degree without her love and support. She has contributed many hours to my project, whether riding along to sites to help sample fish or painting the original fish ladders, she has been there for it all and I am blessed to have her by my side.
I would also like to especially thank my Mom and Dad for their continuous love and belief in me. I have always had lofty dreams, but I know that they have never doubted me for a second. My Dad had an important role in this project. He helped design and build the fish ladders for the field. We used his shop and equipment, and without his expertise, this project would not have been nearly as successful. I thank both my parents for giving me their creativity and ability to work with both my mind and hands. This has shaped the way I look at things and has helped me overcome challenges throughout life.
I thank my advisors Dr. Katie Bertrand and Dr. Brian Graeb. They both saw something in me and gave me a shot. If I needed anything, they were usually able to pull strings to accommodate me, making my job easier. They gave me the free will to pursue research in whatever I wanted and because of that I was able to form a creative project that I was interested in. I also thank Dr. Michael Brown for advice, mentorship, and good humor throughout my time at SDSU.
I thank the Bertrand/Graeb lab group for the countless times spent editing my presentations. I have become friends with many fellow students at SDSU and I would like to thank Matthew Wagner, Cassidy Gerdes, David Schumann, Eli Felts, Jason
Breeggemann, Chad Kaiser, Seth Fopma, Jeremy Kientz, Stephen Jones, Alex Rosburg,
v
Brad Smith, Nicholas Kludt, and Morgan Kauth for directly helping me with my project throughout my time here. I would also like to thank my friends Matthew Neff for being there for me and putting up with me as a roommate my first year of graduate school and
John Harris for helping me with Autodesk software.
I thank my technicians and now friends David Ahrens and Ryan Hansen for helping me with me my field season. Without the hard work of these two men, I could not have done anything. They worked hard, did not complain, and put up with me throughout the summer. Other technicians that I would like to thank that have helped me with either sampling or otolith preparation were Jared Roiland and Ryan Johnston.
I thank Chelsey Pasbrig for helping my project get funded. I thank Jake Davis for inviting me to the Black Hills to research and helping me throughout my time there. I thank Steve Hirtzel for allowing me to conduct research on forest service land and I thank
Greg Simpson for taking time out of his schedule to help me out when our shocker broke down.
I thank the many private landowners who graciously allowed me to conduct research on their land. Some of the best memories I have from the field were from the conversations with these diverse and friendly people.
Funding for this project was provided by South Dakota State Wildlife Grant T-67-
R-1, South Dakota State University Department of Natural Resource Management, South
Dakota Agricultural Experiment Station, and South Dakota Department of Game, Fish, and Parks.
vi
TABLE OF CONTENTS
ABSTRACT ...... viii
Chapter 1: Introduction ...... 1
Importance of stream connectivity ...... 1
Climate change ...... 2
Importance of stream connectivity in a changing climate ...... 3
Barriers to fishes ...... 4
Culverts as barriers to fishes ...... 5
Project overview ...... 7
Literature Cited ...... 10
Chapter 2: Small stream fish ladders for drop culverts ...... 17
Abstract ...... 18
Introduction ...... 19
Methods...... 23
Results ...... 31
Discussion ...... 34
Literature Cited ...... 42
Tables ...... 50
Figures...... 53
Chapter 3: Phenology and population dynamics of Central Stoneroller Campostoma anomalum across a latitudinal gradient...... 67
Abstract ...... 68
Introduction ...... 69
vii
Methods...... 72
Results ...... 75
Discussion ...... 76
Literature Cited ...... 81
Figures...... 88
Chapter 4: Conclusions ...... 96
Summary ...... 96
Management recommendations ...... 98
Research needs...... 99
viii
ABSTRACT
FISH LADDERS DESIGNED FOR DROP CULVERTS AND CENTRAL
STONEROLLER ECOLOGY ACROSS A LATITUDINAL GRADIENT
JOHN ARTHUR LORENZEN
2016
Stream connectivity is essential for the persistence of fish populations and maintenance of fish assemblage structure. Stream-road crossings can become barriers resulting in a loss of connectivity. Creating innovative tools for fisheries managers to allow fish passage through barrier stream-road crossings is important worldwide as climate change may force range shifts to stay within thermal requirements.
Understanding how populations and metapopulations respond to climate is important as fisheries managers can use this information to make informed management decisions in regards to climate change and possible range shifts. Thus, we developed and tested a fish ladder to retrofit barrier stream-road crossings and quantified recruitment, growth, mortality, body condition, and phenology of Central Stoneroller across three latitudes.
The final fish ladder design was based on a Denil-type fish ladder and consisted of 14 evenly spaced flat-top baffles every 15.2 cm at 45° angles to the slope. We tested the fish ladder under controlled and field conditions. During the controlled study, all 14 species stocked were able to ascend the ladders in experimental streams with 13 species passing at rates equal to or greater than 50% whereas only one fish was observed passing the control (no ladder). Fish passed at all 19 field sites. Overall, 1,213 fishes representing
23 species passed fish ladders throughout the summer with a mean passage rate (± SE) in
ix the Black Hills of 1.22 (0.38) fish/day and 28.05 (19.60) fish/day in the Northern
Glaciated Plains.
To understand climatic effects on Central Stoneroller, fish were sampled from three streams in each of three latitudes: 44.5° (South Dakota), 39.1° (Kansas), and 32.0°
(Mississippi). Recruitment was consistent and growth and mortality were similar among latitudes. Maximum age observed was age-3 though age-3 fish were only found in the northern two latitudes. Body condition was similar among high and mid latitudes but was significantly lower in the southern latitude. Peak gonad development occurred later in the north than in the south; stoneroller peak gonadal somatic index (GSI) in Mississippi was
21% on day 65, whereas South Dakota GSI peaked at 15% on day 134. These results indicate that Central Stoneroller phenology is adapted to temperature and northern fish can compensate for shorter growing season as they showed no differences in growth.
1
Chapter 1: Introduction
Importance of stream connectivity
Connectivity is important for fishes in stream networks as many fishes require different parts of the riverscape for different stages of their life history (Fausch et al.
2002; Kashiwagi and Miranda 2009). Unlike terrestrial organisms or lentic fishes that have the ability to move in all directions, lotic fishes are restricted to longitudinal movements constrained by stream banks. Though stream fishes can move laterally into the riparian floodplain when flow exceeds bank height (Ross 2013), these are brief opportunities in typical years.
Northcote (1978) suggested that there are three different types of fish habitat: refuge, feeding, and spawning habitat. Movement among these habitats is essential for the survival and viability of stream fish assemblages (MacDonald and Davies 2007,
Savage and Brenchley 2013). As a result of the uneven distribution of habitat longitudinally within streams and shifts in habitat requirements among life stages in fishes, the ability to move within stream networks can influence stream fish population viability (Poplar-Jeffers et al. 2009). For reasons such as migration, predator avoidance and reproduction, it is often necessary for stream fishes to disperse and move upstream.
The ability to move unhindered provides access to these types of habitats and in turn, decreases the risk of predation and both interspecific and intraspecific competition.
The importance of connectivity is well documented in large anadromous species such as Striped Bass Morone saxatilis, yet many small-bodied potadromous fishes that sexually mature early undergo migrations upstream from larger systems as well (Perkin and Gido 2012). Small stream fishes have been shown to move long distances. Freeman
2
(1995) found that both darters and sunfish moved between habitat types with some individuals travelling 200 and up to 420 m and Schumann (2012) documented Plains
Topminnow movements of over 1,300 m in Nebraska streams. Movement is how fish take advantage of different habitat types and colonization of upstream reaches that would not be possible without stream connectivity.
Climate change
Climate change may result in increased mean annual air temperatures by over
1.5°C by the end of the 21st century (IPCC 2007), which will alter stream thermal regimes, resource quality and availability, hydrology and species distribution (Perkins et al. 2010; Poff and Allan 1995) along with challenging the physiological tolerances of organisms (Fuller et al. 2010, Somero 2010). Climate change will have a profound effect on fish, particularly stream fishes. Fish are ectothermic, meaning they do not produce their own heat; therefore, body temperature is similar to water temperature. Thus, water temperature dictates metabolism (Brown et al. 2004) and therefore, influences ecological processes such as feeding (Huey and Kingsolver 1989). Phelps et al. (2008) demonstrated that temperature and precipitation influence recruitment in some fishes.
Climate change has also altered the phenology of some species (Parmesan 2006).
In the past 20 to 140 years, 59% of 1,598 species studied by Parmesan and Yohe (2003) had altered phenology and/or range with advancements in phenology up to 2.3 days per decade (Parmesan and Yohe 2003). For example, Inouye et al. (2000) observed yellow- bellied marmots Marmota flaviventris emerging from hibernation by up to 23 days early because of a rise in local temperature of 1.4°C even though date of snowmelt and emergence of flowering plants remained constant.
3
Importance of stream connectivity in a changing climate
Climate is a regional filter that encompasses environmental factors such as temperature and precipitation that influence ranges of fishes (Fischer and Paukert 2008,
Ross 2013). Changes in climate may result in species range shifts and may be hindered by physical barriers. Therefore, both stream fragmentation and climate change pose an enormous conservation challenge for fishes (Perkin et al. 2010). Climate change will exacerbate the effects of stream fragmentation (Perkin et al. 2010) as both temperature and precipitation are predicted to increase in variation thus having direct effects on freshwater environments, including prolonged drought and increased intermittency (Milly et al. 2005). Perkin et al. (2010) found that stream fragment length explained 71% of the variability in persistence of pelagic-spawning cyprinids in Great Plains streams. The remaining 29% of the variability was explained by stream discharge (Perkin et al. 2010).
With some streams of the Great Plains predicted to lose 12% of their discharge by 2060 as a result of climate change, many streams may become more fragmented (Perkin et al.
2010). Reduced discharge will isolate fish populations and reduce habitat availability
(Perkin et al. 2010) making solutions to stream fragmentation imperative for the sustainability of some species in the face of a changing climate. Meixler et al. (2009) suggests that eliminating barriers should be given precedence over habitat enhancement for isolated fish populations which are restricted in range. This is extremely important for peripheral populations that are already experiencing conditions that are on the limits of their physiological tolerances. Climate change may force species distributions to shift, and without stream connectivity, extirpations will continue to occur. Stream
4 fragmentation and an increase in environmental variability will continue to be a risk to native stream fishes (Fischer and Paukert 2008).
Barriers to fishes
Barriers to fish movement can isolate necessary habitats for all or some life stages of fishes and therefore, has the potential to decrease productivity of fish populations
(Poplar-Jeffers et al. 2009) and can result in a loss of genetic variability, lowering relative fitness, which could lead to species loss (MacDonald and Davies 2007, Kemp and
O’hanley 2010, Savage and Brenchley 2013). At current levels of fragmentation, species that persist today may not be found in the future. Long term data sets show a time-lag effect on fish assemblages resulting from fragmentation (Taylor et al. 2008, Perkin and
Bonner 2011). Level of fragmentation may stay constant but as a result of a delayed or prolonged response, extinction debts may exist in streams and species present today in areas may yet be lost (Tilman et al. 1994). Topeka Shiner Notropis topeka, a federally listed endangered species is one of many declining as a result of the loss of stream connectivity, and has been shown to be extirpated from fragmented streams (Perkin and
Gido 2012). Fish communities found in fragmented streams have been shown to have lower species richness relative to streams that have maintained connectivity (Alexandre and Almeida 2010; Perkin and Gido 2012) with decreases in species richness of up to 50 percent in fragmented streams (Nislow et al. 2011). Morita and Yamamoto (2002) found that dams in southwestern Japan have caused the extirpation of White-Spotted Char
Salvelinus leucomaenis and Kashiwagi and Miranda (2009) attributed extirpations of
Blackside Darter Percina maculata above small impoundments in their study sites to the loss of downstream connectivity. Barriers also are detrimental to entire guilds. For
5 example, pelagic spawning fishes release semi-buoyant eggs that must drift downstream until they hatch. Barriers then prevent the upstream movement of these fishes once they reach a free-swimming phase (Dudley and Platania 2007).
Barriers not only disrupt fish movement, but can alter upstream and downstream habitat and flow characteristics (Kashiwagi and Miranda 2009). Reservoirs create lentic habitat (Gido et al. 2002) resulting in large areas of standing water that promote high non-native piscivore abundance (Luttrell et al. 1999, Gido et al. 2002). Large reservoirs also release cold water downstream making temperatures unsuitable to stenothermal fishes. Other barriers, such as culverts, constrict the channel and increase velocity (Briggs and Galarowicz 2013) which scours the downstream streambed resulting in a waterfall feature that can become a barrier. Because traits promote local persistence and change longitudinally throughout stream networks, highly variable systems segmented by barriers may result in fish assemblages reduced to tolerant, cosmopolitan fishes (Poff and
Allan 1995).
Culverts as barriers to fishes
Stream-road crossings can become barriers to lotic fishes (Warren and Pardew
1998, Benton et al. 2008, Kemp and O’hanley 2010). A stream-road crossing becomes a barrier to a fish when fast water velocity, high outlet drop, dense suspended debris, excess turbulence, or shallow water depth occurs within or surrounding the stream-road crossing (Wargo and Weisman 2006, Hotchkiss and Frei 2007). Stream-road crossings are prevalent across many landscapes, ranging from flat agricultural lands with stream- road crossings every square mile to mountainous regions where roads are adjacent to and cross streams in valleys (Roni et al. 2002; Poplar-Jeffers et al. 2009). With over 6.4
6 million km of roads in the United States (Norman et al. 2009) and road construction increasing in most countries (Clay 1995), barriers from stream-road crossings have been and will remain a problem for lotic fishes worldwide.
Stream-road crossings may take the form of bridges, box culverts, low water fords, or cylindrical concrete or galvanized steel culverts (hereafter steel culverts). Steel culverts are often preferred at stream-road crossings (Clay 1995; Warren and Pardew
1998; Gibson et al. 2005), and have the most negative effect on overall stream fish movement (Warren and Pardew 1998). Steel culverts typically act as one-way barriers, allowing fish to pass downstream but not upstream. This can contribute to a disproportionate loss of headwater and higher-elevation habitat (Sheer and Steel 2006). If steel culverts do not act as a complete barrier to a species, they may act as a barrier at certain times or for certain life stages within a species. For example, steel culverts designed for passage of adult game fish often do not allow passage of juveniles, eliminating juveniles from important rearing habitat (Roni et al. 2002).
Although there are many situations in which a culvert can become a barrier
(Wargo and Weisman 2006, Hotchkiss and Frei 2007), a physical drop at the outlet has been shown to be the most detrimental (Burford et al. 2009). Though a few centimeters of drop from a culvert may look insignificant, it may be a complete barrier for weakly swimming or benthic oriented fish (MacPherson et al. 2012). Small species such as darters and shiners are weaker jumpers and swimmers than larger bodied roving or potadromous species, and as such, may not be able to pass even the smallest vertical drops. Some larger bodied species such as salmon may not be able to pass vertical drops if the plunge pool on the downstream side is too shallow (Lauritzen 2002, Meixler et al.
7
2009). Prenosil et al. (2016) found that of the 10 game and non-game fishes that they evaluated for jumping ability, only Green Sunfish was able to leap more than 10 cm whereas the other nine species were limited to jumps of 7 cm or less. MacPherson et al.
(2012) found that culvert drop height, slope, and outlet water velocity influenced the distribution of non-game fish with lower densities found upstream of culverts. Culverts with any drop height were found to be impassable barriers to Burbot Lota lota and partial barriers to cyprinids and catostomids (MacPherson et al. 2012). Because of the magnitude of drop culverts on the landscape, Meixler et al. (2009) suggests that culvert removal or fish ladder installation should be given initial precedence over habitat enhancement, particularly in the case of isolated fish populations with range restrictions.
Project overview
Drop culverts are major barriers for stream fishes. More alarming is the vast number of them in existence. Along the western coast, on federal land in Oregon, many of the 10,000 stream-road crossings pose an obstacle to fish passage (Sheer and Steel
2006) and according to the Alaska Department of Fish and Game, within Tongass
National Forest, 66% of the stream-road crossings in salmon streams and 85% of the stream-road crossings in trout streams may prevent fish movement (Flanders and Cariello
2000). Though the consequences of drop culverts are known, limited resources are usually the reason they are not replaced (Norman et al. 2009), and as a result of cost, steel culverts will continue to be the main stream-road crossing structure used in the future
(Gibson et al. 2005; Poplar-Jeffers et al. 2009). Solutions to eliminate drop culverts have been focused on installing large, expensive, permanent cement weirs at intervals across streams below culverts to backwater and eliminate drop (Clay 1995). This solution is
8 very expensive, time consuming, disturbs the streambed, and has historically focused only on game species needs. Despite many studies describing the negative effects that drop culverts have on fish passage, little attention has been given to possible solutions for existing drop culverts to mitigate the passage issue (Gibson et al. 2005).
A principle goal of this project was to design a fish ladder that would allow passage of small stream fishes and be able to attach to most drop culverts. This project did result in a fish ladder design that was able to pass all species experimentally tested and most species encountered in the field, providing state and federal agencies with a valuable tool to enhance stream connectivity.
Alterations in temperature associated with climate change will profoundly affect ecosystems by simultaneously altering the timing and availability of resources and challenging the physiological tolerances of organisms (Fuller et al. 2010, Somero 2010).
Long-term climatic data and general circulation models suggest that global mean annual air temperature is likely to increase over 1.5°C by the end of the 21st century (IPCC
2007), which will alter stream thermal regimes, resource quality and availability, hydrology and species distribution (Perkins et al. 2010; Poff and Allan 1995). Grazing species (e.g., bison) have been shown to control primary producers (Fahnestock and
Knapp 1993). However, it is unclear how climate change will affect grazer performance or interact with diurnal and seasonal patterns of solar energy inputs (photoperiod will remain constant). The match-mismatch hypothesis (Cushing 1969) predicts climate change will cause a mismatch between the metabolic efficiencies of consumers (i.e., grazers) and the availability of resources (i.e., producers). Assessing the dynamic rate functions (i.e., recruitment, growth, and mortality) along with body condition and
9 phenology (timing of peak reproductive condition) of a grazing stream fish, the Central
Stoneroller Campostoma anomalum across a latitudinal gradient will capture a natural transition of climate and allow us to evaluate future responses to the overall performance of this fish and how it may interact with its environment with warmer climates with longer growing seasons.
A principle goal of this project was to quantify the effects of warming on population dynamics (i.e., recruitment, growth, and mortality) along with body condition and reproductive phenology across a latitudinal gradient to predict the effects of warming on Central Stonerollers. Species distributions may need to shift to accommodate changes in temperature and precipitation and may be hindered by barriers. Understanding how populations respond to climate, coupled with innovative tools to enhance stream connectivity will improve the potential for stream conservation and restoration.
10
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connectivity, and salmon persistence in the Willamette and lower Columbia River
basins. Transactions of the American Fisheries Society 135:1654-1669.
16
Somero, G. N. 2010. The physiology of climate change: how potentials for
acclimatization and genetic adaptation will determine ‘winners’ and ‘losers’. The
Journal of Experimental Biology 213:912-920.
Taylor, C. M., D. S. Millican, M. E. Roberts, and W. T. Slack. 2008. Long-term change
to fish assemblages and the flow regime in a southeastern U.S. river system after
extensive ecosystem fragmentation. Ecography 31:787-797.
Tilman, D., R. M. May, C. L. Lehman, and M. A. Nowak. 1994. Habitat destruction and
the extinction debt. Nature 371:65-66.
Wargo, R. S., and R. N. Weisman. 2006. A comparison of single-cell and multi-cell
culverts for stream crossings. Journal of the American Water Resources
Association 42(4):989-995.
Warren, M. L., and M. G. Pardew. 1998. Road crossings as barriers to small-stream fish
movement. Transactions of the American Fisheries Society 127:637-644.
17
Chapter 2: Small Stream Fish Ladders for Drop Culverts
This chapter is being prepared for submission to North American Journal of Fisheries Management and was co-authored by Brian D. S. Graeb, Chelsey A. Pasbrig, and Katie N. Bertrand.
18
Abstract
Stream connectivity is important for fishes as it allows longitudinal migration within watersheds. Stream- road crossings (i.e., steel culverts, box culverts, etc.) can block fishes from migrating upstream hindering their life cycle and ability to recolonize.
Fish ladders designed to retrofit culverts and eliminate stream fragmentation, specifically for small stream fishes, do not exist. We tested five fish ladder designs in replicated stream mesocosms on a mixed assemblage of fishes found in the Northern Glaciated
Plains of eastern South Dakota to evaluate the effect of baffle profile and width on passage rate. Using the design with the highest passage rate of mixed assemblages, we evaluated passage rate of 15 lotic species individually within the same systems. Fish were stocked in experimental streams below the ladders and controls. Trials ran for 48 hours per species and fish that passed were removed every 24 hours for preservation. All 15 species were able to pass our fish ladders as opposed to only one individual of one species passing the control. We then tested fish ladders on culverts in 10 streams in the
Black Hills ecoregion and 9 streams in the Northern Glaciated Plains ecoregion. Fish that ascended ladders were captured in funnel traps to estimate passage rate. Drop height, ladder slope, water velocity, depth, discharge, fish density below culvert, and plunge pool depth were measured at each site to assess influential passage variables. Overall, 1,213 fishes representing 23 species passed fish ladders throughout the summer with a mean passage rate (± SE) in the Black Hills of 1.22 (0.38) fish/day and 28.05 (19.60) fish/day in the Northern Glaciated Plains. Among sites we found no relationship between passage rate and our independent variables though daily discharge influenced passage rate within
19 some sites. Culverts retrofitted with passable fish ladders have the potential to reverse the negative effects of habitat fragmentation on lotic fishes.
Introduction
Fishes use different types of habitat for basic survival needs and for the persistence of populations. Northcote (1978) suggested that there are three different types of fish habitat: refuge, feeding, and spawning habitat. For stream-dwelling fish, connectivity among the three habitat types is essential for persistence of fish populations
(Poplar-Jeffers et al. 2009) and maintenance of fish assemblage structure (MacDonald and Davies 2007, Savage and Brenchley 2013). Stream fishes often must migrate or move upstream to reduce predation risk, reproduce, and feed (MacDonald and Davies
2007). Upstream dispersal also can decrease the intensity of interspecific and intraspecific competition. The importance of connectivity is well documented for economically valuable, large bodied, anadromous species such as Sockeye Salmon
Oncorhynchus nerka, yet many small-bodied potadromous fishes also migrate upstream from larger rivers, lakes, and reservoirs to access refuge, feeding, and spawning habitats
(Perkin and Gido 2012).
Fragmentation can isolate necessary habitats for all or some life stages of fishes and therefore, have the potential to decrease recruitment and growth of fish populations
(Poplar-Jeffers et al. 2009) and can result in a loss of genetic variability, lowering relative fitness, which could lead to population or species loss (MacDonald and Davies 2007,
Kemp and O’hanley 2010, Savage and Brenchley 2013). At current levels of fragmentation, species that persist today may not be found in the future. Long term data sets show a time-lag effect on fish assemblages resulting from fragmentation (Taylor et
20 al. 2008, Perkin and Bonner 2011). Level of fragmentation may stay constant but because of a delayed or prolonged response, extinction debts may exist in streams and species present today in areas may yet be lost (Tilman et al. 1994). Topeka Shiner Notropis topeka, a federally listed endangered species is one of many declining due to the loss of stream connectivity, and has been shown to be extirpated from fragmented streams
(Perkin and Gido 2012). Because traits promote local persistence and change longitudinally throughout lotic systems, highly variable systems segmented by barriers may result in persistence of upstream fish assemblages composed of only fishes with certain traits, such as those with the greatest swimming or jumping ability (Poff and
Allan 1995).
Road crossings fragment stream fish habitats (Warren and Pardew 1998, Benton et al. 2008, Kemp and O’hanley 2010). Road construction is increasing in most countries
(Clay 1995) and in the United States, we have over 6.4 million km of roads (Norman et al. 2009). In much of the Midwest, roads intersect streams about once every square mile.
Road-stream crossings also are abundant in mountainous regions as a result of steep grades and the adjacency of roads and streams in valleys (Roni et al. 2002; Poplar-Jeffers et al. 2009). Hundreds of thousands of road-stream crossings exist in forested drainages of the eastern United States (Nislow et al. 2011). Along the western coast, on federal land in Oregon, many of the 10,000 road-stream crossings pose an obstacle to fish passage
(Sheer and Steel 2006). According to the Alaska Department of Fish and Game, within
Tongass National Forest, 66% of the road-stream crossings in salmon streams and 85% of the road-stream crossings in trout streams may prevent fish movement (Flanders and
Cariello 2000). Fish communities found in streams fragmented by road-stream crossings
21 have been shown to have lower species richness relative to streams that have maintained connectivity (Alexandre and Almeida 2010; Perkin and Gido 2012) with decreases in species richness by up to 50 percent (Nislow et al. 2011).
Road-stream crossings can vary from bridges to box culverts, low water fords to cylindrical concrete or galvanized steel culverts (hereafter steel culverts) though steel culverts are most economical and thus, most preferred (Clay 1995; Warren and Pardew
1998; Gibson et al. 2005). A steel culvert can block fish movement as a result of fast water velocity, high outlet drop, dense suspended debris, excess turbulence, or shallow water depth within or immediately up- or downstream of the culvert (Wargo and
Weisman 2006, Hotchkiss and Frei 2007). Steel culverts cause the greatest reduction in stream fish movement of any road-stream crossing type (Warren and Pardew 1998). Steel culverts typically form one-way barriers, passing fish in the downstream direction but inhibiting upstream passage. This can contribute to a disproportionate loss of headwater and higher-elevation habitat consisting of 1st and 2nd order streams (Sheer and Steel
2006). If steel culverts do not result in a complete barrier to a species, they may still function as a barrier at certain times or for certain age classes. For example, steel culverts designed for passage of adult game fish often block passage of juveniles, decreasing overall recruitment by excluding juveniles from important rearing areas (Roni et al.
2002).
Steel culverts also change stream channel morphology through erosional scouring.
The outflow from the culvert can continually scour the downstream streambed, increasing the distance from the water’s surface to the lip of the culvert, resulting in a waterfall feature that impedes fish movement (Boubee et al. 1999, Wellman et al. 2000, Wargo and
22
Weisman 2006, Olsen and Tullis 2012). Waterfalls are barriers if the height exceeds the jumping capability of the fish, and many fishes are not capable of jumping more than a few inches (Clay 1995). Ficke and Myrick (2011) assessed the jumping capability of three small lotic species. They found that a drop of 10 cm dramatically decreased passage in Brassy Minnow Hybognathus hankinsoni, Common Shiner Luxilus cornutus, and
Arkansas Darter Etheostoma cragini with no fish passing drops of more than 15 cm
(Ficke and Myrick 2011). Drops in water level are detrimental to fish passage though steel culverts that have formed a waterfall on the downstream end (hereafter drop culverts) may be seasonally passable, allowing fish dispersal during periodic floods, for example (Norman et al. 2009; Perkin and Gido 2012).
There is currently no inexpensive solution to mitigate drop culverts (Norman et al.
2009). Furthermore, steel culverts will likely continue to be installed at road crossings, because they’re more cost effective to manufacture and install than bridges (Gibson et al.
2005; Poplar-Jeffers et al. 2009). A fish ladder designed to retrofit drop culverts has been proposed as a possible solution for this issue (Ficke et al. 2011) though none exists.
Fish ladders have been around for hundreds of years (Clay 1995) primarily for the circumventing of large dams by anadromous fishes, but since the advent of automobiles and the expansion of roads, no fish ladders have been specifically designed for road- stream crossings. Fish ladders have historically focused on species with high commercial or recreational value. Though smaller fishes have been observed attempting to traverse these fish ladders, little research has been done on movement of small stream fishes past barriers (Mallen-Cooper and Stuart 2007, Savage and Brenchley 2013). Thus, the objectives of this study were to: 1) compare mean passage rates of small stream fishes
23 among three fish ladder baffle designs in experimental streams, 2) evaluate the effect of baffle slot width in experimental streams on mean passage rate of fishes using the most effective baffle design from objective one, and two additional widths of that baffle design, 3) using the most effective design from objectives one and two, test mean passage rate of 14 lotic species in individual trials in experimental streams, and 4) field test the most effective design on extant fish assemblages in stream-road crossings found in the
Black Hills and Northern Glaciated Plains ecoregions.
Methods
Study area
Black Hills
The South Dakota Black Hills are a domed-shaped uplift situated on the western edge of South Dakota surrounded by short and midgrass prairie (DeWitt et al. 1989). The
Black Hills are an island of mountains and streams within them provide cold water habitat for terrestrial and aquatic species (Breeggemann et al. 2014). Included in the
Level III Middle Rockies Ecoregion, the Black Hills are dominated by ponderosa pine and cold water perennial streams are spring-fed (Omernik 1987) with riparian areas consisting of grazed open meadows, forest, or willow-dominated banks (Koth 2007). The climate is considered continental, highly variable, and characterized by low precipitation, cold winters, and hot summers (Carter et al. 2003). Locally within the Black Hills, climate is driven by elevation, with higher elevations having colder temperatures and greater precipitation (Carter et al. 2003). Geologically, Precambrian metamorphic and intrusive rocks compose the center of the formation and are completely surrounded by
Cretaceous sedimentary rocks (DeWitt et al. 1989). Porous limestone formations of the
24
Northern and Eastern Black Hills cause surface discharge to decrease to little or no flow and are referred to as Loss Zones, creating dispersal barriers to fishes in dry years
(Williamson and Carter 2001). Other barriers to fish movement in the Black Hills include dams and road-stream crossings.
Historically, the Black Hills native fish fauna consisted of Mountain Sucker
Catostomus platyrhynchus and Longnose Dace Rhinichthys cataractae with a few populations of Creek Chub Semotilus atromaculatus, White Sucker Catostomus commersonii, and Fathead Minnow Pimephales promelas in the lower drainages (Koth
2007; Breeggemann et al. 2014). Finescale Dace Chrosomus neogaeus and Longnose
Sucker Catostomus catostomus are also present in the Black Hills though rare and only found in a few isolated areas (Schultz et al. 2012). Three non-native salmonids (i.e.,
Brook Trout Salvelinus fontinalis, Brown Trout Salmo trutta, and Rainbow Trout
Oncorhynchus mykiss) now represent the majority of fish biomass and provide a recreational fishery in Black Hills streams (Schultz and Bertrand 2012).
Northern Glaciated Plains
The Northern Glaciated Plains of eastern South Dakota is a level III ecoregion 46
(Omernik and Gallant 1988). Our study area within this ecoregion consists of three drainages: the Big Sioux, Minnesota, and Red River of the North. These drainages are considered coteaus, consisting of rolling hills with prairie pothole lakes scattered throughout (Bryce et al. 1998). Historically, native short and tallgrass prairie covered the landscape which is now dominated by cattle grazing and rowcrop agriculture (Krause et al. 2013). Climate within this region is considered subhumid with average annual precipication accumulations of 432 to 559 mm per year (Bryce et al. 1998). Fish
25 assemblages can be quite diverse and are composed of warm water species. Species are typically small-bodied non-game fishes.
Experimental streams
Sixteen experimental streams located at South Dakota State University’s Wildlife and Fisheries Sciences Farm at the Agricultural Experiment Station in Brookings, South
Dakota were used to assess the fish ladder designs. Individual experimental streams were comprised of a circular 2.54 m² pool attached to a rectangular 0.84 m² riffle (Matthews et al. 2006) and substrate consisted of gravel. Throughout the trials, well water was continually added and excess water was discharged onto the ground. Atop the riffle section of stream, higher elevation holding tanks were constructed using the same material as the fish ladders. Holding tanks placed above the experimental streams allowed us to create a drop in water level and provide an attachment for the fish ladders.
A sidewall section of holding tank was removed 36 cm above the water level to create a drop from the upstream holding tank to the experimental stream pool below at this height.
The area removed was large enough to fit a fish ladder. All fish ladders were set at this drop with a 27% slope. Preferably, ladders would have been set at a flatter grade (Clay
1995), but limitations with the system did not allow for less slope. Controls were constructed using identical upstream holding tanks. Control holding tanks were fitted with a 50 cm long 61 cm diameter segment of culvert overhanging the pool below with a drop height the same as the fish ladders (36 cm). Immediately at the top of the fish ladder or control, a mesh funnel was placed at the entrance to the upstream holding tank so that any fish that passed either the ladder or control would not be able to leave the holding tank and descend into the pool below. A utility pump (Wayne RUP160) was used to
26 move water from the lower riffle section to the upper holding tank at a rate of 164 l/min.
In ladder treatments, water would then flow down the fish ladder into the plunge pool where it would circulate back into the riffle. In controls, instead of flowing through fish ladders into the pool, water would flow through the section of culvert and drop vertically into the pool to mimic a drop culvert found in the field. Both fish ladders and controls had 0.23 m³ net holding pens below the ladder or vertical drop of the control with a plunge pool depth of 61cm. Holding pens prevented fish from escaping and directed their movement in a way that only upstream was an option. Mesh coverings were placed over the downstream holding pens and the upstream holding tanks to prevent fish from jumping out of the system. The experimental streams were not shaded and were exposed to natural fluctuations in temperature. Figure 2-1 depicts the study system.
Ladder prototypes
Three fish ladder prototypes were fabricated using 7/16” oriented strand board lined with plastic construction film. Ladder designs differed in baffle profile including baffles with a wide flat-top profile (Design 1; Figure 2-2), v-top profile (Design 2; Figure
2-3), and no-baffles present (Design 3; Figure 2-4). Two additional prototypes were fabricated using the same material and baffle profile as design 1, however, these two additional prototypes differed in slot width in order to assess the effects of water velocity without altering discharge or slope. These additional two prototypes included an intermediate slot width (Design 4; Figure 2-5) and a narrow slot width (Design 5; Figure
2-6) with the wide slot width being the original design 1.
Fish collection
27
The fish community of Medary Creek (44.268101, -96.508357) was subsampled on the 25th of June 2014 and the fish community of Peg Munky Run (44.583642, -
96.783672) was subsampled on the 10th of July 2014 using pulsed DC current from a backpack electrofisher. Sufficient individuals of a mixed assemblage of cyprinids were collected to replicate the assemblage in the experimental streams. Species counts from
Medary Creek included: Central Stoneroller Campostoma anomalum, 21 fish/stream;
Sand Shiner Notropis stramineus, 17 fish/stream; Blacknose Dace Rhinichthys atratulus,
13 fish/stream; Creek Chub Semotilus atromaculatus, 5 fish/stream; Common Shiner
Luxilus cornutus, 4 fish/stream; Bluntnose Minnow Pimephales notatus, 2 fish/stream; and Red Shiner Cyprinella lutrensis, 1 fish/stream. Species from Peg Munky Run included: Central Stoneroller, 29 fish/stream; Blacknose Dace, 15 fish/stream; Common
Shiner, 10 fish/stream; Sand Shiner, 8 fish/stream; Tadpole Madtom Noturus gyrinus, 5 fish/stream; Bigmouth Shiner Notropis dorsalis, 3 fish/stream; Bluntnose Minnow, 3 fish/stream; and Creek Chub, 2 fish/stream.
Collected fish were then transported in coolers with stream water and battery powered aerators to the South Dakota State University Agricultural Experiment Station
Wildlife and Fisheries Research Farm. Fish were acclimated to experimental streams in a
1m2 flow-through net pen for 5 days. Fish were fed daily.
During all trials, funnel traps were checked daily to identify and count successful fish. Successful fish were removed daily and all fish were removed at the end of their respective trial and euthanized with tricaine methane sulfonate (MS-222). Fish were preserved in 10% formalin solution and measured for total length (mm TL), standard length (mm SL), body depth (mm BD), and mass (0.1 g). During the trial, water depth
28
(mm) and velocity (m/s) measurements were taken (Flo-Mate 2000) in the center of every baffle or in the place where a baffle would be (in the case of the no-baffle design).
Study design
Experimental trial 1 designs
Four replicates of designs (1-3) in 12 experimental streams were used to assess fish passage among designs. Equitable mixed assemblages of cyprinids, representing the relative abundances sampled in Medary Creek were stocked into 12 experimental streams. Stocking rates were based on the total number of each species sampled equally distributed among treatments in proportion to the number caught in Medary Creek (24
Bluntnose Minnows sampled and distributed in 12 experimental streams = 2 fish/stream).
The trial began on 3rd July 2014 and ran for three days.
We calculated mean percent passage for each replicate. We used an analysis of variance (ANOVA) to test the null hypothesis that there were no differences in percent passage among ladder designs.
Experimental trial 2 baffle notch width
Four replicates of designs 1, 4, and 5 in 12 experimental streams were used to assess the effect of baffle slot width on fish passage. Equitable mixed assemblages of cyprinids, representing the relative abundances sampled in Peg Munky Run were stocked into 12 experimental streams. Stocking rate criteria was the same as trial 1 except based on the Peg Munky Run fish assemblage. The trial began on 17th of July 2014 and ran for three days.
We calculated mean percent passage for each treatment. We tested the assumption of normality with a Shapiro-Wilk test. For data that were not normally distributed we
29 assessed heteroscedasticity and applied variance stabilizing transformations. Where transformations were ineffective, we used non-parametric tests of main effects.
Experimental trial 3 species
Four controls and four replicates of design 1 (Figure 2-2) in 8 experimental streams were used to estimate passage rate on 15 fishes (Figure 2-7). Species were stocked individually into experimental streams at natural densities similar to where they are found in South Dakota. Fish were stocked evenly among experimental streams within each species trial. Trials ran for two days for each species.
We calculated mean percent passage for each species. We used an analysis of covariance (ANCOVA) to test the null hypothesis that there were no differences in percent passage among ladder designs with density as a covariate. If differences in percent passage were detected among species, we used Tukey’s honestly significant difference (HSD) test for multiple comparisons to assess differences in mean percent passage among species.
Experimental trial 3 ecoregion
We grouped fishes geographically by ecoregion they are found in South Dakota
(Black Hills, Northern Glaciated Plains, and both). We calculated mean percent passage for each ecoregion. We used an analysis of variance (ANOVA) to test the null hypothesis that there were no differences in percent passage among fishes found in differing ecoregions. If differences in percent passage were detected among them, we used
Tukey’s (HSD) test to assess differences in mean percent passage among ecoregions.
Experimental trial 3 water movement guild
30
We grouped fishes by water movement guild (Poff and Allan 1995): fast, moderate, slow/none, and generalist water movement guild. We used an analysis of variance (ANOVA) to test the null hypothesis that there were no differences in percent passage among guilds. If differences in percent passage were detected among guilds, we used Tukey’s (HSD) to assess pairwise differences in mean percent passage among guilds.
Field study
Three replicate fish ladders (Figure 2-8) were constructed using steel. Large funnel traps were constructed out of steel and wire mesh, and were attached to the top of the fish ladder to catch all fish that ascended the ladder. We installed block nets 100 meters downstream from the culvert, and used pulsed DC backpack electrofishing (ETS
ABP-3-300, Madison, WI) to triple pass sample fish to depletion prior to installing the fish ladder at each road-stream crossing. We also recorded culvert shape, material, width, height, length, slope, elevation above plunge pool, and plunge pool depth. We noted installation time at each culvert. After fish ladders were installed, we measured slope, and during each day of deployment, three velocity and depth measurements were taken across the bottom, middle, and top ladder baffles.
Fish ladders were deployed for five days at each road-stream crossing. For road- stream crossings with more than one culvert pipe, the ladder was attached to the culvert with the lowest elevation above the plunge pool (Anderson et al. 2012) as the higher elevation culvert pipe is a more difficult barrier to overcome thus reducing the chance of fishes moving upstream around the fish ladder. Water temperature was measured hourly by a data logger (HOBO Pendant; Onset model UA-002-08 Bourne, MA). Funnel traps
31 were monitored daily, and species and total length were recorded prior to releasing fish upstream of the culvert.
Mean passage rate was calculated for each site. Passage rate included all fish by study site. Black Hills and Northern Glaciated Plains data sets were analyzed separately.
Variables included in the analysis were: Julian dates, temperature, drop height, slope of ladder, mean velocity flowing through ladder, mean depth flowing through ladder, discharge through ladder, fish density below culvert, and plunge pool depth. Pearson correlation analysis was performed on variables to assess collinearity. If collinearity existed, we eliminated one of the paired collinear variables. Multiple linear regression was conducted on the remaining variables with passage rate as the dependent variable to see if passage rate was related to any of the independent variables among sites. Within sites, Pearson correlation analysis was used to assess whether daily passage rate was related to fluctuations in discharge which we obtained from the nearest United States
Geological Survey stream gage station.
Results
Experimental trial 1 designs
Mean water velocity in meters/second (± SE) was fastest 1.2 (0.05) in the no- baffle design, intermediate 0.47 (0.02) in the v-top baffle design, and slowest 0.38 (0.01) in the flat-top baffle design. Mean depth (mm) over baffles (± SE) was shallowest 11.5
(0.70) in the no-baffle design, intermediate 20.8 (0.42) in the flat-top baffle design, and deepest 37.73 (0.52) in the v-top baffle design. Zero fish passed the no-baffle design; so therefore, it was not included in further analyses. A Shapiro-Wilk test on percent passage in the flat-top baffle design (W = 0.97, P = 0.78) and the v-top baffle design (W = 0.96, P
32
= 0.64) indicated that the data was normally distributed and a Levene’s test indicated that the data had equal variance (F = 0.52, p = 0.48). There was no significant interaction between design and species (F₁ ₄̦ = 1.13, P = 0.36), so main effects were assessed. Mean percent passage (± SE) was higher 0.53 (0.05) in the flat-top baffle design fish ladders than the v-top baffle design fish ladders (0.42 ± .06; F₁ ₄̦ = 4.49, P = 0.04; Figure 2-9).
Experimental trial 2 baffle notch width
Differences in baffle notch width among the three designs altered water velocity and depth, ultimately affecting passage rates among designs. Mean water velocity in meters/second (± SE) was significantly different among baffle notch widths (F2,11 =
31.71, P < 0.0001) with the narrow baffle ladder velocity 0.75 (0.02) nearly twice as fast as the wide baffle 0.38 (0.01) and intermediate baffle ladders 0.44 (0.01). Because of a non-normal distribution of data, we used a non-parametric Kruskal-Wallis test to test the null hypothesis that there were no differences in median percent passage among water velocities within the flat-top baffle design. Percent passage was similar among water velocity treatments (x²₂ = 1.912, P = 0.384; Figure 2-10). No Tadpole Madtoms were observed passing any ladder, regardless of notch width.
Experimental trial 3 species
The wide flat-top baffle design (Figure 2-2) was chosen for this trial due to having the highest mean passage rate among the designs in trial 1. Shapiro-Wilk tests on percent passage by species indicated that the data was normally distributed and a Levene’s test indicated that the data had equal variance (F = 0.25, P = 0.87). There was no significant interaction between stocking density and species percent passage (F₁₁ ₁̦ = 1.88, P = 0.08); therefore, density was eliminated as a contributing factor. Percent passage was
33 significantly different among species (F₁₃ ₄₂̦ = 8.42, P < 0.0001) and ranged from over
90% for Flathead Chub and Mountain Sucker to under 10% for Plains Topminnow
(Figure 2-11).
Experimental trial 3 ecoregion
A Shapiro-Wilk test was done on percent passage of fishes found in the Black
Hills (W = 0.88, P = 0.15), Northern Glaciated Plains (W = 0.98, P = 0.99), and of fishes found in both ecoregions (W = 0.95, P = 0.24) and the data was normally distributed.
Percent passage was significantly different among ecoregions (F₂ ₅₃̦ = 10.33, P = 0.0002)
(Figure 2-12). According to Tukey’s HSD, Black Hills fishes passed at a significantly higher rate than fishes found in the Northern Glaciated Plains and fishes found in both ecoregions. No differences in passage rate were observed between Northern Glaciated
Plains fishes and fishes found in both ecoregions.
Experimental trial 3 water movement guild
A Shapiro-Wilk test on percent passage of fishes found in the fast water movement guild (W = 0.95, P = 0.70), moderate water movement guild (W = 0.98, P =
0.98), slow to no moving water guild (W = 0.90, P = 0.18), and generalist water moving guild (W = 0.96, P = 0.83) indicated that the data was normally distributed. Percent passage was significantly different among water movement groupings (F3,52 = 8.69, P <
.0001) (Figure 2-13). According to Tukey’s HSD, fishes associated with fast water velocities and moderate water velocities passed at significantly higher rates than fishes associated with slow water velocity. The generalist guild, typically found among all water velocity habitats, passed at rates that were not significantly different from fast, moderate, or slow water movement guilds.
34
Field study
Throughout summer 2015, we installed fish ladders on drop culverts at 10 sites in the Black Hills ecoregion in western South Dakota and 9 sites in the Northern Glaciated
Plains ecoregion in eastern South Dakota. We were able to attach ladders to all culverts encountered in the field which consisted of varying shapes, sizes, and materials and with ladders attached, fish of many sizes, representing 23 species were able to ascend ladders overcoming the drop culverts (Table 2-1). Overall, 1,213 fishes representing 23 species passed fish ladders throughout the summer (Table 2-2) with a mean passage rate (± SE) in the Black Hills of 1.22 (0.38) fish/day and 28.05 (19.60) fish/day in the Northern
Glaciated Plains (Table 2-1).
The Bois de Sioux site passage rate of 184.34 fish/day was an outlier as a result of schooling species passing at much higher rates than observed elseware and was eliminated from further analyses. Regression analysis showed that no relationship existed between passage rate and variables measured in both the Black Hills (F4,6=0.15, P=0.95) and the Northern Glaciated Plains (F6,1=0.28, P=0.89).
Within Black Hills sites, passage rate was not correlated with discharge, but most sites in Eastern South Dakota displayed some correlation (r > 0.60) with the sites: tributary to Brule Creek (n = 5, r = 0.97, p = 0.01), Mckay Fork of the Hidewood (n = 5, r
= 0.88, p = 0.05), and tributary to North Fork Yellow Bank River (n = 5, r = 0.94, p =
0.02) having daily passage rate significantly correlated to daily discharge.
Discussion
A flat-top baffled ladder was identified as the best design for stream-road crossings among those tested and was easily attached to culverts. This fish ladder design
35 was able to dissipate energy efficiently and operate among a wide range of velocities and depths, thus allowing it to pass a larger volume of water without adverse affects on fish passage, much like the basis of this design, a Denil fish ladder (Clay 1995). The fish ladder we tested in the field was slightly over two meters in length making it economically feasible to build and would allow managers to install them efficiently. By directing water flow back onto itself, this fish ladder was able to reduce water velocity near the top of each baffle, thus allowing fish to pass even at steep grades (Bunt et al.
2001). Bunt et al. (2012) suggests that a Denil fish ladder like our design may struggle with attraction efficiency. This is true on large systems such as rivers where discharge through the fish ladder is a fraction of the volume going through the dam; however, culverts on small streams are not on the same scale and the majority of the discharge flows through the ladder. The versatility of a Denil fish ladder along with the challenges presented by drop culverts is why we based our design on this type of fish ladder.
The results of our study show that our fish ladder design allows passage of small- bodied fishes over drop culverts acting as barriers, and is thus a solution to what research has suggested is an issue for nonsalmonid species (Ficke and Myrick 2009).
Experimentally the design was able to pass all 15 species we ran individual trials on, whereas only one Creek Chub was observed passing the control (vertical drop height
35.56 cm). Ten of 19 field sites had vertical drop heights greater than the experimental vertical drop height of 35.56 cm, and vertical drop heights were greater than 15 cm at all sites, which was the maximum jumping height of small stream fishes measured experimentally by Ficke et al. (2011) and Prenosil et al. (2016). Wall and Berry (2004) considered a vertical drop height of more than 6 cm to be a complete barrier to the
36 federally listed Topeka Shiner, and vertical drops ranging from 0.24 m to 0.6 m are presumably impassable to adult salmonids depending on flow characteristics (Anderson et al. 2012). Therefore, depending on the target species, a large portion of drop culverts would be considered barriers for at least part of the year. Bates et al. (2003) recommends culverts to allow passage for all sizes of fish for at least 90% of the passage season.
Because our fieldwork at sites was conducted during relatively short periods of time (5 days/site), it is possible that during other times of the year, increased discharge may inundate culverts, eliminating the barrier at some sites; however, all sites were clearly barriers when fieldwork was conducted and would be considered a complete barrier at that time for most if not all fishes and would remain that way for the vast majority of the year.
Though perched culverts may be semi-passable during flood pulses, upstream fish communities have been shown to be resistant to colonization (Hitt and Roberts 2012) and therefore, periodic connectivity from flood pulses may not be sufficient for fishes to recolonize segments of stream they once occupied prior to culvert perching (Perkin and
Gido 2012). Though semi-passable and not a complete barrier year round, drop culverts have been shown time and again to mimic larger, more permanent barriers resulting in negative consequences on stream fishes (Perkin and Gido 2012). Norman et al. (2009) observed water column species passing drop culverts only during times of high flow when culverts were no longer perched. During base flow when culverts where perched above water level, they observed no upstream movement of water column species or benthic species even though upstream movement probabilities were above 5% at control culvert free sites. Throughout summer 2015, none of our drop culvert sites became
37 inundated and we observed both benthic and water column species passing throughout the summer. Bunt et al. (2001) did not observe either Johnny Darter Etheostoma nigrum or Brook Stickleback Culaea inconstans passing the Denil fish ladders they researched; however, we recorded both Johnny Darter and Brook Stickleback passing our fish ladder design. We found no relationship between any of the variables we recorded and fish passage among sites in the Black Hills and among sites in the Northern Glaciated Plains.
Denil-type fish ladder gradients have been recommended to range from 16 to 20% for trout and salmon (Clay 1995), and would be likely lower for non-game fishes.
However, we observed Creek Chub, Central Stoneroller, Blacknose Dace, and White
Sucker passing our ladder design at slopes up to 60%. This suggests that this fish ladder design can be attached to nearly any culvert at slopes within the range we observed this summer (up to 60 %) and fish assemblages will be able to pass similarly among sites. We did notice a positive relationship between daily discharge and passage rate within eastern
South Dakota sites, suggesting that increased discharge may increase passage rates in some individual streams for some species. We attribute increases in passage rates after rainfall events within some sites in eastern South Dakota to movement behavior, with rainfall triggering the movement. Increases in discharge from rainfall events do not however affect the fishes ability to pass the fish ladders, as we evaluated discharge among sites and it had no affect on passage, but as suggested before, it is anthropocentric, or behavioral affect. No relationship between daily discharge and passage rate within sites in the Black Hills existed. During the time we were in the Black Hills installing and evaluating fish ladders, stream discharge was far above average and never receded during sampling. Because discharge did not spike or recede, but rather stayed consistently high
38 during our time in the Black Hills, we attribute this as the reason no trends exist between passage rate and daily discharge in that ecoregion.
Water velocity that exceeds the critical swimming velocity of a fish will decrease passage rate (Rodriguez et al. 2006). Engineering to minimize water velocities within culverts and fish ladders may increase passage rates. Powers et al. (1997) recommended design velocities of 0.34 m/s in order to allow passage of salmon fry (≤ 60 mm). Powers et al. (1997) and Bouska and Paukert (2009) suggest that prairie stream fishes are able to pass road crossing structures up to 1.1 m/s whereas Ward et al. (2003) suggested that passage of prairie fishes would cease before 0.80 m/s. Some velocities in this study were
86% higher than what Ward et al. (2003) suggests as a maximum velocity; therefore, eliminating the drop at these sites does not completely mitigate that barrier.
Lack of fish ladder use by some species encountered in the field (Black Crappie
Pomoxis nigromaculatus, Blackside Darter Percina maculata, Bluegill Lepomis macrochirus, Brown Trout Salmo trutta, Freshwater Drum Aplodinotus grunniens,
Hornyhead Chub Nocomis biguttatus, Mountain Sucker, Northern Pike Esox lucius,
Rainbow Trout, Red Shiner, Stonecat, Tadpole Madtom, and Walleye Sander vitreus) may be attributed to passage efficiency, rarity of species below the stream-road crossing, physical limitations, or behavioral limitations that affect movement cues (Bunt et al.
2001). It is worth noting, however, that Mountain Sucker, Rainbow Trout, Red Shiner, and Stonecat passed the fish ladders during the experimental stream trials in summer
2014. Furthermore, none of the 13 species that did not pass the ladders in the field occurred at more than two stream-road crossing sites.
39
The ladder deployed at the Bois De Sioux site passed fish at the high mean rate of
184.34 fish/day. We attribute this abnormally high passage rate to schooling species that would pass as a group rather than individuals. Carmine Shiner Notropis percobromus were observed schooling below the drop culvert prior to ladder installation and the majority of the fish that passed the fish ladder at that site were Carmine Shiners on the first day of installation.
Upon arriving in the Black Hills, snowfall was common the first two weeks of the field season and on average, stream temperatures in the Black Hills were 10°C colder than Northern Glaciated Plains streams. Though not significant, temperature or seasonality played some role in passage in the Black Hills. White Suckers have been observed using Denil-type fish ladder designs if water temperatures were consistently above 8°C (Bunt et al. 2001). In the Black Hills we observed similar trends in passage with only one White Sucker passing below 8°C. White Sucker passage did increase as water temperature rose above that threshold. We also observed no Brook Trout passing when water temperatures were below 5°C or above 14°C.
Solutions to eliminate the barrier that a drop culvert presents has been focused on installing large, expensive, and permanent cement weirs at intervals across the stream below the culvert in order to backwater the culvert and eliminate the drop (Clay 1995).
This solution is very expensive, time consuming, disturbs the streambed, and has historically focused only on game species needs. Because removing culverts is typically not feasible, designing new culverts to allow for fish movement and retrofitting existing culverts with fish ladders are the two primary options in which to restore stream connectivity (Ficke et al. 2011). Poplar-Jeffers et al. (2009) surveyed 120 culverts within
40 the Cheat River basin West Virginia and within just these culverts, 61% were perched, 83 of the 120 culverts were considered complete barriers, 34 were considered partial barriers, and only 3 posed no barrier. The cost to replace the culverts that pose an issue just from this study alone was estimated to cost between $5 and $10 million dollars. Even if only the 20 highest priority culverts were to be replaced from this study, it would cost almost $1 million dollars (Poplar-Jeffers et al. 2009). Our fish ladder design not only functioned well, but was very cost effective. If the 73 perched culverts found by Poplar-
Jeffers et al. (2009) were to have our fish ladder design attached, the cost would be much less than replacing the culverts and most likely be under $200,000 total. Our approach of designing a fish ladder to directly attach to most existing culverts has been shown to be successful both biologically and mechanically. Biologically it has passed 23 species including fishes associated with a wide variety of habitats, water velocities, swimming performance, size, and morphology among a wide variety of environmental conditions.
Mechanically it was able to attach to culverts of varying material, size, shape, drop height, and plunge pool depths. The results of this study show that this fish ladder can be a versatile management tool for fisheries managers who are attempting to promote stream connectivity for all fishes in watersheds fragmented by drop culverts.
Other research points to fish ladders as the solution to drop culverts (Hotchkiss and Frei 2007; Ficke et al. 2011). Successful fish ladders can lead to large increases in fish recruitment (Kemp and O’hanley 2010), as habitat connectivity is a vital determinant for the distribution of fish species. Instead of investing time and energy into increasing habitat to increase production, restoring passage through culverts allows access to habitat
41 already available (Roni et al. 2002), thus the potential to increase fish abundance and distribution can be achieved through the nominal cost of installing fish ladders.
Mitigating stream fragmentation across the landscape is vital for the persistence of lotic fishes as fragmentation threatens biodiversity worldwide (Dudgeon et al. 2006).
Lotic fishes are highly succeptible to fragmentation as access to resources is limited to longitudinal dispersal within streams (Perkin et al. 2013). Perkin et al. (2013) found that decreases in barrier permeability led to greater extirpations. Fish ladders retrofitted to drop culverts would increase permeability and allow passage for a greater amount of time throughout the year. This may have a profound effect on fish populations worldwide as extirpations of some endangered species such as the Topeka Shiner has been attributed to habitat fragmentation (Perkin and Gido 2012). Habitat fragmentation can lead to species extinction (Tilman et al. 1994) but with innovative solutions such as this fish ladder design, mitigation of barriers such as drop culverts is possible.
42
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Tables
Table 2-1. Culvert measurements taken at 19 field sites where fish ladders were installed within the Black Hills and Northern Glaciated Plains ecoregions in the spring and summer of 2015. Slate Creek through Swede Gulch are located in the Black Hills ecoregion and Brule Creek tributary through Slip-up Creek are located in the Northern Glaciated Plains ecoregion. Corrugated galvanized pipe is abbreviated as CGP.
Culvert Site Latitude Longitude Shape Material Width (cm) Height (cm) Length (m) Slope (%) Plunge Depth (cm) Slate Creek 43.959283 -103.724515 Circle CGP 70 70 15 3.49 50 Upper Spring Creek 43.905533 -103.746569 Circle CGP 62 62 7.2 5.24 30 Buskala Creek 44.207032 -103.801759 Circle CGP 110 110 12.5 6.99 60 Yellow Creek 44.323525 -103.747173 Circle CGP 166 156 30 4.37 56 Whitewood Creek 44.296519 -103.784266 Circle CGP 125 125 14.2 4.37 84 Cold Spring Creek 44.155644 -104.049601 Circle Steel Piping 82 82 8 2.62 60 Gold Run 44.005678 -103.791039 Circle CGP 170 210 26 2.62 46 Lower Spring Creek 43.898401 -103.725419 Circle Cement 111 111 4.15 1.75 86 North Fork Castle Creek 44.086612 -103.799767 Circle CGP 135 135 14.5 1.75 60 Swede Gulch 44.177556 -103.777293 Oval CGP 296 203 14.6 5.24 80 Brule Creek Tributary 42.982284 -96.756591 Circle CGP 185 185 15 1.75 53 East Union Creek 42.937864 -96.594671 Circle CGP 380 380 24.5 1.75 22 Beaver Creek Tributary 43.601489 -96.539016 Circle CGP 203 170 18.2 1.75 30 Bois De Sioux 45.862398 -96.584818 Oval CGP 270 190 17 1.75 152.4 Mckay Fork, Hidewood Creek 44.68152 -96.842152 Circle CGP 190 190 18.2 4.37 110 Willow Creek 44.896932 -97.066045 Oval CGP 300 210 16.3 1.75 25 North Fork Yellow Bank Tributary 45.135959 -96.811987 Circle CGP 300 300 36 6.12 20 Slip-up Creek Tributary 43.68861 -96.65478 Semi-Circle CGP 145 100 15.6 3.49 36 Slip-up Creek 43.67883 -96.651193 Circle CGP 300 300 24.5 5.24 25
50
Table 2-1. Continued.
Ladder Site Slope (%) Mean Velocity (m/s) Mean Depth (cm) Installation (minutes) Passage Rate (fish/day) Slate Creek 17.63 0.81 13.33 80 2.16 Upper Spring Creek 15.84 0.74 5.44 111 0.11 Buskala Creek 26.79 0.61 6.89 93 0.41 Yellow Creek 36.40 0.52 5.89 90 0.42 Whitewood Creek 17.63 0.59 6.33 100 0.45 Cold Spring Creek 17.63 1.49 15.33 N/A 3.42 Gold Run 26.79 0.76 12.89 100 3.29 Lower Spring Creek 15.84 1.09 14.44 105 0.42 North Fork Castle Creek 26.79 0.84 7.78 240 0.24 Swede Gulch 15.84 0.71 11.11 210 0.60 Brule Creek Tributary 28.67 0.42 3.11 80 14.25 East Union Creek 36.40 0.68 3.33 150 0.64 Beaver Creek Tributary 21.26 0.34 1.89 210 10.62 Bois De Sioux 36.40 0.37 5.00 150 184.34 Mckay Fork, Hidewood Creek 14.05 0.73 7.67 195 9.29 Willow Creek 17.63 0.82 5.89 60 4.66 North Fork Yellow Bank Tributary 60.08 0.21 3.56 225 3.68 Slip-up Creek Tributary 28.67 0.32 1.72 75 14.50 Slip-up Creek 19.44 0.56 2.51 165 10.46
51
52
Table 2-2. Number, mean total length (± SE), minimum length, and maximum length of fish that passed fish ladders at field sites in the Black Hills and Northern Glaciated Plains ecoregions during the summer 2015 field study.
Length Species Passed (#) Mean (mm ± SE) Minimum (mm) Maximum (mm) Bigmouth Shiner 26 60.2 ± 0.8 53 66 Black Bullhead 99 214.1 ± 2.9 106 285 Blacknose Dace 1 90 90 90 Bluntnose Minnow 1 66 66 66 Brook Stickleback 5 48.2 ± 4.8 33 59 Brook Trout 49 167.5 ± 5.0 103 232 Carmine Shiner 609 69.3 ± 0.7 52 96 Central Stoneroller 7 106.0 ± 10.3 64 132 Channel Catfish 1 381 381 381 Common Carp 2 119.0 ± 2.0 117 121 Common Shiner 29 90.4 ± 3.0 60 150 Creek Chub 108 95.0 ± 3.0 44 186 Fathead Minnow 171 54.8 ± 0.6 42 84 Green Sunfish 6 70.2 ± 6.1 56 97 Iowa Darter 1 55 55 55 Johnny Darter 2 37.5 ± 7.5 30 45 Largemouth Bass 3 67.0 ± 4.0 63 75 Longnose Dace 4 99.3 ± 8.1 80 115 Orange Spotted Sunfish 1 67 67 67 Sand Shiner 2 60.0 ± 2.0 58 62 Spotfin Shiner 57 60.0 ± 1.1 48 83 White Sucker 28 114.6 ± 11.8 57 240 Yellow Bullhead 1 136 136 136
53
Figures
Figure 2-1. Control (top) and ladder (bottom) experimental stream setup at South Dakota
State University’s Agricultural Experiment Station in Brookings, South Dakota in summer 2014. Arrows represent direction of flow.
Figure 2-2. Schematic of experimental fish ladder design 1 (wide flat-top) used in experimental trials 1, 2, and 3 in experimental streams at South Dakota State University’s Agricultural Experiment Station in Brookings, South Dakota in summer 2014.
54
Figure 2-3. Schematic of experimental fish ladder design 2 (v-top) used in experimental trial 1 in experimental streams at South
55
Dakota State University’s Agricultural Experiment Station in Brookings, South Dakota in summer 2014.
Figure 2-4. Schematic of experimental fish ladder design 3 (no-baffles) used in experimental trial 1 in experimental streams at South
56
Dakota State University’s Agricultural Experiment Station in Brookings, South Dakota in summer 2014.
Figure 2-5. Schematic of experimental fish ladder design 4 (intermediate slot width) used in experimental trial 2 in experimental
57
streams at South Dakota State University’s Agricultural Experiment Station in Brookings, South Dakota in summer 2014.
Figure 2-6. Schematic of experimental fish ladder design 5 (narrow slot width) used in experimental trial 2 in experimental streams at
58
South Dakota State University’s Agricultural Experiment Station in Brookings, South Dakota in summer 2014.
59
180 Fish ladder
160 Control 12 9
140
120 15 15 100 13 11 80 29 25
15 13 13 60 13 12
Mean totalMean length (mm) 18 40
20
0
Creek Chub Red ShinerSand Shiner Johnny Darter Rainbow Trout White Sucker Flathead ChubLongnose Dace Blacknose Dace Common Shiner Mountain Sucker BluntnoseCentral Minnow Stoneroller Plains Topminnow
Species
Figure 2-7. Mean total length (mm ± SE) of species stocked in fish ladders and controls for experimental trial 3 conducted in eight experimental streams at South Dakota State University’s Agriculture Experiment Station in Brookings, South Dakota in summer 2014. Numbers above bars denote number of fish stocked in each stream.
Figure 2-8. Schematic of fish ladder used for the field study in the Black Hills and Northern Glaciated Plains ecoregions in summer of
60
2015. (1 of 2).
Figure 2-8 continued. Schematic of fish ladder used for the field study in the Black Hills and Northern Glaciated Plains ecoregions in summer of 2015. (2 of 2).
61
62
60 A
50 B
40
30
20
Percent (%) passage
10
0 Flat-top V-top No-baffle Ladder design
Figure 2-9. Mean percent passage (± 1 SE) of fishes among three fish ladder designs (designs 1-3) from trial 1 of the experimental study. Letters depict results of Tukey’s HSD among fish ladder designs. Means with the same letter are not significantly different. Study was conducted in summer 2014 at South Dakota State University’s Agricultural Experiment Station in Brookings, South Dakota.
63
50 A A
40 A
30
20
Percent passage (%) Percent passage
10
0 Wide Intermediate Narrow
Baffle slot width
Figure 2-10. Mean percent passage (± 1 SE) of fishes among three baffle notch widths (designs 1, 4, and 5) from trial 2 of the experimental study. Letters depict results of Tukey’s HSD among baffle notch widths. Means with the same letter are not significantly different. Study was conducted in summer 2014 at South Dakota State University’s Agricultural Experiment Station in Brookings, South Dakota.
64
100 A A A B A B B C C A A B B C C A 80 A B B C A A C B B B C C C C
60 C
40
Percent passage (%)
20
D
0
Creek Chub Red Shiner Sand Shiner White Sucker Flathead Chub Rainbow Trout Johnny Darter MountainLongnose Sucker DaceBlacknose Dace Common Shiner Central StonerollerBluntnose Minnow Plains Topminnow
Species
Figure 2-11. Mean percent passage (± 1 SE) of 14 lotic species in fish ladder design 1 from trial 3 of the experimental study. Letters depict results of Tukey’s HSD among individual species. Means with the same letter are not significantly different. Study was conducted in summer 2014 at South Dakota State University’s Agricultural Experiment Station in Brookings, South Dakota.
65
100
A
80
B 60 B
40
Percent passage (%)
20
0
Both
Black Hills
Northern Glaciated Plains
Ecoregion
Figure 2-12. Mean percent passage (± 1 SE) of fishes in trial 3 of the experimental study grouped by South Dakota ecoregion (Black Hills, Northern Glaciated Plains, and both). Letters depict results of Tukey’s HSD among ecoregion. Means with the same letter are not significantly different. Study was conducted in summer 2014 at South Dakota State University’s Agricultural Experiment Station in Brookings, South Dakota.
66
100
A
80 A A
B
60
B
40
Percent passage (%)
20
0 Fast Moderate Slow/none General
Water movement guild
Figure 2-13. Mean percent passage (± 1 SE) of fishes in trial 3 of the experimental study grouped by water movement guild (Poff and Alan 1995). Letters depict results of Tukey’s HSD among water movement guilds. Means with the same letter are not significantly different. Study was conducted in summer 2014 at South Dakota State University’s Agricultural Experiment Station in Brookings, South Dakota.
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Chapter 3: Phenology and population dynamics of Central Stoneroller Campostoma anomalum across a latitudinal gradient This chapter is being prepared for submission to Transactions of the American Fisheries Society and was co-authored by Brian D. S. Graeb, Keith B. Gido, Jacob F. Schaefer, and Katie N. Bertrand.
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Abstract
Increases in temperature associated with climate change will alter the timing and availability of resources and challenge the physiological tolerances of organisms resulting in species distribution shifts, altered phenology, behavior, and performance. Grazers can control primary producers yet it is unclear how climate change will affect grazer performance or interact with photoperiod. Using latitude as a surrogate for climate, we evaluated the dynamic rate functions, body condition, and timing of peak reproductive condition (GSI) of Central Stoneroller from three latitudes. We collected 50 fish from three reaches in each of the approximate latitudes: 44.5° (South Dakota), 39.1° (Kansas), and 32.0° (Mississippi). Recruitment was consistent along the latitudinal gradient, and mortality was similar at all latitudes with all fish dying after age 2 with the exception of 3 fish in SD and 1 fish in Kansas that lived past age 3. Growth also was similar among latitudes. Body condition was similar between South Dakota and Kansas; however,
Central Stonerollers in Mississippi had significantly lower condition. Peak GSI occurs later in the north than in the south; Stoneroller GSI peaked in MS at 21% on Julian day
65, whereas SD GSI peaked at 15% on Julian day 134. This research enhances the understanding of Central Stoneroller across their range and provides insight into climate change effects on a strong interactor in stream ecosystems, with important implications for other grazer-producer interactions.
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Introduction
Alterations in temperature associated with climate change will profoundly affect ecosystems by simultaneously altering the timing and availability of resources and challenging the physiological tolerances of organisms (Fuller et al. 2010, Somero 2010).
Changes in thermal regimes will shift species distributions, and alter phenology (e.g., flowering, breeding, migration; Parmesan 2006), behavior, and performance (e.g., forage efficiency resulting in growth) of resident species (Angilletta et al. 2002). Single-site experiments have established that grazers (e.g., bison) control primary producers via direct and indirect pathways (e.g., Fahnestock and Knapp 1993). The metabolic theory of ecology predicts that grazers in warmer environments could have greater effects on an ecosystem than grazers in cold environments (Brown et al. 2004). Alternatively, grazer thermal performance curves may be locally adapted (Richter-Boix et al. 2015) through shifts in thermal optima and sensitivity that normalize grazer effects on local environments. Under warming scenarios, local adaptation (in contrast to metabolic theory of ecology) predicts that warming will cause grazers at high latitudes with lower thermal optima, increased growth, and delayed maturation, to become less efficient at regulating primary production.
Long-term climatic data and general circulation models suggest that global mean annual air temperature is likely to increase over 1.5°C by the end of the 21st century
(IPCC 2007), which will alter stream thermal regimes, resource quality and availability, hydrology and species distribution (Perkins et al. 2010; Poff and Allan 1995). Locally adapted thermal optima or sensitivities or constraints to plastic responses (Fuller et al.
70
2010) may disrupt phenology and have cascading effects through food webs (Terblanche et al. 2009; Crowther and Bradford 2013).
Climate change has altered the phenology of some species and has resulted in a match-mismatch between primary producers and those that feed on them (Parmesan
2006). For example, Winder and Schindler (2004) observed a developing asynchrony between peak zooplankton and peak phytoplankton production as a result of climate change increasing temperatures with photoperiod remaining constant. Therefore, reproductive condition is important to evaluate as fecundity may change as a result of climate change and produce earlier year-classes that are not in synchrony with availability of food sources.
Dynamic rate functions (i.e., recruitment, growth, and mortality) affect the ecology and management of fishes (Isley and Grabowski 2007; Maceina and Pereira
2007; Miranda and Bettoli 2007); however, these processes are rarely studied in small lotic systems on non-game species as a result of the cost and time associated with estimating ages from hard structures (DeVries and Frie 1996; Quist and Guy 2001).
Climate change may a profound effect on the dynamic rate functions as temperature and precipitation have been shown to strongly influence recruitment in some species (Phelps et al. 2008) and temperature drives metabolism which ultimately affects growth (Brown et al. 2004). An understanding of factors influencing the population dynamics of non- game species across their range is essential for successful management and conservation as we are faced with a growing number of threatened and endangered species (Durham and Wilde 2008) along with environmental change (i.e., climate change). Without such information, our ability to forecast climate change effects on grazers, and grazer effects
71
on ecosystems remains limited. Range-wide understanding of population dynamics in a broadly distributed species could help predict future responses to climate change.
The Central Stoneroller Campostoma anomalum is distributed throughout central and eastern North America and Canada (Lee et al. 1980). The Central Stoneroller is a benthic grazer feeding primarily on algae and is typically found in small streams with substrates of rock and sand (Miller 1962; Power et al. 1985; Quist and Guy 2001). They have been shown to be extremely important to the structure and function of food webs in their ability to dramatically reduce algal biomass and alter algae composition (Power and
Matthews 1983; Stewart 1987; Gelwick and Matthews 1992). Though shown to be a driver of ecosystem processes (i.e., nutrient dynamics, and benthic community structure;
Power et al. 1988, Matthews 1998), little research on population dynamics exists
(Bisping et al. 2010). With such a large range, species such as Central Stoneroller are found in a variety of environments that span multiple climates (Power and McKinley
1997). Climate has been shown to be a driver in the population structure of Brassy
Minnow Hybognathus hankinsoni, another algivorous minnow species (Falke et al.
2010), and with the impact Central Stonerollers have on their surrounding environment, it is imperative to understand the effect climate has on this species.
Understanding Central Stoneroller dynamic rate functions across a latitudinal gradient is needed for the management, conservation, and the prediction of climate change effects on fisheries. Our overarching goal was to quantify the effects of warming on population dynamics (i.e., recruitment, growth, and mortality) along with body condition and reproductive phenology (timing of peak reproductive condition) across a latitudinal gradient to predict the effects of warming on Central Stonerollers. The
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objective of this study is to assess Central Stoneroller population dynamics in three streams at three latitudes using age structure analysis, growth analysis, length-weight relations to evaluate body condition, and the gonadal somatic index to evaluate reproductive condition and timing of peak spawn.
Methods
Study area
To quantify variation in Central Stoneroller population dynamics (i.e., recruitment, growth, and mortality), body condition, and reproductive condition across latitudes, we collected stonerollers from each of 3 sites within each of three latitudes in spring 2014. We sampled three reaches in each of the approximate latitudes: 44.5°
(hereafter South Dakota), 39.1° (hereafter Kansas), and 32.0° (hereafter Mississippi).
Two reaches in South Dakota were in tributaries to the Big Sioux River (Medary Creek and Peg Munky Run) and one reach was a tributary to the Minnesota River (South Fork
Whetstone River) (Figure 3-1). All three streams consisted of sand or gravel substrate.
The Big Sioux and Minnesota drainages are considered coteaus, consisting of rolling hills with prairie pothole lakes scattered throughout (Bryce et al. 1998). Historically, native short and tallgrass prairie covered the landscape which is now dominated by cattle grazing and rowcrop agriculture (Krause et al. 2013). Mean ambient temperatures by month range from -5.1°C in January to 27.6°C in July. All three reaches in Kansas
(Carnehan Creek, Deep Creek, and Kings Creek) are tributaries in the Kansas River basin
(Figure 3-1) and are located on Konza Prairie. Konza Prairie is grassland with topography of ridges and valleys overlaying limestone bedrock (Gray 1997). The area has never been plowed though it was historically used for grazing (Gray 1997). Streams on Konza Prairie
73
are dominated by cobble, pebble, and gravel substrate (Bertrand and Gido 2007). Mean ambient temperatures by month range from -2.7°C in January to 26.6°C in July (Gray
1997). Of the reaches in Mississippi, two were within James Creek and one was in
Widows Creek all of which are in the Bayou Pierre drainage in southwestern Mississippi
(Figure 3-1). Mississippi streams were characterized by firm gravel substrata and surrounding landscape was historically forested, though land use change is influencing the area and increasing erosion (Ross et al. 2001). Mean ambient temperatures by month range from 14.7°C in January to 32.7°C in July and August.
Fish collection
In South Dakota, fish were collected in a single electrofishing pass using a backpack electrofisher (Engineering Technical Services model ABP-3-300, University of
Wisconsin, Madison) while sampling in an upstream direction. Using dip nets, two netters collected stunned fish. In Kansas and Mississippi, fish were collected using both electrofishers and seines. Seines were used in areas where fish abundance was low and shocking was inadequate. For age structure analysis to assess recruitment, growth, and mortality, 50 Central Stonerollers were collected during April 2014 at 3 sample reaches
(Peg Munky, Medary, and South Fork Whetstone) in South Dakota, 30 to 60 Central
Stonerollers were collected during March 2014 at 3 sample reaches (Kings, Carnehan, and Deep) in Kansas, and 50 Central Stonerollers were collected in February 2014 at 3 sample reaches (Widows and 2 locations on James) in Mississippi. Fish at all sites in all states were euthanized with tricaine methane sulfonate (MS-222) and frozen. For reproductive condition (gonadal somatic index, hereafter GSI) analysis, 20 Central
Stonerollers were collected every two weeks at all sites from mid-February (in
74
Mississippi) through June (in South Dakota). All fish were euthanized with MS-222 and preserved in 10% formalin.
Laboratory
Fish were measured (mm standard length; (SL), and weighed (0.01 g). For age structure analysis, lapillus and asteriscus otoliths were removed and stored in vials until dry. Once dry, otoliths were mounted on microscope slides with super glue. Using wetted
1000 grit sandpaper, we polished mounted otoliths. As a result of poor annuli formation on asteriscus otoliths, we used lapillus otoliths for aging. Two readers independently examined and estimated ages. If disagreements existed, otoliths were re-examined until an agreement was reached. Consensus ages were used for all analyses. For GSI analysis, gonads were removed and weighed (0.01 g) and organs were removed from fish and fish were weighed (0.01 g) to obtain an eviscerated (somatic) mass. Eviscerated mass was used in calculating GSI.
Recruitment, Growth, and Mortality
Age frequency histograms were developed to assess Central Stoneroller age structure. We calculated mean length-at-age for each latitude (state) and each reach. We evaluated mean length-at-age by calculating 95% confidence intervals for each state and reach. Means were considered different if confidence intervals were not overlapping. We did not analyze age-3 fish because of low numbers.
Body Condition
Analysis of covariance (ANCOVA) was used to test for differences in slope of log10 transformed length and weight among states. Weight was used as the dependent variable and length and state were covariates.
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Reproductive Condition
Internal organs were removed and we measured eviscerated body mass to reduce variability as a result of stomach contents. Reproductive condition was evaluated by calculating GSI (gonad mass/eviscerated body mass*100). GSI was plotted by Julian date to assess peak gonad development.
Results
Central Stoneroller age ranged from 1 to 3. No missing year classes were found within states among observed ages; however, individuals older than age 2 were only found at Peg Munky Run and Kings Creek in South Dakota and Kansas, respectively
(Figure 3-2). Between 87 and 89% of the Kansas and Mississippi fish collected were age-
1 and between 11 and 12% were age-2 whereas 66% of fish collected were age-1 fish and
32% were age-2 fish in South Dakota (Figure 3-2). Maximum age of Central Stoneroller across all sample sites was 3, though very few individuals survived past 2 (n = 4) and no fish collected in Mississippi were estimated to be older than 2 years of age (Figure 3-2).
There was no difference in Central Stoneroller mean length-at-age among states
(Figure 3-3). Though not statistically significant, trends did exist in mean length-at-age.
In general, mean length at age was greater in the north than in the south. The two populations with the highest mean length-at-age were in South Dakota and both populations in Mississippi had the lowest mean length-at-age, with the exception of
Kings Creek in Kansas, which was the lowest (Figure 3-4). There was some variability in mean length at age-1 among creeks within latitudes. Age-1 fish were approximately 18% longer in the Whetstone River than in Medary Creek in South Dakota, and age-1 fish
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were approximately 20% longer in Deep Creek than Kings Creek in Kansas. There were no differences among creeks in mean length at age in Mississippi (Figure 3-5).
We found variability among latitudes in the effect of length on weight. There was a significant interaction between the effects of length and state on weight (ANCOVA;
F2,420 = 12.21, P < 0.01). South Dakota and Kansas fish weights responded similarly to length however, Mississippi fish increased less in weight to length than Kansas or South
Dakota fish resulting in a flatter regression slope (Figure 3-6) and indicating poorer condition.
GSI peaked when air temperatures reached 12°C, regardless of latitude (Figure 3-
7), and thus peak GSI occurred 69 days later in South Dakota than in Mississippi.
Stoneroller peak GSI in MS was 22% of body mass on day 65, peak GSI in KS was 13% on day 100, and SD GSI peaked at 15% on day 134 (Figure 3-8).
Discussion
Results from our study show that the population dynamics of Central Stoneroller are similar across latitudes and body condition is greater in higher-latitude populations.
Results from our study also demonstrate that the temperature for peak reproduction is fixed across latitudes for stonerollers, a dominant stream herbivore (Power et al. 1985).
Gonad mass as a function of body mass (GSI) peaks when air temperatures reach approximately 12°C, regardless of Julian date (Figure 3-7). The differences in magnitude of GSI are consistent with a latitudinal cline in energy allocation to growth vs. reproduction, producing larger and longer lived individuals at high latitudes
(temperature-size rule: Arendt 2011).
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An ectotherm’s body temperature affects physiological reaction rates (Huey and
Kingsolver 1989) making ambient temperature arguably the most important ecophysiological variable. Climate change will alter thermal regimes; therefore, higher mean annual air temperatures and earlier warming will lead to changes in phenology (i.e., spawning time; Parmesan 2006) and performance (i.e., growth; Angilletta et al. 2002).
Theoretically, because temperature for peak reproduction is fixed, increases in mean annual air temperature leading to shorter winters will cause stonerollers to reproduce earlier which may lead to a mismatch between larval stonerollers and necessary resources. Mismatched timing of yearly events has been observed for some species
(Parmesan 2006). For example, Inouye et al. (2000) observed yellow-bellied marmots
Marmota flaviventris emerge from hibernation by up to 23 days sooner because of a rise in local temperature of 1.4°C even though date of snowmelt and emergence of flowering plants stayed constant. Additionally, in the past 20 to 140 years, 59% of 1,598 species studied by Parmesan and Yohe (2003) had altered phenology and/or range with advancements in phenology up to 2.3 days per decade (Parmesan and Yohe 2003).
Central Stoneroller recruitment and mortality were similar among latitudes, although no Mississippi fish were found over age two. Bisping et al. (2010) found the maximum age of Central Stoneroller in Iowa streams to be four years of age, Quist and
Guy (2001) found the maximum age of Central Stonerollers in Kansas streams to be three years of age with 97% of the fish age-2 or less, and Gunning and Lewis (1956) found the maximum age of Central Stoneroller in Illinois to be three years with 77% of the fish age-
2 or less. For our northern two latitudes, our results are most similar to Quist and Guy
(2001) as we only found four age-3 individuals. Cold climates tend to lower mortality in
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some species (Angilletta et al. 2004; Yamahira et al. 2007). We did find that age-3
Central Stoneroller were only found in our northern two latitudes suggesting that this species does have lower mortality in northern areas of its range, however, because of low sample size, age-3 stonerollers may exist at other locations in southern latitudes and local factors within latitude may have biased mortality results.
We found no difference in growth among latitudes. Along with Central
Stoneroller in our study, many other fishes show no significant relationship between latitude and body size, thus suggesting a mechanism to compensate for shorter growing season exists in some fishes (Belk and Hourston 2002). Conover and Present (1990) found that Atlantic Silversides Menidia menidia in northern latitudes had higher growth rates than those in southern latitudes. Similar results were found by Jensen and Johnsen
(1986) for juvenile Atlantic Salmon Salmo salar, Power and McKinley (1997) for Lake
Sturgeon Acipenser fulvescens, and Jonassen et al. (2000) for juvenile Atlantic Halibut
Hippoglossus hippoglossus. These relationships demonstrate what has been termed
“countergradient variation” by Conover (1990) which suggests that an inverse relationship between length of growing season and growth may exist in some species
(Conover 1990). We can only speculate about the mechanism behind similar growth rates across the latitudinal gradient in our study. First, it is possible that metabolic rates of high-latitude populations are adapted to be higher at a given temperature than low- latitude (warmer climate) populations (Terblanche et al. 2009). Clarke (1991) explained
‘metabolic cold adaptation’ as compensation by ectotherms growing in colder climates at rates similar to warmer climates. Second, similar growth rates could be the result of adaptations to winter mortality. Conover (1990) suggests that faster growth rates in
79
higher latitudes may reduce size-selective overwinter mortality. Higher mortality of smaller fish may then favor faster growing fish with increasing latitude (Jonassen et al.
2000). Third, northern latitude fish may also remain closer to optimal temperature for
Central Stoneroller growth for longer periods of time thus resulting in increases in season-long growth relative to southern latitude fish (Conover 1990). Two models of latitudinal growth compensation have been proposed by Yamahira and Conover (2002).
Models include a species adaptation to either temperature or growing season. An adaptation to temperature would imply similar growth among latitudes with maximum size being the same; however, optimal growth would be locally adapted to a temperature typically found in their native environment so that colder temperatures would allow for faster growth in northern latitude fishes and vice versa for southern latitude fishes
(Yamahira and Conover 2002). An adaptation to growing season would imply that a species grows over the same temperature range, but high latitude fishes can grow faster within that same temperature range to compensate for a shorter growing season
(Yamahira and Conover 2002). A more straightforward reason has been put forward by
Power and McKinley (1997) who suggests that food quality or quantity may be higher per individual in northern latitudes when summer growth is taking place.
Growth is not only latitude based and therefore local factors within latitude among streams are affecting growth. We did observe this as age-1 Whetstone River and
Medary Creek Central Stonerollers in South Dakota had significantly different lengths.
This was also the case for Deep Creek and Kings Creek in Kansas. Felts et al. (2014) found growth differences in Northern Pearl Dace Margariscus nachtriebi among four southwestern South Dakota streams and suggested that differences could exist due to
80
local differences in food supply, habitat quality, biotic interactions, or genetics. Despite similar growth, we found differences in length-weight relationships among latitudes.
Northern and mid-latitude fish appear to be in better condition having added more weight relative to length than southern-latitude fish.
We found temperature for peak reproduction to be fixed across latitudes in
Central Stoneroller suggesting that climate change will advance spawning time by earlier spring-time warming. Though no differences in recruitment, growth, or mortality were found for Central Stoneroller among latitudes, age-3 fish only present in the northern two latitudes and lower body condition of southern latitude populations suggests that climate change may negatively impact southern populations more than northern populations as they are already on the southern extent of their range. Adaptations to colder climate for northern populations of Central Stoneroller seem to exist as similar growth was observed among latitude despite vast differences in growing season. Additional research with larger sample size on the latitudinal differences within species would expand the knowledge and implications of climate change.
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Yamahira, K., and D. O. Conover. 2002. Intra- vs. interspecific latitudinal variation in
growth: adaptation to temperature or seasonality? Ecology 83(5):1252-1262.
Yamahira, K., M. Kawajiri, K. Takeshi, and T. Irie. 2007. Inter- and intrapopulation
variation in thermal reaction norms for growth rate: evolution of latitudinal
compensation in ectotherms with a genetic constraint. Evolution 61(7):1577-1589.
Figures
Figure 3-1. Map of states and basins Central Stoneroller were collected in 2014. Basins included: Minnesota River basin (northern
basin in South Dakota), Big Sioux River basin (southern basin in South Dakota), Kansas River basin (Kansas), and Bayou Pierre River 88
basin (Mississippi).
140
120
100
80 SD KS 60 Numberof fish MS
40
20
0 1 2 3 Age (years)
Figure 3-2. Age frequency of Central Stoneroller collected in South Dakota, Kansas, and Mississippi, 2014.
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120
SD 110 KS 100 MS
90
80
70
60 Standardlength(mm)
50
40
30
20 1 2 3 Age (years)
Figure 3-3. Mean length-at-age (mm ± 95% CI) of Central Stoneroller collected from South Dakota, Kansas, and Mississippi.
90
100 Medary Creek
Peg Munky Run
90 Whetstone River
Carnehan Creek
Deep Creek 80 Kings Creek
James Creek
70 Widows Creek Standardlength(mm) 60
50
40 1 2 3 Age (years)
Figure 3-4. Mean length-at-age of Central Stoneroller from South Dakota (Medary Creek, Peg Munky Run, and Whetstone River),
Kansas (Carnehan Creek, Deep Creek, and Kings Creek), and Mississippi (James Creek and Widows Creek). 91
110 South Dakota Kansas Mississippi 100
90
80
70
60
Standard length (mm) length Standard Medary Creek Carnehan Creek 50 Peg Munky Run Deep Creek James Creek Whetstone River Kings Creek Widows Creek 40 1 2 3 1 2 3 1 2 3 Age (years)
Figure 3-5. Mean length-at-age (mm ± 95% CI) of Central Stoneroller collected from sites within South Dakota, Kansas, and Mississippi.
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Figure 3-6. Central stoneroller Log10 weight by log10 standard length for three latitudes (states).
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Figure 3-7. Mean daily air temperature (°C) by Julian date for three latitudes (states). Location of arrows represents Julian date on which peak female GSI occurred in each state [note similar temperature (horizontal line) of peak female GSI].
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30 Mississippi Kansas 25 South Dakota
20
15
10
Female GSI
5
0
40 60 80 100 120 140 160 180
Julian date Figure 3-8. Mean female GSI (± SE) estimated semi-weekly in 3 streams in each of 3 latitudes (states). Data were pooled at streams within latitudes.
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Chapter 4: Conclusion
Summary
Increases in temperature associated with climate change will alter the timing and availability of resources and challenge the physiological tolerances of organisms resulting in species distribution shifts, altered phenology, behavior, and performance. Because climate forms a gradient of cold to warm as latitude decreases, latitude can be used to evaluate differences among populations from different climates and make future predictions about populations under warming scenarios.
We evaluated the dynamic rate functions, body condition, and peak reproductive condition (GSI) of Central Stoneroller Campostoma anomalum collected from three latitudes. We sampled three reaches in each of the approximate latitudes: 44.5° (South
Dakota), 39.1° (Kansas), and 32.0° (Mississippi). All latitudes contained no missing year classes demonstrating consistent recruitment and all but three individuals in South
Dakota and one individual in Kansas died before age three resulting in similar mortality.
We found growth similar among latitudes. Differences in growth within latitude clouded the comparison among latitudes; therefore, biologically, growth may be faster for Central
Stoneroller in northern latitudes though we did not have the sample size necessary to elucidate this. Body condition was similar among the upper two latitudes, however, stonerollers in southern latitudes increased in weight per unit length at a slower rate suggesting that southern populations of Central Stonerollers are in poorer condition which may be attributed to thermal stress from being located on the southern extent of their range. Peak GSI occurred later in the north than in the south; stoneroller peak GSI in
MS was 21% on day 65, whereas SD GSI peaked at 15% on day 134. This suggests that
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stoneroller gonad development and spawning is adapted to temperature as opposed to day length. This may lead to advances in spawning time and create a mismatch between larval stonerollers and their environment.
As climate change becomes more evident, species distributions may have to shift in order to survive. Stream-road crossings that form barriers will exacerbate the effects of climate change as they will not allow range shifts to accommodate for changing climates and may lead to declines in fish populations and possible species loss. Stream-road crossings are present worldwide and are very abundant in the United States and other developed countries. Many existing stream-road crossings are barriers to fish movement and as human population increases; roads will expand and add more potential barriers to stream networks. Cost of stream-road crossings will always favor culverts which are the most prone to become barriers, and in many situations, improper installation without addressing biological needs will lead to the continuous addition of barriers to fish movement.
One of the main obstacles to fish movement past a culvert stream-road crossing is a result of perching. Perching of the downstream side of the culvert forms a waterfall
(drop culvert), and becomes a barrier when a fish cannot leap high enough to overcome the drop. It is therefore important to create solutions for existing and future drop culverts.
The solution for drop culverts we developed and researched was a fish ladder designed to retrofit culverts, eliminating the drop and allowing passage.
We conducted lab and field studies to evaluate a fish ladder designed to retrofit perched culverts. In the lab, when altering baffle profile (flat-top, v-top, and no baffles) we found that the flat-top baffle design passed lotic fishes found in South Dakota at a
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higher mean passage rate than the other two designs. When evaluating passage rate of stream fishes found in South Dakota at three different baffle slot widths (wide, moderate, and narrow) within the flat-top baffle fish ladder we found no difference in passage rate.
This suggested that this fish ladder design would be able to function and pass fish in a variety of velocity conditions (as velocity increased with decreasing baffle slot width).
All species were able to pass the flat-top baffle fish ladder though species were observed passing at different rates. We do not know if fish were attempting and failing at passing or not attempting at all, therefore differences in passage rate could be a result of physiological abilities or behavior. In the field, ladders were able to attach to any culvert design or material we encountered. Fish were observed passing ladders at all sites though passage rates differed among sites. We measured factors that would influence passage and found no relationship between any of the factors and passage rate among sites suggesting that this ladder design was able to function similarly under all conditions. We did observe a correlation in daily discharge and daily passage rate suggesting that rainfall events may trigger upstream movement for fishes. We speculate that changes in daily discharge did not affect fish ability to pass ladders but rather it was behavioral, providing a motivation to move upstream.
Management recommendations
Central Stoneroller may become extirpated from the southern extent of their range as a result of climate change. Southern populations will need to disperse and therefore, stream connectivity is vital. We recommend mitigating stream fragmentation in watersheds where stonerollers are present. In addition, we recommend continued monitoring of abundance and size structure of stonerollers which will inform future
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managers of population health. We do not know if advancements in spawning time will negatively impact stoneroller populations or their surroundings through match- mismatched interactions, however, it is something managers should be aware of as future management actions such as habitat enhancement may need to take place.
Habitat fragmentation from stream-road crossings is detrimental to fishes and will exacerbate the effects of climate change. Our fish ladder design provides a tool for fisheries managers to mitigate this fragmentation and allow passage over drop culverts so fishes have access to habitat located longitudinally in streams. Because many stream-road crossings are constructed and installed improperly resulting in barriers, we recommend that proper inspection of newly installed stream-road crossings take place so that they allow fish passage. Because our fish ladder was successful both in experimental and field trials, we recommend it to be used on existing and future drop culverts to allow fish passage. We also recommend stream-road crossings surveys to determine high-priority sites for fish ladder installation (i.e., presence of species of greatest conservation need, suitable upstream habitat). Finally, until ladder durability is determined, we recommend that ladders be used as a temporary tool to be deployed during times when fish are moving upstream and taken out during winter months, or times of year when upstream passage is low.
Research needs
This research contributed to the understanding of the dynamic rate functions, body condition, and phenology of peak gonad development in an ecologically important stream fish, the Central Stoneroller along with contributing to the understanding of climate change on this species. Additional research would enhance the understanding of
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the effects of climate change on lotic species and provide more clarity to fully grasp what the future holds. More research is required to fully understand the differences in the dynamic rate functions of Central Stoneroller across their range. A larger sample size is required to reduce variation among sites within latitude. If a larger sample size is used, differences in these metrics may be found to exist.
This research also developed a management tool for fisheries professionals to eliminate habitat fragmentation from drop culverts. Additional research and design on future prototypes of this fish ladder may result in increased passage or an understanding of why some fish were not observed using our design in the field. Future researchers could explore the use of passive integrated transponder (PIT) tags to evaluate fish ladder attraction, and actual passage efficiency of fish ladders. Future researchers also need to evaluate durability as we do not know for sure the life span on these structures. Ladder strength may need to increase if they are left to overwinter in streams.