1. Introduction
River or stream flooding (i.e., out-of-banks flow) is a common phenomenon worldwide, brought on naturally by heavy rainfall or extensive snowmelt [
1], or artificially by managed releases from reservoirs [
2]. Flooding can occur in a wide range of systems, from tropical rivers [
3] to desert streams [
4], from high mountain streams [
5] to lowland coastal rivers [
3,
6,
7], and from high-gradient low-order headwaters [
5,
8,
9] to high-order wide-floodplain systems [
10,
11]. Some rivers and streams are prone to frequent, recurring flooding [
5,
8], whereas others rarely experience floods. Floods also exhibit extreme variability, in both magnitude (total discharge and/or elevation change relative to baseflow) and duration (time spent out-of-banks) [
1,
12].
Floods can dramatically alter the environments of both rivers [
13] and their riparia [
14], impacting both physical and biological components of aquatic habitats and their adjoining terrestrial corridors [
3,
8,
9,
15]. These effects can be either negative or positive, depending on flood variables, location within the stream or river system [
11,
16], and the physical or biological stream/riparian characteristic being examined [
12,
17,
18]. From a human perspective, small floods may have neutral to positive effects on many of the ecosystem services (the benefits humans obtain from the natural functioning of the system) provided by a river or stream, but extreme floods cause losses in virtually every ecosystem service [
19].
Extreme (also termed catastrophic or historic) floods can have truly devastating impacts on river and stream environments [
1,
5,
9,
16]. These once-in-100-years to once-in-2000-years events wreak havoc on instream and riparian physical features [
3,
9] and may have long-lasting, negative effects on aquatic biota [
1,
5,
9]. However, some biota may be resistant and/or resilient to even these drastic floods, demonstrating amazing abilities to survive scouring flows and returning to pre-flood abundances only weeks or months after floodwaters subside [
5,
8,
9].
Although many aquatic organisms can recover quickly from extreme floods, others may not. For example, species that are poor recolonizers, those that inhabit vulnerable habitats, or those that are in more sensitive life stages at the time of flooding are likely to be severely impacted by extreme floods [
8]. Such species may include benthic fishes like sculpins (Cottidae) and longnose dace (
Rhinichthys cataractae), which are considered poor recolonizers [
8,
20] and may be vulnerable to crushing by bedload movement of their cobble/boulder habitats during catastrophic flooding [
8,
21]. Young-of-year or juvenile life stages of these and other species also may be at greater risk than larger or adult fish during flood events, due to both small size and poor swimming ability [
22].
Sculpin are common fish of marine and freshwater systems, especially in circumpolar regions of the Northern Hemisphere [
23,
24]. Many species inhabit cold and coolwater rivers and streams, where they can numerically dominate fish communities [
20,
25]. The slimy sculpin (
Uranidea cognatus), with the widest geographical distribution of any sculpin species in North America [
26,
27], has been suggested as an excellent species for environmental effects monitoring due to its small size, limited mobility, early maturity, fast growth rate, high reproductive output, and high abundance [
20,
28,
29].
A once-in-2000-years flood impacted many coldwater streams and rivers across southeastern Minnesota in August 2007 [
9,
30], scouring stream habitats and reducing aquatic macroinvertebrate densities by up to 95% [
9]. Post-flood surveys on several regional streams indicated that salmonids (brook trout,
Salvelinus fontinalis and brown trout,
Salmo trutta) and slimy sculpin had survived the floods, although numbers were greatly reduced (N. Mundahl, unpublished data). However, there was concern that reduced aquatic prey resources might result in poorer fish condition, which ultimately could impact subsequent spawning success. Consequently, this study was undertaken to assess the impact of flooding on autumn diets and condition of slimy sculpin in two area streams that differed in severity of flooding and where, fortuitously, sculpin had been collected one to eight weeks prior to flooding for examination of fish condition and diets. It was hypothesized that sculpin would exhibit poorer condition and reduced diets (fewer prey and reduced prey biomass) post- versus pre-flood in both streams, with reductions greater in the stream with more severe flooding. In addition, sculpin in these two streams, plus one additional stream that also had experienced catastrophic flooding, were examined the following spring (seven months post-flood) just prior to spawning to assess diets, fish condition, and reproductive fitness. It was expected that sculpin in the two streams that had the worst flooding the previous summer would display poorer diets and condition and reduced reproductive fitness relative to sculpin in the stream that had less severe flooding.
5. Discussion
Flooding during August 2007 had severe effects on streams, rivers, and the surrounding landscape throughout southeastern Minnesota and west-central Wisconsin. Damage to human structures was extensive (
$272 million US in losses) across a broad geographic area, and flooding resulted in 10 human fatalities. Streams and rivers crested many meters above flood stage [
31], flowing with enough power to destroy bridges and roadways, lift houses off their foundations, sweep cars off roads, and obliterate agricultural crops [
37]. Rock and mud landslides blocked some streams, forcing the creation of new channels. In the aftermath of such devastation, concern eventually shifted from anthropogenic losses to the status of fish that had inhabited these streams prior to flooding [
30].
The impacts of extreme flooding on lotic fish communities can be highly variable and complicated to understand or explain [
1]. Although juvenile or small-bodied fishes may be highly susceptible to heavy losses during extreme floods in some systems [
1], in other systems this same size group of fish may be unaffected or actually increase in abundance post-flood [
16]. Habitat complexity may play a crucial role in reducing flood impacts on fish [
8]. In addition, many fish species display remarkable resistance and resilience to even extreme flooding [
2,
5,
8,
16], in fact even depending on both regular seasonal and periodic catastrophic floods to clean spawning substrates, deepen channels and pools, reduce non-native fish abundance, and reset food webs [
1,
5,
8,
16,
38]. Long-term exposure of many fish species to the variable abiotic environments of lotic systems has allowed them to evolve and/or adapt unique sets of characteristics (e.g., morphological, physiological, behavioral) that allow them to better cope with flood conditions [
1].
Despite the catastrophic flooding, surveys after the flooding in southeastern Minnesota indicated that many native brook trout and introduced brown trout had survived [
30]. Most juvenile trout (5–8 cm total length) that had hatched out five months prior to the floods were lost, but all sizes of older fish remained in the streams, although in reduced numbers [
30]. The resilience of those survivors was impressive, as they spawned two to three months after flooding and produced a record hatch of young trout during spring 2008 [
30]. The same positive, post-flood response to late summer flooding has been observed previously for brown trout, attributed to improved fall spawning conditions (larger and higher-quality gravel beds) and enhanced environments for incubating trout eggs and, later, emerging fry (due to reduced fine sediments within spawning gravels) [
5].
The presence of slimy sculpin in all streams after catastrophic flooding was encouraging, especially since small benthic species like sculpins are considered highly vulnerable to such destructive events [
8,
21,
22]. Powerful flooding in Garvin Brook and Trout Run produced severe channel scouring and displacement of boulders >40 cm in diameter, simultaneously destroying sculpin habitat and food resources. The presence of sculpin in Gilmore Creek and Garvin Brook only two weeks post-flood suggest that either fish were able to find refuges [
5,
8] that enabled them to survive mass bedload rearrangement, or they were able to quickly recolonize stream reaches from nearby, less-impacted systems [
39,
40]. Although slimy sculpin often are considered as sedentary and unlikely to exhibit more than small-scale movements [
29], there is increasing evidence that they are capable of both rapid and long-range movements, especially when recolonizing defaunated stream reaches [
39,
40,
41]. An apparent lack of less-impacted reaches in Garvin Brook (severe flooding affected the entire stream, extending from headwater springs downstream to the confluence with the Mississippi River) suggests that survival in refuges was most likely, especially since even young-of-year sculpin were large enough by August to significantly reduce their vulnerability to elevated current velocities [
5,
22]. Slimy sculpin also have the ability to squeeze into extremely tight spaces by compressing their skull widths by up to 20% [
42], which would permit them to retreat into very small openings beneath large boulders or into cracks between layers of sedimentary limestone bedrock that are common in regional streams [
43], allowing them to wait out catastrophic flows in protected spaces. A study conducted in Gilmore Creek reported that slimy sculpin were more abundant a short time after the flood than they were prior to flooding [
31].
Based on examination of slimy sculpin condition and diets before and after catastrophic flooding, it was apparent that differing severities of flooding did not impact these variables exactly as expected. Pre- and post-flood diets differed in Garvin Brook where flooding was most severe, but not in Gilmore Creek where flooding was less intense. However, sculpin in both streams consumed lower prey biomass after the flood, even though fish in Garvin Brook significantly increased the numbers of prey consumed. Condition of sculpin declined post-flood in Garvin Brook as expected, but sculpin condition improved after flooding in Gilmore Creek. Before spring spawning, sculpin condition had improved in Garvin Brook, but declined in Gilmore Creek. Just prior to spawning, sculpin from the two most severely flood-impacted streams were in better condition than those in the less-impacted stream, but no measure of reproductive fitness other than oocyte size differed among streams.
Reduced prey biomass in sculpin stomachs after flooding suggests that prey availability was limited [
9]. The dominance of sculpin diets by Chironomidae (midge) larvae was not surprising, as these organisms were the most abundant aquatic invertebrates after the flood and comprised a large proportion of the invertebrate assemblage for many months thereafter [
9]. Sculpin are opportunistic feeders, taking advantage of whatever prey are available and abundant [
36], so their ability to find and consume prey after a flood should be adequate. Differing pre- versus post-flood diets in Garvin Brook, but not in Gilmore Creek, follows expectations that more severe flooding would produce greater impacts on sculpin diets, but the increased numbers of prey in Garvin Brook sculpin stomachs relative to the pre-flood survey appears counterintuitive. Reduced condition of sculpin post-flood in Garvin Brook suggest that fish may have experienced a one-to-two-week period of food deprivation during and after flooding, followed by compensatory increases in feeding rate and prey consumption [
44]. However, the small size of midge larvae still resulted in reductions in ingested biomass post-flood relative to pre-flood feeding. Reduced body condition of sculpin has been reported previously when fish consumed smaller, lower-energy prey [
45].
As expected, condition of sculpin in flood-damaged Garvin Brook declined significantly after the flood, likely attributable to a combination of reduced prey resources and increased stress post-flood [
9,
32,
33]. In contrast, sculpin in Gilmore Creek exhibited improved condition post-flood, despite reductions in numbers and dry mass of prey in stomachs. Condition is a responsive indicator of exposure of sculpin to a variety of stressors [
29,
46], so significant reductions in condition post-flood in Garvin Brook are suggestive of stressful environmental conditions in the few weeks following floods. Environmental conditions apparently were better in less-impacted Gilmore Creek, since fish displayed improved condition after flooding. This result is difficult to explain from a diet perspective, since Gilmore Creek sculpin contained fewer prey items in their stomachs and reduced dry weight prey biomass after flooding compared to before. Perhaps the more diverse diet of Gilmore Creek sculpin, containing organisms considerably larger than midge larvae (especially caddisfly larvae, a variety of crustaceans, and small fish), may have provided nutrients sufficient to maintain and even improve overall sculpin body condition post-flood [
45].
Prior to spring spawning, temperate zone fishes in general, and sculpin in particular, direct energy resources to their reproductive organs to maximize their reproductive fitness [
47,
48]. Prey resources often are abundant and diverse during this time period [
9], although territorial behaviors among fish may result in reduced food intake in those individuals relegated to poorer habitats [
47,
49]. As cold-adapted species capable of feeding effectively at winter temperatures [
48], sculpin should lose little of their endogenous energy reserves and maintain condition through the winter months. However, reduced food (energy) intake during winter and early spring due to reduced prey resources post-flooding [
9], while reproductive organs were enlarging in preparation for spawning, could have resulted in the reduced condition factors observed for sculpin in Gilmore Creek (both males and females).
The lack of any significant differences in GSI or HSI indices, or in oocyte numbers of sculpin among the three streams examined indicates that there were no observable effects of differing flood magnitude on sculpin reproduction seven months post-flood. Although differing availabilities of prey in the various systems may have resulted in altered food intake and possible declines in overall fish condition, this was not translated into reduced reproductive fitness. Even under adverse environmental conditions, reproductive tradeoffs in fish are common [
48]. Many fish species, apparently including sculpin, can allocate significant energy reserves toward maximizing their reproductive output, even if that allocation may compromise post-spawning survival of some fish [
48]. Such allocations toward reproduction have been reported in other fish species deprived of food prior to spawning [
48].