Complex to simple: Fish growth along the Illinois River network
Introduction
Many environmental drivers influence fish growth within riverine landscapes (Brett, 1979). This includes biological and physical factors (e.g., Neuheimer and Taggart, 2007; Korman and Campana, 2009; Zeug et al., 2019), as well as variations in resources and stress (e.g., Johnston et al., 1990; Adams et al., 1996; Morrongiello et al., 2013), all of which vary spatially within a river network (Fig. 1). Riverine fish populations commonly exhibit spatial structure, because vital rates including survival, growth, and reproduction differ among local and regional patches (Thomas and Kunin, 1999; Tsuboi et al., 2020); which can be enhanced or constrained by local movement and migration. As a result, a diverse portfolio of life-history traits – including growth rates (cf. Schindler et al., 2010) – exists along river networks.
Globally, river ecosystems are subject to a multitude of anthropogenic stressors including channelization, hydrological modifications, land-use changes, and urbanization (Best, 2019; Sabater et al., 2019). These stressors can act individually or collectively to influence fish growth in river ecosystems. Large-scale anthropogenic stressors also have the potential to ‘flip’ river ecosystems into another regime or state (sensu Resilience Thinking, Holling, 1973) from which recovery cannot occur (Erős et al., 2019). Anthropocene Rivers (sensu Kelly et al., 2018) exhibit different ecosystem structure and function compared to natural rivers (Thoms et al., 2018). Studies on the highly modified Illinois River (Illinois, USA), for example, show pronounced and persistent changes in ecosystem structure (i.e., Shannon diversity of fish; DeBoer et al., 2019) and function (i.e., trophic status and food web character; DeBoer et al., 2020) in response to many human-induced stressors over 100 years. These ecosystem changes are indicative of the new Anthropocene regime. Anthropocene Rivers are inferred to also exhibit a loss of heterogeneity across many scales (Padial et al., 2020) resulting in a homogenization of ecosystem function, including life-history traits like growth (Moore et al., 2010). A diverse portfolio of life-history traits confers population stability (Schindler et al., 2010) and resilience to environmental change (Greene et al., 2010). This is increasingly important in the Anthropocene (sensu Reed et al., 2010), when large-scale and irreversible changes to river ecosystems are becoming common (Kelly et al., 2018; Erős et al., 2019). At present, we know very little about the character of life-history traits like fish growth along Anthropocene Rivers.
Heterogeneity of ecosystem structure and function is an inherent feature of riverine landscapes, driven, in part, by a diverse physical template (sensu Southwood, 1977; 1988). Heterogeneity is spatial variation in the environment (White and Brown, 2005). The River Continuum Concept (RCC, Vannote et al., 1980) suggests fish growth should differ systematically downstream (e.g., Tedesco et al., 2009; Kelly et al., 2016), whereas the Flood Pulse Concept (FPC, Junk et al., 1989) hypothesizes fish growth should differ within river networks as result of differential lateral connections (e.g., Gutreuter et al., 1999; Phelps et al., 2015). The Riverine Ecosystem Synthesis (RES, (Thorp et al., 2010)) infers fish growth should differ among functional process zones (FPZs) - large-scale lengths of a river network with a similar hydro-geomorphological character. FPZs have distinct riverbed habitats (Collins et al., 2014), fish communities (Boys and Thoms, 2006; Elgueta et al., 2019), benthic macroinvertebrate communities (Bellmore and Baxter 2014; Elgueta et al., press), and food web character (Thoms and Delong, 2018). Moreover, according to the RES, heterogeneity in fish growth should occur along Anthropocene Rivers, because FPZs have different capacities to absorb disturbances (i.e., different resilience; Walker and Salt, 2012).
Modifications to the physical environment of river ecosystems can result in evolutionary changes to the character of important life-history traits (Stearns, 1992; Wootton, 2012). These modifications can also change interactions between the physical environment and ecosystem function (cf. Thoms et al., 2017; Thoms and Delong, in review). Anthropogenic stressors can simplify riverine landscapes, homogenize riverine communities, and favor nonnative fishes (Olden et al., 2004; Padial et al., 2020). However, the effect of physical simplification on life-history traits like fish growth in Anthropocene river ecosystems remains unclear (Pyron et al., 2019; Sturrock et al., 2020). Our aim is to examine the character of fish growth along the entire main channel of an Anthropocene River. We predict: 1) a diverse portfolio of fish growth will be maintained along the Anthropocene Illinois River, 2) fish growth in the Anthropocene Illinois River will differ from other river ecosystems, and 3) a positive association between FPZ physical heterogeneity and fish growth will be maintained in the Anthropocene Illinois River.
Section snippets
Study area
The Illinois River is a multi-use waterway connecting the Great Lakes at Chicago, Illinois with the Mississippi River at Grafton, Illinois (Lian et al., 2012) (Fig. 2).
The main stem of the Illinois River is approximately 439 km long, and has a watershed area of 72,701 km2, nearly 44% of the land area of Illinois (Talkington, 1991). Four major stressor phases have been identified for the Illinois River Basin: the Settlement, Degradation, Recovery, and Biological Invasion phases (
Methods
Four species of potamodromous fish were collected along the length of the Illinois River, from each FPZ (Fig. 2). These fishes are commercially and recreationally important: bluegill (Lepomis macrochirus), white bass (Morone chrysops), channel catfish (Ictalurus punctatus), and bigmouth buffalo (Ictiobus cyprinellus). They are also representative of four functional feeding guilds (FFGs): bluegill for invertivores, white bass for piscivores, channel catfish for omnivores, and bigmouth buffalo
Results
The majority (7 of 12) of species-growth metric combinations examined did not differ among FPZs (Table 1). k and L∞ for bluegill and white bass did not differ among FPZs (Table 1, Fig. 3, Fig. 4), and L∞ for channel catfish and bigmouth buffalo did not differ among FPZs (Table 1, Figs. 3 and 4). Only channel catfish GI did not differ among FPZs; GI of bluegill, white bass, and bigmouth buffalo differed among FPZs (Table 1, Fig. 5). Of the five species-growth metric combinations that differed
Discussion
The portfolio of growth metrics recorded for the Anthropocene Illinois River display a limited diversity. Fishes from all four FFGs exhibited a similar single growth character: faster growth rates (k) and smaller maximum sizes (L∞) (cf. Table 2). This limited portfolio for the different FFGs likely indicates a stressful environment for fishes that constrains life-history expression (sensu Fox and Keast, 1991) and one that constrains the ecosystem's capacity to maintain viable populations (sensu
Conclusions
Human alteration of river ecosystems has simplified their natural physical heterogeneity and homogenized ecosystem function, thus undermining their resilience (Nilsson et al., 2005). Moreover, the influence of heterogeneity on ecosystem resilience is an emerging issue in river science and management. Heterogeneity is especially important in the context of mitigation and rehabilitation (not “restoration”, given the irreversibility of certain anthropogenic changes; Erős et al., 2019). Enhanced
Author statement
Jason DeBoer – Conceptualization, Methodology, Investigation, Formal analysis, Writing - Original Draft, Writing - Review & Editing
Martin Thoms – Conceptualization, Methodology, Formal analysis, Writing – Original Draft, Writing - Review & Editing
James Lamer - Formal analysis, Writing - Original Draft
Andrew Casper – Conceptualization, Writing - Original Draft
Michael Delong – Methodology, Writing - Original Draft
Declaration of Competing Interest
The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.
Acknowledgements
We thank Sabina Berry for processing channel catfish spines, Andrew Weiland for processing otoliths, Levi Solomon for acting as second reader for age estimation, and Travis Brenden and Derek Ogle for statistical consultation and code refinement. JDB was supported by an Australian Government Research Training Program (RTP) Scholarship from the University of New England. The authors declare that they have no known competing financial interests or personal relationships that could have appeared to
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