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Intercontinental Comparison of Fish Life History Strategies Along a Gradient of Hydrologic Variability Julian D

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Intercontinental Comparison of Fish Life History Strategies Along a Gradient of Hydrologic Variability Julian D American Fisheries Society Symposium 73:83–107, 2010 © 2010 by the American Fisheries Society Intercontinental Comparison of Fish Life History Strategies along a Gradient of Hydrologic Variability Julian D. Olden* School of Aquatic and Fishery Sciences, Box 355020, University of Washington Seattle, Washington 98195, USA Mark J. Kennard Australian Rivers Institute, Griffith University, Nathan, Queensland 4111, Australia Abstract.—The flow regime is considered the primary driver of physical processes in riverine ecosystems; thus we expect that the trait composition of fish assemblages might respond similarly to hydrologic variability, even at broad spatial scales. Here, we test the hypothesis that freshwater fish life history strategies on two continents (south- ern United States and eastern Australia) converge along gradients of hydrologic vari- ability and primary productivity at the drainage scale. Our results show that the fishes of the United States and Australia conform to the three-dimensional adaptive space arising from the trade-offs among three basic demographic parameters of survival, fecundity, and onset and duration of reproductive life. Species from both continents represent the endpoints in adaptive space defining the periodic (19% versus 33% for the United States and Australia, respectively), opportunistic (69% versus 52%), and equilibrium life history strategies (12% versus 15%). We found evidence that fish life history composition of drainage basins in the two continents have converged across similar gradients of hydrologic variability and productivity despite phylogenetic and historical differences. Moreover, these relationships were largely consistent with pre- dictions from life history theory. Increasing hydrologic variability has promoted the greater prevalence of opportunistic strategists (a strategy that should maximize fit- ness in environmental settings dominated by unpredictable environmental change) while concurrently minimizing the persistence of periodic-type species (a strategy typically inhabits seasonal, periodically suitable environments). Our study provides a conceptual framework of management options for species in regulated rivers be- cause life history strategies are the underlying determinants for population responses to environmental change and therefore can be used to classify typical population re- sponses to flow alteration or mitigation via environmental flow prescriptions. Introduction nized (Naiman et al. 2008). Flow has been suggested to be the “master variable” that deter- The importance of hydrologic variability for mines pattern and process in rivers, thus limit- shaping the biophysical attributes and func- ing the distribution and abundance of species tioning of riverine ecosystems is well recog- and regulating ecological integrity (Poff et al. * Corresponding author: [email protected] 1997). Natural spatial variation in the hydro- 83 84 olden and kennard logic regime is influenced by variations in cli- Poff et al. 2006; Verberk et al. 2008; Frimpong mate and basin geology, topography, and veg- and Angermeier 2010, this volume). Species etation, which interact at multiple spatial and traits may be intercorrelated through physio- temporal scales to shape the physical template logical constraints, trade-offs (i.e., investments upon which ecological and evolutionary pro- in one trait leaving fewer resources available cesses operate in river ecosystems (Poff and for investment in another), or spin-offs (i.e., in- Ward 1990; Bunn and Arthington 2002; Lake vestments in one trait reduce costs or increase 2008). the benefits of investment in another trait), The natural flow-regime paradigm postu- resulting in the creation of life history strate- lates that the structure and function of riverine gies or tactics represented as sets of coevolved ecosystems, and the adaptations of their con- traits that enable a species to cope with a range stituent riparian and aquatic species, are dictat- of ecological problems (Stearns 1992). Com- ed by patterns of intra- and interannual varia- parative studies from a diverse array of fishes tion in river flows (Poff et al. 1997). A rich body in marine and freshwater systems have inde- of literature has demonstrated that the long- pendently identified three primary life his- term physical characteristics of flow variability tory strategies that represent the endpoints of have strong consequences at local to regional a triangular continuum arising from essential scales and at time intervals ranging from days trade-offs among the basic demographic pa- (ecological effects) to millennia (evolution- rameters of survival, fecundity, and onset and ary effects). Flow variability, including flood duration of reproduction (Winemiller 1989, and drought events, interacts with the under- 1992; Winemiller and Rose 1992; Vila-Gispert lying geology to shape the river’s physical and and Moreno-Amich 2002; Vila-Gispert et al. chemical templates and constrain assemblage 2002; King and McFarlane 2003; see Figure structure for fish (e.g., Lamouroux et al. 2002; 1). Winemiller and Rose (1992) synthesized Hoeinghaus et al. 2007; Kennard et al. 2007), these life history trade-offs and proposed the stream invertebrates (e.g., Poff and Ward 1989; following characteristic biological and habitat Vieira et al. 2004; Dewson et al. 2007), ripari- environmental attributes associated with the an plants (e.g., Nilsson et al. 1993; Naiman and three life history strategies. Periodic strategists Décamps 1997; Pettit et al. 2001), and riparian are large-bodied fishes with late maturation, invertebrates (e.g., Wenninger and Fagan 2000; high fecundity per spawning event, and low ju- Lambeets et al. 2008). Adaptations to natural venile survivorship (i.e., no parental care) and flow regimes include behaviors that enable in- typically inhabit seasonal, periodically suitable sects to avoid desiccation by droughts, fish life environments with large-space spatial (patchi- history strategies that are synchronized to take ness) and temporal (seasonality) heterogene- advantage of floodplain inundation, and plant ity. Opportunistic strategists are small-bodied morphologies that protect roots by jettisoning fishes with early maturation, low fecundity per seasonal biomass during floods (reviewed in spawning event, and low juvenile survivorship Lytle and Poff 2004). and typically inhabit habitats subjected to fre- There has been considerable interest in the quent and intense disturbances. Equilibrium identification of major axes of ecological strat- strategists are small to medium-bodied fishes egy variation in freshwater ecosystems based with moderate maturation age, low fecundity on trait correlations across large numbers of per spawning event, and high juvenile survivor- species (Winemiller 2005; Olden et al. 2006; ship (i.e., provides parental care) and typically intercontinental comparison of fish life histories 85 Figure 1. Triangular life history model depicting environmental gradients selecting for endpoint strate- gies defined by optimization of demographic parameters generation time, fecundity, and juvenile sur- vivorship (modified from Winemiller 2005). Example species from the study regions and representative hydrologic regimes expected to favor each life history strategy are illustrated. Australian fish illustrations by B. J. Pusey; U.S. fish illustrations freely available on the Internet. inhabit environments with low variation in odic fish species in rivers of the Cote d’Ivoire habitat quality and strong biotic interactions. (Africa) compared to equilibrium species. By The demographic parameters discussed comparing trends in native and nonnative spe- above are direct reflections of the ways in cies distributions among fish life history strate- which fish allocate energy to reproduction, gies in the lower Colorado River basin (USA), and the three life history strategies of the con- Olden et al. (2006) found that century-long tinuum can be interpreted as being adaptive modifications in flow variability have likely with respect to relative variability and predict- promoted the spread of nonnative equilibrium ability of temporal and spatial variation in abi- strategies (favored in constant environments) otic environmental conditions, food availabil- while leading to greater distributional declines ity, and predation pressure (Winemiller 2005). of native species located along the periodic- Hydrological variability plays a dominant role opportunistic continuum (strategies favored in shaping physical processes in riverine eco- in more unpredictable and variable environ- systems, and a number of recent studies have ments). Tedesco et al. (2008) found higher supported the association between hydrology proportions of periodic species in highly sea- and fish life history strategies. Tedesco and sonal drainage basins of western Africa (e.g., Hugueny (2006) found that regional climate rivers with short and predictably favorable (presumably related to hydrologic variability) seasons), whereas more hydrologically stable induced greater population synchrony of peri- basins with a wet season of several months 86 olden and kennard were dominated by equilibrium strategists. Col- tunistic-periodic-equilibrium trichotomy. The lectively, these studies (and others) suggest that freshwater fish faunas differ strikingly between the distributions of freshwater fishes are shaped, the two regions, with the U.S. fauna being ex- at least in part, by interactions between life his- tremely diverse and with low
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