Michael D. Yard

Food‐web dynamics in a large river discontinuum

Nearly all ecosystems have been altered by human activities, and most communities are now composed of interacting species that have not co-evolved. These changes may modify species interactions, energy and material flows, and food-web stability. Although structural changes to ecosystems have been widely reported, few studies have linked such changes to dynamic food-web attributes and patterns of energy flow. Moreover, there have been few tests of food-web stability theory in highly disturbed and intensely managed freshwater ecosystems. Such synthetic approaches are needed for predicting the future trajectory of ecosystems, including how they may respond to natural or anthropogenic perturbations. We constructed flow food webs at six locations along a 386-km segment of the Colorado River in Grand Canyon (Arizona, USA) for three years. We characterized food-web structure and production, trophic basis of production, energy efficiencies, and interaction-strength distributions across a spatial gradient of pertu...

Trout Piscivory in the Colorado River, Grand Canyon: Effects of Turbidity, Temperature, and Fish Prey Availability

Abstract Introductions of nonnative salmonids, such as rainbow trout Oncorhynchus mykiss and brown trout Salmo trutta, have affected native fishes worldwide in unforeseen and undesirable ways. Predation and other interactions with nonnative rainbow trout and brown trout have been hypothesized as contributing to the decline of native fishes (including the endangered humpback chub Gila cypha) in the Colorado River, Grand Canyon. A multiyear study was conducted to remove nonnative fish from a 15-km segment of the Colorado River near the Little Colorado River confluence. We evaluated how sediment, temperature, fish prey availability, and predator abundance influenced the incidence of piscivory (IP) by nonnative salmonids. Study objectives were addressed through spatial (upstream and downstream of the Little Colorado River confluence) and temporal (seasonal and annual) comparisons of prey availability and predator abundance. Data were then evaluated by modeling the quantity of fish prey ingested by trout durin...

Nonnative Fish Control in the Colorado River in Grand Canyon, Arizona: An Effective Program or Serendipitous Timing?

Abstract The federally endangered humpback chub Gila cypha in the Colorado River within Grand Canyon is currently the focus of a multiyear program of ecosystem-level experimentation designed to improve native fish survival and promote population recovery as part of the Glen Canyon Dam Adaptive Management Program. A key element of this experiment was a 4-year effort to remove nonnative fishes from critical humpback chub habitat, thereby reducing potentially negative interactions between native and nonnative fishes. Over 36,500 fish from 15 species were captured in the mechanical removal reach during 2003–2006. The majority (64%) of the catch consisted of nonnative fish, including rainbow trout Oncorhynchus mykiss (19,020), fathead minnow Pimephales promelas (2,569), common carp Cyprinus carpio (802), and brown trout Salmo trutta (479). Native fish (13,268) constituted 36% of the total catch and included flannelmouth suckers Catostomus latipinnis (7,347), humpback chub (2,606), bluehead suckers Catostomus d...

A quantitative life history of endangered humpback chub that spawn in the Little Colorado River: variation in movement, growth, and survival.

While the ecology and evolution of partial migratory systems (defined broadly to include skip spawning) have been well studied, we are only beginning to understand how partial migratory populations are responding to ongoing environmental change. Environmental change can lead to differences in the fitness of residents and migrants, which could eventually lead to changes in the frequency of the strategies in the overall population. Here, we address questions concerning the life history of the endangered Gila cypha (humpback chub) in the regulated Colorado River and the unregulated tributary and primary spawning area, the Little Colorado River. We develop eight multistate models for the population based on three movement hypotheses, in which states are defined in terms of fish size classes and river locations. We fit these models to mark–recapture data collected in 2009–2012. We compare survival and growth estimates between the Colorado River and Little Colorado River and calculate abundances for all size classes. The best model supports the hypotheses that larger adults spawn more frequently than smaller adults, that there are residents in the spawning grounds, and that juveniles move out of the Little Colorado River in large numbers during the monsoon season (July–September). Monthly survival rates for G. cypha in the Colorado River are higher than in the Little Colorado River in all size classes; however, growth is slower. While the hypothetical life histories of life-long residents in the Little Colorado River and partial migrants spending most of its time in the Colorado River are very different, they lead to roughly similar fitness expectations when we used expected number of spawns as a proxy. However, more research is needed because our study period covers a period of years when conditions in the Colorado River for G. cypha are likely to have been better than has been typical over the last few decades.

Of Travertine and Time: Otolith Chemistry and Microstructure Detect Provenance and Demography of Endangered Humpback Chub in Grand Canyon, USA

We developed a geochemical atlas of the Colorado River in Grand Canyon and in its tributary, the Little Colorado River, and used it to identify provenance and habitat use by Federally Endangered humpback chub, Gila cypha. Carbon stable isotope ratios (δ13C) discriminate best between the two rivers, but fine scale analysis in otoliths requires rare, expensive instrumentation. We therefore correlated other tracers (SrSr, Ba, and Se in ratio to Ca) to δ13C that are easier to quantify in otoliths with other microchemical techniques. Although the Little Colorado River’s water chemistry varies with major storm events, at base flow or near base flow (conditions occurring 84% of the time in our study) its chemistry differs sufficiently from the mainstem to discriminate one from the other. Additionally, when fish egress from the natal Little Colorado River to the mainstem, they encounter cold water which causes the otolith daily growth increments to decrease in size markedly. Combining otolith growth increment analysis and microchemistry permitted estimation of size and age at first egress; size at first birthday was also estimated. Emigrants < 1 year old averaged 51.2 ± 4.4 (SE) days and 35.5 ± 3.6 mm at egress; older fish that had recruited to the population averaged 100 ± 7.8 days old and 51.0 ± 2.2 mm at egress, suggesting that larger, older emigrants recruit better. Back-calculated size at age 1 was unimodal and large (78.2 ± 3.3 mm) in Little Colorado caught fish but was bimodally distributed in Colorado mainstem caught fish (49.9 ± 3.6 and 79 ± 4.9 mm) suggesting that humpback chub can also rear in the mainstem. The study demonstrates the coupled usage of the two rivers by this fish and highlights the need to consider both rivers when making management decisions for humpback chub recovery.

Assessing Predation Risks for Small Fish in a Large River Ecosystem between Contrasting Habitats and Turbidity Conditions

Abstract This study examined predation risk for juvenile native fish between two riverine shoreline habitats, backwater and debris fan, across three discrete turbidity levels (low, intermediate, high) to understand environmental risks associated with habitat use in a section of the Colorado River in Grand Canyon, AZ. Inferences are particularly important to juvenile native fish, including the federally endangered humpback chub Gila cypha. This species uses a variety of habitats including backwaters which are often considered important rearing areas. Densities of two likely predators, adult rainbow trout Oncorhynchus mykiss and adult humpback chub, were estimated between habitats using binomial mixture models to examine whether higher predator density was associated with patterns of predation risk. Tethering experiments were used to quantify relative predation risk between habitats and turbidity conditions. Under low and intermediate turbidity conditions, debris fan habitat showed higher relative predation risk compared to backwaters. In both habitats the highest predation risk was observed during intermediate turbidity conditions. Density of likely predators did not significantly differ between these habitats. This information can help managers in Grand Canyon weigh flow policy options designed to increase backwater availability or extant turbidity conditions.

Inferring species interactions through joint mark–recapture analysis

Introduced species are frequently implicated in declines of native species. In many cases, however, evidence linking introduced species to native declines is weak. Failure to make strong inferences regarding the role of introduced species can hamper attempts to predict population viability and delay effective management responses. For many species, mark-recapture analysis is the more rigorous form of demographic analysis. However, to our knowledge, there are no mark-recapture models that allow for joint modeling of interacting species. Here, we introduce a two-species mark-recapture population model in which the vital rates (and capture probabilities) of one species are allowed to vary in response to the abundance of the other species. We use a simulation study to explore bias and choose an approach to model selection. We then use the model to investigate species interactions between endangered humpback chub (Gila cypha) and introduced rainbow trout (Oncorhynchus mykiss) in the Colorado River between 2009 and 2016. In particular, we test hypotheses about how two environmental factors (turbidity and temperature), intraspecific density dependence, and rainbow trout abundance are related to survival, growth, and capture of juvenile humpback chub. We also project the long-term effects of different rainbow trout abundances on adult humpback chub abundances. Our simulation study suggests this approach has minimal bias under potentially challenging circumstances (i.e., low capture probabilities) that characterized our application and that model selection using indicator variables could reliably identify the true generating model even when process error was high. When the model was applied to rainbow trout and humpback chub, we identified negative relationships between rainbow trout abundance and the survival, growth, and capture probability of juvenile humpback chub. Effects on interspecific interactions on survival and capture probability were strongly supported, whereas support for the growth effect was weaker. Environmental factors were also identified to be important and in many cases stronger than interspecific interactions, and there was still substantial unexplained variation in growth and survival rates. The general approach presented here for combining mark-recapture data for two species is applicable in many other systems and could be modified to model abundance of the invader via other modeling approaches.

Trends in Rainbow Trout Recruitment, Abundance, Survival, and Growth during a Boom-and-Bust Cycle in a Tailwater Fishery

AbstractData from a large-scale mark–recapture study were used in an open-population model to determine the cause for long-term trends in growth and abundance of a Rainbow Trout Oncorhynchus mykiss population in the tailwater of Glen Canyon Dam, Arizona. Reduced growth affected multiple life stages and processes, causing negative feedbacks that regulated the abundance of the population, including higher mortality of larger fish; lower rates of recruitment (young of the year) during years when growth was reduced; and lower rates of sexual maturation in the following year. High and steady flows during spring and summer 2011 resulted in a very large recruitment event. The population had declined tenfold by 2016 due to a combination of lower recruitment and reduced survival of larger trout. Survival rates for 225-mm and larger Rainbow Trout in 2014, 2015, and 2016 were 11, 21, and 22% lower, respectively, than average survival rates between 2012 and 2013. Abundance at the end of the study would have been thre...

Does Bioelectrical Impedance Analysis Accurately Estimate the Physiological Condition of Threatened and Endangered Desert Fish Species

AbstractBioelectrical impedance analysis (BIA) is a nonlethal tool with which to estimate the physiological condition of animals that has potential value in research on endangered species. However, the effectiveness of BIA varies by species, the methodology continues to be refined, and incidental mortality rates are unknown. Under laboratory conditions we tested the value of using BIA in addition to morphological measurements such as total length and wet mass to estimate proximate composition (lipid, protein, ash, water, dry mass, energy density) in the endangered Humpback Chub Gila cypha and Bonytail G. elegans and the species of concern Roundtail Chub G. robusta and conducted separate trials to estimate the mortality rates of these sensitive species. Although Humpback and Roundtail Chub exhibited no or low mortality in response to taking BIA measurements versus handling for length and wet-mass measurements, Bonytails exhibited 14% and 47% mortality in the BIA and handling experiments, respectively, indi...