As the prey thickens: rainbow trout select prey based upon width not length

Drift-feeding fish are typically considered size-selective predators. Yet, few studies have explicitly tested which aspect of prey “size” best explains size selection by drift-foraging fish. Here, we develop a Bayesian discrete choice model to evaluate how attributes of both prey and predator simultaneously influence size-selective foraging. We apply the model to a large dataset of paired invertebrate drift (n = 784) and rainbow trout (Oncorhynchus mykiss) diets (n = 1028). We characterized prey “size” using six metrics (length, width, area, hemispherical area, volume, mass) and used pseudo-R2 to determine which metric best explained observed prey selection across seven taxa. We found that rainbow trout are positively size-selective, they are selecting prey based upon differences in prey width, and size-selectivity increases with fish length. Rainbow trout demonstrated strong selection for the adult and pupae stages of aquatic insects relative to their larval stages. Our study provides strong empirical evidence for size-selective foraging in rainbow trout and demonstrates prey selection is based primarily upon width, not length or area as has been widely reported.

Freshwater reefs as mesohabitats for the assessment of diel invertebrate drift patterns

Freshwater reefs (known as tufa barriers) are special karst features recognized for highly heterogeneous habitat structures, complex hydrogeological features, and unique macrozoobenthos drift (downstream dispersal) patterns. Our study objective was to investigate diel and seasonal drift patterns between barriers and pools, both composed of moss-rich and fishless mesohabitat types, aligned on a small spatial scale within the karst, tufa-precipitating Plitvice Lakes hydrosystem. We monthly sampled drift at the two mesohabitat types (barriers and pools) during midday and dusk and examined quantitative and qualitative drift compositions, including drifting invertebrates, moss, and associated particulate organic and inorganic matter (APOIM). Barriers displayed higher invertebrate drift densities than those of pools. The same pattern was observed for moss and APOIM. At both mesohabitat types, invertebrate drift showed peak but highly variable densities during late spring and summer (mean >100 individuals m-3), whereas during late winter and early spring the drift densities were 5-fold lower than those densities. The nonmetric multidimensional scaling analysis revealed that invertebrate drift seasonality was influenced by seasonal drift patterns of aquatic moss and moss-dwelling invertebrate taxa. Adult and/or larval Riolus spp. and larval Hemerodromia spp. were found to be the most significant for the separation of midday and dusk within the NMDS ordination of barriers and pools. At barriers, midday drift densities of invertebrates, moss, and APOIM were higher than the respective dusk records. Within pools, invertebrate drift was largely aperiodic. We suggest that increased midday and/or aperiodic drift are a consequence of the lack of fish between barrier- and pool-mesohabitats. Our results further indicated that aquatic invertebrates inhabiting fast-flowing barriers and slow-flowing pools mostly exhibit “passive drift” mediated by transport agents such as water flow and dislodged aquatic vegetation. The observed spatio-temporal drift patterns are also likely influenced by ontogenetic shifts in drift periodicity (i.e., shifts depending on the development stage and morphological characteristics of the individual taxa) as well as benthic distribution of moss-dwelling invertebrate taxa. We can conclude that biotic (vs. abiotic) controls of drift are likely minimized in the fishless case of the freshwater reefs and associated barrier–pool sequences within Plitvice Lakes hydrosystem.

Foraging modes and movements of Oncorhynchus mykiss as flow and invertebrate drift recede in a California stream

Salmonids frequently adapt their feeding and movement strategies to cope with seasonally fluctuating stream environments. Oncorhynchus mykiss tend to drift-forage in higher velocity habitat than other salmonids, yet their presence in streams with seasonally low velocity and drift suggests behavioral flexibility. We combined 3-D videogrammetry with measurements of invertebrate drift and stream hydraulics to investigate the drivers of O. mykiss foraging mode and movement during the seasonal recession in a California stream. From May to July (2016), foraging movement rate increased as prey concentration and velocity declined; however, movement decreased in August as pools became low and still. In May, 80% of O. mykiss were drift-foraging, while by July, over 70% used search or benthic-foraging modes. Velocity and riffle crest depth were significant predictors of foraging mode, while drift concentration was a poor univariate predictor. However top ranked additive models included both hydraulic variables and drift concentration. A drift-foraging bioenergetic model was a poor predictor of foraging mode. We suggest that infall and benthic prey, as well as risk aversion, may influence late-summer foraging decisions.

Rheotaxis revisited: a multi-behavioral and multisensory perspective on how fish orient to flow

ABSTRACTHere, we review fish rheotaxis (orientation to flow) with the goal of placing it within a larger behavioral and multisensory context. Rheotaxis is a flexible behavior that is used by fish in a variety of circumstances: to search for upstream sources of current-borne odors, to intercept invertebrate drift and, in general, to conserve energy while preventing downstream displacement. Sensory information available for rheotaxis includes water-motion cues to the lateral line and body-motion cues to visual, vestibular or tactile senses when fish are swept downstream. Although rheotaxis can be mediated by a single sense, each sense has its own limitations. For example, lateral line cues are limited by the spatial characteristics of flow, visual cues by water visibility, and vestibular and other body-motion cues by the ability of fish to withstand downstream displacement. The ability of multiple senses to compensate for any single-sense limitation enables rheotaxis to persist over a wide range of sensory and flow conditions. Here, we propose a mechanism of rheotaxis that can be activated in parallel by one or more senses; a major component of this mechanism is directional selectivity of central neurons to broad patterns of water and/or body motions. A review of central mechanisms for vertebrate orienting behaviors and optomotor reflexes reveals several motorsensory integration sites in the CNS that could be involved in rheotaxis. As such, rheotaxis provides an excellent opportunity for understanding the multisensory control of a simple vertebrate behavior and how a simple motor act is integrated with others to form complex behaviors.

Evaluating a pattern of ecological character displacement: charr jaw morphology and diet diverge in sympatry versus allopatry across catchments in Hokkaido, Japan

Similar species that overlap in sympatry may diverge in characters related to resource use as a result of evolution or phenotypic plasticity. Dolly Varden charr (Salvelinus malma) and whitespotted charr (S. leucomaenis) overlap along streams in Hokkaido, Japan, and compete by interference for invertebrate drift-foraging positions. Previous research has shown that as drift declines during summer, Dolly Varden shift foraging modes to capture benthic prey, a behaviour facilitated by their subterminal jaw morphology. We compare body and jaw morphology of Dolly Varden in sympatry vs. allopatry in two locations to test for character displacement. Statistical analysis showed significant divergence in characters related to foraging, which was correlated with variation in individual charr diets. Dolly Varden in sympatry had shorter heads and lower jaws than in allopatry, and even within sites charr with these characteristics fed less on drifting terrestrial invertebrates but more on benthic aquatic invertebrates. Those in allopatry had longer heads and lower jaws, and fed more on terrestrial invertebrates. The close proximity of sites in one stream suggests that Dolly Varden may display phenotypic plasticity similar to other charr, allowing rapid responses in morphology to the presence of competitors. These morphological shifts probably help them maintain positive fitness when competing with whitespotted charr in Hokkaido streams.

Relationship between background invertebrate drift concentration and flow over natural flow recession and prediction with a drift transport model

This study advances understanding of the flow dependency of invertebrate drift in rivers and its relevance to drift-feeding fish. Background drift concentration varied spatially and with flow over natural flow recession (lower mid-range to low flow) in a reach of a New Zealand river, largely consistent with passive entrainment. Seven taxonomic groups (dominated by Leptophlebiidae and Chironomidae) exhibited positive drift concentration–flow relationships, and one (sandy/stony-cased caddisflies (Conoesucidae)) exhibited negative relationships. A mechanistic drift transport model accurately predicted the slope, but not y intercept, of the drift concentration–flow relationship for the total drift community that positively responded to flow but performed more poorly at the taxon or size-class level. Partitioning the relative influence of drift entry and dilution revealed that positive drift concentration–flow relationships arose from entry overwhelming dilution with increasing flow. Drift transport models have potential for predicting relative (%) effects of flow change on concentration and rate of drift-prone invertebrates. This paves the way for drift transport models to inform inputs to net rate of energy intake models for drift-feeding fish.

Spatial linkages between geomorphic and hydraulic conditions and invertebrate drift characteristics in a small mountain stream

Although invertebrate drift plays a fundamental role in river ecosystems, spatial linkages between its characteristics and hydrogeomorphic conditions remain poorly understood. This study, conducted in a small stream in British Columbia, provides insight into such relationships. Most notably, drift concentration was positively correlated with bed scour, and mean body size was negatively correlated with bed fill. Although correlational design precludes inference regarding mechanisms, these results are consistent with protracted effects of flood-related bed disturbances on spatial pattern of drift characteristics under low flow conditions. We detected no robust associations between drift and flow characteristics that could not be explained by bed disturbances. Because of high spatial variability, drift characteristics were statistically distinct only in few habitat types. Compared with middepth, invertebrate drift concentration was higher near water surface, and mean body size was larger near the bed. Mean body size was also statistically smaller in riffle–pool transitions, which experienced largest bed fill. Possible legacy effects of bed disturbances could have broader implications for drift-related energy and nutrient fluxes in river corridors and for quantitative drift models.