Mathematical description of the stimuli to the lateral line system of fish derived from a three-dimensional flow field analysis

The three-dimensional potential flow field around a moving fish with an axially symmetric body is investigated mathematically. The case of a moving fish in open water and that of one approaching a plane surface in front of it are considered. For these cases the spatial and temporal distributions of the stimuli to the lateral line system are derived from the flow field analysis. This is done in dependence on the shape and the size of the fish.     

Shape variation in a benthic stream fish across flow regimes

Evolution of fish body shapes in flowing and non-flowing waters have been examined for several species. Flowing water can select for fish body shapes that increase steady swimming efficiency, whereas non-flowing water can favor shapes that increase unsteady swimming efficiency. Benthic stream fishes often use areas near the substrate that exhibit reduced or turbulent flow, thus it is unclear which swimming forms would be favored in such environments, and how shape might change across flow regimes. To test the relationship between fish body shape and flow regime in a benthic stream fish, we used geometric morphometric techniques to characterize lateral body shape in mountain sucker (Catostomus platyrhynchus) across flow rates, using stream gradient as an indicator of stream flow. Mountain suckers from low-flow environments were more streamlined, consistent with steady swimming body shapes, whereas mountain suckers from high flows had deeper bodies, consistent with unsteady swimming body shapes. In addition, smaller individuals tended to have more robust body shapes. These patterns are opposite to those predicted for stream fishes in the mid-water column. The benthic stream environment represents a distinct selective environment for fish shape that does not appear to conform to the simple dichotomy of flowing versus non-flowing water.     

Morphological plasticity in a native freshwater fish from semiarid Australia in response to variable water flows

In fishes, alterations to the natural flow regime are associated with divergence in body shape morphology compared with individuals from unaltered habitats. However, it is unclear whether this morphological divergence is attributable to evolutionary responses to modified flows, or is a result of phenotypic plasticity. Fishes inhabiting arid regions are ideal candidates for studying morphological plasticity as they are frequently exposed to extreme natural hydrological variability. We examined the effect of early exposure to flows on the development of body shape morphology in the western rainbowfish (Melanotaenia australis), a freshwater fish that is native to semiarid northwest Australia. Wild fish were collected from a region (the Hamersley Ranges) where fish in some habitats are subject to altered water flows due to mining activity. The offspring of wild‐caught fish were reared in replicated fast‐flow or slow‐flow channels, and geometric morphometric analyses were used to evaluate variation in fish body shape following 3, 6, 9, and 12 months of exposure. Water flows influenced fish morphology after 6 and 9 months of flow exposure, with fish in fast‐flow environments displaying a more robust body shape than those in slow‐flow habitats. No effect of flow exposure was observed at 3 and 12 months. Fishes also showed significant morphological variation within flow treatments, perhaps due to subtle differences in water flow among the replicate channels. Our findings suggest that early exposure to water flows can induce shifts in body shape morphology in arid zone freshwater fishes. Morphological plasticity may act to buffer arid zone populations from the impacts of anthropogenic activities, but further studies are required to link body shape plasticity with behavioral performance in habitats with modified flows.     

Mathematical model of flow in the vitreous humor induced by saccadic eye rotations: effect of geometry

Saccadic eye rotations induce a flow in the vitreous humor of the eye. Any such flow is likely to have a significant influence on the dispersion of drugs injected into the vitreous chamber. The shape of this chamber deviates from a perfect sphere by up to 10–20% of the radius, which is predominantly due to an indentation caused by the lens. In this paper we investigate theoretically the effect of the domain shape upon the flow field generated by saccades by considering an idealized model. The posterior chamber geometry is assumed to be a sphere with a small indentation, undergoing prescribed small-amplitude sinusoidal torsional oscillations, and, as an initial step towards understanding the problem, we treat the vitreous humor as a Newtonian fluid filling the chamber. The latter assumption applies best in the case of a liquefied vitreous or a tamponade fluid introduced in the vitreous chamber after vitrectomy. We find the flow field in terms of vector spherical harmonics, focusing on the deviation from the flow that would be obtained in a perfect sphere. The flow induced by the departure of the domain geometry from the spherical shape has an oscillating component at leading order and a smaller-amplitude steady streaming flow. The oscillating component includes a circulation cell formed every half-period, which migrates from the indentation towards the center of the domain where it disappears. The steady component has two counter-rotating circulations in the anterior part of the domain. These findings are in good qualitative agreement with the experimental results of Stocchino et al. (Phys Med Biol 52:2021–2034, 2007). Our results predict a significant reduction in the expected time for drug dispersal across the eye compared with the situation in which there is no fluid flow present.     

Mechanism of vestibular adaptation of fish under microgravity.

In a space experiment, the adaptation of goldfish behavior during flight and readaptation after landing were investigated. Six goldfish (1 normal, 1 with otoliths removed on both sides, 4 with otoliths removed on one side) were flown in a fish package (F/P) of Aquatic Animal Experiment Unit (AAEU). The dorsal light responses (DLRs) of fish with otoliths removed were recorded after operation until launch and after landing. The behaviors of the fish were recorded with a video camera on Mission Elapsed Time (MET) Day-00, 02, 05, 08, 12. On MET Day-00, two fish with otoliths removed on one side showed flexion of body toward the operated side. These fish also showed rolling behavior toward the operated side. However, the body flexion disappeared on MET Day-05 or MET Day-08. No rolling behaviors were observed after that time. Five fish showed backward looping behaviors during the mission. Although the frequency of looping episodes decreased after MET Day-08, five fish still showed looping behavior on MET Day-12, that was the last day of video recording on orbit. In microgravity, visual system of fish did not seem to provide sufficient cues to prevent them from looping or rolling. After landing, no looping and rolling behavior was observed. However, the tilt angle of the DLR increased in the fish with otolith removed 5 month before launch but not in normal fish and those with otoliths removed 2 weeks before launch. These results suggest that the behavioral dysfunction and the adaptational process in space are dependent on vestibular inputs.     

Maneuvering hydrodynamics of fish and small underwater vehicles

The understanding of fish maneuvering and its application tounderwater rigid bodies are considered. The goal is to gaininsight into stealth. The recent progress made in NUWC is reviewed.Fish morphology suggests that control fins for maneuverabilityhave unique scalar relationships irrespective of their speedtype. Maneuvering experiments are carried out with fish thatare fast yet maneuverable. The gap in maneuverability betweenfish and small underwater vehicles is quantified. The hydrodynamicsof a dorsal fin based brisk maneuvering device and a dual flappingfoil device, as applied to rigid cylindrical bodies, are described.The role of pectoral wings in maneuvering and station keepingnear surface waves is discussed. A pendulum model of dolphinswimming is presented to show that body length and tail flappingfrequency are related. For nearly neutrally buoyant bodies,Froude number and maneuverability are related. Analysis of measurementsindicates that the Strouhal number of dolphins is a constant.The mechanism of discrete and deterministic vortex sheddingfrom oscillating control surfaces has the property of largeamplitude unsteady forcing and an exquisite phase dependence,which makes it inherently amenable to active control for precisionmaneuvering. Theoretical control studies are carried out todemonstrate the feasibility of maneuverability of biologicallyinspired bodies under surface waves. The application of fishhydrodynamics to the silencing of propulsors is considered.Two strategies for the reduction of radiated noise are developed.The effects of a reduction of rotational rate are modeled. Theactive cambering of blades made of digitally programmable artificialmuscles, and their thrust enhancement, are demonstrated. Next,wake momentum filling is carried out by artificial muscles atthe trailing edge of a stator blade of an upstream stator propulsor,and articulating them like a fish tail. A reduction of radiatednoise, called blade tonals, is demonstrated theoretically.     

鱼体(去鳃)和鱼鳃对不同形态铜的积累特征

在实验室条件下研究实验鱼Ｐａｒａｃｈｅｉｒｏｄｏｎ对人工河水中不同形态的积累特征,对比了鱼体（去鳃）和鱼鳃对铜吸收量的差异,并探讨了鱼对铜的吸收机理。研究结果表明,实验鱼鳃部和体内铜积累量均随水相游离铜浓度增高,暴露时间增长而增加,但鳃部积累浓度较鱼体其余部分高一个数量级,其从水相富集铜的速率显著高于鱼体。     

Coping with flow: behavior, neurophysiology and modeling of the fish lateral line system

With the mechanosensory lateral line fish perceive water motions relative to their body surface and local pressure gradients. The lateral line plays an important role in many fish behaviors including the detection and localization of dipole sources and the tracking of prey fish. The sensory units of the lateral line are the neuromasts which are distributed across the surface of the animal. Water motions are received and transduced into neuronal signals by the neuromasts. These signals are conveyed by afferent nerve fibers to the fish brain and processed by lateral line neurons in parts of the brainstem, cerebellum, midbrain, and forebrain. In the cerebellum, midbrain, and forebrain, lateral line information is integrated with sensory information from other modalities. The present review introduces the peripheral morphology of the lateral line, and describes our understanding of lateral line physiology and behavior. It focuses on recent studies that have investigated: how fish behave in unsteady flow; what kind of sensory information is provided by flow; and how fish use and process this information. Finally, it reports new theoretical and biomimetic approaches to understand lateral line function.     

Conforming to the status flow: The influence of altered habitat on fish body-shape characteristics

1. Anthropogenic stream impoundments often result in altered flow regimes and reservoir formations. Functional fish characteristics may change in response to altered flow conditions, and understanding the underlying mechanisms at work (e.g. natural selection and phenotypic plasticity) is of great importance in both evolutionary and conservation contexts. Cyprinella lutrensis is a widely distributed minnow native to stream systems in Missouri, U.S.A. Following impoundment of the Osage River, C. lutrensis currently occurs in the resultant Truman Reservoir and surrounding tributaries. To examine the impacts of this habitat alteration on functional fish characteristics, we hypothesised that C. lutrensis populations in altered systems with no-flow would show an overall less streamlined body shape as compared to those in flowing systems, assuming body streamlining is beneficial for locomotion in flowing environments.
2. We conducted field and laboratory studies to test effects of flow and no-flow on C. lutrensis body shape. Geometric morphometric techniques were used to quantify body shape in field and laboratory individuals, and non-structural lipid extractions were conducted to assess whether fat storage contributes to body shape under flow and no-flow treatments.
3. Morphological comparisons between C. lutrensis from Truman Reservoir and surrounding streams yielded significant differences in body shape, with greater body streamlining in stream populations. A laboratory experiment using artificial stream units revealed similar significant differences in body streamlining between flow and no-flow treatments within a single generation. In addition, individuals under no-flow treatment had significantly higher levels of non-structural lipids compared to flow treatment individuals.
4. Our results indicate that changes in flow regime can alter the evolutionary trajectories (i.e. related to morphology) of fish populations. We suggest that body shape alterations are likely to occur within and across generations in a cumulative manner, therefore, it is plausible that both natural selection and plastic response mechanisms play roles in defining minnow body shape following flow alteration.

     

Hydrodynamics of suction feeding of fish in motion

Forward motion was shown to increase the efficiency of suction feeding by causing the ingested volume to be predominantly in front of the fish, instead of being a sphere centred on the fish mouth. This increases the distance from which a prey, located in front of the predator, can be ingested by suction.
A general hydrodynamic model of the external effects of the suction process is presented, showing that this is dependent upon a single non-dimensional parameter, the ratio of a characteristic mouth dimension, divided by the forward swimming speed and the suction time. Data for large-mouth bass, Micropterus salmoides show that the observed average forward velocity while feeding, of 3.1 body length/s, serves to increase the ingestion range by more than 60% over suction while not moving.     

The influence of viscous hydrodynamics on the fish lateral-line system

Fish exhibit many behaviors that involve sensing water flows with their lateral-line system. In many situations, viscosity affects how the flow interacts with the body of the fish and the neuromasts of the lateral line. Here we discuss how viscosity influences the stimulus to the fish lateral-line system. The movement of a fish's body creates flows that can interfere with the detection of external signals, but these flows can also serve as a source of information about nearby obstacles and the fish's own hydrodynamic performance. The viscous boundary layer on the surface of the skin alters external signals by attenuating the low-frequency components of stimuli. The stimulus to each neuromast depends on the interaction of the fluid surrounding the neuromast and the structural properties of that neuromast, including the number of mechanosensory hair cells it contains. A consideration of the influences of viscosity on flow, at both the whole-body and receptor levels, offers the promise of a more comprehensive understanding of the signals involved in behaviors mediated by the lateral-line system.     

Effects of linear acceleration on fish behavior and eye movements.

Behavioral responses and eye movements of fish during linear acceleration were reviewed. It is known that displacement of otoliths in the inner ear leads to body movements and/or eye movements. On the ground, the utriculus of the vestibular system is stimulated by otolith displacement caused by gravitational and inertial forces during horizontal acceleration of whole body. When the acceleration is imposed on the fish's longitudinal axis, the fish showed nose-down and nose-up posture for tailward and noseward displacement of otolith respectively. These responses were understood that the fish aligned his longitudinal body axis in a plane perpendicular to the direction of resultant force vector acting on the otoliths. When the acceleration was sideward, the fish rolled around his longitudinal body axis so that his back was tilted against the direction in which the inertial force acted on the otoliths. Linear acceleration applied to fish's longitudinal body axis evoked torsional eye movement. Direction of torsion coincided with the direction of acceleration, which compensate the change of resultant force vector produced by linear acceleration and gravity. Torsional movement of left and right eye coordinated with each other. In normal fish, both sinusoidal and rectangular acceleration of 0.1G could evoke clear eye torsion. Though the amplitude of response increased with increasing magnitude of acceleration up to 0.5 G, the torsion angle did not fully compensate the angle calculated from gravity and linear acceleration. Removal of the otolith on one side reduced the response amplitude of both eyes. The torsion angle evoked by rectangular acceleration was smaller than that evoked by sinusoidal acceleration in both normal and unilaterally labyrinthectomized fish. These results suggest that eye torsion of fish include both static and dynamic components.     

Morphological selection in an extreme flow environment: body shape and waterfall-climbing success in the Hawaiian stream fish Sicyopterus stimpsoni

Flow characteristics are a prominent factor determining body shapes in aquatic organisms, and correlations between body shape and ambient flow regimes have been established for many fish species. In this study, we investigated the potential for a brief period of extreme flow to exert selection on the body shape of juvenile climbing Hawaiian gobiid fishes. Because of an amphidromous life history, juvenile gobies that complete an oceanic larval phase return to freshwater habitats, where they become adults. Returning juveniles often must scale waterfalls (typically with the use of a ventral sucker) in order to reach the habitats they will use as adults, thereby exposing these animals to brief periods of extreme velocities of flow. Hydrodynamic theory predicts that bodies with larger suckers and with lower heights that reduce drag would have improved climbing success and, thus, be well suited to meet the demands of the flows in waterfalls. To test the potential for the flow environment of waterfalls to impose selection that could contribute to differences in body shape between islands, we subjected juvenile Sicyopterus stimpsoni to climbing trials up artificial waterfalls (～100 body lengths) and measured differences in body shape between successful and unsuccessful climbers. Waterfalls appear to represent a significant selective barrier to these fishes, as nearly 30% failed our climbing test. However, the effects of selection on morphology were not straightforward, as significant differences in shape between successful and unsuccessful climbers did not always match hydrodynamic predictions. In both selection experiments and in adult fish collected from habitats with different prevailing conditions of flow (the islands of Hawai'i versus Kaua'i), lower head heights were associated with exposure to high-flow regimes, as predicted by hydrodynamic theory. Thus, a premium appears to be placed on the reduction of drag via head morphology throughout the ontogeny of this species. The congruence of phenotypic selection patterns observed in our experiments, with morphological character divergence documented among adult fish from Hawai'i and Kaua'i, suggests that differences in morphology between subpopulations of adult climbing gobies may result, at least in part, from the selective pressures of high-velocity flows encountered by migrating juveniles.     

Resource allocation in yolk-feeding fish

An insight is made into the main processes that occur in fish during endogenous feeding period. The ways in which yolk absorption rate can be measured are evaluated. Essential amino acids and polyunsaturated fatty acids are preferentially retained for incorporation into body tissue. Profound physiological and anatomical changes in yolk and a sequence of slow, fast, and a second period of slow absorption occur during the endogenous feeding period. Attempts to quantify the ontogenetic sequence are reviewed. Various methods of body size assessment are compared, and sources of bias in individual and population growth estimates are discussed. Several calorimetric methods are compared of which direct calorimetry using an oxygen bomb is the reference method. An advanced elemental analysis (CHNS) is a reliable technique that is adequate for early stages. Indices of growth potential are reviewed including a comparison of different measures, models and approaches used to estimate growth. Changes in body hydration, caloric value, content of lipids, protein, free amino acids (FAA) and minerals, and in content of RNA and DNA occur in early ontogeny. Ways to quantify metabolic rate are identified. Mean relative respiration rate of initial egg before activation is very low, about 20 mm³ g⁻¹ h⁻¹. Ontogenetic sequence in absolute metabolic rate of fish embryos and yolk-feeding larvae involves an increase through hatching to a peak at the time of first feeding ability, and a decrease under starvation. Models predicting the relationship between oxygen consumption and age in yolk-feeding fish are reviewed. Sequence of metabolic fuels begins with use of small molecules as carbohydrates, soon switched to FAA. Later lipids are progressively used, they provide energy for swimming activity. After yolk depletion body protein-bound amino acids are mobilised. In this review I focused on the major environmental variables as temperature, oxygen, salinity, pH, toxic xenobiotics, light, UV radiation, magnetic field and substrate, along with intrinsic factors as egg or body size, sex and genetic factors. A question was posed on how the extrinsic and intrinsic factors determine yolk absorption, growth and metabolic rates in yolk-feeding fish. Special attention is devoted to fish body size attained exclusively on yolk. A considerable variety of body size responses to temperature was found, for which several explanations are forwarded. Methodological progress made recently is characterised and the most conspicious advances in understanding of fish early life history are highlighted. Information derived from these studies can be used in management of fish populations in the field and to optimise activities in aquaculture.     

Water temperature and fish growth: otoliths predict growth patterns of a marine fish in a changing climate

Ecological modeling shows that even small, gradual changes in body size in a fish population can have large effects on natural mortality, biomass, and catch. However, efforts to model the impact of climate change on fish growth have been hampered by a lack of long‐term (multidecadal) data needed to understand the effects of temperature on growth rates in natural environments. We used a combination of dendrochronology techniques and additive mixed‐effects modeling to examine the sensitivity of growth in a long‐lived (up to 70 years), endemic marine fish, the western blue groper (Achoerodus gouldii), to changes in water temperature. A multi‐decadal biochronology (1952–2003) of growth was constructed from the otoliths of 56 fish collected off the southwestern coast of Western Australia, and we tested for correlations between the mean index chronology and a range of potential environmental drivers. The chronology was significantly correlated with sea surface temperature in the region, but common variance among individuals was low. This suggests that this species has been relatively insensitive to past variations in climate. Growth increment and age data were also used in an additive mixed model to predict otolith growth and body size later this century. Although growth was relatively insensitive to changes in temperature, the model results suggested that a fish aged 20 in 2099 would have an otolith about 10% larger and a body size about 5% larger than a fish aged 20 in 1977. Our study shows that species or populations regarded as relatively insensitive to climate change could still undergo significant changes in growth rate and body size that are likely to have important effects on the productivity and yield of fisheries.     

Ecomorphology of solitary chemosensory cell systems in fish: a review

Synopsis Solitary chemosensory cells (SCCs) are present in the skin of a wide spectrum of lower vertebrates, such as lampreys, elasmobranchs, teleost fishes and some amphibians (Kotrschal 1991, Whitear 1992). However, due to the difficulties studying them, virtually all our present knowledge on SCCs stems from the anterior dorsal fin of two species of rocklings (Gadidae). This fin is a peculiar chemosensory organ, carrying approximately 5 million SCCs (Kotrschal et al. 1984, Kotrschal & Whitear 1988). The evidence derived from this model on the structure of SCCs, on their innervation and brain representation, on the flow dynamics at the receptors, on their electrophysiological responses and behavioral relevance indicates that this fin is actively sampling for substances leaked from other fish, such as body mucus and bile components. Possibly, the rockling anterior dorsal fin aids in predators avoidance. To generate hypotheses on the functions and biological roles of the generalized., scattered SCC systems present in most fishes, their structural parameters are put in perspective to taste bud structure and function and to the rockling results. Ecomorphological reasoning serves to establish testable hypotheses: in essence, SCC systems spread over the body surface may be designed as general water samplers, but not for the exact localization of a stimulus source. If the function of the latter is equally dependent on water flow, as the rockling fin organ, fish would have to rely either on the ambient water flow, or speed up their own swimming to optimize SCC input. If SCCs are indeed evolved in the context of predator avoidance, a comparison between life history intervals and between species should reveal, that the system varies in accordance with predation pressure. It is concluded, that in fish, SCCs are certainly an important source of environmental information. If we do not understand functions and biological roles of SCCs, it will not be possible to explain fish behavior and ecology. Evidently, further investigations are urgently needed.     

Reproductive ecology of cultured fish in the wild

Domestication has been shown to have an effect on morphology and behaviour of Atlantic salmon ( Salmo salar ). We compared swimming costs of three groups of juvenile Atlantic salmon subject to different levels of domestication: (1) wild fish; (2) first generation farmed fish origination from wild genitors; and (2) seventh generation farmed fish originating from Norwegian aquaculture stocks. We assessed swimming costs under two types of turbulent flow (one mean flow velocity of 23 cm s⁻¹ and two standard deviations of flow velocity of 5 and 8 cm s⁻¹). Respirometry experiments were conducted with fish in a mass range of 5–15 g wet at a water temperature of 15° C. Our results confirm (1) that net swimming costs are affected by different levels of turbulence such that, for a given mean flow velocity, fish spent 1·5‐times more energy as turbulence increased, (2) that domesticated fish differed in their morphology (having deeper bodies and smaller fins) and in their net swimming costs (being up to 30·3% higher than for wild fish) and (3) that swimming cost models developed for farmed fish may be also be applied to wild fish in turbulent environments.     

The intestinal loops of bluegill fish, Lepomis macrochirus

The structural configurations of the intestinal loops of bluegill fish (Lepomis macrochirus) and a correlation of their probable functional significance have been investigated. The bluegill fish accommodates its relatively large intestine in its smaller body cavity by developing two intestinal loops. Regional variations in the intestinal wall exist due to the presence of the intestinal loops. The structural complexity of the intestine by loop formation has apparently created a problem by posing an obstruction as well as deviation to the smooth flow of food. In order to solve this problem structural modifications in the walls of the intestinal loops are needed, as is observed by the variation in muscle thickness of greater and lesser curvature of both the intestinal loops.     

The scaling of locomotor performance in predator-prey encounters: from fish to killer whales.

During predator-prey encounters, a high locomotor performance in unsteady manoeuvres (i.e. acceleration, turning) is desirable for both predators and prey. While speed increases with size in fish and other aquatic vertebrates in continuous swimming, the speed achieved within a given time, a relevant parameter in predator-prey encounters, is size independent. In addition, most parameters indicating high performance in unsteady swimming decrease with size. Both theoretical considerations and data on acceleration suggest a decrease with body size. Small turning radii and high turning rates are indices of maneuverability in space and in time, respectively. Maneuverability decreases with body length, as minimum turning radii and maximum turning rates increase and decrease with body length, respectively. In addition, the scaling of linear performance in fish locomotion may be modulated by turning behaviour, which is an essential component of the escape response. In angelfish, for example, the speed of large fish is inversely related to their turning angle, i.e. fish escaping at large turning angles show lower speed than fish escaping at small turning angles. The scaling of unsteady locomotor performance makes it difficult for large aquatic vertebrates to capture elusive prey by using whole-body attacks, since the overall maneuverability and acceleration of small prey is likely to be superior to that of large predators. Feeding strategies in vertebrate predators can be related to the predator-prey length ratios. At prey-predator ratios higher than approximately 10(-2), vertebrate predators are particulate feeders, while at smaller ratios, they tend to be filter feeders. At intermediate ratios, large aquatic predators may use a variety of feeding methods that aid, or do not involve, whole body attacks. Among these are bubble curtains used by humpback whales to trap fish schools, and tail-slapping of fish by delphinids. Tail slapping by killer whales is discussed as an example of these strategies. The speed and acceleration achieved by the flukes of killer whales during tail slaps are higher and comparable, respectively, to those that can be expected in their prey, making tail-slapping an effective predator behaviour.     

Size-assortativeness in multi-species fish shoals

Theory predicts that fish should assort in shoals on the basis of similar phenotypic traits to minimize predation risk and to maximize foraging efficiency. A single phenotypic character, body size, was considered and the hypothesis tested that free-ranging fish shoals are sizeassorted. Furthermore, a second test investigated whether fish within multi-species shoals are more strongly size-assorted with conspecifics than with heterospecifics. Twelve fish shoals, each comprising two different species (golden shiner Notemigonus crysoleucas , and banded killifish Fundulus diaphanus ) were caught in the littoral zone of a north temperate lake using a beach seine. Shoal membership size ranged from 36 to 776 fish, and mean standard body length of members ranged from 18 to 34 mm. Fish were assorted by body size at two different levels, namely, between shoals and at the level of species within shoals. Body sizes of shiners and killifish within shoals were significantly different in seven out of 12 shoals, with killifish being the larger species in five cases and shiners being the larger one in two cases. Because there is considerable overlap in body size between the two species in the population, the observed species-related size-assortativeness within shoals was not just the by-product of a directional size difference (between species) in the population. These findings provide strong quantitative evidence for size-sorting in free-ranging fish shoals and raise questions concerning the formation of multi-species fish shoals.