Dependency and independency among fish density and electivity indices in a stream fish assemblage

Habitat preferences can be identified from population density and electivity indices, which can be derived from either abundance or occurrence data. This study examined the preference detectability and independency of density and electivity indices from different data sources -- abundance and occurrence. Eight scenarios comprising four fish species along two environmental gradients (flow velocity and water depth) in an undisturbed mountain stream were used in this study. Fish density and seven electivity indices were calculated from abundance and occurrence data for the eight scenarios. The correlation analyses illustrated that abundance electivity indices were positively correlated with density in all eight scenarios while occurrence electivity indices were positively correlated with density in only three scenarios. Electivity indices were positively correlated with each other within the same data quality (abundance or occurrence) but not between. The Chesson’s L index was less correlated with other indices and should be used cautiously. Natural historical traits, such as social interactions among organisms, may affect the preference performance of density and electivity indices. In conclusion, density is a good preference indicator. Abundance electivity indices can represent density while occurrence electivity density should be used cautiously for fast assessment. Three electivity indices, Ivlev’s E and E′ and Jacobs’ D, were recommended for their ease of understanding.     

The reliability of palatability estimates obtained from rumen contents analysis and a field-based index of diet selection

Understanding herbivore foraging behaviour is often limited by not knowing the palatability of food items. Electivity indices can overcome this challenge by relating the consumption of food items to their relative abundance in the environment. We compare eight widely used electivity indices to relate the level of consumption of 48 food items in the rumen contents of 402 red deer Cervus elaphus in New Zealand to the environmental availability of food items. Additionally, we derive a field-based index of palatability from observed browse damage recorded during vegetation surveys at our site. Electivity indices were positively correlated with each other and with the browse survey index, which was also positively correlated with electivity indices from other studies. Thus, the choice of electivity index is unimportant when making comparisons of relative palatability among food items. Our analyses suggest that quantitative measures of palatability may be reliably inferred from observed browse damage in the field, which is quicker and less destructive to estimate than electivity indices. As a result, we demonstrate a useful technique for predicting herbivore diet selection.     

A comparison of methods for quantitative analysis of feeding selection of fishes

Synopsis Predator-prey data were obtained in a study of feeding ecology of land-locked alewife,Alosa pseudoharengus, in Claytor Lake, Virginia, and subjected to two indices of electivity as well as a nonparametric paired-comparisons statistical test. Values obtained from the two indices were often contradictory with each other and the statistical test. Attributes of the statistical procedure render it more appropriate than electivity indices to describe feeding selectivity in fish populations. Results of the study also demonstrated the importance of considering the size composition of zooplankton populations in prey preference studies; measures of selection were often positive for zooplankters 1.0 mm while negative for those of the same prey group < 1.0 mm length. Because numerous problems are inherent with in situ feeding selectivity studies of planktivorous fishes, selectivity determinations, regardless of methodology, should be construed in a relative rather than absolute context.     

Selective feeding behaviour of the common carp Esomus danricus (Ham.) in its natural habitat

Food selectivity by the common freshwater carp, Esomus danricus (Ham.) was studied through Ivlev's equation of electivity. The selective feeding seemed to be influenced by many factors, including the accessibility, abundance and taste, of the food items, the mechanical and physiological adaptations on the part of the fish to capture and digest those food organisms, as also the inherent and instinctive property of the fish to prefer certain types of food over the others. The absence of strong specific selectivity by the fish for a variety of planktonic food organisms was found to be a phenomenon compensatory to stenophagism, in order to make use of whichever food items remained in the habitat at the time when the preferred genera of plankton were not available, due to the process of succession.     

Food and food selection of cisco (Coregonus albula L.) in a dysoligotrophic lake

With minor exceptions, cisco (C. albula) in Lake Suomunjärvi fed on cladocerans and copepods. The food habits were different among two size classes of fish. The composition of cisco diet changed at different depths and times of the day, but the distribution of zooplankton was usually very similar in each period irrespective of the depth or diel period. Daphnia, Bosmina, Cyclops scutifer and Heterocope appendiculata were the main diet of cisco. Calculations of fish electivity indices showed that cisco selected usually large species of cladocerans and copepods and a small cladoceran, Bosmina coregoni. Body-size selection was clear for Daphnia and Bosmina.     

The food and feeding relationships of larval and 0+ year juvenile fishes in lowland rivers and connected waterbodies. II. Prey selection and the influence of gape

The relative importance of taxa- and size-specific prey selection, and the influence of gape on the prey consumed by the larvae and 0+ year juveniles of four fish species were investigated in 'main river', 'marina' and 'pond' macrohabitats in the lower River Trent, England. A general sequence of ontogenetic shifts in food consumption was reflected in the electivity indices of particular prey taxa, partly due to the restrictions imposed by the gape of 0+ year fishes. Certain taxa, however, were consistently selected over others, irrespective of size, suggesting that taxa-specific, as well as size-specific, prey characteristics may be important in the selection process. There were significant, positive relationships between maximum prey (zooplankton) length and maximum gape height for larvae, but not for 0+ year juveniles. The majority of fishes, however, consumed prey substantially smaller than the maximum theoretically possible inferred from their gape. The greater size ranges of zooplankton in connected waterbodies compared with main river channels provide suitable prey for a range of developmental steps and fish species, and may, thus, enhance recruitment success.     

The food and feeding habits of young perch, Perca fluviatilis, in Windermere

SUMMARY. The food of 0+ perch, Perca fluviatilis L. has been investigated in Windermere, the largest lake in the English Lake District. Whole guts and stomachs from 800 fish were investigated during four 24-h periods between June and September in 1975 and in 1976. Ciliates, algae, rotifers and cyclopoid nauplii constituted the first food of larval perch in Windermere. Vertical samples were taken from the zooplankton, at the same time and in the same location where the fish were sampled, and the concentration of the different food organisms studied. Electivity of the different food organisms was studied from lvlev's index, which showed positive electivity, or selection for some organisms, e.g. Bosmina obtusirostris (Sars.) and copepods (both cyclopoid and calanoid), and negative for others such as Daphnia hyalina var. galeata (Sars.), Leptodora kindti (Focke) and Bythotrephes longimanus (Leydig). In studying predatorprey relationships, Daphnia hyalina was chosen as the 'standard prey', being the most abundant of all food organisms in the diet of young perch, also because its maximum body-depth (carapace width (mm)) was among the largest of the cladocera and copepods in Windermere. Comparison of the maximum carapace width of Daphnia eaten with that of the plankton revealed that smaller specimens were taken by juvenile perch in July, but they tended to select larger organisms during August and September.     

Basis of selective predation by the aquatic larvae of the salamander, Ambystoma tigrinum

SUMMARY. 1. The effect of several factors on feeding selectivity by three size classes of larvae of Ambystoma tigrinum (Green) was examined. Five types of invertebrate prey were used: Chaoborus, Daphnia , damselflies, Hyalella and water boatmen.
2. Five variables were examined in relation to the electivity indices associated with each prey type: ease of prey consumption, ease of prey capture, relative prey size, prey activity, and microhabitat overlap. No single variable explained the observed electivity.
3. For small larvae (snout-vent length, SVL, <2.5cm) relative prey size was correlated with electivity. For large larvae (SVL >6.5cm), ease of prey consumption, prey activity, and relative prey size predicted electivity values. However, none of the measured variables was related to electivity values found for medium sized larvae (SVL >3.5 and <5.5cm).     

Food of an endangered cyprinodont (Aphanius iberus): ontogenetic diet shift and prey electivity

Synopsis We studied the ontogenetic diet shift and prey electivity of an endangered cyprinodontid fish endemic to the Iberian Peninsula, the Spanish toothcarp (Aphanius iberus). The toothcarp’s diet was omnivorous, dominated by harpacticoid copepods (Mesochra lilljeborgi and Tisbe longicornis), copepod nauplii and detritus. Diet composition varied greatly among habitats, depending on prey availability. In a rarely inundated habitat (glasswort), there was more consumption of the isopod Protracheoniscus occidentalis and the harpacticoid copepod Mesochra lilljeborgi, while in algal mats another harpacticoid (Tisbe longicornis), chironomid dipterans and invertebrate eggs were more important in diet. Although a benthic feeding habitat has previously been suggested, in our study the diet was based rather on water column organisms for both glasswort and algal mat habitats. There was also an ontogenetic diet shift, with an increase of mean prey length with fish length, clearly linked to a microhabitat change. Smaller fish showed positive electivity and greater reliance on planktonic prey (e.g. copepod nauplii, the harpacticoid copepods Mesochra lilljeborgi and Tisbe longicornis, the rotifer Brachionus plicatilis, and ostracods), while larger fish elected and preyed on more benthic organisms (e.g. Canuella perplexa, Mesochra rapiens, and ephydrid dipterans).     

Prey selection by flounder, Platichthys flesus, in the Douro estuary, Portugal

Prey selection by the flounder, Platichthys flesus (Linnaeus, 1758), in an estuarine nursery was investigated and the major factors influencing food choice by this species were assessed. Diet breadth was narrow, reflecting the low prey diversity observed in the benthos. A gradual ontogenetic shift from small prey such as amphipods to larger prey like polychaetes and bivalves was observed. Amphipods had positive electivity values in the upper estuary and negative values in the lower estuary.Polychaetes showed the inverse pattern. Bivalve electivity values were always positive. Differential selectivity throughout the estuary was mainly related to spatial segregation of flounders according to size, with the smaller individuals concentrating in the upper estuary and larger individuals concentrating in the lower estuary. Amphipods such as Corophium spp. play a crucial role in the flounder diet because of their small size, low mobility and diel activity pattern. As prey, the polychaete value increases throughout flounder ontogeny since the flounder size range is compatible with the larger mouth gapes and detection ability of larger fish. Bivalve electivity values for flounder are mainly related to high calorific values. The absence of Crangon crangon (Linnaeus, 1758) in the diet may be due to low water temperature since the cost–benefit involved in the capture of highly mobile prey is too high at low temperatures. It was concluded that flounder must use several sensory features to detect and capture prey in turbid estuarine waters and that field studies provide important background information on the actual predator preferences under natural conditions.     

Selective predation by carp (Cyprinus carpio L.) on benthic molluscs in Skadar Lake, Yugoslavia

Analysis of predation by carp ( Cyprinus carpio L.) on mollusc species in Skadar Lake, Yugoslavia, indicated both generalized and specific features of a predator-prey interaction. Vertical distribution of molluscs in sediments seemed unrelated to predation rates by carp. Although size selective predation may occur, electivity indices appeared most related to shell shape and thickness. Of the three most abundant species, electivity was high for Valvata piscinalis but low for Pyrgula annulata and Dreissencia polymorpha. Valvata has a more readily crushed shell, a higher proportion of occupied shells and a greater percentage of organic content than Pyrgula. Although a potentially preferred prey item, Dreissencia probably escaped intensive predation by being attached to larger objects. Carp appeared to utilize a sequence of tactile and gustatory cues in the prey selection process.     

Prey selection and functional response of juvenile lemon sharks Negaprion brevirostris

Prey preference of juvenile lemon sharks Negaprion brevirostris was examined in experiments conducted in field-based pens where the fish were offered meals of live teleosts: grey snapper Lutjanus griseus and yellow fin majorra Gerres cinereus in ratios of L. griseus : G. cinereus varying from 10:0 to 2:8. Relative electivity indices for each treatment showed that N. brevirostris preyed on G. cinereus rather than L. griseus . They showed a prey preference relating to density of G. cinereus (type II functional response) but not to L. griseus .     

Effects of prey size and foraging mode on the ontogenetic change in feeding niche ofColostethus stepheni (Anura: Dendrobatidae)

The feeding niche ofColostethus stepheni changes during ontogeny. Small individuals eat small arthropods, principally mites and collembolans, and larger frogs eat bigger prey of other types. The shift in prey types is not a passive effect of selection for bigger prey. There is a strong relationship between electivity for prey types and frog size, independent of electivity for prey size. Four indices of general activity during foraging (number of movements, velocity, total area utilized and time spent moving), which are associated with electivity for prey types in adult frogs and lizards, did not predict the ontogenetic change in the diet ofC. stepheni. Apparently, the behavioral changes that cause the ontogenetic change inC. stepheni are more subtle than shifts in general activity during foraging. Studies of niche partitioning in communities of anurans that do not take into consideration ontogenetic changes in diet and seasonal changes in the size structures of populations present a partial and possibly erroneous picture of the potential interactions among species.     

The sampling characteristics of electivity indices

Summary Electivity indices measure the utilization of food types (r) in relation to their abundance or availability in the environment (p). Foods that constitute a larger proportion of the diet than of the available foods are considered preferred; conversely those proportionately underrepresented in the diet are avoided. A food is eaten at random if its proportion in the diet equals its proportion in the environment. A family of electivity indices stemming from Ivlev's (1961) classic monograph exist and differ only in the particular algorithm used to calculate electivity from r and p.For each available index I graphed the values of electivity as contours for all combinations of r and p. These graphs are compared to illustrate the strengths and weaknesses of each index on the basis of the following criteria: 1) the value of the index when r=p for a food, 2) the symmetry of the electivity value as feeding deviates from random, 3) the possible range of index values, 4) the linearity of changes in electivity over the full range of r and p, 5) the sensitivity of the index to sampling errors, 6) the statistical testability of the electivity, and 7) the stability of the electivity value for a food type that changes relative abundance or occurs in combination with different food types. No one index ideally satisfies all the criteria.The host preferences of gypsy moth, Lymantria dispar, feeding on tree foliage in an undisturbed deciduous forest in southwestern Quebec, Canada were used to compare the available indices: Ivlev's electivity, E; Ivlev's forage ratio, E'; Jacob's modified electivity, D; Jacob's modified forage ratio, log Q; Chesson's alpha; Strauss' linear index, L; and Vanderploeg and Scavia's relativized electivity, E ^*. The electivity values calculated by each index differ one from another; host trees shown as preferred by one index will frequently appear avoided according to an alternative index. The rank order electivities for the 19 available host trees, however, are remarkably similar for all but Strauss' linear index, L. Populus grandidentata, Quercus rubra, Ostrya virginiana, and Amelanchier were the most preferred host trees in the sampled forest; Prunus serotina, Acer pensylvanicum, A. rubrum, Betula lutea, and Fraxinus americana were most avoided. The use of Vanderploeg and Scavia's E ^* index is recommended.     

Early dry-season community structure and habitat use of young fish in tributaries of the River Sinnamary (French Guiana, South America) before and after hydrodam operation

We examined fish community structure and habitat use at the start of the dry seasons: (1) in 10 tributaries of the River Sinnamary (French Guiana) before and after the start of dam operation, and (2) in 10 upstream tributaries and at 10 littoral sites in the newly-created reservoir after the start of operation to assess the impact on fish juveniles of a hydroelectric dam built on the river's lower section. After the first year of dam operation, juvenile fish communities downstream of the dam showed an important decrease of the relative abundance of Characiformes, and Perciformes dominated. Principal components analysis revealed a distinct upstream-to-downstream progression in the juvenile fish communities with post-reservoir downstream and reservoir sites representing transitions between the upstream and pre-reservoir downstream sites. Canonical correspondence analysis and electivity indices of fish-habitat associations revealed three relatively distinct groups of sites, corresponding to the downstream, reservoir and upstream taxa. The proportion of juveniles presenting higher-than expected frequencies (Fisher's exact test) towards local environmental variables was higher for taxa more often caught in upstream sites. Inversely, juvenile taxa more frequently observed in downstream and reservoir sites appeared less selective towards local environmental characteristics. In the downstream reaches of the river, hydrodam operation is expected to drive the fish community towards a new biologically accommodated state where tolerant species will dominate and sensitive species will be lacking. This revised version was published online in July 2006 with corrections to the Cover Date.     

On the diurnal rhythm and feeding electivity in kilka (<Emphasis Type="Italic">Clupeonella cultriventris</Emphasis>) in Rybinsk Reservoir

Diurnal rhythms, feeding electivity, and feeding resources were studied for the new for Rybinsk Reservoir fish species, kilka (Clupeonella cultriventris). It is revealed that, in Rybinsk Reservoir, kilka actively feeds during day time. Selective feeding is a characteristic of this fish. Yearlings prefer small Cyclopoida and their young specimens dominating in zooplankton. Two-year-old fish actively select innumerous but large crustaceans of genera Heterocope, Bythotrephes, Leptodora, and Cyclops. Possible factors determining selection of food items by kilka are discussed. These factors are as follows: zooplankters’ biomasses, the extent of plankters’ aggregations, body size, and visibility of some prey species for kilka (as determined by movement pattern, transparency and cuticle color, presence of eggs, or large eye).     

Habitat use of juvenile white sturgeon in the Kootenai River, Idaho and British Columbia

Ultrasonic telemetry was used to assess habitat features utilized by 36 endangered juvenile white sturgeon, Acipenser transmontanus, in the lower 120 km of the Kootenai River of Idaho, USA and British Columbia, Canada during the summer and early fall of 1999 and 2000. All 36 fish were initially captured in pools using gillnets and released there, but most of the subsequent telemetry contacts were in glides, indicating these fish moved freely between the two macro-habitats. The low electivity indices indicated little preference between glides and pools. Most contacts were in glides, in the outside bend of the river channel (50), and in or near a visually defined thalweg. Contacts were most often associated with sand substrates and no cover. Physical habitat characteristics (nose [bottom] water velocity, depth, substrate, and cover) were recorded at 168 contact locations. The combination of significantly greater velocities and depths at contact sites vs. non-contact sites (p < 0.01) indicated these fish actively found and used areas of higher velocity and greater depth within the Kootenai River. There was little cover found for fish in the river other than large sand dunes and depth. The combination of depth and nose velocity data supported the idea that large sand dunes are providing refugia in the form of velocity breaks.     

Assessing feeding electivity in Acanthaster planci: a null model analysis

Feeding electivity was investigated in a non-outbreaking population of the crown-of-thorns starfish Acanthaster planci (L.) from North Sulawesi, Indonesia. A null model-based approach was used to assess the feeding pattern of Acanthaster in relation to the availability of coral prey in the field. Of a total of 70 species of corals recorded as prey, massive species, particularly of Faviidae, tended to be more frequently consumed than would be expected under the assumption of random feeding by A. planci. Branched and encrusting/laminar forms of corals that occurred in relatively exposed sites were apparently not preferred, pointing to the importance of non-acroporan massive species of corals in cryptic habitats as prey for A. planci. The null model-based electivity index Z introduced here directly measured the deviation from random feeding, while two common indices (Ivlev’s and Vanderploeg and Scavia’s) only partially reflected such deviations (hence, prey selection cannot be accurately demonstrated by these). Electivity values (Z) for poritid species and Acropora palifera, the most common Acropora species in the study site, were significantly negative, indicating apparent avoidance of them by Acanthaster. Our results indicate that accessibility to different coral species and the choice/avoidance of certain species are the important elements of feeding in non-outbreaking populations of Acanthaster inhabiting spatially variable reef environments. A similar consideration may apply to the feeding patterns of other corallivores that possess superior/inferior mobility to Acanthaster. The present study emphasizes the merit of testing the observed patterns, using null models for a rigorous assessment of feeding preferences.     

The diet of breeding Dippers Cinclus cinclus and their nestlings in the catchment of the River Wye, mid-Wales: a preliminary study by faecal analysis

A preliminary study by faecal analysis into the diet of seven pairs of breeding Dippers and their nestlings was undertaken in the catchment of the River Wye in spring 1983. Macro-invertebrate items in faeces could be identified to family and quantified on the basis of mouthparts whilst the macro-invertebrate prey available to each pair of Dippers was assessed quantitatively by riffle-sampling.
In terms of numerical abundance, ephemeropteran nymphs, particularly Baetidae, and trichopteran larvae, particularly Hydropsychidae, were more important to adults and nestlings, respectively. When prey weights were taken into account, trichopteran larvae were the most important macro-invertebrate group to both, but especially to nestlings.
Some macro-invertebrate groups were selected in proportion to their abundance in the environment although indices of electivity indicated that prey selection for adults and nestlings, respectively, was negatively and positively correlated with prey weight. Significant changes in diet were also apparent with nestling age: smaller items, such as ephemeropteran nymphs, were eaten by younger nestlings whilst larger items, such as trichopteran larvae, became more frequent up to Day 15 after hatching.     

Deep Vision: An In-Trawl Stereo Camera Makes a Step Forward in Monitoring the Pelagic Community

Ecosystem surveys are carried out annually in the Barents Sea by Russia and Norway to monitor the spatial distribution of ecosystem components and to study population dynamics. One component of the survey is mapping the upper pelagic zone using a trawl towed at several depths. However, the current technique with a single codend does not provide fine-scale spatial data needed to directly study species overlaps. An in-trawl camera system, Deep Vision, was mounted in front of the codend in order to acquire continuous images of all organisms passing. It was possible to identify and quantify of most young-of-the-year fish (e.g. Gadus morhua, Boreogadus saida and Reinhardtius hippoglossoides) and zooplankton, including Ctenophora, which are usually damaged in the codend. The system showed potential for measuring the length of small organisms and also recorded the vertical and horizontal positions where individuals were imaged. Young-of-the-year fish were difficult to identify when passing the camera at maximum range and to quantify during high densities. In addition, a large number of fish with damaged opercula were observed passing the Deep Vision camera during heaving; suggesting individuals had become entangled in meshes farther forward in the trawl. This indicates that unknown numbers of fish are probably lost in forward sections of the trawl and that the heaving procedure may influence the number of fish entering the codend, with implications for abundance indices and understanding population dynamics. This study suggests modifications to the Deep Vision and the trawl to increase our understanding of the population dynamics.