Null model analysis of species associations using abundance data

The influence of negative species interactions has dominated much of the literature on community assembly rules. Patterns of negative covariation among species are typically documented through null model analyses of binary presence/absence matrices in which rows designate species, columns designate sites, and the matrix entries indicate the presence (1) or absence (0) of a particular species in a particular site. However, the outcome of species interactions ultimately depends on population-level processes. Therefore, patterns of species segregation and aggregation might be more clearly expressed in abundance matrices, in which the matrix entries indicate the abundance or density of a species in a particular site. We conducted a series of benchmark tests to evaluate the performance of 14 candidate null model algorithms and six covariation metrics that can be used with abundance matrices. We first created a series of random test matrices by sampling a metacommunity from a lognormal species abundance distribution. We also created a series of structured matrices by altering the random matrices to incorporate patterns of pairwise species segregation and aggregation. We next screened each algorithm-index combination with the random and structured matrices to determine which tests had low Type I error rates and good power for detecting segregated and aggregated species distributions. In our benchmark tests, the best-performing null model does not constrain species richness, but assigns individuals to matrix cells proportional to the observed row and column marginal distributions until, for each row and column, total abundances are reached. Using this null model algorithm with a set of four covariance metrics, we tested for patterns of species segregation and aggregation in a collection of 149 empirical abundance matrices and 36 interaction matrices collated from published papers and posted data sets. More than 80% of the matrices were significantly segregated, which reinforces a previous meta-analysis of presence/absence matrices. However, using two of the metrics we detected a significant pattern of aggregation for plants and for the interaction matrices (which include plant-pollinator data sets). These results suggest that abundance matrices, analyzed with an appropriate null model, may be a powerful tool for quantifying patterns of species segregation and aggregation.     

The effects of castration, sex ratio and population density on social segregation and habitat use in Soay sheep

We analysed 16 years of census data gathered on the island of Hirta (archipelago of St. Kilda) to investigate the effects of castration, population density, sex ratio, season and group type on habitat use and social segregation of Soay sheep. From 1978 to 1980, 72 male lambs were castrated. We used this experiment to study how a change in reproductive status could affect sociality and habitat choice of these males. Males, females and castrates were all segregated outside the rutting season in autumn. Castrates were the least segregated from females in spring and summer but were most segregated from them during the pre-rut. The more equal the sex ratios, the higher was the degree of social segregation. The three sex classes used similar habitat types, namely, Holcus agrostis, Agrostis festuca and Calluna habitats. Holcus agrostis and Agrostis festuca were top- and second-ranked in female and castrate habitat use, while Holcus agrostis and Calluna were the two top habitat types used by rams. It is unclear why males included Calluna heath habitats, but it cannot be excluded that they might have shifted their use depending on forage availability. A lack in differences in habitat use between castrates and females suggests that body size differences alone cannot be the driving factor for habitat segregation in male and female Soay sheep and that there are reasons other than body size that could motivate reproductive males to use additional habitat types, such as Calluna heath. Although habitat use shifted from one habitat type to the next between low- and high-population-density years and between seasons, there was no clear link between population density and how different groups (male, female or castrate) used these areas. We discuss effects of reproductive status, population density and sex ratio on social segregation and habitat use and suggest that these factors need to be taken into account when investigating causes of sexual segregation in ungulates.     

Role of competition and predation in determining habitat occupancy of Cerithiidae (Gastropoda: Prosobranchia) on the rocky,intertidal, Red Sea coasts of Sinai

Six cerithiid snail species occur on rocky intertidal flats along the Sinai coasts of the Red Sea: Clypeomorus moniliferum, Cerithium caeruleum, C. scabridum, C. columna, Clypeomorus tuberculatum and Cerithium echinatum. The present study, conducted at 22 stations covering almost the whole length of Sinai, covered the 4 yr period from October 1972 to August 1976, and describes the habitat occupancy of 5 of these species (C. echinatum is excluded for lack of data). Several of these species (sometimes all) often occur together, and in such cases are distincly segregated by habitat. However all species considerably overlap in their distribution along the axes of four major interrelated abiotic gradients, thus excluding the possibility that habitat segregation is determined by larval settlement preferences. Other distributional patterns observed at some sites, such as lack of overlap or contact between belts of the various species and the relative abundance of food available to all species, make postlarval competitive interactions unlikely. The existence and the degree of proximity of a coral reef with its associated predatory fishes, influence the cerithiids' distributional patterns. Differences between the cerithiid species in their vulnerability to fish predation, associated with differences between sites in the abundance and the accessibility of predatory fishes, and in the availability of refuges for each cerithiid species, can satisfactorily explain the observed distributional patterns including co-occurrence with habitat segregation. It is proposed that habitat segregation is caused by predation on young stages by generalist fishes which may totally eliminate a certain species at a given site; the same site may provide refuges for recruits of another species, allowing these to survive to an advanced age. In general, the flat's structural complexity is associated with its diversity of refuges from predation, and hence with the number of co-existing species. This mechanism for co-existence and habitat segregation in tropical Cerithiidae may also be instrumental in maintaining the high species diversity of other tropical benthic communities.Paper No. 12 in the series Colonization of the Eastern Mediterrancan by Red Sea species immigrating through the Suez Canal     

Ecology,feeding competition and social structure in baboons

Predictions of the model of van Schaik (1989) of female-bonding in primates are tested by systematically comparing the ecology, level of within-group contest competition for food (WGC), and patterns of social behaviour found in two contrasting baboon populations. Significant differences were found in food distribution (percentage of the diet from clumped sources), feeding supplant rates and grooming patterns. In accord with the model, the tendencies of females to affiliate and form coalitions with one another, and to be philopatric, were strongest where ecological conditions promoted WGC. Group fission in the population with strong WGC was “horizontal” with respect to female dominance rank, and associated with female-female aggression during a period of elevated feeding competition. In contrast, where WGC was low, females’ grooming was focused on adult males rather than other females. Recent evidence suggests that group fission here is initiated by males, tends to result in the formation of one-male groups, and is not related to feeding competition but to male-male competition for mates. An ecological model of baboon social structure is presented which incorporates the effects of female-female competition, male-male competition, and predation pressure. The model potentially accounts for wide variability in group size, group structure and social relationships within the genus Papio. Socio-ecological convergence between common baboons and hamadryas baboons, however, may be limited in some respects by phylogenetic inertia. Received: 22 April 1994/Accepted after revision: 9 December 1995     

Invasional interference due to similar inter- and intraspecific competition between invaders may affect management

As the number of biological invasions increases, the potential for invader-invader interactions also rises. The effect of multiple invaders can be superadditive (invasional meltdown), additive, or subadditive (invasional interference); which of these situations occurs has critical implications for prioritization of management efforts. Carduus nutans and C. acanthoides, two congeneric invasive weeds, have a striking, segregated distribution in central Pennsylvania, U.S.A. Possible hypotheses for this pattern include invasion history and chance, direct competition, or negative interactions mediated by other species, such as shared pollinators. To explore the role of resource competition in generating this pattern, we conducted three related experiments using a response-surface design throughout the life cycles of two cohorts. Although these species have similar niche requirements, we found no differential response to competition between conspecifics vs. congeners. The response to combined density was relatively weak for both species. While direct competitive interactions do not explain the segregated distributional patterns of these two species, we predict that invasions of either species singly, or both species together, would have similar impacts. When prioritizing which areas to target to prevent the spread of one of the species, it is better to focus on areas as yet unaffected by its congener; where the congener is already present, invasional interference makes it unlikely that the net effect will change.     

Testing sexual segregation and aggregation: old ways are best

The study of sexual segregation has received increasing attention over the last two decades. Several hypotheses have been proposed to explain the existence of sexual segregation, such as the "predation risk hypothesis," the "forage selection hypothesis," and the "activity budget hypothesis." Testing which hypothesis drives sexual segregation is hampered, however, by the lack of consensus regarding a formal measurement of sexual segregation. By using a derivation of the well-known chi-square (here called the sexual segregation and aggregation statistic [SSAS]) instead of existent segregation coefficients, we offer a reliable way to test for temporal variation in the occurrence of sexual segregation and aggregation, even in cases where a large proportion of animals are observed alone. A randomization procedure provides a test for the null hypothesis of independence of the distributions of males and females among the groups. The usefulness of SSAS in the study of sexual segregation is demonstrated with three case studies on ungulate populations belonging to species with contrasting life histories and annual grouping patterns (isard, red deer, and roe deer). The existent segregation coefficients were unreliable since, for a given value, sexual segregation could or could not occur. Similarly, the existent segregation coefficients performed badly when males and females aggregated. The new SSAS was not prone to such limitations and allowed clear conclusions regarding whether males and females segregate, aggregate, or simply mix at random applicable to all species.     

Honeybees use a Lévy flight search strategy and odour-mediated anemotaxis to relocate food sources

The availability of food resources changes over time and space, and foraging animals are constantly faced with choices about how to respond when a resource becomes depleted. We hypothesise that flying insects like bees discover new food sources using an optimal Lévy flight searching strategy and odour-mediated anemotaxis, as well as visual cues. To study these searching patterns, foraging honeybees were trained to a scented feeder which was then removed. Two new unrewarding feeders, or ‘targets’, were then positioned up- and downwind of the original location of the training feeder. The subsequent flight patterns of the bees were recorded over several hundred metres using harmonic radar. We show that the flight patterns constitute an optimal Lévy flight searching strategy for the location of the training feeder, a strategy that is also optimal for the location of alternative food sources when patchily distributed. Scented targets that were positioned upwind of the original training feeder were investigated most with the numbers of investigations declining with increasing distance from the original feeder. Scented targets in downwind locations were rarely investigated and unscented targets were largely ignored, despite having the same visual appearance as the rewarding training feeder.     

Mortality is associated with social rank in the clown anemonefish (<Emphasis Type="Italic">Amphiprion percula</Emphasis>)

Elucidating the causes of post-recruitment mortality is a vital step toward understanding the population dynamics of coral reef fishes. Predation is often considered to be the primary proximate cause of mortality. It has, however, proven difficult to discern the relative contributions of predation and other processes, such as competition for food, shelter, or mates, to patterns of mortality. To determine which other processes might be important drivers of mortality patterns, factors related to mortality in the clown anemonefish Amphiprion percula (Lacepède, 1802) were examined. Patterns of mortality will not be driven by predation in A. percula, because these fish are well protected from predators by their close association with sea anemones. Mortality rates were based on the disappearance of known individuals from a population of 201, in 57 groups, during a 1-year field study (in 1997), in Madang Lagoon, Papua New Guinea. Mortality rate of A. percula was low (14% per annum) compared to other coral reef fish, probably due to the protection from predators afforded by the anemone. Six factors (reef, depth, anemone diameter, number of individuals, density, and standard length) showed no association with the probability of mortality (P>0.05). Rank was the only factor associated with the probability of mortality (P<0.03); low-rank individuals (ranks 4–6) suffered a higher mortality rate than high-rank individuals (ranks 1–3) (P<0.01). The most likely explanation for this pattern was that competition for rank, amongst individuals within an anemone, resulted in some individuals evicting their subordinates. Individuals probably competed for rank because it conferred access to reproduction, and not because it conferred access to food or shelter. Such competition for reproduction will be intense whenever some individuals obtain a greater share of reproduction than others do, and it may be an important process influencing the dynamics of coral reef fish populations.Electronic Supplementary Material Supplementary material is available for this article if you access the article at . A link in the frame on the left on that page takes you directly to the supplementary material.Communicated by J.P. Grassle, New Brunswick     

Interspecific comparisons of growth and diet among late larvae of three co-occurring clupeoid species in the Kii Channel,Japan

Growth and diet were compared among larvae of Japanese anchovy Engraulis japonicus, Japanese sardine Sardinops melanostictus and Pacific round herring Etrumeus teres. Compositions of prey items of the three species in the same month showed greater similarity than for the same species in different months. Prey size as well as prey taxa of the three species overlapped considerably with one another. Therefore, interspecific prey competition is likely in the case of limited food availability. The most abundant species tended to change from anchovy to round herring in early winter, from round herring to sardine in late winter and from sardine to anchovy in early spring, indicating a temporal segregation in use of the nursery grounds. Similar seasonal changes in growth rates were observed for the three species. Although interspecific prey competition is likely, the temporal segregation and similar temporal changes in growth rates could favor their coexistence.     

Food abundance affects energy intake and reproduction in frugivorous female Assamese macaques

In most mammals, female fertility and reproduction are strongly influenced by nutritional status and, therefore, by foraging conditions. Here, we investigate the relationship between food resources, feeding competition, energy intake and reproduction in a group of wild female Assamese macaques (Macaca assamensis) in northeastern Thailand. Over 2,100 h of data on feeding behaviour, energy intake and activity budgets were combined with data on resource characteristics, female reproduction and physical condition. We found that an increase in food availability had a positive effect on female energy intake and conception rates. In addition, it appeared that females incurred energetic costs during lactation and that females with a better physical condition during the mating season were more likely to conceive. The annual birth season occurred a few months before the annual peak in food availability, causing peak lactation to coincide with a period of high food availability. This suggests that females use the increased food abundance to compensate for the energetic costs of lactation. Neither energy intake rates nor activity budgets were influenced by female dominance rank, even during periods when the levels of contest competition were predicted to be high. In line with this, we found no evidence for rank-related differences in reproduction. The apparently limited influence of feeding competition in female Assamese macaques adds to the debate on the extent to which patterns in feeding competition and fitness can reliably be predicted based on ecological conditions. We suggest that this may partially be resolved by including potential competition-reducing mechanisms into the predictive framework.     

Aggresive competition and individual food consumption in wild brown capuchin monkeys (Cebus apella)

Summary The impact of aggresive competition on food intake at all the resources used is analysed for every member of a group of brown capuchin monkeys (Cebus apella) in the Manu National Park, Peru, where they live in groups of 8–14 animals. An individual's food intake at a given tree was affected independently both by its domirance rank (Fig. 1) and by how much aggression it received (Table 5). Food intake was not strongly affected by body size when dominance rank was held constant by partial correlation. At food sources where high rates of fighting occurred, an individual's food intake depended more on its domirance status than on the rate of aggression it received (Fig. 2). However, food intake at resources where rates of fighting were low depended mostly on the rate of aggression received. When aggression over food was absent, the food intakes of dominants and subordinates were equal. Dominants had significantly greater total energy intake (20.5% more) than did subordinates, even though more than one third of their diet came from food sources where little or no fighting occurred (Fig. 3). Energy intake was also significantly greater for individuals that received little aggression. The only adult that emigrated from the main study group was the individua with the lowest energy intake. Competition for food within groups was more than ten times as intense as competition between brown capuchin groups.     

Environmental and genetic influences on mating strategies along a replicated food availability gradient in guppies (Poecilia reticulata)

Food availability is expected to influence the relative cost of different mating tactics, but little attention has been paid to this potential source of adaptive geographic variation in behavior. Associations between the frequency of different mating tactics and resource availability could arise because tactic use responds directly to food intake (phenotypic plasticity), because populations exposed to different average levels of food availability have diverged genetically in tactic use, or both. Different populations of guppies (Poecilia reticulata) in Trinidad experience different average levels of food availability. We combined field observations with laboratory “common garden” and diet experiments to examine how this environmental gradient has influenced the evolution of male mating tactics. Three independent components of variation in male behavior were found in the field: courtship versus foraging, dominance interactions, and interference competition versus searching for mates. Compared with low-food-availability sites, males at high-food-availability sites devoted more effort to interference competition. This difference disappeared in the common garden experiment, which suggests that it was caused by phenotypic plasticity and not genetic divergence. In the diet experiment, interference competition was more frequent and intense among males raised on the greater of two food levels, but this was only true for fish descended from sites with low food availability. Thus, the association between interference competition and food availability in the field can be attributed to a genetically variable norm of reaction. Genetically variable norms of reaction with respect to food intake were found for the other two behavioral components as well and are discussed in relation to the patterns observed in the field. Our results indicate that food availability gradients are an important, albeit complex, source of geographic variation in male mating strategies.     

Nonlinear effects of food aggregation on interference competition in mallards

Previous studies of interference competition have shown an asymmetric effect on intake rate of foragers on clumped resources, with only subordinate individuals suffering. However, the food distributions in these studies were uniform or highly clumped, whereas in many field situations, food aggregation is intermediate. Here we investigated whether food distribution (i.e., uniform, slightly clumped, and highly clumped) affects the behavioral response of mallards foraging alone or competing with another. Although the amount of food was the same in all distributions, the mallards reached higher intake rates, visited fewer patches, and showed longer average feeding times in the highly clumped distribution. Competing mallards had lower intake rates on the slightly clumped than on the uniform or highly clumped food distributions. Subordinates generally visited more patches and had shorter feeding times per patch, but their intake rates were not significantly lower than those of dominants. Therefore, we propose that subordinates do not necessarily suffer from interference competition in terms of intake rate, but do suffer higher search costs. In addition, although dominants had significantly higher average feeding times on the best quality patches of the highly clumped food distribution, such an effect was not found in the slightly clumped distribution. These findings indicate that in environments where food is aggregated to a lesser extent, monopolization is not the best strategy for dominants. Our results suggest that interference experiments should use food distributions that resemble the natural situation animals are faced with in the field.     

Competitive regimes in forest-dwelling Hanuman langur females (Semnopithecus entellus)

Van Schaik’s socioecological model predicts interrelations among food distribution, competitive regimes, and female social relationships. To test the internal consistency of the model, feeding competition was examined in three differently sized groups of a forest-dwelling population of Hanuman langurs (Semnopithecus entellus). The nutritional condition of females was used as a direct indicator of feeding competition and related to the seasonal variation in resource distribution and abundance. Female dominance hierarchies were characterized by displacements. Dominance hierarchies were significantly linear and relatively stable, but less so with increasing group size. Physical condition correlated with dominance rank and high-ranking females were in the best condition, indicating within-group contest competition. The strength of this relationship became less pronounced with increasing group size. The females of the medium-sized group were in the best physical condition indicating between-group contest plus within-group scramble competition. Closer examination revealed variable costs and benefits of group foraging with a predominance of within-group scramble competition when food was more abundant. The results support some basic predictions of the model. Limiting food abundance was bound to ubiquitous within-group scramble competition. The use of clumped resources translated into differences in net energy gain based on dominance. In contrast to the predictions, group-size-related costs and benefits were related to food abundance instead of food distribution. As predicted, within-group contest competition was linked to a linear dominance hierarchy. The absence of nepotism and coalitions in Hanuman langurs may be attributed to dominance hierarchies that are unstable through time, probably minimizing fitness gain via kin support. Received: 25 May 1999 / Received in revised form: 18 February 2000 / Accepted: 25 February 2000     

Observations on the diurnal vertical migrations of an oceanic animal community

Diurnal changes in abundance caused by vertical migrations have been examined in populations of copepods, ostracods, euphausiids, amphipods, decapods, chaetognaths, siphonophores and fish. The animals were taken in a series of hauls made over a 24 h period with an opening-closing midwater trawl system (RMT 1+8), consisting of a net of 1 m² mouth area combined in the same frame as one of 8 m² mouth area. The samples were taken at 250 m depth in a position 30°N; 23°W on 7/8 April 1972. The specific composition of the community and the numbers of individuals changed continuously with time. The numbers of fish, decapods and chaetognaths increased at night, but those of copepods, ostracods and euphausiids decreased. More species of fish, decapods and copepods were present by night than by day, whereas the numbers of species per haul for other groups remained fairly constant. The relative abundances of groups caught by the RMT 1 have been analysed, but similar treatment of the RMT 8 samples was impossible as only 3 groups were taken from this net. Non-migrants were a minority in every group except chaetognaths. Migrant species have been put into one of 6 transitory categories according to their patterns of abundance and hence migrations. Within each category, migratory behaviour varied both inter- and intraspecifically. The patterns of abundance of many species were smooth and continuous, suggesting slow migratory cycles of small amplitude. Conversely, extensive migrants had discontinuous patterns and presumably more rapid movements. Few migrants had a steady numerical plateau between their upward and downward migrations, and most apparently moved up or down continuously. The presence of migratory species in the sampled layer depended upon the time of day or night. It is concluded that, in a vertical series of hauls, the depths of occurrence of migrants will vary with the sampling time. Further-more, a vertical series will show a species minimum migration range but not necessarily its maximum. Individuals of some species were out of phase with the migrations of their main populations. There is evidence that the distributions and migrations of some species of decapods, euphausiids, copepods and fish could be related to the distribution of underwater light. Three pairs of congeneric copepod species were both spatially and temporally segregated for at least part of their diurnal cycles. Such an orderly arrangement could provide a means of reducing competition between species. Some species, however, overtook others on their migrations and the pattern of underwater light cannot, therefore, regulate the distribution of all species in the same way.     

Competition affects risk-sensitivity in foraging shrews

Summary Studies of risk-sensitive foraging have so far focused only on the effect of food demand on choice of feeding site. We suggest that competition is likely to be another factor influencing risksensitivity. A choice experiment with common shrews showed that, in the absence of competition, risk-aversion increased with increasing food intake relative to requirement. When apparent competitors were present, however, shrews were risk-indifferent regardless of their estimated requirement. The switch to risk-indifference in the presence of competitors appears to be an all-or-nothing rule of thumb which is not modified by experience with reward probability distributions.     

Effects of food particle concentration on feeding current velocity in sex species of marine Bryozoa

Experiments were carried out using established methods to measure feeding current velocity in six species of marine Bryozoa collected in 1982 and 1983 around low-tide mark at Port St. Mary, Isle of Man (Flustrellidra hispida, Alcyonidium gelatinosum, A. hirsutum, Electra pilosa, Membranipora membranacea and Bowerbankia gracilis feeding on Tetraselmis suecica). It was found that there was a positive correlation between feeding current velocity and the concentration of food (phytoplankton) in the environment. It was also found that, at a fixed particle concentration, feeding current velocity was strongly correlated with lophophore height and (less strongly) with tentacle number. The competitive and evolutionary implications of these relationships are considered. It is tentatively suggested that interspecific competition for food may be of some significance in marine Bryozoa and that the removal of food supplies to competing colonies may play a role in colony overgrowth and competition for space.     

Predation, scramble competition, and the vigilance group size effect in dark-eyed juncos (Junco hyemalis)

In socially feeding birds and mammals, as group size increases, individuals devote less time to scanning their environment and more time to feeding. This vigilance “group size effect” has long been attributed to the anti-predatory benefits of group living, but many investigators have suggested that this effect may be driven by scramble competition for limited food. We addressed this issue of causation by focusing on the way in which the scan durations of free-living dark-eyed juncos (Junco hyemalis) decrease with group size. We were particularly interested in vigilance scanning concomitant with the handling of food items, since a decrease in food handling times (i.e. scan durations) with increasing group size could theoretically be driven by scramble competition for limited food resources. However, we showed that food-handling scan durations decrease with group size in an environment with an effectively unlimited food supply. Furthermore, this food-handling effect was qualitatively similar to that observed in the duration of standard vigilance scans (scanning exclusive of food ingestion), and both responded to changes in the risk of predation (proximity of a refuge) as one might expect based upon anti-predator considerations. The group size effects in both food-handling and standard scan durations may reflect a lesser need for personal information about risk as group size increases. Scramble competition may influence vigilance in some circumstances, but demonstrating an effect of competition beyond that of predation may prove challenging. Received: 22 September 1998 / Received in revised form: 1 February 1999 / Accepted: 14 February 1999     

Effects of temperature and food availability on growth and reproduction in the neustonic pedunculate barnacle Lepas anserifera

To elucidate the life history of neustonic animals, growth and reproductive patterns were investigated in the hermaphroditic pedunculate barnacle Lepas anserifera in field and laboratory experiments in Wakayama, western Japan from 2006 to 2008. The effects of temperature (19, 24 or 29°C) and food availability (once or twice a week) on growth and reproduction were also studied in the laboratory. The barnacles grew and matured rapidly, especially in the field: individuals on the average grew from 3 mm to more than 12 mm in capitulum length within 15 days and some were brooding. High temperature and high food availability resulted in greater growth. High temperature also resulted in earlier maturation of both testes and ovaries, whereas the effect of food availability was less clear. The rapid growth and maturation, together with earlier maturation at higher temperatures, may be an adaptation to ephemeral floating objects to which they attach.     

Spatial and temporal feeding patterns of copepods from the North Pacific centra gyre

Spatial and temporal feeding patterns (determined from an index of gut fullness) are described for 10 typical species of calanoid copepods collected from the North Pacific central gyre (September 1968 to June 1977), an area where the zooplankton is food limited and there were a-priori reasons to suspect that feeding and competition for food were important in regulating zooplankton community structure. Over 100 samples from 11 cruises to the eastern part of the gyre were examined, and patterns of gut fullness were related to environmental variables and the copepod species structure. The copepods studied all tended to be omnivores and food generalists. Males had lower indices of gut fullness than females but both males and females of a species had similar spatial and temporal feeding patterns. Guts were usually fuller at night than during the day, even in nonmigrating species; however, within nighttime depth distributions, no depths were preferred for feeding. There were also differences between species in mean gut fullness, but different species tended to have similar spatial and temporal feeding patterns. There was considerable spatial variability, and locales could be identified in which most species had higher indices of gut fullness. The copepods were not necessarily more abundant in these locales, nor did these tend to be areas of above average chlorophyll concentration. These patterns were consistent with relatively nonselective feeding, and there was no evidence that these species separate their niches by feeding at differing places or times.