Habitat structure directly affects aggression in convict cichlids Archocentrus nigrofasciatus

Aggressive behavior can be an important factor in determining how animals use and divide space and resources. Previous studies have shown that aggression in fishes can be influenced by a variety of factors, including water temperature and resource levels. In this study, we tested if the amount of habitat structure in the environment affected aggression levels in female convict cichlids Archocentrus nigrofasciatus. We performed a laboratory experiment in which we placed female convict cichlids into an aquarium with low or high amounts of habitat structure and monitored the dominant female's behavior toward the subordinate female. Aggressive behavior in convict cichlids primarily consists of chases and bites. We found that the total time the dominant female spent chasing the subordinate female was greater when there was a low amount of habitat structure as compared to when there was a high amount of habitat structure. We also found that both the average duration of a chasing bout and the number of bites directed at the subordinate fish increased when there was a low amount of structure, but the number of chases did not. These results indicate that increased habitat structural complexity decreases aggressive behavior in convict cichlids [Current Zoology 56 (1): 52-56, 2010].     

Disturbance chemical cues determine changes in spatial occupation by the convict cichlid Archocentrus nigrofasciatus

I studied the effect of disturbance chemical cues on fish that make trade-offs between foraging in an open area and remaining in a safe refuge. I used convict cichlids Archocentrus nigrofasciatus that were either visually exposed to a predator (n = 8) or exposed to water conditioned by chemical cues from disturbed conspecifics (n = 8). Fish visually exposed to a predator decreased their ingestion rate and spent more time in the refuge than in the foraging area, while fish receiving water from frightened conspecifics did not alter their ingestion rate or time spent in the refuge and foraging site, but increased their spatial occupation (i.e., motion). These results suggest that convict cichlids recognized the predator by visual cues. Moreover, disturbance cues are a form of threatening public information that may increase fish spatial occupation due to increased exploring behaviour; but is not sufficiently alarming to alter feeding or increase refuge use.     

Density dependence in foraging habitat preference of eastern grey kangaroos

For a free‐ranging forager, the suitability of a patch is dependent on population density, resource supply, resource quality, and the costs of foraging or dispersal. We quantified differences among three foraging habitats and compared this variation to temporal patterns of habitat preference by free‐ranging eastern grey kangaroos, Macropus giganteus. We investigated selection on a fine‐grained spatial scale, and asked whether habitat preference is constrained by density‐dependent mechanisms. Variation in the quantity and quality of resources among habitats was greatest during spring, when biomass and quality were highest, and differences among habitats were most pronounced. However, consistent and discernable differences among habitats were not obtained, indicating that the system fluctuated around an equilibrium state. Using isodar regressions to examine the consumer‐density relationships among habitats, open‐woodland habitat was favoured over the two open‐forest habitats for foraging. Seasonal isodars indicated that density dependence regulated preference between the three foraging habitats during autumn, spring and summer, but not during winter, when variability in resources among habitats was lowest.     

An anthropogenic habitat within a suboptimal colonized ecosystem provides improved conditions for a range‐shifting species

Many species are shifting their ranges in response to the changing climate. In cases where such shifts lead to the colonization of a new ecosystem, it is critical to establish how the shifting species itself is impacted by novel environmental and biological interactions. Anthropogenic habitats that are analogous to the historic habitat of a shifting species may play a crucial role in the ability of that species to expand or persist in suboptimal colonized ecosystems. We tested if the anthropogenic habitat of docks, a likely mangrove analog, provides improved conditions for the range‐shifting mangrove tree crab Aratus pisonii within the colonized suboptimal salt marsh ecosystem. To test if docks provided an improved habitat, we compared the impact of the salt marsh and dock habitats on ecological and life history traits that influence the ability of this species to persist and expand into the salt marsh and compared these back to baselines in the historic mangrove ecosystem. Specifically, we examined behavior, physiology, foraging, and the thermal conditions of A. pisonii in each habitat. We found that docks provide a more favorable thermal and foraging habitat than the surrounding salt marsh, while their ability to provide conditions which improved behavior and physiology was mixed. Our study shows that anthropogenic habitats can act as analogs to historic ecosystems and enhance the habitat quality for range‐shifting species in colonized suboptimal ecosystems. If the patterns that we document are general across systems, then anthropogenic habitats may play an important facilitative role in the range shifts of species with continued climate change.     

Aggression in closely related Malawi cichlids varies inversely with habitat complexity

Within the past 2 million years, the cichlids of Lake Malawi have diversified into well over 500 species resulting in one of the worlds largest lacustrine fish radiations. As a result, many of the habitats within the lake support a high diversity of species. In these highly species rich communities, male cichlids must acquire and defend a territory to successfully reproduce. Within the rock-dwelling cichlids of Lake Malawi (mbuna), this has resulted in the formation of polyspecific leks on the heterogeneous rocky benthos. Aggression is fairly common in these leks and has been tied not only to individual reproduction but to the larger phenomenon of community assembly and the maintenance of biological diversity. In this study, I examined the patterns of aggressive acts of four species within the mbuna genus Maylandia at two locations in the southern Lake Malawi. The number of aggressive acts of two sympatric species was examined at each location. At each site, one species defends territories over bedrock and the other over cobble. The number of aggressive acts across the four species was compared. The influence of habitat type on male aggression was examined and the targets of male aggression were identified to evaluate several hypotheses concerning the evolution of male aggression. The results show that aggression quantitatively varied among species, was largely directed towards heterospecifics, and was strongly influenced by habitat type. The aggressive behavior of one sympatric species pair, Maylandia benetos and Maylandia zebra, was observed under controlled laboratory conditions. Laboratory results support field observations: the bedrock associated species performed more aggressive acts and aggressive acts were directed equally at con- and heterospecifics. The results of this study suggest that habitat complexity plays a larger role in shaping aggressive behavior than other suggested factors such as competition for resources.     

Antipredator Behaviour and Suppressed Aggression by Convict Cichlids in response to Injury-released Chemical Cues of Conspecifics but not to those of an Allopatric Heterospecific

In aquatic environments, chemical cues serve as an important source of information for the detection of predation risk. Here, we investigate the response of convict cichlids, Cichlasoma nigrofasciatum, to injury-released chemical cues. We exposed pairs of juvenile convict cichlids first to dechlorinated tap water (control), then later to one of two test stimuli: 1. chemical cues from injured convict cichlids; or 2. chemical cues from injured mosquito fish, Gambusia affinis. Gambusia are allopatric and phylogenetically unrelated to convict cichlids. Gambusia skin was used to control for a general response to injured fish. In response to conspecific cues, convict cichlids significantly increased time spent near the bottom of test aquaria and time under a shelter object. In response to Gambusia skin, convict cichlids tended to increase time spent near the tank bottom but did not increase use of the shelter object. There was a trade-off between antipredator and agonistic behaviours. In response to convict cichlid cues, there was a significant reduction in the frequency of approaches and bites. Gambusia skin extract had no significant effect on aggressive behaviour. These data suggest a species-specific antipredator response to conspecific alarm pheromones in a New World cichlid fish and demonstrate a trade-off between predator avoidance and intraspecific aggression. Further, the presence of an alarm response in this model species sets the stage for the use of chemical cues as a research tool to manipulate predation risk in studies of the interaction between predation risk and reproductive behaviour.     

Time and travelling costs during chick‐rearing in relation to habitat quality in Little Owls Athene noctua

In many bird species, parents adjust their home‐ranges during chick‐rearing to the availability and distribution of food resources, balancing the benefits of energy intake against the costs of travelling. Over recent decades, European agricultural landscapes have changed radically, resulting in the degradation of habitats and reductions in food resources for farmland birds. Lower foraging success and longer foraging trip distances that result from these changes are often assumed to reduce the reproductive performance of parents, although the mechanisms are not well understood. We tested the behavioural response of chick‐rearing Little Owls Athene noctua to variation in habitat diversity in an agricultural landscape. We equipped females with GPS loggers and received adequate range‐use data for 19 individuals (6063–14 439 locations per bird). In habitats dominated by homogeneous cropland habitats, home‐ranges were over 12 ha in size, whereas in highly diverse habitats they were below 2 ha. Large home‐ranges were associated with increased flight activity (117% of that of birds in small home‐ranges) and distances travelled per night (152%), increased duration of foraging trips (169%) covering larger distances (246%), and reduced nest visiting rates (81%). The study therefore provides strong correlative evidence that Little Owls breeding in monotonous farmland habitats expend more time and energy for a lower benefit in terms of feeding rates than do birds in more heterogeneous landscapes. As nestling food supply is the main determinant of chick survival, these results suggest a strong impact of farmland characteristics on local demographic rates. We suggest that preserving and creating islands of high habitat diversity within uniform open agricultural landscapes should be a key target in the conservation of Little Owl populations.     

Diet and habitat overlap in two sympatric primate species, the Tana crested mangabey Cercocebus galeritus and yellow baboon Papio cynocephalus

Diet and habitat overlap was studied in two sympatric primate species sharing two neighbouring patches of fragmented gallery forest in Tana River, Kenya. Systematic data on feeding and ranging behaviour was collected on one group each of the Tana crested mangabey Cercocebus galeritus and yellow baboon Papio cynocephalus between August 1992 and February 1993. When rainfall was low and fruit resources scarce, yellow baboons spent most of their foraging time in the open woodlands while mangabeys foraged within the forest. At this period, diet and habitat overlaps between the two species were low. As rainfall increased, followed by a gradual rise in fruit availability, yellow baboons shifted their foraging range and both species became confined to the forest habitat. Consequently, both diet and habitat overlaps increased, peaking at the end of the rainy season. Mangabeys showed a reduced within-group dispersal and also spent significantly less time foraging in a given forest patch when yellow baboons were also present within the same forest patch.
Increased habitat and diet overlaps during the wet season need not have resulted in increased interspecific competition for food because at this period, fruits were readily available in the forest.     

Foraging and habitat use of common duikers,Sylvicapra grimmia,in a heterogeneous environment within the Soutpansberg,South Africa

We investigated aspects of the foraging behaviour and activity patterns of free‐ranging common duikers (Sylvicapra grimmia) within the Soutpansberg, South Africa. We used giving‐up densities (GUD) and camera traps to test for habitat selection and patch‐use behaviour by common duikers inhabiting a grassland containing distinct ‘islands' of woody vegetation. Foraging in or around a wooded island was affected by its surrounding vegetation. GUDs were significantly lower in portions dominated by tall grass and scattered ferns and highest in areas with open short grass and thick fern. Using grids of 5 × 5 stations, we mapped the duikers' foraging on a larger scale that incorporated neighbouring rocky hillsides. The duikers preferred feeding in areas with tall grass and scattered fern (sufficient cover and escape routes), followed by the wooded islands and thick fern (lack of sightlines/escape routes and presence of predator‐ambush sites), whereas little foraging occurred at the edges and rocky areas (hard substrate that impede escape potential). Photos (total 873) revealed solitary activity, highest in the late afternoon. Photos of vigilant individuals were mostly from rocky and fern habitats. Our results suggest that the duikers allocated their feeding efforts, activity and vigilance patterns to attune to their perceived risk of predation within their heterogeneous environment.     

High interindividual variability in habitat selection and functional habitat relationships in European nightjars over a period of habitat change

An animal's choice of foraging habitat reflects its response to environmental cues and is likely to vary among individuals in a population. Analyzing the magnitude of individual habitat selection can indicate how resilient populations may be to anthropogenic habitat change, where individually varying, broadly generalist populations have the potential to adjust their behavior. We collected GPS point data from 39 European nightjars (Caprimulgus europaeus) at a UK breeding site where restoration measures have altered large areas of habitat between breeding seasons. We calculated individual habitat selection over four breeding seasons to observe changes that might align with change in habitat. We also analyzed change in home range size in line with change in habitat availability, to examine functional relationships that can represent trade‐offs made by the birds related to performance of the habitat. Individual explained more of the variation in population habitat selection than year for most habitat types. Individuals differed in the magnitude of their selection for different habitat types, which created a generalist population composed of both generalist and specialist individuals. Selection also changed over time but only significantly for scrub habitat (60% decrease in selection over 4 years). Across the population, individual home range size was 2% smaller where availability of cleared habitat within the home range was greater, but size increased by 2% where the amount of open water was higher, indicating the presence of trade‐offs related to habitat availability. These results highlight that using individual resource selection and specialization measures, in conjunction with functional responses to change, can lead to better understanding of the needs of a population. Pooling specialist and generalist individuals for analysis could hide divergent responses to change and consequently obscure information that could be important in developing effective conservation strategies.     

Home‐range size and habitat use of European Nightjars Caprimulgus europaeus nesting in a complex plantation‐forest landscape

In Europe, the consequences of commercial plantation management for birds of conservation concern are poorly understood. The European Nightjar Caprimulgus europaeus is a species of conservation concern across Europe due to population depletion through habitat loss. Pine plantation‐forest is now a key Nightjar nesting habitat, particularly in northwestern Europe, and increased understanding of foraging habitat selection is required. We radiotracked 31 Nightjars in an extensive (185‐km²) complex conifer plantation landscape in 2009 and 2010. Home‐range 95% kernels for females, paired males and unpaired males were an order of magnitude larger than song territories of paired males, emphasizing the importance of habitats beyond the song territory. Nightjars travelled a mean maximum distance of 747 m from the territory centre each night. Home‐range placement relative to landscape composition was examined by compositional analysis. Pre‐closure canopy forest (aged 5–10 years) was selected at all scales (MCP, 95% and 50% kernels), with newly planted forest (aged 0–4 years) also selected within 50% kernels. For telemetry fixes relative to habitat composition within 2 km of their territory centre, individuals again selected pre‐closure and newly planted forest, and also grazed grass heath. Open ungrazed habitat was not selected, with implications for open habitat planning for biodiversity conservation within public‐owned forests. Despite the Nightjars’ selection for younger growth, moth biomass was greater in older forest stands, suggesting that foraging site selection reflects ease of prey capture rather than prey abundance. Within large plantation‐forest landscapes, a variety of growth stages is important for this species and our results suggest that grazing of open habitats within and adjacent to forest will additionally benefit the European Nightjar.     

Interaction between seed crypsis and habitat structure influence patch choice in a granivorous bird, the chaffinch Fringilla coelebs

Food abundance is an important determinant in habitat and patch selection but food accessibility and detectability is less often considered. Foraging on more cryptic seeds may increase predation risk by increasing the length of head down periods. Habitat structure may interact with this as birds are less able to detect predators with their head lowered in riskier obstructed habitats. We investigated patch choice in chaffinches Fringilla coelebs foraging in obstructed and open habitats and artificially manipulated the search times of seeds by colouring them either yellow or black. One trial consisted of a choice between the conspicuous seed in the open patch, and the cryptic seed in the obstructed patch; in the second trial the treatments were reversed. Individuals were more willing to forage in the obstructed habitat when the yellow seeds were present (43% of pecks made in the obstructed patch) than when the black seeds were present (18% of pecks in the obstructed patch). Differences in search time are likely to explain this result: yellow seeds were located almost twice as fast (1.26±0.60 seconds) as black ones (2.36±0.88 seconds). This experiment shows that individual foraging decisions may be influenced not only by food abundance but by the properties of individual food items (in this case seed crypsis) and the structure of the habitat they are present in.     

Scales and costs of habitat selection in heterogeneous landscapes

Summary Two scales of habitat selection are likely to influence patterns of animal density in heterogeneous landscapes. At one scale, habitat selection is determined by the differential use of foraging locations within a home range. At a larger scale, habitat selection is determined by dispersal and the ability to relocate the home range. The limits of both scales must be known for accurate assessments of habitat selection and its role in effecting spatial patterns in abundance. Isodars, which specify the relationships between population density in two habitats such that the expected reproductive success of an individual is the same in both, allow us to distinguish the two scales of habitat selection because each scale has different costs. In a two-habitat environment, the cost of rejecting one of the habitats within a home range can be expressed as a devaluation of the other, because, for example, fine-grained foragers must travel through both. At the dispersal scale, the cost of accepting a new home range in a different habitat has the opposite effect of inflating the value of the original habitat to compensate for lost evolutionary potential associated with relocating the home range. These costs produce isodars at the foraging scale with a lower intercept and slope than those at the dispersal scale.Empirical data on deer mice occupying prairie and badland habitats in southern Alberta confirm the ability of isodar analysis to differentiate between foraging and dispersal scales. The data suggest a foraging range of approximately 60 m, and an effective dispersal distance near 140 m. The relatively short dispersal distance implies that recent theories may have over-emphasized the role of habitat selection on local population dynamics. But the exchange of individuals between habitats sharing irregular borders may be substantial. Dispersal distance may thus give a false impression of the inability of habitat selection to help regulate population density.     

Optimal foraging by firemouth cichlids, Cichlasoma meeki, in a social context

Pairs of juvenile firemouth cichlids, Cichlasoma meeki, were allowed to feed on Tubifex worms for 5 min. In the experiment, the worms were distributed so as to simulate a patchy resource with two levels of average abundance. The dominant member of each ichlid pair exhibited feeding behaviour consistent with expectations from optimal foraging theory, i.e. it fed primarily in the most profitable patch initially. When food was relatively abundant, the subordinate fish fed predominantly in the second most profitable patch. In this way it minimized the costs associated with agonistic activity. This study illustrates the usefulness of a multivariate approach in the study of foraging behaviour.     

Chainsawing for conservation: Ecologically informed tree removal for habitat management

Summary In many ecosystems, increases in vegetation density and the resulting closure of forest canopies are threatening the viability of species that depend upon open, sunlight‐exposed habitats. Consequently, we need to develop management strategies that recreate open habitats while minimizing the impacts on non‐target areas. Selective logging creates canopy gaps, but may result in undesirable effects in other respects. Thus, chainsaws have not been a popular tool for conservation. We conducted a landscape‐scale experiment to test whether selective tree removal can restore patch‐level habitat quality for Australia’s most endangered snake (Hoplocephalus bungaroides) and its main prey (the lizard Oedura lesueurii). We selectively removed canopy trees surrounding 25 overgrown rock outcrops and compared the resultant habitat structure and abiotic conditions to 30 overgrown, shady outcrops and 20 open, sunny outcrops. Removing vegetation decreased canopy cover by 19% in experimental plots and increased incident radiation and thermal regimes. These changes increased the availability of suitable shelter sites for our target species by 131%. At the landscape scale, our manipulations had a trivial effect on forest habitat; by increasing the area of sun‐exposed outcrops, we decreased forest cover by <0.1%. Our results show that targeted canopy removal can increase the availability of sun‐exposed habitat patches for endangered species in biologically meaningful ways. Thus, selective tree felling may be an effective conservation tool for open‐habitat specialists threatened by vegetation overgrowth.     

Slow, fast and in between: habitat distribution and behaviour of larvae in nine species of libellulid dragonfly

1. Activity and microhabitat use are important factors determining species performance in habitats that differ in permanence and species composition of top predators. This study examined the relationship between the distribution across a gradient of habitat permanence and an associated transition in the composition of top predators and the behaviour of species of larval dragonflies. It also assessed the relationship between larval behaviour, body size and the duration of the larval stage. In laboratory mesocosms the mobility of the different species was measured, as was the extent to which they associated with artificial vegetation. 2. Species mobility was positively related to their natural occurrence in habitats in which invertebrates or small‐bodied fish were the top predators, and negatively related with the frequency with which species co‐existed with large‐bodied fish, the permanence of the habitat and the length of the larval stage. 3. Rather than falling into strict low and high mobility categories, habitat generalists that occurred across the habitat gradient, co‐existing with different top predators, had variable mobility levels. In these generalists, mobility was positively related to how frequently they were found in natural habitats in which invertebrates were the top predators. 4. The extent to which species utilized the artificial vegetation in mesocosms was associated with the length of the larval period but was not associated with mobility or species habitat distribution in the field.     

Short‐term Costs and Benefits of Habitat Complexity for a Territorial Fish

An increase in habitat complexity is thought to decrease visibility and the territory size of visually oriented animals. Hence, the addition of physical structure has been viewed as a useful restoration technique to increase the density of territorial species, particularly in stream fishes. However, a decrease in territory size may have a negative effect on the fitness of individual organisms. We attempted to quantify some of the positive and negative effects of increasing habitat structure on the behaviour and growth rate of wild young‐of‐the‐year (YOY) Atlantic salmon. Fish were exposed to one of two habitat treatments in mesh enclosures in Catamaran Brook, New Brunswick: a gravel substrate (low complexity) or a gravel substrate with boulders added (high complexity). Wild‐caught individuals were tagged, weighed and measured before being stocked at densities of 1·m^?2 for 7‐d trials. While fish from high‐complexity treatments benefited from lower rates of aggression, they also had lower foraging rates and smaller territories compared to those in low‐complexity treatments. Specific growth rate, however, did not differ significantly between treatments. While the addition of structure to a habitat may be beneficial at the population level in terms of an increase in population density, our results suggested that individual fish may pay some short‐term costs in these environments. Further research is needed to evaluate the longer term costs and benefits of adding structure to improve the habitat quality for stream salmonids.     

Designing mosaic landscapes for Black Grouse Tetrao tetrix using multi‐scaled models

With increasing pressures on land for human use, it is important to identify the habitat requirements of key species, not just in terms of a correlation with a given habitat feature, but also the relationship between species presence and its coverage, proximity to other habitat types, and importance at different spatial scales. We used maximum entropy to estimate the optimal proportions of 18 habitat types, plus elevation and habitat richness associated with the presence of leks of Black Grouse Tetrao tetrix within an 800‐km² study area in Perthshire, Scotland. We repeated the analysis at several radii (0.2–3 km) to assess how the importance of different habitats changed with proximity to lek and scale. We then examined habitat features or combinations of features that were associated with large leks or positive lek growth. Models at all radii had satisfactory predictive power. Using response curves from maxent , we constructed ideal habitat mixes for leks at each radius. At the 2‐km radius, suitability was highest with around 20% each of three moorland types and open/mixed forestry, whereas close to leks (0.2 km), higher proportions of grouse moor and lower proportions of closed‐canopy woodland were optimal. The relationship between habitat and lek size or direction of lek growth was complex, indicating that a landscape containing large or productive leks can be the result of more than one combination of habitats. This demonstrates a degree of flexibility in designing landscapes for Black Grouse conservation, so landowners can prioritize combinations of habitats that are the most practical and/or economical, while still serving the requirements of the target species.     

Habitat preferences and habitat restoration options for small‐bodied and juvenile fish species in the northern Murray–Darling Basin

Degradation of instream habitats in the northern Murray–Darling Basin has occurred through numerous stressors, including siltation, clearing of bankside vegetation, intrusion of livestock and impacts of pest species. A better understanding of habitat preferences of native fish species could help guide future instream habitat restoration actions. The habitat choices of seven native fish species, juvenile Murray Cod (Maccullochella peelii), juvenile Golden Perch (Macquaria ambigua ambigua), juvenile Silver Perch (Bidyanus bidyanus), adult Murray–Darling Rainbowfish (Melanotaenia fluviatilis), adult Olive Perchlet (Ambassis agassizii), adult Un‐specked Hardyhead (Craterocephalus stercusmuscarum fulvus) and adult carp gudgeons (Hypseleotris spp.) were tested in preference troughs to help inform potential habitat restoration actions in the Condamine catchment. Each species was given a choice between pair combinations of open sandy habitat, submerged macrophytes, emergent plants and rocky rubble. Habitat preferences varied between species. Murray Cod, Golden Perch, carp gudgeons and Olive Perchlets preferred structure over open sandy habitat, whilst juvenile Silver Perch, Un‐specked Hardyhead and Murray–Darling Rainbowfish did not avoid open sandy habitats. Juvenile Murray Cod preferred rocky rubble habitat over all other habitat choices. Use of complex rock piles to provide nursery habitat for Murray Cod populations is a potential restoration option. Introduction of rock could also benefit Golden Perch and carp gudgeons. Use of emergent plants, submerged macrophytes and rocky rubble for habitat restoration all appear to have merit for one or more species of small‐bodied fishes or juvenile stages of larger sized fishes. Rocky rubble or floating attached macrophytes could be viable restoration options in areas too turbid to establish submerged macrophytes. These habitat interventions would complement existing actions such as re‐snagging and provision of fish passage to assist with sustainable management of native fish populations.     

Divergence in social foraging among morphs of the three‐spined stickleback,Gasterosteus aculeatus

Foraging in social groups has a number of benefits but can also increase the risk of exploitation. High tendency to shoal may be correlated with groups foraging, although facultatively social fish adjust both shoaling decisions and food resource defence based on intrinsic and extrinsic factors. The main aim of this study was to examine the relationships between shoaling, solitary foraging and aggression, forager tolerance of conspecifics joining at a discovered food patch and forager exploitation of resources discovered by others. We used two intra‐lacustrine three‐spined stickleback morph pairs, lava and mud, and monomorphic morphs from each of lava and mud habitats. The lava morph formed less cohesive shoals, was bolder during solitary foraging, approached and entered an occupied food patch less frequently than the mud morph, suggesting a link between shoaling and the propensity for social foraging. However, shoaling tendency and joiner tolerance were not correlated at a population level. Intralacustrine lava and mud morphs differed more markedly in joiner tolerance than morphs from single habitat lakes, whereas the opposite was true for shoaling tendency. We conclude that, in addition to differentiation in shoaling tendency, the lava and mud morphs differ in social foraging and these variations may act to promote population divergence. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 194–203.