Habitat-specific chemical cues influence association preferences and shoal cohesion in fish

The structure of social animal groups can be dynamic, characterized by high rates of group fission and fusion. Despite this, group composition is often well ordered by factors such as species, body size and by numerous other phenotypic traits. Research in shoaling fishes has revealed that individuals refine group membership decisions still further and are capable of assimilating chemical cues pertaining to recent habitat and prey use by prospective group mates, preferring to associate with others whose recent resource use history closely matches their own. In this study, we firstly examined the dynamics of the formation and breakdown of these preferences, revealing that they can be acquired and replaced in a matter of just a few hours. Using such cues enables individuals to accurately assess the resource use of conspecifics, allowing them to indirectly sample the local environment while reducing the chances of acquiring outdated information that can precipitate maladaptive behaviors. Secondly, we found that shoals composed of individuals with shared recent habitat use history were more cohesive compared to those where the constituent individuals differed in recent habitat use. Increased shoal cohesion may reduce predation risk, and could enhance the ability of individuals to detect and use social information.     

A risk-balancing trade off between foraging rewards and predation hazard in a shoaling fish

Summary In controlled experiments, shoaling European minnows fed in equal numbers on two equally-rewarded food patches, and likewise on unequal patches the numbers feeding reflected food levels. For equal food, minnows did not feed at the patch where they had encountered a simulated diving avian predator, whereas they chose to feed at this risky patch when it was more valuable. By avoiding hazard for equal food, but accepting predator risk for higher food rewards, the minnows performed a risk-balancing trade-off.     

Size-assorted fish shoals and the majority's choice

Similarity among group members may serve as a defence against visually hunting predators that preferentially attack individuals who are phenotypically different from the group majority. The presence of such odd individuals in an otherwise homogeneous group may, however, increase the vulnerability of the other group members as well. Individuals might thus be expected to form uniform groups in order to decrease predation risk, not only in trying to avoid being odd in a group, but also when attempting to avoid being accompanied by odd individuals. This hypothesis was tested with small and large three-spined sticklebacks, Gasterosteus aculeatus. Focal fish were offered the choice between a shoal consisting of conspecifics that were all similar in body length to the focal fish (matching shoal) and a shoal in which one or a few individuals differed in body length from the focal fish (non-matching shoal). In the control experiment, all individuals in the non-matching shoal differed in size from the focal fish. The control confirmed that individuals preferably joined the matching shoal when the alternative option was to be odd in another one. However, when the alternative for size-assortative shoaling was to belong to the majority in a mixed shoal, the shoal choice of individuals appeared on average to be random. Visual contact with a live pike, Esox lucius, did not affect the shoal choice pattern. Furthermore, despite the frequency-dependent nature of the oddity effect, varying the number of odd individuals in the non-matching shoal did not have a significant effect on individual's shoaling decisions. These results suggest that size-assortativeness in fish shoals is not a result of individuals avoiding being among the majority in a mixed group. Received: 2 September 1998 / Received in revised form: 12 May 1999 / Accepted: 29 May 1999     

Bigamy and mate choice in the biparental cichlid fish Cichlasoma nigrofasciatum

Summary Sixty percent of C. nigrofasciatum males showed bigamous behaviour when held in small, outdoor ponds at an adult sex ratio of 5 males to 10 females. No male held more than two pair bonds simultaneously. Bigamy had reproductive benefits; bigamous males, on average, fathered more broods that survived well into the free-swimming fry stage than did monogamous males. In ponds where adult size varied, no positive correlations were found, for either sex, between adult size and frequency of spawning or success of broodrearing, despite predictions that larger size would confer reproductive benefits on both sexes. A significant positive correlation was found between the sizes of mated individuals in the pond where both sexes varied in size. On theoretical grounds this relationship is likely to be more strongly influenced by female than by male choice of spawning partner.     

Social information,conformity and the opportunity costs paid by foraging fish

Animals pay opportunity costs when pursuing one of several mutually exclusive courses of action. We quantified the opportunity costs of conforming to the behaviour of others in foraging sticklebacks (Pungitius pungitius), using an arena in which they were given the option of shoaling in one area or searching for food in another. Fish foraging in the absence of stimulus conspecifics found the prey patch sooner and spent longer exploiting it than those in trials where a stimulus shoal was present. Furthermore, in trials where the stimulus shoal exhibited feeding cues, subjects approached them sooner and spent more time shoaling with them, exploring less of the arena than in trials where the stimulus shoal exhibited no such cues. This suggests sensitivity not only to the mere presence of conspecifics, but also to the social information that they produce. We also saw that groups of focal fish, compared to single individuals, were less influenced by the stimulus shoal and explored more of the arena, a behaviour that may be attributed to facilitation, competition or both. Such opportunity costs are likely to be offset by benefits such as reduced predation risk, and we discuss this in terms of the trade-offs associated with living in groups.     

Influence of fish on habitat choice of water birds: a whole system experiment

It is notoriously difficult to study population interactions among highly mobile animals that cannot be meaningfully confined to experimental plots of limited size. For example, migratory water birds are believed to suffer from competition with resident fish populations for shared food resources. While observational evidence in support of this hypothesis is accumulating, replicated experiments addressing this issue at the proper spatial scale are lacking. Here, we report from a replicated whole-system experiment in which we stocked large (0.07 km2), shallow (< or =2.5 m deep), highly eutrophic ponds in the bird protection area "Ismaninger Speichersee mit Fischteichen" with different densities of carp and assessed the responses of water birds and their food resources during summer over several years. In all years, the biomasses of benthic macroinvertebrates, macroalgae, and macrophytes as well as the densities of herbivorous, carnivorous, and omnivorous water birds were reduced in carp ponds compared to fishless ponds. The negative effects of carp on food resources and on the numbers of water birds feeding in carp ponds increased over the season (May-September) and were stronger at high than at low stocking densities of carp. Consequently, differences in resource densities between ponds with and without carp increased, and the ranking of ponds with respect to resource densities became more predictable over the season. These factors may have contributed to a seasonal improvement of the birds' abilities to track resource densities across ponds, as suggested by tight correlations of bird numbers on ponds with resource densities late, but not early, in the season.     

The indirect effects of eutrophication on habitat choice and survival of fish larvae in the Baltic Sea

The structure of the habitat is usually crucial for growth and survival of young life stages. Presently, some nursery areas of fish larvae are changing due to eutrophication, e.g. due to enhanced growth of ephemeral filamentous algae at the expense of perennial species. We studied the influence of two habitats, one with filamentous algae (Cladophora glomerata) and the other with bladder wrack (Fucus vesiculosus), on habitat choice of pike larvae (Esox lucius) in the absence/presence of a predator or a competitor. We further tested whether the habitat choice is adaptive in increasing survival under predation threat. In contrast to expectations, pike larvae preferred the habitat with ephemeral filamentous algae to the bladder wrack, thriving in clean waters, independent of the presence/absence of both predator/competitor. In addition, the survival of the larvae was higher in the filamentous algae in the presence of predators, which suggested that the habitat preference of the larvae was adaptive. The structure of the bladderwrack habitat was probably too open for newly hatched larvae, which implies that F. vesiculosus and other large brown algae are not as important refuges for young larvae as previously thought.     

Kin recognition in rainbowfish (Melanotaenia eachamensis): sex, sibs and shoaling

Living with relatives can be beneficial to individuals via the evolution of kin-directed altruism, but this is tempered by the increased risk of inbreeding. Therefore, in social species, the ability to recognise relatives can be highly advantageous. This study focuses on kin discrimination in the Lake Eacham rainbowfish, Melanotaenia eachamensis, an endangered freshwater species from north-east Queensland, Australia. First, I examined kin recognition abilities when a combination of both chemical and visual recognition cues was available. When given a choice of shoaling with same-sex groups, females spent significantly longer with full-sibs rather than half-sibs, full-sibs rather than non-relatives and half-sibs rather than non-relatives. Males spent significantly longer shoaling with full-brothers versus half-brothers, but showed no other shoalmate preferences. Second, in the presence of only chemical cues, females did not discriminate among groups of different levels of relatedness, but males showed a non-significant tendency to associate with full-sibs rather than non-relatives. Male shoaling behaviour seemed to be more influenced by factors other than relatedness, e.g. intra-sexual competition. Finally, I found that the shoaling preferences of females changed when exposed to groups of males. Females preferred to associate with non-relatives rather than half-brothers and non-relatives rather than full-brothers. There was no significant difference in the time spent with half-brothers versus full-brothers. Taken together, my results suggest that females have very good kin recognition abilities. They prefer to shoal with female relatives but avoid male relatives, and so are able to balance the benefits of nepotism and the costs of incest. Received: 2 May 2000 / Revised: 26 July 2000 / Accepted: 30 July 2000     

Experimental assessment of sensory modalities of coral-reef fish larvae in the recognition of their settlement habitat

One of the great mysteries of coral-reef fish ecology is how larvae locate the relatively rare patches of coral-reef habitat on which they settle. The present study aimed to estimate, by experiments in aquaria, the sensory modalities of coral-reef fish larvae for senses used in searching for their species settlement habitat. Larval recognition of settlement habitat can be based on the detection of conspecifics and/or of characteristics of coral habitat using visual, chemical and mechanical cues. For this study, larvae were captured with crest nets and were then introduced into experimental tanks that allowed testing of each type of cue separately (visual, chemical or mechanical cues). Among the 18 species studied, 13 chose their settlement habitat due to the presence of conspecifics and not based on the characteristics of coral habitat, and 5 species did not move toward their settlement habitat (e.g. Scorpaenodes parvipinnis, Apogon novemfasciatus). Among the different sensory cues tested, two species used the three types of cues (Parupeneus barberinus and Ctenochaetus striatus: visual, chemical and mechanical cues), six used two types (e.g. Myripristis pralinia: visual and chemical cues; Naso unicornis: visual and mechanical cues), and five used one type (e.g. Chrysiptera leucopoma: visual cues; Pomacentrus pavo: chemical cues). These results demonstrate that many coral-reef fish larvae could in practice use sensory cues for effective habitat selection at settlement, and have the ability to discriminate species-specific sensory cues.Communicated by J. Krause     

Altered responses to female odors in parasitized male mice: neuromodulatory mechanisms and relations to female choice

There is accumulating evidence that females may preferentially select parasite-free or -resistant males. Minimal attention has, however, been paid to the mate preferences and responses of the parasitized male hosts themselves. Here, we considered the effects of parasitic infection on male host mate responses, the neuromodulatory correlates of these responses, and the relations of these responses to female mate choice. Using an odor “preference” test, we examined the effects of different stages of an acute, sub-clinical infection with the naturally occurring, enteric, single host, protozoan parasite, Eimeria vermiformis, on the responses of male mice, Mus musculus domesticus, to the odors of estrous females along with the responses of uninfected females to the parasitized males. At 4 days post-infection (non-infective, pre-patent stage) E. vermiformis-infected male mice showed a significantly decreased preference for the odors of estrous females, whereas at 10 days post-infection (infective, patent stage) infected males showed a significantly increased preference for the odors of estrous females. Parasitized males displayed no significant changes in their responses to the odors of non-estrous females, supporting effects on the reproductively related responses of the host. In parallel, estrous females displayed a reduced interest in the odors of infected males. Least interest was expressed in the odors of the patent, infective males, consistent with the avoidance of contagion. Using selective opioid peptide receptor agonists and antagonists we found evidence that enhanced kappa opioid peptide (e.g., dynorphin) activity was related to the decreased sexual interest of the pre-infective males, while augmented delta opioid peptide (e.g., enkephalin) activity was associated with the enhanced responses of the infective males to females. We further showed that acute kappa opiate administration reduced the responses of uninfected males to females and that uninfected females displayed modified responses to the odors of uninfected males subject to acute modifications of opioid activity. We suggest that these differential shifts in endogenous opioid activity in the parasitized males are associated with and, or related to alterations in neuro-immune and endocrine functions. These findings show that parasitic infection can have, depending on the stage of infection and associated neuromodulatory changes, either significant facilitatory or inhibitory effects on male host preferences for and responses to females. Received: 22 April 1996 / Accepted after revision: 15 March 1997     

The costs and benefits of territory sharing for the Caribbean coral reef fish,Scarus iserti

Summary Along the Caribbean coast of Panama, groups of unrelated female striped parrotfish, Scarus iserti, co-defend a common feeding territory. Field manipulations of group size and composition were performed to examine the benefits and costs accrued by dominant fish within these shared territories. Dominant fish benefit from the presence of relatively large subordinates because they share in the defense of the territory. Removals of these fish caused increases in defense time and decreases in feeding time for dominant group members. Dominants benefit from the presence of small subordinates because they increase the foraging efficiency of dominants. Removals of smaller subordinates caused reductions in the feeding time of dominant fish, although no changes in defense time occurred. Concurrently, dominant fish reduce costs of resource depletion by displacing subordinate group members from good food patches. Dominance interactions within a group reduce the amount of time subordinates spend feeding (subordinate individuals fed at higher rates following the removal of a dominant) and limit a subordinate's access to high quality resources. This combination of benefits and reduced costs ultimately contributes to the economic defensibility of a striped parrotfish territory and has led to the evolution of group territorial behavior in the absence of kin selection and cooperative parental care.     

Predator shoaling moderates the confusion effect in blue-green chromis,Chromis viridis

Summary In experiments, blue-green chromis [Chromis viridis (Cuvier 1830)] were fed on either scattered or aggregated swarms of brine shrimp (Artemia sp.). Ten runs with each prey dispersion treatment were performed with shoals of one, two, five and ten chromis. The mean lag in reaching peak feeding rate for fish fed on aggregated prey was significantly shorter in the larger chromis shoals. In contrast, with the scattered treatment all such lags were similar and very short. As foraging proceeded, higher feeding rates were observed in the larger feeding shoals, regardless of prey dispersion. Prey capture success (i.e. the rate of retention of intercepted prey) declined with time, but was significantly higher in groups of ten fish. Two main conclusions emerge. Firstly, grouping facilitated initiation of feeding by individuals preying on concentrated swarms and reduced the delay in reaching a maximum feeding level. This may have been due to a suppression of the confusion effect through reduced reliance upon vigilance. Secondly, reduced vigilance allowed larger shoals of chromis to feed effectively over more extended periods. Trends of increasing shoal cohesion and decreasing prey retention rate with time were consistent with a postulated increase in antipredator vigilance with declining feeding motivation.     

The effects of mating system on male mate choice in a coral reef fish

Summary We show how mate limitation appears to be critical in determining whether or not males exercise mate choice among available females. Thalassoma bifasciatum is a Caribbean reef fish with two distinct mating patterns: group-spawning and pair-spawning. In both mating systems, female fecundity is variable and size dependent, and female availability is high. However, sperm competition among group-spawning males apparently limits the number of effective matings in which a male may engage, whereas territorial pair-spawning males have little or no such limitation. Group-spawning males should be discriminating in their choice of mates and our data confirm this: there is strong evidence for assortative mating in group-spawns, with more large males joining in mating groups around large females. In contrast, pair-spawning males show no indication of mate preferences, and spawn with all females who arrive at their territories.     

Reproductive rates,operational sex ratios and mate choice in St. Peter’s fish

Operational sex ratio (OSR) theory predicts that sexual differences in potential reproductive rates (PRRs) create biases in the OSR and thus determine the relative strength of sexual selection (competition and choice) operating on each sex. Although this theory is well accepted, empirical studies that quantify it are still lacking. This paper presents such a study. I measured the natural OSR of Galilee St. Peter’s fish (Sarotherodon galilaeus) in the field (Lake Kinneret) and examined the direction of mate choice in the laboratory. The OSR in Lake Kinneret was male biased. Both a male-biased sex ratio and higher male reproductive rates (twice as fast as females) contributed to the skew in the OSR, but the sexual differences in PRR were shown to be the main factor causing variation in the OSR. Females, the sex with the lower PRR, were more selective for mates. The faster male reproductive rate may explain why females are more selective for mates despite varying less in quality. Received: 29 May 1995/Accepted after revision: 13 April 1996     

Application of sensory and microbial analysis to assess quality of fish in Siliguri city of West Bengal, India

Sensory and certain microbial analyses were applied to assess the quality of raw fish sold at a market in Siliguri cityof West Bengal, India. In regular surveys undertaken during June to August 2008, a particular fish species was randomly selected, its source was noted and a sensory analysis, the quality index method (QIM) was applied to assess its quality Raw fish samples were also collected and a small quantity (about 1 g) of scales oran upper layer of the skin surface (forscale-less fish samples), gill, liverand a portion of gut with gut-contents were aseptically removed for enumeration of the total aerobic heterotrophic bacteria, Aeromonas spp., Pseudomonas spp., Salmonella spp. and coliform counts. Oreochromis mossambicus and Tenulosa ilisha recorded significantly higher QIM scores, compared to other species (p<0.05). Riverine fish, Lepidocephalichthys guntea and Channa punctatus scored the lowest QIM scores (0) while scores for Puntius ticto and Mystus vittatus and pond cultured species like Cirrhinus mrigala, Catla catla, Labeo rohita, Labeo bata and Cyprinus carpio were very marginal (p<0.05). Aeromonas spp., Salmonella spp. and total coliforms were recorded from all the studied species while Pseudomonas spp. was isolated from only seven species. Among the tissues examined, the lowest counts of total heterotrophic bacteria, Aeromonas spp., Pseudomonas spp., Salmonella spp. and total coliforms were recorded from the skin in every fish species. Highest counts of pathogenic bacteria (except Pseudomonas spp.) were recorded in Tenulosa ilisha for all the tissues except liver. Since fish are properly cooked in Bengali households, the risk of disease from fish consumption is relatively less. However, some tribes residing in the region are known to consume undercooked fish and proper cooking methods should be followed in view of the present findings to avoid health risks. Besides, utmost care should be taken while handling fish.     

Time and recruitment costs as currencies in manipulation studies on the costs of reproduction

Life history theory predicts that parents will have lower Darwinian fitness if they tend clutches that are above or below the size they naturally produce. We experimentally tested for relationships between fitness and clutch size in Tree Swallow (Tachycineta bicolor) offspring and parents. Over 130 trios of nests initiated on the same day were randomly divided among reduce (-3 eggs), control (3 eggs picked up and replaced), or add (+3 eggs) manipulations. Pre-manipulation modal clutch size was six eggs (range before manipulations was 1-10; afterwards, it was 1-11). Hatching took longer in larger clutches, but the proportion of eggs hatching and fledging was similar for clutches from 4 to 10, so that clutches of 10 produced the maximum number of fledgling. Parental feeding rates were higher for larger broods, but per capita feeds to nestlings were fewer, and nestlings were smaller. Nonetheless, survival of both young and adults, based on recaptures in subsequent years, was not significantly affected by manipulations. Manipulations also had no significant effect on subsequent reproduction, including the number of fledglings produced by either local recruits or returning breeders. Collectively, our results failed to detect fitness costs associated with tending larger clutches for either parents or the offspring reared and suggested directional selection for larger clutch size. However, because clutches that hatch later produce fewer recruits, the extra days required to lay more eggs and to fledge extra young may eliminate a large part of the advantage that would accrue to parents producing enlarged clutches. For example, our data suggest that there may be less than a 16% benefit to producing nine instead of six eggs, rather than 50%, as is suggested by experimentally manipulated egg numbers alone. Thus, time, rather than costs of reproduction, may be the crucial constraint selecting against Tree Swallows laying larger clutches.     

Female choice in sage grouse: the roles of attraction and active comparison

Previous studies of female choice in sage grouse Centrocercus urophasianus have implicated both the acoustic quality and repetition rate of the stereotyped strut display as putative cues for female choice. Stages in the choice process at which specific components of male courtship display influence female decisions were investigated using field observations of female pre-mating behavior. Females visited a subset of territorial males and then actively chose one of these as a mate. The order in which males were visited suggested that females searched until an acceptable mate was found, rather than employing a “best-of-n” tactic. Numbers of females visiting a male were related to differences in an acoustical component of display (inter-pop interval) whereas the probability that a visiting female mated was related to display rate (Table 3), indicating that initial attraction and active choice are influenced by different components of display. In addition, inter-pop interval and display rate tended to covary inversely (Fig. 1), suggesting that attraction and active choice may impose conflicting selection pressures on display performance. Received: 11 November 1995/Accepted after revision: 16 March 1996