Changing tactics: dominance, territoriality, and the responses of "primary" males to competition from conditional breeders in the variegated pupfish (Cyprinodon variegatus)

Using variegated pupfish, we examined the flexibility in "primary" male tactics when dealing with the aggressive costs of competition. Analogous to conditional mating strategies, we expected primary males to exhibit one of two interrelated tactics (i.e. dominance or territoriality) in response to different numbers of competitors. In the field, competitors influenced aggression. Primary males defended territories; residents facing more intruders engaged in more chases and obtained fewer spawns per female. In the laboratory, primary males showed dominance at low density, controlling most of the aquarium. With increasing competitor numbers, primary males reduced the area controlled and defended territories. Territories occurred with intermediate to high competitor numbers and only under male-biased sex ratios. During these interactions, aggression was highest and competitors were too many for dominant males to suppress the assertions made by each subordinate to increase its rank. Relinquished control of the entire domain enabled a previously subordinate male to establish a territory in the undefended portion of the aquarium. Reduction in defended area related to a reduction in territorial males' spawning success relative to dominants. These results suggested that primary males, like conditional breeders, would adopt the tactic that enabled them to spawn despite the constraints of competition.     

Agonistic behavior in a schooling fish: form,function and ontogeny

Synopsis Although juvenile chum salmon,Oncorhynchus keta, are generally regarded as a schooling fish, when presented with a defensible point-source of food, some individuals abandon schooling, aggressively subordinate competitors, and monopolize food. When food is removed, fish gradually abandon solitary agonistic behaviors and return to schooling behavior. Agonism increases in frequency and intensity as juveniles age. The ability to alternate facultatively between schooling and solitary agonistic behavior may enable juvenile chum to respond to local patterns of food distribution and predation risk. The ontogenetic increase in agonism may result in school dispersal as fish move from the estuary into coastal waters, and may well reflect a shift in the costs versus the benefits of schooling as fish mature and become less vulnerable to predation.     

Facultative schooling behavior in the spottail shiner (Notropis hudsonius): possible costs and benefits

Synopsis If an individual is less susceptible to predation in a group it should be able to reduce its vigilance for predators and perhaps also its tendency to flee from predators, thereby gaining time and energy for feeding and other activities. These predictions were examined in the field (Veronica Lake, northwestern Ontario) using a facultative schooling species, the spottail shiner (Notropis hudsonius). Schools of various sizes attacked with a fish predator model had a significantly lower reaction distance to the predator than solitary shiners. Solitary shiners must therefore invest more time and energy in defense but may gain a feeding advantage because their visual field does not overlap with conspecific competitors.     

Male tactics and reproductive success in the harem polygynous bat Saccopteryx bilineata

Alternative tactics in reproductive behavior enable individualsto maximize their fitness in relation to competitors in thesame population. In many taxa, territoriality is a common tacticof males to increase their reproductive success. In the batSaccopteryx bilineata, territorial males defend roosting areasfor females against other males and court females throughout the year. Peripheral males in the same colonies do not defendterritories but compete with territorial males for reproductionwith females. In this study, we monitored the behavior of themales in a natural colony over three reproductive seasons.We compared morphological and age data and measured the reproductiveoutput of males adopting the territorial or peripheral tactic.No differences in body size or weight were detected betweenmale types, but the probability of adopting a tactic seemedto be age dependent. Peripherals were often young males andreplaced territorials in several cases, whereas the oppositecase was not observed. Peripherals were not excluded from reproduction,but territorials were more likely to reproduce. Variation in reproductive success was high within both male tactics, andthe reproductive success of some peripherals was comparableto territorials, but, on average, the reproductive successof territorials was more than twice as high. Therefore, behavioraltactics do not seem to be equally profitable in general butmay represent different phases in the reproductive life of manyS. bilineata males.     

Producer–Scrounger Games in a Spatially Explicit World: Tactic Use Influences Flock Geometry of Spice Finches

Group-foraging animals can either search for their food (producer) or search for opportunities to join the food discoveries of others (scrounger). To maximize food returns, producers should distance themselves from potential competitors whereas scroungers should increase proximity to potential producers. We investigated the extent to which playing one or the other tactic affected an individual's location in captive flocks of ground-feeding spice finches ( Lonchura punctulata ) as they foraged for hidden clumps of food on an aviary floor. We constrained some individuals to use the producer tactic by pre-training them to find food hidden under lids. Constrained producers foraged significantly further from the center of flocks than constrained scroungers. Flocks with many scroungers were significantly more compact than flocks with fewer scroungers. The results are consistent with published simulations of spatially explicit producer–scrounger models and suggest that the use of producer and scrounger foraging tactics be included as a factor that affects an individual's position within foraging groups.     

Elevated corticosterone levels elicit non-calling mating tactics in male toads independently of changes in circulating androgens

Competition among males for a limited number of females may result in the expression of condition-dependent alternative mating tactics. In such cases, decision rules mediating mating tactic expression are likely to be influenced by a male's external as well as internal environment. For example, experimental studies with anuran amphibians (frogs and toads) indicate that changes in the social-acoustic environment alter the probability that an individual adopts a calling versus non-calling "satellite" mating tactic. However, there is considerable variation in the behavioral responses of individuals in such studies, suggesting that physiological differences among individuals play an important role in tactic expression. For instance, recent models predict that natural elevation in adrenal glucocorticoids during vocal production alters androgen production and/or neural activity to mediate transitions between reproductive tactic expression in anuran amphibians. Using corticosteroid injections, we show that elevation in circulating corticosterone levels significantly increase the probability that free-ranging male Great Plains toads (Bufo cognatus) and Woodhouse's toads (B. woodhousii) adopt a non-calling satellite tactic. Corticosterone-induced behavioral transitions occurred rapidly (<1 h) and independently of fluctuations in circulating androgen levels, suggesting a direct effect of glucocorticoids on brain centers controlling vocal production. We discuss our findings in the context of behavioral studies that alter the social-acoustic environment to examine its influence on tactic expression.     

Experience does not alter alternative mating tactics in the burying beetle Nicrophorus vespilloides

Alternative male mating tactics are widespread, but the cuesthat determine which tactic is adopted remain unclear. Sizeis commonly associated with alternative mating tactics, butit is not known how individuals gauge their size effectively,especially given that size is relative and frequency dependent.One possibility is that interactions with conspecifics are usedto assess size, relative to potential competitors, and thusfine-tune tactics. Success in mating might also influence matingtactics given that this should indicate the potential availabilityof mates in the population. We tested these ideas in the buryingbeetle Nicrophorus vespilloides, examining whether individualsuse the outcome of larval or adult interactions as cues to adjustthe tactics used to acquire mates. Male N. vespilloides employ2 tactics; search for a carcass, a resource required for reproduction,or release a pheromone (call) to attract a mate. Males are plasticin the amount of time they invest in each tactic, and in a relatedspecies (Nicrophorus orbicollis), male size influences the tacticadopted. We examine the potential effects of parental care,sibling competition, relative size within a brood, and adultexperience of agonistic interactions and mating on tactic adoption.Absolute size was consistently the best predictor of callingrate, with smaller males calling more often than larger males.We suggest that the lack of a response to adult cues may reflectunpredictability in the occurrence of social interactions orstable size distributions in this population.     

Alternative reproductive tactics and status-dependent selection

The status-dependent selection model on alternative reproductivetactics predicts a single switch-point in status: usually allplayers above some status (e.g., competitive ability) shouldpractice the tactic with the higher average payoff, while thosebelow that point should make the "best of a bad job" by practicingthe alternative, lower payoff tactic. Many empirical studiesindeed show a relationship between status and tactic choice,but they do not conform to this single switch-point prediction.I modify the status-dependent selection model by consideringstatus-dependent fitness that is mediated, at least in part,by resource acquisition (e.g., status-based competition forterritories or nuptial gifts). With variation in resource quality,predicted tactic-choice distributions change: a high-statusmale may be territorial on a high-quality territory, a lowerstatus male may practice an alternative tactic, and an evenlower status male may be territorial on a low-quality territory.Tactic choice thus alternates as in many empirical studies andcan appear to be but is not actually stochastic. As the numberof theoretically predicted switch-points increases, however,mixed or mixed-conditional strategies should become more prevalent.While alternative tactics will likely usually differ in meanpayoff, viewing alternative reproductive tactics as inherently"better" or "worse" (e.g., viewing cuckoldry as "worse"—thebest of a bad job) is misleading if not tempered with awarenessthat payoff can vary greatly within tactics and overlap betweentactics.     

Increasing vulnerability to predation increases preference for the scrounger foraging tactic

When animals forage in groups, individuals can search for foodthemselves (producer tactic) or they can search for and joinother individuals that have located food (scrounger tactic).The scrounger tactic may provide greater antipredator benefitsthan the producer tactic because "scroungers" hop with theirheads up and tend to occupy central positions in a group, whereas"producers" hop with their heads down and tend to occupy edgepositions. We tested whether increasing an individual's vulnerabilityto predation (using wing-loading manipulations) causes an increasedpreference for the scrounger tactic in zebra finches (Taeniopygiaguttata). Wing-loading manipulations were effective at increasingfocal individuals' perception of vulnerability to predation;treatment individuals increased their total time allocated tovigilance, whereas control individuals did not. Treatment individualsalso increased their use of the scrounger tactic (proportionof hops with head up) and scrounged a greater proportion ofpatches, whereas control individuals exhibited no changes. Ourresults are consistent with the hypothesis that the scroungertactic confers greater antipredator benefits than the producertactic, although whether antipredator benefits are achievedthrough differences in head orientation, spatial position, orboth, remains unclear. Our finding that individuals adjust theiruse of the scrounger tactic according to changes in their phenotypeprovides evidence for phenotype-limited allocation strategiesin producer–scrounger games.     

Urea is a dynamic pool of bioavailable nitrogen in coral reefs

Urea may be an important source of nitrogen in low nutrient coral reef environments because corals and other organisms can assimilate it easily and it is found throughout ocean waters. We measured the distribution and concentrations of urea in seagrass beds, areas of schooling fish, coral formations and bottom sediments in the Upper Florida Keys Reef Tract. The flux of urea from bottom sediments was also measured. Ambient concentrations of urea in the offshore reefs were similar to the concentrations of nitrate and ammonium. Seagrass beds, areas of schooling fish and coral formations had elevated concentrations of urea that were up to eight times higher than nitrate in the system. Numerous ephemeral hotspots of urea that were 8–20 times the ambient urea concentration existed in seagrass beds, areas of schooling fish, and above sediments. Coastal areas and inland canals had high urea concentrations where urban runoff and septic effluents were prevalent, but there was no anthropogenic influence in the offshore habitats. Urea concentrations above bottom sediments were not different from ambient concentrations and benthic flux chamber incubations showed biological activity in carbonaceous sediments but no net urea production. The decrease in urea concentrations from coasts and inland waterways to a consistent ambient concentration in the offshore reef system and ephemeral hotspots of high urea concentration suggest that urea is a dynamic pool of bioavailable nitrogen in the reefs of the Upper Florida Keys.     

Phytoplankton competition in deep biomass maximum

Resource competition in heterogeneous environments is still an unresolved problem of theoretical ecology. In this article, I analyze competition between two phytoplankton species in a deep water column, where the distributions of main resources (light and a limiting nutrient) have opposing gradients and co-limitation by both resources causes a deep biomass maximum. Assuming that the species have a trade-off in resource requirements and the water column is weakly mixed, I apply the invasion threshold analysis (Ryabov and Blasius, Ecol Lett 14:220–228, 2011) to determine relations between environmental conditions and phytoplankton composition. Although species deplete resources in the interior of the water column, the resource levels at the bottom and surface remain high. As a result, the slope of resources gradients becomes a new crucial factor which, rather than the local resource values, determines the outcome of competition. The value of resource gradients nonlinearly depend on the density of consumers. This leads to complex relationships between environmental parameters and species composition. In particular, it is shown that an increase of both the incident light intensity or bottom nutrient concentrations favors the best light competitors, while an increase of the turbulent mixing or background turbidity favors the best nutrient competitors. These results might be important for prediction of species composition in deep ocean.     

Paternal inheritance of growth in fish pursuing alternative reproductive tactics

In species with indeterminate growth, age‐related size variation of reproductive competitors within each sex is often high. This selects for divergence in reproductive tactics of same‐sex competitors, particularly in males. Where alternative tactics are fixed for life, the causality of tactic choice is often unclear. In the African cichlid Lamprologus callipterus, large nest males collect and present empty snail shells to females that use these shells for egg deposition and brood care. Small dwarf males attempt to fertilize eggs by entering shells in which females are spawning. The bourgeois nest males exceed parasitic dwarf males in size by nearly two orders of magnitude, which is likely to result from greatly diverging growth patterns. Here, we ask whether growth patterns are heritable in this species, or whether and to which extent they are determined by environmental factors. Standardized breeding experiments using unrelated offspring and maternal half‐sibs revealed highly divergent growth patterns of male young sired by nest or dwarf males, whereas the growth of female offspring of both male types did not differ. As expected, food had a significant modifying effect on growth, but neither the quantity of breeding substrate in the environment nor ambient temperature affected growth. None of the environmental factors tested influenced the choice of male life histories. We conclude that in L. callipterus growth rates of bourgeois and parasitic males are paternally inherited, and that male and female growth is phenotypically plastic to only a small degree.     

Laboratory experiments on the schooling and feeding behaviour of Trichogaster fasciatus (Pisces: Perciformes)

Laboratory experiments were performed on the schooling and feeding behaviour of Trichogaster fasciatus . There was a tendency on the part of individuals of similar size to school together; the instinct being stronger in younger specimens. Unavailability of similar-sized individuals led to schooling by members of different size-groups. Column feeding was exhibited by the fish. While feeding at the bottom, the fishes formed angles varying from 30°90°. Larger individuals formed a right angle more often than younger members of the species. Optimum degree of crowding seemed necessary for active feeding as confinement of single individual, or overcrowded condition, both resulted in a decline of feeding activity.     

Complex behaviour by coral-reef fish larvae in open-water and near-reef pelagic environments

We present the first in situ observations of the pelagic larvae of coral-reef fishes feeding, schooling and being preyed upon. In addition, we report on their behavioural interactions with adult and juvenile fishes. Observations on over 500 larvae of over 50 species (mostly from four families) near the end of their pelagic interval were made in both open water (> 1 km offshore) and near-reef environments. Nearly 10% of larvae were seen to feed in open water, but < 1% fed near the reef. Presettlement schooling was observed in five species of four families. We observed no predation upon larvae in open water except near the bottom. Near the reef, 8.5% of larvae were eaten. The main predators near and on the reef were a species of wrasse and lizardfishes. Rates of predation seem to differ among genera of pomacentrids, perhaps related to differences in behaviour when settling. When confronted with adult fishes, which happened largely near the reef, larvae reacted with a limited range of behaviours, including sheltering near the observer, swimming to the surface, slowing or stopping, or swimming offshore. The frequency of these behaviours differed among larvae of three pomacentrid genera. Interactions with reef residents, particularly pomacentrids, were common, and usually involved aggression by the resident toward settling larvae. This may act to discourage settlement during the day when such residents are active. These data show that behaviour of late larvae of coral-reef fishes is complex and can greatly influence survival and recruitment. Further, behaviour differs among taxa, showing that not only are larvae not passive, but also that a generalised behaviour of larvae does not exist.     

Social network predicts loss of fertilizations in nesting males of a fish with alternative reproductive tactics

Alternative reproductive tactics (ARTs) evolve when there is strong intra-sexual competition between conspecifics for access to mates. Typically, larger “bourgeois” males reproduce by securing the access to reproductive resources while smaller “parasitic” males reproduce by stealing fertilizations from larger males. A number of factors can influence the reproductive success of each tactic, including intrinsic (e.g. size) and extrinsic (e.g. tactic relative frequency) variables. An example where plastic ARTs occur is the peacock blenny Salaria pavo, with large males reproducing by defending nests and attracting females (bourgeois tactic) and small males reproducing by achieving sneaked fertilizations (parasitic tactic). In this study, we conducted field observations on individually tagged animals to determine their social network and collected eggs from 11 nests to determine the fertilization success of each male tactic. Paternity estimates for 550 offspring indicated an average fertilization success for nest-holder males of 95%. Nest-holder male morphological traits and social network parameters were tested as predictors of fertilization success, but only the number of sneakers present in the nest-holder’s social networks was found to be a predictor of paternity loss. Although male morphological traits had been previously found to be strongly correlated with reproductive success of nest-holder males, as measured by the number of eggs collected in the male’s nest, no correlation was found between any of the measured morphological traits and fertilization success for these males. The results suggest a stronger influence of the social environment than of morphological variables in the proportion of lost fertilizations by nest-holder males of this species.     

Schooling as a strategy for taxis in a noisy environment

Many aquatic animals face a fundamental problem during foraging and migratory movements: while their resources commonly vary at large spatial scales, they can only sample and assess their environment at relatively small, local spatial scales. Thus, they are unable to choose movement directions by directly sampling distant parts of their environment. A common strategy to overcome this problem is taxis, a behaviour in which an animal performs a biased random walk by changing direction more rapidly when local conditions are getting worse. Such an animal spends more time moving in right directions than wrong ones, and eventually gets to a favourable area. Taxis is inefficient, however, when environmental gradients are weak or overlain by noisy small-scale fluctuations. In this paper, I show that schooling behaviour can improve the ability of animals performing taxis to climb gradients, even under conditions when asocial taxis would be ineffective. Schooling is a social behaviour incorporating tendencies to remain close to and align with fellow members of a group. It enhances taxis because the alignment tendency produces tight angular distributions within groups, and dampens the stochastic effects of individual sampling errors. As a result, more school members orient up-gradient than in the comparable asocial case. However, overly strong schooling behaviour makes the school slow in responding to changing gradient directions. This trade-off suggests an optimal level of schooling behaviour for given spatio-temporal scales of environmental variations. Social taxis may enhance the selective value of schooling in pelagic grazers such as herrings, anchovies and Antarctic krill. Furthermore, the degree of aggregation in a population of schooling animals may affect directly the rate and direction of migration and foraging movements.     

Effects of Social Structure on the Behaviour and Performance of Alternative Reproductive Phenotypes in Male Rock Shrimp, Rhynchocinetes typus

Males that adopt alternative mating tactics within a conditional strategy often undergo costly morphological changes when switching to the next phenotype during ontogeny. Whether costs of changing to a subsequent reproductive phenotype are outweighed by a higher mating probability may depend on the frequencies of different phenotypes in a group of competitors. Benefits and costs associated with different phenotype frequencies depend on interactions within and between alternative phenotypes, but the underlying behavioural mechanisms have rarely been studied. Herein, we used the rock shrimp Rhynchocinetes typus as a model: ontogenetic male stages of this species differ in morphological and behavioural traits that indicate alternative reproductive phenotypes. The small, subordinate, male stage (typus) develops via several intermediate stages (intermedius) to the dominant male stage (robustus): in competitive interactions the typus males usually employ the sneaking tactic, while the robustus males invariably employ the monopolizing fighter tactic. In laboratory experiments, we manipulated phenotype frequencies to examine whether there are frequency‐dependent effects on searching behaviour, aggressiveness and mating probability. With increasing frequency of robustus males, the rate of aggressive interactions among them increased. Furthermore, robustus males increased walking velocity when more than one robustus male was present. In contrast, typus males did not adjust their searching or aggressive behaviour. The increase of aggressive interactions among robustus males provided more opportunities for typus males to seize a temporarily unguarded female. While typus males exploit fights among robustus males that produce mating opportunities for them, robustus males benefit from typus males, which reveal the presence of receptive females. We suggest that each phenotype benefits from the presence of the other phenotype and suffers costly interference among individuals of the same phenotype. Whether frequency‐dependent effects on the mating probability of subordinates also affect their ontogenetic switchpoint should be examined in future studies.     

Universal reserve constants and selection criteria in the periodically changing environment

The authors analyse different approaches (geometrical and analytical) to investigate the dynamic of competing populations in changing environment. For the broad range of models a sufficient character of selection is determined: in changing environment one population excludes the others if its productivity is higher with some reserve. In competition models "everybody against everybody" there is a universal reserve constant that is not depended on the number of populations in community. In the models "everybody against one" reserve constant can increase without limits in increasing the number of competitors.     

Functions of Scales and Photophores in Mesopelagic Luminescent Sharks

In sharks bioluminescence is only known from the family Squalidae. It evolved independently in two out of six squalid subfamilies, Dalatiinae and Etmopterinae. The distribution of photophores was mapped in several species. It is suggested that in the Dalatiinae, which do not school, but migrate vertically, luminescence serves as ventral countershading. The Etmopterinae school and feed close to the bottom. Their luminescence is an aid in schooling. Four different placoid scale patterns are found in luminescent sharks and they allow to accommodation the photophores in the skin.     

Using behavior and ecology to exploit schooling fishes

By the late 1980's, humans were removing 76 million metric tons (MMT) of marine fishes annually. The potential sustainable catch is somewhere between 69 and 96 MMT. As a result, major fisheries have collapsed or are in danger of collapsing. Many of these species school. Schooling is effective against gape-limited predators because of dilution and confusion. However, larger predators may exploit schooling behavior to sequester and consume a non-trivial fraction of the group. This is the strategy of fishers. Both gear and fisher behavior have evolved to take advantage of the seemingly canalized response of schooling species. This paper examines the ways artisanal and western fishers have exploited knowledge of the behavior and ecology of schooling species to aid in fish capture. Topics include object association; use of light, sound, and chemicals; perceived barriers; predator-prey and other trophic interactions; inherent cyclical rhythms such as diel migration, lunar spawning, and seasonality; and correlations with the physical environment. Exploiting schooling allows fishers to increase efficiency through knowledge of when and where fish aggregate, or by extending the conditions under which aggregation occurs. However, knowledge of behavioral ecology can also be used to conserve schooling stocks. Gear selectivity, group size and population dynamics, and fisher efficiency are all potential areas of integration between behavioral ecology and fishery management. However, no amount of integration of behavioral ecology into fishery management will have the intended conservation effects if fishing effort is not limited to at least numerical if not behaviorally-sustainable levels.