Nestmate recognition in the unicolonial ant Formica paralugubris

In unicolonial populations of ants, individuals can mix freelywithin large networks of nests that contain many queens. Ithas been proposed that the absence of aggression in unicolonialpopulations stems from a loss of nest mate recognition, butfew studies have tested this hypothesis. We investigated patternsof aggression and nest mate recognition in the unicolonial woodant, Formica paralugubris. Little aggression occurred, evenbetween workers from nests separated by up to 5 km. However,when aggression took place, it was directed toward non–nestmates rather than nest mates. Trophallaxis (exchange of liquidfood) occurred very frequently, and surprisingly, workers performedsignificantly more trophallaxis with non–nest mates thanwith nest mates (bias 2.4:1). Hence, workers are able to discriminatenest mates from non–nest mates. Higher rates of trophallaxisbetween non–nest mates may serve to homogenize the colonyodor or may be an appeasement mechanism. Trophallaxis rate andaggression level were not correlated with geographical distanceand did not differ within and between two populations separatedby several kilometers. Hence, these populations do not representdifferentiated supercolonies with clear-cut behavioral boundaries.Overall, the data demonstrate that unicoloniality can evolvedespite well-developed nest mate recognition. Reduced levelsof aggression might have been favored by the low rate of interactionswith foreign workers, high cost of erroneously rejecting nestmates, and low cost of accepting foreign workers.     

Genetic monogamy in blue-headed vireos and a comparison with a sympatric vireo with extrapair paternity

Based on the breeding synchrony hypothesis, we predicted, intwo congeners that nest in simiilar habitat but differ in nestingsynchrony, that blue-headed vireos (Vireo solitarius) wouldhave fewer extrapair fertilizations (EPFs) thaii red-eyed vireos(V. olivaceus EPFs were rare in blue-headed vireos (1/37 nestlings),but common in red-eyed vireos (11/19 nestlings). We studiedthe behavior of blue-headed vireos to determine what factorscould promote genetic monogamy. We found no evidence that malesmate guarded to prevent extrapair copulations from occurring.Males did not follow fertile mates closely when mates left thenest (14–25% of female departures) and, during the egg-layingperiod, males were often alone on the nest (22.3 mm/h). Femaleblue-headed vireos, but not red-eyed vireos, obtain direct benefitsfrom social mates such as nest building and incubation (49.1%of the total), and they assess male quality long before becomingfertile. Female blue-headed vireos spent more time incubatingwhen their mates had low incubation effort. Furthermore, maleincubation effort was positively correlated with nest survivalduring incubation. We discuss the evolution of genetic monogamyand sex role convergence in blue-headed vireos in relation toasynchronous breeding.     

Conditional strategies in territorial defense: do Carolina wrens play tit-for-tat?

Neighboring territorial animals are viewed primarily as intenserivals, but there are also opportunities for cooperation amongcompetitors. Many animals respond less aggressively towardneighbors than to strangers. This phenomenon, termed the "dearenemy" effect, should be stable only when the reduced aggressionis reciprocal. Territory owners should use conditional strategiesin territorial defense, showing reduced aggression toward neighbors who cooperate by respecting territorial boundaries but increasingaggression toward invading neighbors. In this study I examinedthe response of territory owners to playbacks of neighborsat shared boundaries before and after intrusions by that neighboror by strangers. Results showed that territory owners did notincrease their aggression toward defecting neighbors but did increase their aggression toward neighbors after a simulatedintrusion by a stranger. This surprising result might reflectthe long-term relationship between neighboring Carolina wrensand the threat posed by rare intruding strangers.     

Quorum sensing by encounter rates in the ant Temnothorax albipennis

Emigrating colonies of the ant Temnothorax (formerly Leptothorax)albipennis can choose the best of several nest sites, even whenthe active ants organizing the move do not compare sites. Thiscollective ability depends on a quorum rule used by ants assessinga candidate site. Only when the site's population has surpasseda threshold do they switch from slow recruitment of fellow activeants by tandem runs to rapid transport of the majority of thecolony. Here, I show that ants perceive the achievement of aquorum through their rate of direct encounters with nest matesat the site. When ants in a crowded site were prevented fromtactile contact with nest mates, they recruited by tandem runs,as though to an empty nest. Furthermore, when the encounterrate was raised independent of population, by reducing the sizeof the candidate nest, ants started to transport at a significantlylower population. The switch occurred at the same encounterrate regardless of nest size, whether the rate was measuredas the mean over the entire visit or as the inverse of the latencyuntil the first encounter. Because encounter rate reflects thedensity of nest mates and thus varies with nest size as wellas population, the ants' collective decision-making algorithmmay be robust to the exact population at which the switch totransport occurs. Ants cease monitoring quorum presence afterswitching to transport, coincident with an abrupt shorteningof visit duration by approximately 2 min, which may be interpretedas the time required for quorum detection.     

Regulation of worker activity in a primitively eusocial wasp, Ropalidia marginata

Ropalidia marginala, a tropical, primitively eusocial, polistinewasp, is unusual in that the queen (the sole egg-layer) is neitherthe most behaviorally dominant nor the most active individualin the colony. The queen by herself rarely ever initiates interactionstoward her nest mates or unloads returning foragers. There arealways a few workers in the colony who are more dominant andactive than the queen. Absence of the queen from her colonydoes not affect colony maintenance activities such as foragingor brood care, but it always results in one individual becomingvery aggressive and dominant. The dominant worker becomes thenext queen if the original queen does not return. The queendoes not appear to play any significant role in colony activityregulation. Instead, colony activities appear to be regulatedby several mechanisms including dominance behavior toward foragers,feeding of larvae, and the unloading of returning foragers,all mediated by workers themselves. Regulation of colony maintenanceappears to be based on direct evaluation of the needs of thecolony by the workers themselves. The queen however has perfectreproductive control over all workers; workers never lay eggsin the presence of the queen. It appears therefore that themechanisms involved in regulation of worker activity and workerreproduction are separate in R marginata. These findings contrastwith other primitively eusocial species where the queen actsas a "central pacemaker" and controls both worker activity andworker reproduction.     

Worker biting interactions and task performance in a swarm-founding eusocial wasp (Polybia occidentalis, Hymenoptera: Vespidae)

Workers in many insect societies interact via body contact withtheir nest mates, and social biting and other forms of contactmay play a general role in regulating task performance. HereI present evidence that social biting affects task performancewithout direct reproductive conflict in Polybia occidentalis,a swarm-founding eusocial wasp. Polybia occidentalis workersengaged in social biting with nest mates. Most workers thatwere active on the nest surface participated in biting interactions,but individuals differed significantly in their rates of biting and of being bitten. Rates of being bitten corresponded withnonreproductive task performance: more biting was directedat foragers than nonforagers, and foraging rates were correlatedwith rates of being bitten. Furthermore, some on-nest workersinitiated foraging activity immediately after they were bitten.Together these patterns suggest that social biting influencesforaging rates by increasing workers' probabilities of leavingthe nest. Variation in biting rates did not correspond withdifferences in reproductive physiology: highly active bitersand recipients did not differ in body size or in ovary development.In P. occidentalis and in other eusocial insects with largeworker forces, biting and other types of social contact amongworkers may regulate task performance independently of directreproductive competition.     

Spatial relatedness and brood parasitism in a female-philopatric bird population

The spatial structure of relatedness between individuals ina population can be crucial for social selection and evolution.Here we analyze a female alternative reproductive tactic, conspecificbrood parasitism, in relation to spatial relatedness among femalesin a Baltic Sea population of the common eider Somateria mollissima.The role of relatedness in brood parasitism is debated: somemodels predict parasite avoidance of related hosts, others predicthost–parasite relatedness. We estimate pairwise relatednessfrom protein fingerprinting of egg albumen in 156 nests, withpairwise nest distances ranging from 1 to 6 km. Relatednessincreases significantly from the longest distances to an averageof r 0.09 below 20 m. Brood parasitism is common, and averagepairwise relatedness between host and parasite is estimatedat 0.18–0.21. Parasites thus do not avoid relatives, andcombined with the findings of a similar study in another eiderpopulation, the results show that mean host–parasite relatednessis higher than that among close neighbors. High host–parasiterelatedness is therefore not an effect of natal philopatry alone;some other form of kin bias is also involved. Recognition andassociation between birth nest mates is a candidate mechanismfor further study.     

Seasonal Variation in Response to Neighbors and Strangers by a Territorial Songbird

Reduced aggression toward territorial neighbors, termed the ‘dear‐enemy’ effect, is thought to arise because territorial animals benefit by avoiding contests with neighbors with whom they have already established relationships. The dear‐enemy effect has been described in many taxa, but few studies have considered whether or not neighbors’ relationships are affected by changes in the social environment. In this study, I tested whether Carolina wrens, Thryothorus ludovicianus, behaviorally discriminate between neighbors and strangers in two different social environments: in spring when territories have been established for several months, and in fall when an influx of new birds claiming territories might de‐stabilize wren neighborhoods. Comparisons of responses of territorial males to playbacks of songs from neighbors and strangers showed that Carolina wrens show the dear‐enemy effect in spring, but not in fall in this design. The apparent lack of a differential response to neighbors and strangers in fall might be due to a reduction in aggression toward strangers. This study provides evidence that seasonal changes in the dear‐enemy effect coincide with seasonal changes in the social environment.     

Temporal Patterns of Territorial Behavior and Circulating Testosterone in the Lapland Longspur and Other Arctic Passerines

We studied territorial aggression in relation to circulatingtestosterone levels in free-living birds of four species innorthern Alaska. The Lapland longspur, Calcarius lapponicus,is an abundant breeding passerine on the arctic tundra. Unlikemany passerines at lower latitudes, male Lapland longspurs donot defend a "multiple-purpose territory" that serves to providenest sites, food and shelter. Rather, after arrival on the breedinggrounds, they perform aerial display flights over a looselydenned "nest area" for a very brief period of two days or so,showing tolerance of other males. This song display may be involvedin courtship. During this phase, male longspurs show a briefand pronounced peak in circulating testosterone levels, andare not aggressive toward simulated territorial intrusions (STIs).Males then "guard" their sexually receptive mates for aboutten days, during which they are highly aggressive toward STIs,but do not sing as much. During the next phase, incubation,the males become very tolerant of conspecific males. Their circulatingtestosterone levels decline to baseline levels, and they generallydo not sing or display aggression in response to STIs. Threeother passerines, the white-crowned sparrow, Zonotrichia leucophrysgambelii, American tree sparrow, Spizella arborea, and savannahsparrow, Passerculus sandwichensis, show patterns of territorialaggression typical of species studied at lower latitudes. Welldefinedterritories are defended for several weeks, during which thereis a prolonged peak in plasma concentrations of testosterone.These three species continue to sing and display aggressioneven late in the season, unlike the longspurs. The peak of testosteronein the longspurs occurs simultaneously with the peak in songdisplay, while in mid-latitude species it occurs with the peakin reproductive aggression. These data suggest that the interrelationshipof testosterone and aggression in Lapland longspurs may be differentfrom that of passerines with multiple-purpose territories, andmay be related to the constraints of breeding in the open arctictundra.     

Deceptive use of alarm calls by male swallows, Hirundo rustica: a new paternity guard

Male swallows (Hirundo rustica) frequently gave alarm callswhen their fertile mates left the nest vicinity without themale during egg laying. Females engaged in extrapair copulationsduring such absences when they were not guarded intensely bytheir mates. However, males were sometimes able to disrupt extrapaircopulation attempts toward their mates by giving alarm calls,causing all nearby swallows to take flight. I experimentallytested the idea that male swallows gave alarm calls when theircertainty of paternity was at stake. When colonially breedingfemale swallows were temporarily chased away from their nestduring the egglaying period in the absence of their mates, malesnearly always gave alarm calls when they returned to the emptynest. In contrast, colonially breeding males rarely gave alarmcalls when their mates were absent during the prelaying or theincubation periods. Solitarily breeding males rarely gave alarmcalls in the absence of their mates, irrespective of the timein their nesting cycle. However, solitarily breeding males didgive alarm calls when their mates were absent from the nestif a male mount previously had been presented near their nestduring the egg-laying and the prelaying periods, but not duringthe incubation period. Deceptive use of alarm calls (in theabsence of any predators) by male swallows apparently was aneffective paternity guard.     

Neighbor-stranger discrimination by song in a suboscine bird, the alder flycatcher, Empidonax alnorum

Bird song and its functions have been studied extensively formore than 50 years, but almost entirely in oscine passerines.Few studies have investigated any aspect of song in suboscinepasserines. This is significant because song development andthe extent of individual variation in song differs greatly betweenthese groups. Learning and auditory feedback play major rolesin song development in all oscines studied, but apparently nopart in song ontogeny in suboscines. The ability of territorialoscine males to discriminate between songs of neighbors andstrangers has received considerable attention, but this phenomenonis virtually unstudied in suboscines. We tested whether a suboscinebird, the alder flycatcher (Empidonax alnorum), was able todiscriminate between songs of neighbors and strangers despitelimited individual variation in song. We performed playbackexperiments to measure responses of males to songs of neighborsand strangers broadcast from the territory boundary shared bythe subject and the neighbor. Subjects responded more aggressivelyto songs of strangers than to songs of neighbors. These resultsfurther our understanding of the evolution of song and its functionsin suboscines by demonstrating that, similar to their oscinerelatives, they can discriminate between the songs of neighborsand strangers.     

Nocturnal Mate Recognition and Nest guarding by Female Convict Cichlids (Pisces,Cichlidae: Cichlasoma mgrofasciatum)

This study investigated the ability of nest-guarding convict cichlid (Cichlasoma mgrofasciatum) females to recognize their own mate and to defend their brood effectively against strangers at night. Cichlids in the laboratory bred in nesting boxes made of plexiglas. At the fry stage, various conspecific males were introduced, at night, into the nest box of brood-guarding females. Female reaction was observed with infrared equipment. If the introduced male was their own mate, females showed little reaction. If the introduced male was a stranger, females reacted by directing head shakes, tail beats, and pushes or bites at the intruder. The frequency of these acts was significantly higher in the presence of strangers than in the presence of mates; the frequency also rose significantly as the size of strangers increased. The two largest classes of strangers fought with the females. These fights included circling behaviour, but not mouth-locking. Some aspects of female behaviour suggested that mate recognition is based on short-range chemical cues and not on sound, size, or rapidly diffusing odours. Mate recognition in the nest, at night, is an example of non-visual communication in fish, and active nest defense at night shows that the convict cichlid, normally considered to be diurnal, can engage in coordinated and effective behaviour in the absence of light.     

Honeybee guards do not use food-derived odors to recognize non-nest mates: a test of the Odor Convergence hypothesis

Honeybee (Apis mellifera) colonies rob honey from each other during periods of nectar shortage. Persistent robbing can killweak colonies. Primarily responsible for preventing robbingare guard bees. Previous research has shown that the probabilityof both nest mate and non-nest mate workers being acceptedby guards at the nest entrance increases as nectar availability increases. The mechanism responsible for this change in guardacceptance can be explained by two competing hypotheses: OdorConvergence and Adaptive Threshold Shift. In this study wetested the Odor Convergence hypothesis. The acceptance by guardsat the nest entrance of workers transferred between four coloniesthat had been fed either odorless sucrose syrup (two colonies)or diluted heather honey (Calluna vulgaris) (two colonies)was measured for 3 days before feeding and during 2 weeks offeeding. Despite the large sample sizes, the probability ofguards accepting non-nest mates was not affected by the similaritiesor dissimilarities in food odor between guards' and non-nestmates' colonies. This finding contrasts with the accepted wisdom that food odors are important in nest mate recognition in honeybeesand the data, therefore, strongly reject the Odor Convergencehypothesis.     

Sexual conflict among polygynous pied flycatchers feeding young

In the polygynous pied flycatcher, Ficedula hypoleuca, reproductivesuccess of females is constrained by male food provisioningduring the nestling period. Hence, there will be conflictinginterests among the male and each of his mates as to how malefeeding effort should be shared among broods. This paper describesthree experiments designed to examine the parental behaviorof the members of a bigynous trio, i.e., the male and his twomates, in light of these conflicts. In all experiments, primaryand secondary broods were manipulated to hatch on the same dayto reduce the difference in brood-reproductive value due toage. Males divided their effort equally when the two broodswere the same size. However, males did not allocate their investmentin proportion to brood size when brood sizes differed, but investedmore heavily per young in the larger broods. This finding suggeststhat males tried to optimize the joint effort of their two mates.Males and females showed similar responses to experimental reductionin brood demands, which indicates no difference in their willingnessto invest in offspring. When one of the male’s mates wasremoved temporarily, the male increased his total feeding rateand provided proportionately more food to the "motherless" brood.Through flexible allocation of parental investment, males seemable to optimize their reproductive interests in the two broods.The only way a polygynously mated female might successfullyincrease the amount of male assistance at her nest is to makeher own brood more valuable for the male, relative to the otherbroods he might have. We discuss some ways this might be achieved.[Behav Ecol 1991;2:106–115]     

Nests as ornaments: revealing construction by male sticklebacks

Nests are built by animals from a variety of taxa, and serveas receptacles for eggs and developing offspring. Where nestsare built solely or mainly by one sex, they also have the potentialto serve as extended ornaments, because aspects of constructionpotentially reveal or amplify characteristics of the builderto prospective mates. Here, we develop novel indices to quantifynest structure and examine variation in temporal and structuralaspects of nest construction in relation to morphological,immunological, and physiological traits in male three-spinedsticklebacks, Gasterosteus aculeatus. Wild-caught male sticklebacksthat began construction within 3 days of being transferredto the laboratory built "neater" nests than fish that tooklonger to start, and we present alternative testable hypothesesthat could explain this pattern. Various characteristics ofnest-building males correlated with nest structure. The relativeweight of the building male's kidney—which secretes aglue-like protein used in nest building and whose developmentis androgen-dependent—correlated positively with nest "neatness." We also found males with enlarged spleens (an indicatorof immune stress) to construct less "compact" nests. The structureof a nest may therefore be important not only in determiningits functional capacity, but may also act as a quality-revealingornament. We suggest that females may gain valuable informationregarding male health status and androgen levels from nestinspection.     

Immuno-Gold Localization of Nitrogenase in Root Nodules of Elaeagnus pungens Thunb

The molybdenum-iron component of nitrogenase (Mo-Fe component)was purified from soybean nodule bacteroids and antibody wasraised against it in rabbits. Antibody raised against the 53kDa polypeptide which was the major protein in the Mo-Fe componentfraction of soybean nitrogenase was confirmed to be specificto the nitrogenase by immunodiffusion and immunotitration. Thenitrogenase from root nodules of Elaeagnus pungens cross-reactedwith the antibody and appeared from the results of the immunodiffusionto be partially identical to soybean nitrogenase. Using the antibody, we examined intracellular localization ofnitrogenase in root nodules of Elaeagnus pungens, in which Frankiais present as a symbiont, by immuno-gold labelling. Thin sectionsof nodules of Elaeagnus pungens were first treated with anti-nitrogenasespecific antibody and then with colloidal gold-protein A asa marker. The gold particles were observed to be concentratedin the vesicles of the endophyte Frankia. This provides strongsupport for the existence E of nitrogenase in the vesicles.Furthermore, our results suggested that nitrogenase localizesin the hyphae of the endophyte Frankia in Elaeagnus pungensnodules. ¹Present address: Iwata Experiment Station, Japan Tobacco Inc.,Iwata-gun, Shizuoka 438, Japan. (Received March 9, 1988; Accepted July 28, 1988)     

Behavioral mechanisms preventing filial ovicide in house wrens

Because unmated house wrens (Troglodytes aedon) regularly destroyclutches of conspecifics, we examined mechanisms that preventbreeding individuals from destroying eggs in their own nests.Results of our experimental field study indicate that the egg-destroyingbehavior of males is partially suppressed after mating and thatthe suppression may prevent filial ovicide in some individuals.At least 35% of males continue to destroy conspecific eggs duringthe incubation period of their mates. Because filial ovicideby males is prevented independently of female presence at thenest, we conclude, after eliminating alternative hypotheses,that filial ovicide by males is prevented through their recognitionof nest locations. Egg-destroying behavior of females is alsosuppressed during the incubation. The suppression is strongerthan in males because fewer females than males destroy foreigneggs over a comparable time period. Females respond to experimentallychanged nest location and nest structure, but not egg coloration.We conclude that filial ovicide by females is prevented throughthe suppression of their eggdestroying behavior and throughtheir recognition of nest location and structure.     

The Costs of Neighbors for a Territorial Dragonfly, Perithemis tenera

Past researchers have often considered neighbors to be beneficial to territorial residents, particularly compared with non‐neighbor conspecific competitors. However, neighbors have the potential to be costly to residents in terms of both defensive costs and lost resources. In this study, we assessed the relative costs of defending a mating territory against neighbors and non‐neighbors for the dragonfly Perithemis tenera, comparing across males with different numbers of contiguous neighbors; we also examined the possibility that the presence of contiguous neighbors might reduce the detection of potential mates. When neighbors were present, residents experienced a greater total number of intrusions by males; this increase in intrusions was due to higher numbers of intrusions by neighbors, as the number of intrusions by non‐neighbor males did not differ. Residents with immediately adjacent neighbors also made more sorties toward neighbors than did residents whose nearest neighbors’ territories were not immediately adjacent. Interestingly, although the number of visits by females did not vary with the presence of neighbors, residents with neighbors made fewer sorties toward females than did residents without neighbors. Our results suggest that defensive costs increased when neighbors were present, that residents with neighbors may have missed opportunities to acquire mates, and thus that living with neighbors can be costly in this species.     

Chick begging as a signal: are nestlings honest?

Begging by dependent avian offspring is known to correlate withhunger level, and parents use this as a signal of brood demandto adjust their chick feeding behavior. While there is informationon how each chick adjusts its begging to its own condition,little is known of how chicks adjust to the state of their nestmates. In two experiments we manipulated the competitive environmentof individual European starling (Sturnus vulgaris) chicks byaltering the state of nest mates while holding the state oftarget chicks constant In the first experiment we placed thetarget chick's nest mates in neighboring nests with brood sizesof two, five, or eight chicks. Following the manipulation wereturned them to their own nests and recorded begging behavioron videotape. In the second experiment we separated a targetchick from its siblings and manipulated feeding level in thelaboratory. The siblings were fed at one of three levels; meanwhile,all the target chicks were fed at the intermediate level. Afterthe manipulation we placed the target chicks with their siblingsand recorded their begging in response to an artificial stimulus.In neither experiment was the begging effort of the unmanipulatedtarget chicks affected by the changes in begging behavior oftheir siblings. This result supports the view that begging isa reliable signal of individual chick state and does not involveresponses to the effort of nest mates.     

Bumble bees (Bombus terrestris) store both food and information in honeypots

Social insect foragers often transmit information about foodsources to nest mates. In bumble bees (Bombus terrestris), forexample, successful foragers use excited motor displays anda pheromone as communication signals. In addition, bees couldmake use of an indirect pathway of information flow, via thehoney stores. We show here that, indeed, bees in the nest continuouslymonitor honeypots and sample their contents, thus obtaininginformation on supply and demand of nectar. When there is aninflux of nectar into the nest, the colony deploys more workersfor foraging. The number of new foragers depends on sugar concentration.Foragers returning with high-quality sugar solution displaymore "excited runs" on the nest structure. The recruits' response,however, does not depend on modulated behavior by foragers:more workers start to forage with high quality of incoming nectar,even when this nectar is brought by a pipette. Moreover, weshow that the readiness of bees to respond to recruitment signalsor incoming nectar also depends on colony demand. When colonynectar stores are full, the response of bees to equal amountsof nectar influx is smaller than when stores are empty. Whencolony nectar stores are depleted, foragers spend more timerunning excitedly and less time probing pots in the nest andrun with higher average speed, possibly to disperse the alertingpheromone more efficiently. However, more bees respond to nectarinflux to empty stores, whether or not this is accompanied byforager signals. Thus, honeypots serve to store informationas well as food.