Colony size selection determines adult survival and dispersal preferences: allee effects in a colonial bird

Avian coloniality traditionally has been investigated by examining how breeding success varies with colony size, but other crucial fitness components rarely have been examined. This may lead to wrong conclusions because unmeasured parameters may change the final fitness balance. We used multistate capture-recapture models to investigate adult survival and dispersal in relation to colony size within a long-term monitored population of lesser kestrels (Falco naumanni). Nest predation probability decreases with colony size, and adult survival is predicted to show the same trend because adults are exposed to the same suite of predators. As expected, survival probability was higher in large colonies (0.72+/-0.015; mean+/-SE) than in medium or small colonies (0.65+/-0.02). Additionally, dispersal probabilities were higher going from small to large colonies (0.20+/-0.01) than from large to small (0.08+/-0.01), as predicted by theory of habitat selection shaped by fitness maximization. These asymmetries are likely to generate size-specific colony population dynamics, so they should be taken into account in studies of colonial birds and other metapopulation-like systems. Allee effects, that is, positive density dependence, appear to be the cause of the evolution of dispersal behavior and may explain the maintenance of coloniality in this species.     

Testosterone and group size in cliff swallows: testing the "challenge hypothesis" in a colonial bird

The "challenge hypothesis" states that increases in testosterone levels of male animals during the breeding season are directly related to the extent of intrasexual competition for resources or mates that they experience. Although often tested in territorial species, the challenge hypothesis has not been evaluated for colonial animals that live in groups of different sizes and that thus experience different intensities of intrasexual competition. We measured circulating testosterone levels of male and female cliff swallows (Petrochelidon pyrrhonota) in southwestern Nebraska, where these birds nest in colonies of widely different sizes. Males had significantly higher testosterone levels than females, as expected. For males especially, there was a seasonal rise in testosterone levels early in the nesting cycle, corresponding to the period when birds were establishing nest ownership and egg laying, and then a fall as they switched to parental duties. Testosterone levels varied significantly with colony size; for both sexes, birds in larger colonies had higher levels of testosterone than those in smaller colonies when controlling for date. Age and body mass were not related to testosterone levels. Higher levels of testosterone for birds of both sexes in larger colonies probably reflect greater competition for matings, often extra pair, in the more social nesting situations. The results support the predictions of the challenge hypothesis.     

Effect of colony size on the competitive outcome of encrusting colonial organisms

The relative importance of colony size ratio of interacting species was studied in Tomioka Bay, Japan. Six encrusting colonial species belonging to the following three different taxonomic groups were tested: Ascidia (three species), Bryozoa (two) and Porifera (one). Colonies of these organisms were grown in the community of sessile organisms developed on plastic panels. Logistic regression analysis was carried out to determine the effect of size ratio on the competitive outcome of interacting colonies. The results between all possible combinations among these six species did not show a significant size effect in competitive outcome (i.e. a larger colony size did not always prove important in the success of a competitive interaction with smaller colonies of other species). On the contrary, competitive success depends on the types of species interacting. Certain species such asDidemnum moseleyi (ascidian) andHaliclona sp. (sponge), in spite of being smaller in colony size, won in competitive interactions with larger colonies of other species such asDiplosoma mitsukurii (ascidian) andWatersipora subovoidea (bryozoan). These results contradict the one reported earlier: that the larger the colony size, the more chance the colony will have to win in competitive interactions.     

Rates of brood development in a social wasp: effects of colony size and parasite infection

Summary Colonies of eusocial insect species are most vulnerable during the founding stage. Many species have evolved means to minimize the length of the founding, or pre-emergence, stage by accelerating the rate of development of the first worker offspring. Other things being equal, the sooner a colony can begin producing workers, the less the risk of colony failure, the steeper the growth curve of the colony during the ergonomic stage, and the larger the colony will be at reproductive maturity. Swarm-founding species, whose founding units consist of hundreds or thousands of workers, may face less selection pressure to minimize the duration of the founding stage than independent-founding species. However, swarm size varies within species, and small swarms face greater risk of extinction during the founding stage than large swarms. This consideration predicts that within a species, small swarms should have shorter founding stages than large swarms, likely by rearing a small group of precocious brood. On the other hand, evidence that large social groups organize colony labor more efficiently, gather resources more predictably, and homeostatically maintain physical conditions inside the nest within narrower ranges, predicts that larger groups should rear all brood more rapidly and therefore have shorter founding stages. To test whether small or large swarms have shorter founding stages in colonies of Polybia occidentalis, a Neotropical swarm-founding wasp, we measured brood development rates in colonies collected after 28 days of development, just short of the minimum egg-to-adult development time. We found that as size increased across colonies, pre-emergence times decreased, mean age of pupae in the nest increased, and median age of the brood (larvae + pupae) increased. That is, brood developed significantly faster in large colonies than in small ones. Using these same measures, we also found that infection by a gregarine parasite increased brood development time, independently of colony size.Received 10 April 2003; revised 29 October 2003; accepted 21 November 2003.     

Feather micro-organisms and uropygial antimicrobial defences in a colonial passerine bird

1.  The relationship between the composition of communities of micro-organisms and their hosts remains poorly understood. We conducted extensive field studies of feather-degrading bacteria, other cultivable bacteria, and fungi on the plumage of a migratory bird, the barn swallow Hirundo rustica Linnaeus, to understand the association between micro-organisms, host sociality and host antimicrobial defences, as reflected by the size of the uropygial gland.
2.  The abundance of feather-degrading bacteria, but not other cultivable bacteria or fungi, decreased with increasing size of the uropygial gland.
3.  Females had more feather-degrading bacteria than males.
4.  Barn swallows living in larger colonies had more feather-degrading bacteria than less social conspecifics.
5.  These findings suggest that the uropygial gland plays a specific role in regulating the abundance of feather-degrading bacteria that furthermore depends on the social environment of the host.     

Lifetime fitness correlates of natal dispersal distance in a colonial bird

1. Obtaining empirical evidence of the consequences of dispersal distance on fitness is challenging in wild animals because long-term, unbiased data on reproduction, survival and movement are notoriously difficult to obtain. 2. Lifetime fitness correlates of natal dispersal distance were studied in an isolated population of the facultatively colonial lesser kestrel Falco naumanni (Fleischer) monitored during 8 years at north-eastern Spain, where most birds (83%) dispersed from their natal colony to settle at distances ranging from 112 m to 136.5 km. 3. Neither annual breeding success nor age at recruitment was affected by natal dispersal distance. However, a capture-mark-recapture analysis revealed that survival during the year following recruitment decreased exponentially with dispersal distance, with differences of up to 15% between philopatrics and long-distance dispersers. In subsequent years, it remained similar irrespective of the natal dispersal distance moved. These results did not seem to be biased by long-distance dispersers settling differentially in the periphery of the population (which could emigrate permanently and be considered dead in future occasions) or within-individual consistency in successive dispersal distances, so our results appear to reflect genuine survival differences between dispersal tactics. 4. Average lifetime fledgling production, average lifetime recruitment success and rate-sensitive individual fitness (λ(ind)) also decreased with the distance from the natal to the first-breeding colony, indicating that dispersal decisions early in life affecting immediate survival prospects may translate into long-term fitness costs. 5. Both survival and lifetime fitness models including continuous dispersal distances significantly improved the characterization of the effect on fitness compared with models considering dispersal as a discrete process (i.e. dispersal vs. philopatry at a colony level). 6. Long-distance dispersers were more likely to establish new colonies regardless of whether they recruited in the centre or the periphery of the population, revealing their important role in the colonization of unoccupied patches. Individuals experienced a higher probability of mortality in small and newly funded colonies, so lifetime fitness costs of dispersal seem to be explained by recruitment in sites where average quality is low because of high uncertainty in survival prospects.     

Genome size of chrysophytes varies with cell size and nutritional mode

The cellular content of nuclear DNA varies up to 200,000-fold between eukaryotes. These differences can arise via different mechanisms. In particular, cell size and nutritional mode may influence evolution of the nuclear DNA content. Chrysophytes comprise organisms with different cell organizations and nutritional modes. Heterotrophic clades evolved independently several times from phototrophic or mixotrophic ancestors. Thus, chrysophytes are an ideal model taxon for investigating the effect of the nutritional mode on cellular DNA content. We investigated the genome size of heterotrophic, mixotrophic, and phototrophic chrysophytes. We demonstrate that cell sizes and genome sizes differ significantly between taxa with different nutritional modes. Phototrophic strains tend to have larger cell volumes and larger genomes than heterotrophic strains do. The investigated mixotrophic strains had intermediate cell volumes and small to intermediate genome sizes. Heterotrophic chrysophytes had the smallest genomes and cell volumes compared to other chrysophytes. In general, genome size increased with cell volume, but cell volume only partially explained the variation in genome size. In particular, genome sizes of mixotrophic strains were smaller than expected based on cell sizes.     

Socially-induced variation in physiological mediators of parental care in a colonial bird

Social facilitation of reproduction occurs in humans and animals, and may represent one of the bases of reproduction in groups. However, its underlying physiological mechanisms remain largely unexplored. Here, we found in a colonial bird, the Adélie penguin (Pygoscelis adeliae), that the number of parental interactions (nest relief ceremonies) performed by breeding individuals on the colony was positively related to prolactin levels in other breeding individuals exposed to these interactions (i.e. focal individuals). As prolactin is typically involved in the expression of parental behaviour in birds, this suggests that parental interactions by conspecifics represent social cues that might increase parental motivation in focal individuals. Moreover, parental interactions were not related to corticosterone levels in focal individuals, suggesting that these social cues were not stressful for penguins. However, social stimulation still had a cost for focal individuals, as it was negatively related to their antioxidant defences (a component of self-maintenance). As social stimulation was also positively related to prolactin levels, this highlights the fact that social stimulation acts on the trade-off between reproduction and self-maintenance. For the first time, the results of the current study shed light on the physiological factors potentially underlying social facilitation of parental care. Importantly, they suggest that, even though social facilitation of parental care may increase breeding performance, it can also negatively affect other fitness components.     

Interspecific parasite exchange in a mixed colony of birds

Studies of avian host-parasite interactions rarely include consequences of relationships among hosts for either the host or parasite species. In this study, we examine the ectoparasitic burden of adult and nestling European bee-eaters (Merops apiaster) and rock sparrows (Petronia petronia) in a mixed colony. We found that (1) each bird species had its own species of lice; (2) hematophagous mites parasitized both adults and nestlings of both bird species; (3) Carnus hemapterus, a common parasite of nestling bee-eaters, also infested rock sparrow nestlings, a species not previously described as a host for this dipteran; and (4) whereas C. hemapterus did not show high host specificity within the colony, the emergence of adult flies was synchronized with the start of hatching in bee-eater nests. We suggest that coexistence of these 2 bird species results in parasite exchange, bee-eaters obtaining mites from sparrows and sparrows becoming infested by C. hemapterus. Differences in the detrimental effects of parasite transfer for each host species may result in a process of apparent competition mediated by shared parasites. Interspecific parasite exchange is an important aspect of host-parasite relationships in mixed colonies, which requires further attention.     

Testis size increases with colony size in cliff swallows

By using a sample of over 800 male cliff swallows (Petrochelidonpyrrhonota) that died during a rare climatic event in our Nebraskastudy area in 1996, we investigated how testis size was relatedto body size, age, parasite load, a bird's past colony-sizehistory, and spleen size. Testis volume increased with bodysize. After correcting for body size, testis volume was lowestfor birds age 1 and 2 years but did not vary with age for males3 years old or more. Birds occupying parasite-free (fumigated)colonies had significantly larger testes than did birds at nonfumigatedsites. Testis volume increased significantly with the size ofthe breeding colonies a bird had used in the past. Testis volumehad no relationship with spleen volume after correcting forbody size. The results show within a species that larger testesare favored in more social environments, probably reflectinga response to increased rates of extrapair copulation (and thussperm competition) among cliff swallows in large colonies. Thepresence of ectoparasites, by inflating levels of plasma corticosterone,may in turn reduce testis mass. These data provide no supportfor the hypothesis that large testes, perhaps by producing moretestosterone, are immunosuppressive and thus costly for thatreason.     

How pollinator-mediated mating varies with population size in plants

In most higher plants sexual interactions are mediated by animal pollinators that affect the number and differential reproductive success of mates. The number and sex of breeding individuals in populations are central factors in evolutionary theory, but the quantitative effect of plant population size on pollinator-mediated mating is understudied. We investigated variation in pollen removal (male function) and fruit set (female function) among flowering populations of different size of two bumblebee-and one butterfly-pollinated, rewardless, pollen-limited, hermaphroditic orchid species in Sweden. As the amount of pollen removed from plants by insects (either absolute or proportional) increased, so did the number of pollinations, whereas the proportions of plants with different pollinator-designated functional sex (male, female, hermaphrodite) depended primarily on the ratio between the amount of fruit set and pollen removed within populations. A larger population size was found to have several effects: (1) the total numbers of pollinia removed and fruits set increased; (2) the proportion of pollen removed from plants decreased; (3) the proportion of flowers pollinated decreased in the butterfly-but was not affected in the bumblebee-pollinated species; (4) the ratio between fruits set and pollinia removed increased linearly in the bumblebee-pollinated species but reached a maximum at c. 80 individuals in the butterfly-pollinated species; (5) the numbers of pollinator-designated pure male and hermaphrodite individuals increased; and (6) the variance in pollinium removal, but not fruit set, increased among individuals. These findings empirically verify the basic importance of population size for the mating structure of outcrossing plants, and indicate that selection for female sexual traits is reinforced when population size is smaller while selection for male sexual traits is reinforced when population size is larger.     

Ejaculate size varies with socio-sexual situation in an insect

Abstract.

• 1 Males operate within a finite energy budget and cannot produce limitless supplies of sperm. On the other hand, when a female mates with a second male while still containing fertile sperm from a rival male, selection should favour the male that inseminates more sperm. Optimal strategy should thus be for males to exercise discretion in the allocation of sperm to individual females. Assuming the outcome of sperm competition to be based on either the raffle or kamikaze principles, the sperm competition hypothesis predicts a positive association between the probability that the sperm will find themselves in competition with sperm from rival males and the number of sperm inseminated.
• 2 The beetle, Tenebrio molitor L., behaves in accordance with this hypothesis. Males accompanied by a rival male before and during mating inseminate more sperm per ejaculate than unaccompanied males. Accompanied males are also faster to initiate mating and more likely to show post-copulatory guarding. Adjustment of number of sperm inseminated was shown by males subjected to both long-term (5 days) and short-term (5 min) exposure to potential intrasexual competitors. Individual males exposed to both levels also demonstrated the ability to adjust their ejaculate according to socio-sexual situation.
• 3 We conclude that male T.molitor adjust the number of sperm they inseminate according to some perception of the risk of sperm competition.

     

Singing in the moonlight: dawn song performance of a diurnal bird varies with lunar phase

It is well established that the lunar cycle can affect the behaviour of nocturnal animals, but its potential to have a similar influence on diurnal species has received less research attention. Here, we demonstrate that the dawn song of a cooperative songbird, the white-browed sparrow weaver (Plocepasser mahali), varies with moon phase. When the moon was above the horizon at dawn, males began singing on average 10 min earlier, if there was a full moon compared with a new moon, resulting in a 67% mean increase in performance period and greater total song output. The lack of a difference between full and new moon dawns when the moon was below the horizon suggests that the observed effects were driven by light intensity, rather than driven by other factors associated with moon phase. Effects of the lunar cycle on twilight signalling behaviour have implications for both pure and applied animal communication research.     

Ectoparasites cause increased bilateral asymmetry of naturally selected traits in a colonial bird

Abstract Parasitism has been shown to correlate with levels of bilateral symmetry in some organisms, with more asymmetric individuals often having more parasites. However, few studies have shown experimentally that parasitism directly causes increased asymmetry. By fumigating some cliff swallow (Petrochelidon pyrrhonota) colonies and leaving others untreated, we investigated experimentally whether ectoparasitism by the cimicid swallow bug led to higher levels of asymmetry in length of wings, outer tail feathers, and tarsus among juvenile and adult birds. Juveniles from fumigated colonies measured soon after fledging had significantly less asymmetry in wing and outer tail length than juveniles from nonfumigated colonies; asymmetry in tarsus length was unaffected by parasitism. Adults that had undergone one or more post‐juvenal molts on the wintering grounds showed no differences in asymmetry between those reared in fumigated vs. nonfumigated colonies. These results show that ectoparasitism directly leads to increased feather asymmetry in cliff swallows, probably through parasite‐induced nutritional stress. Because wing and tail asymmetry impair flight performance and reduce foraging efficiency, the increased asymmetry caused by parasites represents a fitness cost to cliff swallows. This is among the few experimental studies to show an effect of parasites on asymmetry of naturally selected characters.     

Brain size,head size and behaviour of a passerine bird

A recent increase in comparative studies of the ecological and evolutionary consequences of brain size in birds and primates in particular have suggested that cognitive abilities constitute a central link. Surprisingly, there are hardly any intraspecific studies investigating how individuals differing in brain size behave, how such individuals are distributed and how brain size is related to life history and fitness components. Brain mass of the barn swallow Hirundo rustica was strongly predicted by external head volume, explaining 99.5% of the variance, allowing for repeatable estimates of head volume as a reflection of brain size. Repeatability of head volume within and between years was high, suggesting that measurement errors were small. In a 2 years study of 501 individual adult barn swallows, I showed that head volume differed between sexes and age classes, with yearlings having smaller and more variable heads than older individuals, and females having smaller and more variable heads than males. Large head volume was not a consequence of large body size, which was a poor predictor of head volume. Birds with large heads arrived early from spring migration, independent of sex and age, indicating that migratory performance may have an important cognitive component. Head volume significantly predicted capture date and recapture probability, suggesting that head volume is related to learning ability, although morphological traits such as wing length, aspect ratio and wing area were unimportant predictors. Intensity of defence of offspring increased with head volume in females, but not in males. Barn swallows with large heads aggregated in large colonies, suggesting that individuals with large heads were more common in socially complex environments. These results suggest that brain size is currently under natural and sexual selection, and that micro‐evolutionary processes affecting brain size can be studied under field conditions.     

Song repertoire size varies with HVC volume and is indicative of male quality in song sparrows (Melospiza melodia)

Complex birdsong is a classic example of a sexually selected ornamental trait. In many species, females prefer males with large song repertoires, possibly because repertoire size is limited by the size of song control nuclei which reflect developmental success. We investigated whether song repertoire size was indicative of brain area and male quality in song sparrows (Melospiza melodia) by determining if repertoire size was related to the volume of song control nucleus HVC, as well as several morphological, immunological and genetic indices of quality. We found that males with large repertoires had larger HVCs and were in better body condition. They also had lower heterophil to lymphocyte ratios, indicating less physiological stress and a robust immune system as measured by the number of lymphocytes per red blood cell. Song repertoire size also tended to increase with neutral-locus genetic diversity, as assessed by mean d2, but was not related to internal relatedness. Our results suggest several mechanisms that might explain the finding of a recent study that song sparrows with large song repertoires have higher lifetime fitness.     

Truncated power laws reveal a link between low-level behavioral processes and grouping patterns in a colonial bird

Background

Departures from power law group size frequency distributions have been proposed as a useful tool to link individual behavior with population patterns and dynamics, although examples are scarce for wild animal populations.

Methodology/Principal Findings

We studied a population of Lesser kestrels (Falco naumanni) breeding in groups (colonies) from one to ca. 40 breeding pairs in 10,000 km² in NE Spain. A 3.5 fold steady population increase occurred during the eight-year study period, accompanied by a geographical expansion from an initial subpopulation which in turn remained stable in numbers. This population instability was mainly driven by first-breeders, which are less competitive at breeding sites, being relegated to breed solitarily or in small colony sizes, and disperse farther than adults. Colony size frequency distributions shifted from an initial power law to a truncated power law mirroring population increase. Thus, we hypothesized that population instability was behind the truncation of the power law. Accordingly, we found a power law distribution through years in the initial subpopulation, and a match between the power law breakpoint (at ca. ten pairs) and those colony sizes from which the despotic behavior of colony owners started to impair the settlement of newcomers. Moreover, the instability hypothesis was further supported by snapshot data from another population of Lesser kestrels in SW Spain suffering a population decline.

Conclusions/Significance

Appropriate analysis of the scaling properties of grouping patterns has unraveled the link between local agonistic processes and large-scale (population) grouping patterns in a wild bird population.