Helpers in colonial cooperatively breeding sociable weavers <Emphasis Type="Italic">Philetairus socius</Emphasis> contribute to buffer the effects of adverse breeding conditions

Some studies on the effects of helpers in cooperatively breeding vertebrates show a positive effect of helper presence on reproductive output whereas others find no effect. One possibility for this discrepancy is that helpers may have a positive effect when breeding conditions are adverse, while their effect might go unnoticed under good conditions. We investigate this hypothesis on sociable weavers Philetairus socius, a colonial cooperatively breeding passerine that inhabits a semi-arid region where breeding conditions vary markedly. We used multivariate mixed models to analyse the effect of helpers on reproduction under contrasting environmental and social conditions while controlling for parental and colony identity. We found that reproductive success in sociable weavers was primarily influenced by nest predation and rainfall. In addition, colony size was negatively associated with hatching and fledging success and number of young fledged per season. Helpers had a less prominent but significant influence on feeding rates and reproductive outcome. In agreement with expectations, the presence of helpers counteracted some of the negative effects of breeding in periods of low rainfall or in large colonies and was also associated with an increased number of young fledged per season. Our results illustrate that the effect of helpers might be detectable mostly under unfavourable conditions, but can contribute to improve reproductive performance in those situations.     

Group-living in cliff swallows as an advantage in avoiding predators

Summary Cliff swallows (Hirundo pyrrhonota) in SW Nebraska, USA, nest in colonies and associate in groups away from their colonies. The degree to which group-living in this species affords advantages in the avoiding of predators was examined. The distance from the colony at which a snake predator was detected increased with colony size. In flocks away from the colonies, group vigilance increased, but the time that each individual spent vigilant decreased, with flock size. As a result, birds in large flocks had more time for preening and mud-gathering. Cliff swallows did not effectively mob predators and thus were unable to deter predators regardless of group size. Nesting within each colony was highly synchronous, but when the effects of ectoparasites on nesting success were removed, individuals nesting during the peak breeding period were no more successful than those nesting before or after the peak. This suggests that swamping of predators is unlikely in cliff swallow colonies. Nests at the edges of colonies were more likely to be preyed upon than nests nearer the center, suggesting that colonial nesting conferred some selfish herd benefits. Overall reproductive success did not vary with colony size. While cliff swallows receive some anti-predator benefits by living in groups, the avoidance of predators is probably not a major selective force for the evolution of coloniality in this species.     

Fitness costs of an immune response in the house martin (<Emphasis Type="Italic">Delichon urbica</Emphasis>)

Immune responses constitute a major way for hosts to defend themselves against parasites. Because hosts do not habitually produce strong responses all the time, immune responses might be costly to produce or maintain. We tested experimentally if the production of a response to a challenge with a novel antigen resulted in a cost in terms of fitness using the highly colonial house martin Delichon urbica as a model system. We injected adult breeding birds during laying of the first clutch with either Newcastle disease virus (NDV) or a control injection, and the clutch was subsequently removed to induce relaying. NDV stimulates the non-specific immune system, causing production of antibodies during a period of more than 2 weeks. Accordingly, we found an increase in leukocyte counts in experimental birds compared to controls. Experimental treatment reduced the frequency of re-laying, caused a delay in timing of relaying and a reduction in brood size. Quality of nestlings in terms of body size, body mass and T-cell-mediated immune response did not differ significantly between treatments. Overall, seasonal reproductive success differed significantly between treatments, showing that the production of an immune response by adult birds is costly in terms of future fecundity.     

Why are larger convict cichlid (Cichlasoma nigrofasciatum) fry sometimes adopted into broods of smaller fry?

Females of Elasmucha grisea defend their eggs and small nymphs against invertebrate predators. Females sometimes guard their clutches side by side on the same birch leaf. We studied benefits of this joint guarding both in the field and in the laboratory. We found that adjacent females had significantly larger clutches than solitary females. In the laboratory, we studied the effectiveness of joint versus single defence against ant (Formica uralensis) predators. We established female pairs from initially singly guarding females by cutting off pieces of leaves with egg clutches and pasting them beside another female guarding her clutch. In the control group the females with their clutches were similarly cut off but these clutches were placed on another leaf without any female. The birch twigs where females guarded their clutches were placed in cages in close proximity to laboratory ant nests. In the experimental treatment, two females guarded their clutches together and at the same nest there was another birch twig without a female. In the control treatment two twigs with one female on each were placed close to another ant nest. Two females defended their clutches significantly more successfully, losing fewer eggs than did the single females. This primitive form of female sociality in parent bugs resembles colonial nesting in birds, where communal defence is also important. However, to our knowledge this is the first experiment where the benefit of joint guarding has been tested directly by manipulating the size of the breeding group rather than by measuring the risk of predation in groups of different size.     

Fieldfare (Turdus pilaris) breeding success in relation to colony size,nest position and association with Merlins (Falco columbarius)

Summary Clutch size, nestling production and breeding success were studied in colonial Fieldfares (Turdus pilaris) in a subalpine birch forest during ten breeding seasons. Reproductive success was highest for central pairs in large colonies; such pairs benefited most from communal defence against nest predators. Fieldfares and Merlins (Falco columbarius) usually bred in association. Fieldfares breeding away from Merlins had lower breeding success than pairs associated with Merlins, which also benefited by reduced nest predation. Fieldfares apparently chose to nest near Merlins, which had already laid eggs when the thrushes started nest-building.     

Group size and ectoparasitism affect daily survival probability in a colonial bird

Little is known in general about how group size or ectoparasitism affect survival in colonial animals. We estimated daily within-season survival probabilities for nesting adult and recently fledged juvenile cliff swallows (Petrochelidon pyrrhonota) at 239 colonies from 1983 to 2003 in southwestern Nebraska, USA. Some colonies had been fumigated to remove ectoparasites. We conducted mark-recapture at each colony site to estimate daily survival. There were no systematic differences between males and females in daily survival. Adults and juveniles occupying parasite-free colonies had, on average, 4.4% and 62.2% greater daily survival, respectively, than their counterparts in naturally infested colonies. Daily survival of all birds increased with colony size for both parasite-free colonies and those under natural conditions, although the effect was stronger for adults at fumigated sites and for juveniles. Average daily survival probability for adults tended to increase during warmer and drier summers. Although daily survival varied at some sites over the course of the nesting cycle, there were no strongly consistent within-year temporal effects on survival. Even small differences in daily survival probability can translate into large effects on mean lifespan. The deleterious effects of ectoparasites on daily survival within the season represent a previously unknown cost of ectoparasitism. The increase in within-season survival with colony size reflects the net effects of many costs and benefits associated with colony size. Ectoparasitism is probably the most important cost that tends to partly balance the positive effects of large colonies. The greater survival of cliff swallows in the larger colonies is a previously unknown advantage of colonial nesting.Communicated by P. Heeb     

Colonial and single breeding in fieldfares,Turdus pilaris L.: a comparison of nesting success in early and late broods

Summary Pairs of fieldfares breed singly or in colonies. Their breeding season was subdivided into an early part, when the trees were still leafless, and a late part, when the trees had leaves. Early colonial pairs has a significantly higher nesting success than single pairs. However, amongst late broods the nesting success of both categories of pairs was the same. Colonies of only two pairs show a weak trend suggesting that it could be advantageous for early broods to have a neighbour nearby, while the nesting success of late broods seems to be highest if the neighbour is farther away. The number of neighbours is more important than the distance to the nearest neighbour when colonies of different sizes are examined. The nesting success of colonial broods is also influenced by the time of breeding. Success increases with colony size, in early colonies, but the opposite applies for the late breeding season.A little owl was placed near single and colonial nests to show how communal defence contributed to the greater success of early colonial nests. Predators entering a colony risk being contaminated by significantly more faeces due to the higher rate of attacks by fieldfares using aimed defecation near colonial nests and these could affect the predators' flying ability.The possibility of fieldfare colonies serving as information centres for food finding is also discussed.     

Nestling size rank in the little egret (Egretta garzetta) influences subsequent breeding success of offspring

Few studies have investigated the long-term fitness consequences of nestling size hierarchies in altricial birds. In this study, we investigated whether or not the size rank order of siblings influences subsequent breeding success in the little egret, Egretta garzetta. From a marking program allowing individual recognition of wild birds, we obtained data on the breeding success of 56 pairs comprising individuals for which the size rank order was known. The breeding success in these pairs was positively influenced by the age of the marked bird but negatively affected by the laying date of the pair and the size rank order of the marked individual. There was also a significant difference between breeding colonies. We suggest two main hypotheses for a link between size rank order of individuals and their breeding success and we discuss our results in relation to current hypotheses on the adaptive value of hatching asynchrony. Received: 10 August 1998 / Accepted after revision: 13 December 1998     

Solitary nesting as an alternative breeding tactic in colonial nesting bluegill sunfish (Lepomis macrochirus)

Colonial breeding can evolve in response to benefits afforded by clumped individuals, such as reduced predation and increased ease of assessing potential mates. However, colonial breeding can also impose costs such as increased disease transmission or increased cuckoldry. Here, we investigate solitary nesting as a potential alternative breeding tactic in colonial breeding bluegill sunfish (Lepomis macrochirus). Most male bluegill, termed parentals, compete for nesting sites in colonies and then court and spawn with females and provide sole care of the eggs. Although nesting in a colony results in reduced predation and fungal infection of broods, it comes at a cost of increased parasitism by specialized cuckolder males that do not nest. We found that 4.5% of parentals forgo spawning in a colony and instead construct nests solitarily. Solitary males were of similar size and age to colonial males, but were in significantly better condition. Solitary males obtained as many eggs as males nesting in the center of colonies, and significantly more than males nesting on the periphery of colonies. Thus, females do not appear to discriminate against solitary males. Solitary males had smaller ear tabs, a presumed sexually selected character used by parental males in intrasexual competition, than colonial males. Tracking data revealed consistency in nesting tactic (but not nest position within the colony) between spawning attempts. We suggest that solitary nesting represents either a facultative decision made by parental males in top condition at the onset of breeding, or a life history decision to forgo spawning in colonies.Communicated by K. Lindström     

A test of the mismatch hypothesis: How is timing of reproduction related to food abundance in an aerial insectivore?

In seasonal environments, vertebrates are generally thought to time their reproduction so offspring are raised during the peak of food abundance. The mismatch hypothesis predicts that reproductive success is maximized when animals synchronize their reproduction with the food supply. Understanding the mechanisms influencing the timing of reproduction has taken on new urgency as climate change is altering environmental conditions during reproduction, and there is concern that species will not be able to synchronize their reproduction with changing food supplies. Using data from five sites over 24 years (37 site-years), we tested the assumptions of the mismatch hypothesis in the Tree Swallow (Tachycineta bicolor), a widespread aerial insectivore, whose timing of egg-laying has shifted earlier by nine days since the 1950s. Contrary to the mismatch hypothesis, the start of egg-laying was strongly related to food abundance (flying insect biomass) during the laying period and not to timing of the seasonal peak in food supply. In fact, food abundance generally continued to increase throughout the breeding season, and there was no evidence of selection based on the mistiming of laying with the seasonal peak of food abundance. In contrast, there was selection for laying earlier, because birds that lay earlier generally have larger clutches and fledge more young. Overall, initial reproductive decisions in this insectivore appear to be based on the food supply during egg formation and not the nestling period. Thus, the mismatch hypothesis may not apply in environments with relatively constant or abundant food throughout the breeding season. Although climate change is often associated with earlier reproduction, our results caution that it is not necessarily driven by selection for synchronized reproduction.     

Edge Effects in the Great Tit: Analyses of Long-term Data with GIS Techniques

Abstract:  In contemporary fragmented landscapes, edges are commonplace, and understanding the effects of edge environments is thus essential for the conservation of forest communities. The reproductive output of forest passerines is often reduced close to forest edges. Possible explanations include overcrowding by conspecifics, elevated rates of predation, and the occurrence of lower-quality habitat and/or individuals at forest edges. We attempted to separate these processes by examining edge effects in the absence of nest predation and by effectively controlling for differences in breeding density and the quality of habitats and individuals. We used an edge distance index (EDI), which accounts for the number and distribution of edges in close proximity to a breeding location, to help explain variation in breeding density, nesting success, and reproductive traits of 8308 pairs of Great Tits ( Parus major ) breeding between 1965 and 2005, in Wytham, near Oxford, United Kingdom. Results from linear mixed modeling confirmed higher breeding density and a higher proportion of immigrant individuals at forest edges. Nevertheless, independently of these effects, we also found that birds laying later, with smaller clutches but larger eggs, were typical of edge environments. The number of offspring recruited to the breeding offspring per breeding attempt was also reduced at edges, both directly and mediated through changes in clutch size and laying date. Edge effects on life histories were detectable within individual females and up to 500 m from the woodland edge. Woodland edges are increasingly common in contemporary fragmented landscapes. Therefore these results, which suggest a pervasive effect of edges on reproduction, are of considerable importance to the management and conservation of forest communities.     

Behavioral tactics of subadult female floaters in the tree swallow

Summary There are large numbers of reproductively mature female tree swallows (Tachycineta bicolor) which do not breed due to limits of suitable nesting cavities. Many of these floaters are one-year-old females that have a distinctive subadult plumage. This study examines the behavioral tactics that these subadult female floaters use to obtain breeding opportunities. Early in the season, subadult floaters tended to intrude briefly (Fig. 4) on many nest sites in succession (Figs. 2, 3), although they rarely gained close access to nest sites (Fig. 5). Subadults responded very quickly to vacant nest sites, where the resident female had been experimentally removed, by entering the nest cavity and defending it from conspecifics. We argue that the early season exploratory behavior increases a subadult's chances of discovering a vacant nest site, rather than increasing its success in evicting resident females or laying eggs in other females' nests. During the nestling period, subadult females intruded on fewer nest sites for longer periods, and often gained close access to the nest site. Late in the season, subadult floaters may be gathering information on the quality of nest sites for the next breeding season, rather than searching for current breeding opportunities. The reproductive tactics of subadult female tree swallows are consistent with the breeding threshold model for the evolution of delayed plumage maturation in passerines.     

Size-dependent reproduction in the Caribbean gorgonian Pseudoplexaura porosa

Reproductive success among clonal taxa is often portrayed as a simple function of clone size, but reproduction in Pseudoplexaura porosa (Houtuyn), a common Caribbean gorgonian, reflects a more complex relationship between size and gamete production. Tagged colonies were sampled at two reefs in the San Blas Islands, Panama during the 1995 spawning season to determine the colony size at first reproduction, size-dependent polyp fecundities, and whole colony gonad production, and to follow the gametogenic cycle. Additional data were obtained during non-spawning months in 1984 and 1985. Of 120 colonies ranging in height from 20 to 250 cm, only colonies taller than 50 cm were reproductive. An average of 4.33 oocytes polyp⁻¹, 560 to 800 μm in diameter, were released during monthly synchronous spawning events from June to September. Oocytes released each month developed from a group of intermediate size oocytes present at the end of the preceding month. Almost all oocytes >560 μm were released during each spawning event. Spermatogenesis was a shorter process than oogenesis. During each spawning month, mature spermaries developed from a group of <125 μm spermaries. Virtually all spermaries >190 μm were released each month, and most of the spermary volume in mature male polyps was generated anew each month of spawning. Among reproductive colonies, polyp fecundity increased with colony height from 50 to 200 cm. Per capita fecundity of >200 cm tall female colonies was lower than for 150 to 200 cm colonies, but whole colony fecundity of large colonies was greater due to the exponential increase in the number of polyps that occurs with increasing colony height. In male colonies, both polyp and colony reproductive output increased with colony height. The large amounts of sperm produced by large male colonies probably contributes to the high rates of in situ fertilization observed in P. porosa. Differences in fecundity as a function of colony size and sex suggest differences in some combination of the cost of reproduction and/or allocation of resources to reproduction. Received: 9 September 1998 / Accepted: 7 June 1999     

Frequency and timing of extrapair fertilisation in the polyandrous red phalarope (Phalaropus fulicarius)

In sequentially polyandrous bird species, where females mate with different males in succession during a single breeding season, sperm stored by females can occasionally lead to extrapair fertilisations (EPFs) in clutches cared for by the second and subsequent males. Thus, we predicted that in red phalaropes (Phalaropus fulicarius) – a sex-role-reversed, sequentially polyandrous, arctic breeding shorebird – EPFs would occur more frequently in clutches laid later in the breeding season. We used multilocus DNA profiling to examine the frequency and timing of EPFs in a population of red phalaropes breeding in the Canadian high arctic. Using a technique to determine parentage without maternal DNA, we inferred that 6 of 70 chicks in 18 broods resulted from EPFs – one extrapair chick in each of 6 broods. These results were supported by a further analysis using microsatellite DNA. As predicted, broods containing EPFs hatched from clutches laid significantly later in the season than did broods containing no EPFs. The difference in median hatch dates between broods with and without EPFs was 9.0 days, or 38% of the entire egg-laying period in that season. For the whole breeding season, we estimated that 6.5% of chicks were sired by extrapair males, which is similar to extrapair paternity estimates for other sex-role-reversed birds, but relatively low compared to the majority of socially monogamous species studied so far. Received: 24 August 1998 / Received in revised form: 18 January 1999 / Accepted: 24 January 1999     

Infanticidal and anti-infanticidal strategies in the swallow Hirundo rustica

Summary Five definite and nine suspected cases of infanticide were documented in the swallow Hirundo rustica. Unmated males visited neighbouring nests and removed entire broods during the first days of the nestling period, preferentially from nests owned by young late-breeding individuals. Infanticide followed the disappearance of the male nest owner in 11 of the 14 cases. The unmated infanticidal male later mated with the female victim although renesting during the same season only took place in 12 of the 14 cases. The fraction of unmated males increased with colony size, and infanticide was relatively more common if unmated males were abundant. Females did not engage in extra-pair copulations in order to avoid infanticide. Intense guarding of the nest and its contents prevented infanticide. Nest guarding intensity was higher in colonial compared with solitarily breeding swallows, and guarding intensity increased with colony size. Experimental removal of male swallows during the early nestling period reduced the nest guarding intensity and increased frequency of visits from unmated males in colonies, but not among solitarily breeding swallows. Colonial nests from which the male was removed suffered from infanticide more often than solitary nests, and nests where infanticide was recorded were guarded significantly less intensely than other nests before the infanticidal incidents.     

Maternal characteristics and the production and recruitment of sons in the eastern kingbird (<Emphasis Type="BoldItalic">Tyrannus tyrannus</Emphasis>)

Sex allocation theory predicts that if variance in reproductive success differs between the sexes, females who are able to produce high-quality young should bias offspring sex ratio towards the sex with the higher potential reproductive success. We tested the hypothesis that high-quality (i.e., heavy) female eastern kingbirds (Tyrannus tyrannus) that bred early in the breeding season would produce male-biased clutches. A significant opportunity for sexual selection also exists in this socially monogamous but cryptically polygamous species, and we predicted that successful extra-pair (EP) sires would be associated with an excess of male offspring. Although population brood sex ratio did not differ from parity, it increased significantly with female body mass and declined with female breeding date, but was independent of the morphology and display (song) behavior (correlates of reproductive success) of social males and EP sires. Male offspring were significantly heavier than female offspring at fledging. Moreover, the probability that male offspring were resighted in subsequent years declined with breeding date, and was greater in replacement clutches, but lower when clutch size was large. Probability of resighting female offspring varied annually, but was independent of all other variables. Given that variance in reproductive success of male kingbirds is much greater than that of females, and that male offspring are more expensive to produce and have a higher probability of recruitment if fledged early in the season, our results support predictions of sex allocation theory: high-quality (heavy) females breeding when conditions were optimal for male recruitment produced an excess of sons.     

Is reduced clutch size a cost of parental care in Eastern Phoebes (Sayornis phoebe)?

What is the cost of parental care in birds? Previous studies using observational and experimental techniques on nest building and clutch sizes in a small migrant flycatcher, the Eastern Phoebe (Sayornis phoebe), led to contradictory results that did not show a consistent cost of current reproductive effort on residual reproductive output. The data presented here indicate that different elements of parental behaviors are indeed costly because they reduce various aspects of phoebes' subsequent reproductive performance. Experimental removal of old nesting structures at previously used breeding sites reduced but did not eliminate the chance of phoebes' settlement in the subsequent year. Comparing sites at which phoebes did and did not build new nests showed that nest builders completed their first clutches later, had lower probabilities of second breeding attempts, and more often lost their nesting attempt due to fallen nest structures than nest reusers. There was, however, no significant effect of nest building on the clutch sizes and rates of cowbird parasitism of first nesting attempts. Overall, sites with newly built nests had lower seasonal reproductive effort than sites with reused nests. I also examined phoebes' relative residual reproductive output in a separate breeding season when nest building was not experimentally manipulated. When controlled for confounding variables this analysis indicated that in those phoebes that did breed for a second time, the relative decrease of the sizes of first to presumed second clutches was greater at sites where first breeding attempts consisted of more total nestlings. These data are consistent with the hypothesis that parental care is costly in Eastern Phoebes and support predictions of trade-offs between the nest building, brood care, and residual egg-investment components of reproduction.     

Reproductive synchrony in the fieldfare (Turdus pilaris) in relation to spring arrival,nest predation and nestling starvation

Summary In SW Sweden, solitary fieldfares (Turdus pilaris) were synchronized to the same degree as those living in colonies. In N Sweden, I investigated the degree of synchronization among colonies of varying size because solitary breeding occurred only occasionally. The degree of synchronization was not related to colony size. Synchronization was strongest in years when breeding started late and weakest in years when breeding commenced early. In N Sweden, the nest predation risk increased slightly as the season progressed. In SW Sweden, nestling mortality due to starvation was highest in early breeders, but their fledgling production exceeded that of late breeders. Increased nest predation penalized late layers.     

Are female boat-tailed grackle colonies neutral assemblages?

Female boat-tailed grackles (Quiscalus major) nest in colonies, and apparently settle independently of male activities. Associations between colony females may be mutually beneficial (cooperative hypothesis) or females may be penalized by associating (competitive hypothesis). Contrary to predictions based on either cooperative or competitive behavior, (1) reproductive success was not related to colony size nor to internest distance; (2) intracolony nesting synchronies were the same as those of the whole population, and within colonies, there was no relationship between the timing and spacing of nearest-neighbor nests; and (3) already established females were only occasionally aggressive toward females attempting to settle near them, and their aggressive response was independent of their stage of nesting. The results suggest that females act independently of each other and do not affect each other's fitness (neutral female hypothesis). Colonies may be neutral aggregations in sites secure from ground predators. Although females benefit by co-occupying predator-free sites, advantages and disadvantages of colomality do not appear to be related to intrasexual association.     

Flexible incubation rhythm in northern fulmars: a comparison between oceanographic zones

Variation in the timing and abundance of marine food resources is known to affect the breeding behaviour of many seabirds, constraining our understanding of the extent to which these behaviours vary in different parts of a species’ range. We studied incubation shifts of northern fulmars (Fulmarus glacialis) breeding at two colonies in Arctic Canada (High Arctic oceanographic zone) and one colony in the UK (Boreal oceanographic zone) between 2001 and 2005. Fulmars in Arctic Canada had longer incubation shifts than previously reported at more southern colonies, presumably because marine productivity is lower early in the breeding season in the Arctic. Shift durations were particularly long at one colony in years with abnormally late, extensive sea-ice cover, although at the other Arctic colony, where sea-ice cover is predictably late every year, the duration of shifts was shorter than expected. At the Boreal colony, incubation shifts were much longer than expected, similar to Arctic colonies, and likely attributable to poor marine food supplies in the North Sea in recent years. Collectively, our data suggest that fulmars can adjust their incubation rhythm to compensate for poor marine feeding conditions, although this may incur a cost to body condition or reproductive success.