Density dependent effects between three competitive bird species

Summary Density and breeding success of the great tit Parus major, blue tit Parus caeruleus and collared flycatcher Ficedula albicollis were studied in nest box colony in oak forest over a period of 19 years.Intraspecific density dependent clutch size reduction was found with blue tit and great tit. In interspecific relation the high density of blue tits reduced the clutch size of great tits.In the hatching period neither intraspecific nor interspecific density dependence were showed between the tits when the third competitive species, collared flycatcher was present. The collared flycatcher significantly reduced the hatching success of both tit species and the number of fledglings of great tit.The effects of the great tits and combined density of the great and blue tits on the hatching failure and number of fledglings of collared flycatcher were found when the density of the tits was high. There were not significant relationships to the single density of blue tits.The temporal variability of the competition of the three bird species is discussed.     

Nest destruction elicits indiscriminate con‐ versus heterospecific brood parasitism in a captive bird

Following nest destruction, the laying of physiologically committed eggs (eggs that are ovulated, yolked, and making their way through the oviduct) in the nests of other birds is considered a viable pathway for the evolution of obligate interspecific brood parasitism. While intraspecific brood parasitism in response to nest predation has been experimentally demonstrated, this pathway has yet to be evaluated in an interspecific context. We studied patterns of egg laying following experimental nest destruction in captive zebra finches, Taeniopygia guttata, a frequent intraspecific brood parasite. We found that zebra finches laid physiologically committed eggs indiscriminately between nests containing conspecific eggs and nests containing heterospecific eggs (of Bengalese finches, Lonchura striata vars. domestica), despite the con‐ and heterospecific eggs differing in both size and coloration. This is the first experimental evidence that nest destruction may provide a pathway for the evolution of interspecific brood parasitism in birds.     

Asymmetric competition between two tit species: a reciprocal removal experiment

1. In a 5-year field experiment, competition for food was tested between great tit ( Parus major L.) and blue tit ( P. caeruleus L.), two common hole-nesters in Central Europe. Experimental ('allopatric') populations of both species were created during the breeding seasons by preventing the nesting or egg laying of one of the competing species.
2. An asymmetric relationship was found between the two tits; blue tits were more successful in the competitive interaction. Detectable effects were found only in nestling condition. Great tits raised lighter nestlings when breeding sympatrically with blue tits.
3. A possible mechanism is suggested that could be responsible for the different competitive abilities of the two species; blue tits are more effective in utilizing the most abundant size categories of caterpillar food supply than great tits.     

On the origin and rarity of interspecific nest parasitism in birds

Interspecific nest parasitism is surprisingly rare in birds given the potential advantages for the parasite of exploiting the parental care of other species. One possibility is that chicks will not thrive with the parental care and food of heterospecifics. I simulated parasitism in nonparasitic congeners by switching eggs between nests of three species of titmice (great tit Parus major, blue tit Parus caeruleus, and coal tit Parus ater). The experiment showed that compatibility of parental care was not a constraint preventing parasitism. I also used the model system to compare fitness consequences of inter- and intraspecific nest parasitism, addressing the problem of which form is ancestral. Fledging success (body mass, survival) was higher when an egg was added to the nest of a smaller species than to the nest of a conspecific and also higher when the parasitic chick hatched early rather than late relative to host chicks. This suggests that interspecific nest parasitism may not require a stage of intraspecific nest parasitism before evolving but may start from a larger species directly exploiting the parental care of a smaller species or a species with shorter incubation period directly exploiting a species with longer incubation period.     

Experimental manipulation of temperature reduce ectoparasites in nests of blue tits Cyanistes caeruleus

Several models predict changes in the distributions and incidences of diseases associated with climate change. However, studies that investigate how microclimatic changes may affect host–parasite relationships are scarce. Here, we experimentally increased the temperature in blue tit Cyanistes caeruleus nest boxes during their breeding season to determine its effects on the parasitic abundance (i.e. of nest‐dwelling ectoparasites, blood‐sucking flying insects and hemoparasites) in nests and the host condition of nestlings and adults. The temperature was increased using heat mats placed underneath the nest material, which resulted in an average temperature increase of 3ºC and a reduction in relative humidity of about six units. The abundance of mites Dermanyssus gallinoides and blowfly pupae Protocalliphora azurea was significantly reduced in heated nest boxes. Although not statistically significant, a lower prevalence of flea larvae Ceratophyllus gallinae was also found in heated nests. However, heat treatment did not affect hemoparasite infection of adult blue tits or the body condition of adult and nestling blue tits. In conclusion, heat treatment in blue tit nests reduced nest‐dwelling ectoparasites yet without any apparent benefit for the host.     

Climate change can alter competitive relationships between resident and migratory birds

Climate change could affect resource competition between resident and migratory bird species by changing the interval between their onsets of breeding or by altering their population densities. We studied interspecific nest-hole competition between resident great tits and migrant pied flycatchers in South-Western Finland over the past five decades (1953-2005). We found that appearance of fatal take-over trials, the cases where a pied flycatcher tried to take over a great tit nest but was killed by the tit, increased with a reduced interspecific laying date interval and with increasing densities of both tits and flycatchers. The probability of pied flycatchers taking over great tit nests increased with the density of pied flycatchers. Laying dates of the great tit and pied flycatcher are affected by the temperatures of different time periods, and divergent changes in these temperatures could consequently modify their competitive interactions. Densities are a result of reproductive success and survival, which can be affected by separate climatic factors in the resident great tit and trans-Saharan migrant pied flycatcher. On these bases we conclude that climate change has a great potential to alter the competitive balance between these two species.     

Sexual imprinting and the origin of obligate brood parasitism in birds

We discuss two pathways along which obligate brood parasitism (OBP) may evolve and examine some of the critical steps that must be passed by letting great tits Parus major be reared by blue tits Parus caeruleus in a field experiment. The cross-fostered chicks survived well in blue tit nests, but their local recruitment and reproductive success was much lower than that of controls. The effect was strongest when great tits grew up with siblings of the host species rather than with conspecific siblings in blue tit nests. The low success seemed to be caused by misimprinting because the cross-fostered birds behaved like blue tits in several aspects (species association, alarm calls, and aggressive response by resident females to caged intruders). Some birds of both sexes were apparently so strongly imprinted that they did not attract or accept a social mate of their own species. We conclude that imprinting may be necessary for OBP to evolve in birds because the parasite must be attracted to the nests of the host species to add eggs and thereby continue the parasitic life cycle. However, strong imprinting may also prevent OBP from occurring if parasitic offspring seek a mate from the host species.     

Bird predation by tawny owls (Strix aluco L.) and its effect on the reproductive performance of tits

The density of great tit Parus major L. and blue tit Parus caeruleus L. was artificially increased by placing nest-box colonies for these species in the vicinity of the nests of breeding tawny owls during 1993–1997. Bird prey composition in the owl nests, the proportion of parents disappearing from the breeding tit populations and the reproductive performance of the widowed parents were analysed. The frequency of predation on tits by tawny owls was greater in areas where tit density had been artificially increased. Owls preyed more on tits during the feeding period of owlets than during the incubation period and more in years when snow covered the ground during the incubation period than when it did not. Mortality due to predation was male biased and more females lost their mates in populations breeding near tawny owl nests. Reproductive performance of the widowed parents was lower and their body weights were lighter at the end of the nestling period than those found in birds rearing youngs with their mates. Predation by owls increased the between-year turnover in the breeding tit population: widowed parents did not return to the nesting site for the next breeding season.     

Rival imprinting: interspecifically cross-fostered tits defend their territories against heterospecific intruders

Failure to recognize conspecifics in social interactions such as mate choice and aggressive encounters will often result in reduced fitness. Studies on mate choice show that the ability to recognize conspecifics as mates is not universally present at birth, but often needs to be learned. In contrast, little is known about the ontogeny of intrasexual species recognition. To test whether learning influences the recognition of sexual rivals, we compared the aggressive response towards intruders of interspecifically cross-fostered individuals and controls reared by conspecific parents. We simulated territorial intrusion by presenting either a caged individual or playback song near the nest of breeding pairs of great tits, Parus major, and blue tits, P. caeruleus. Great tits reared by blue tit parents responded much more to blue tit stimuli than did great tit controls, and furthermore showed stronger responses to blue tit stimuli than to those of their own species. Blue tits reared by great tits responded much more to great tit stimuli than did blue tit controls. In contrast, blue tits cross-fostered to coal tits, P. ater, did not respond more to coal tits than did blue tit controls. There was a species difference in the response to conspecifics: blue tits cross-fostered to great tits responded more to conspecifics than did cross-fostered great tits. The results were similar for males and females. We conclude that learning influences intrasexual species recognition in these tits. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Intraspecific Variation and Interspecific Differences in the Bacterial and Fungal Assemblages of Blue Tit (Cyanistes caeruleus) and Great Tit (Parus major) Nests

Although interest in the relationship between birds and microorganisms is increasing, few studies have compared nest microbial assemblages in wild passerines to determine variation within and between species. Culturing microorganisms from blue tit (Cyanistes caeruleus) and great tit (Parus major) nests from the same study site demonstrated diverse microbial communities with 32 bacterial and 13 fungal species being isolated. Dominant bacteria were Pseudomonas fluorescens, Pseudomonas putida, and Staphylococcus hyicus. Also common in the nests were the keratinolytic bacteria Pseudomonas stutzeri and Bacillus subtilis. Dominant fungi were Cladosporium herbarum and Epicoccum purpurascens. Aspergillus flavous, Microsporum gallinae, and Candida albicans (causative agents of avian aspergillosis, favus, and candidiasis, respectively) were present in 30%, 25%, and 10% of nests, respectively. Although there were no differences in nest mass or materials, bacterial (but not fungal) loads were significantly higher in blue tit nests. Microbial species also differed interspecifically. As regards potential pathogens, the prevalence of Enterobacter cloacae was higher in blue tit nests, while Pseudomonas aeruginosa—present in 30% of blue tit nests—was absent from great tit nests. The allergenic fungus Cladosporium cladosporioides was both more prevalent and abundant in great tit nests. Using discriminant function analysis (DFA), nests were classified to avian species with 100% accuracy using the complete microbial community. Partial DFA models were created using a reduced number of variables and compared using Akaike’s information criterion on the basis of model fit and parsimony. The best models classified unknown nests with 72.5–95% accuracy using a small subset of microbes (n = 1–8), which always included Pseudomonas agarici. This suggests that despite substantial intraspecific variation in nest microflora, there are significant interspecific differences—both in terms of individual microbes and the overall microbial community—even when host species are closely related, ecologically similar, sympatric, and construct very similar nests.     

Multiple responses to increasing spring temperatures in the breeding cycle of blue and great tits (Cyanistes caeruleus,Parus major)

Many bird species start laying their eggs earlier in response to increasing spring temperatures, but the causes of variation between and within species have not been fully explained. Moreover, synchronization of the nestling period with the food supply not only depends on first‐egg dates but also on additional reproductive parameters including laying interruptions, incubation time and nestling growth rate. We studied the breeding cycle of two sympatric and closely related species, the blue tit Cyanistes caeruleus and the great tit Parus major in a rich oak‐beech forest, and found that both advanced their mean first‐egg dates by 11–12 days over the last three decades. In addition, the time from first egg to fledging has shortened by 2–3 days, through a decrease in laying interruptions, incubation time (not statistically significant) and nestling development time. This decrease is correlated with a gradual increase of temperatures during laying, suggesting a major effect of the reduction in laying interruptions. In both species, the occurrence of second clutches has strongly decreased over time. As a consequence, the average time of fledging (all broods combined) has advanced by 15.4 and 18.6 days for blue and great tits, respectively, and variance in fledging dates has decreased by 70–75%. Indirect estimates of the food peak suggest that both species have maintained synchronization with the food supply. We found consistent selection for large clutch size, early laying and short nest time (laying to fledging), but no consistent changes in selection over time. Analyses of within‐individual variation show that most of the change can be explained by individual plasticity in laying date, fledging date and nest time. This study highlights the importance of studying all components of the reproductive cycle, including second clutches, in order to assess how natural populations respond to climate change.     

Effect of an ectoparasite on lay date, nest-site choice, desertion, and hatching success in the great tit (Pants major)

Ectoparasites are common in most bird species, but experimentalevidence of their effects on life-history traits is scarce.We investigated experimentally the effects of the hematophagoushen flea (Ceratophyllus gallinae) on timing of reproduction,nest-site choice, nest desertion, clutch size, and hatchingsuccess in the great tit (Parus major). When great tits wereoffered a choice on their territory between an infested anda parasite-free nest-box, they chose the one without parasites.When there was no choice, the great tits in a territory containingan infested nest-box delayed laying the clutch by 11 days ascompared with the birds that were offered a parasite-free nestingopportunity. The finding that there was no difference in phenotypictraits related to dominance between the birds nesting in infestedboxes and birds nesting in parasite-free boxes suggests thatthe delay is not imposed by social dominance. Nest desertionbetween laying and shortly after hatching was significandy higherin infested nests. There was no difference between infestedand parasite-free nests in clutch size, but hatching successand hence brood size at hatching were significantly smallerin infested nests. Nest-box studies of great tits have beenseminal in the development of evolutionary, ecological, andbehavioral theory, but recently a polemic has arisen in theliterature about the validity of the conclusions drawn fromnest-box studies where the naturally occurring, detrimentalectoparasites are eliminated by the routine removal of old nestsbetween breeding seasons. Our study suggests that this criticismis valid and that the evaluation of the effects of ectoparasitesmay improve our understanding of behavioral traits, life-historytraits, or population dynamics     

Leafing phenology and timing of egg laying in great tits Parus major and blue tits P. caeruleus

To evaluate the importance of tree leafing for the start of laying and clutch size of birds, we compared the breeding phenology of great tits Parus major and blue tits P. caeruleus between one coastal and two inland sites in the same geographical region. Because of the cooling influence of the sea, trees at the coastal site were known to initiate budburst about a week later than at the inland sites. During 5 years, breeding by the tits and the leaf phenology of birch Betula pendula , and oak Quercus robur were monitored. The leaf phenology of birch and oak explained a significant part of the between-year variation in the start of egg laying in blue and great tits, respectively. The tits started laying earlier at the sites with an early budburst, i.e. normally inland. However, leaf phenology was not an absolute cue to the start of laying, since blue tits laid earlier relative to leafing at the inland site than at the coastal site, and both tit species laid eggs earlier relative to leafing during late springs. In neither species was clutch size affected by leafing phenology. However, great tit females at the coastal site consistently produced fewer eggs than did those at the inland site. No such difference was found in the blue tits. Although leafing phenology may predict the start of laying in tits, other factors also influence its timing. These factors might include other cues, or differing life-history trade-offs depending on site or general climatic factors during the spring.     

Egg concealment is an antipredatory strategy in a cavity‐nesting bird

Birds have developed different behavioural strategies to reduce the risk of predation during the breeding period. Bird species that nest in the open often cover their eggs to decrease the risk of predators detecting the clutches. However, in cavity nesters, the potential functions of egg covering have not been explored despite some bird species that nest in cavities also covering their eggs as open nesters do. We analysed whether egg covering is an antipredatory behaviour in the blue tit (Cyanistes caeruleus). We simulated an increase in the perceived risk of predation at experimental nests by adding predator scent inside the nest boxes during the egg‐laying period, whilst adding lemon essence or water to control nest boxes. Birds exposed to predator chemical cues in the nest of experimental pairs more frequently covered their eggs than birds exposed to an odorous control. These results suggest that egg covering may have evolved as an antipredatory behaviour also in cavity nesters to reduce the risk of egg predation and thus increase reproductive success in birds.     

Blue tits (Cyanistes caeruleus) respond to an experimental change in the aromatic plant odour composition of their nest

Although the use of olfaction by birds is now widely recognised, the olfactory abilities of passerine birds remain poorly explored, for historical reasons. Several studies however suggest that passerines can perceive volatile compounds in several biologically relevant contexts. In Corsica, recent findings suggest that cavity-nesting blue tits may use volatile compounds in the context of nest building and maintenance. Although they build their nests mainly from moss, female blue tits also frequently incorporate fragments of several species of aromatic plants in the nest cup. In field experiments, breeding female blue tits altered their nest maintenance behaviour in response to experimental addition of aromatic plants in their nest. In aviary experiments, captive male blue tits could be trained to detect lavender odour from a distance. Here I report results from a field study aimed to test whether adult blue tits altered their chick-feeding behaviour after an experimental change in nest odour composition. I experimentally added fragments of aromatic plant species that differed from those brought in the nests before the start of the experiment in a set of experimental nests and added moss, the basic nest material, in a set of control nests. Both male and female blue tits hesitated significantly longer entering the nest cavity after addition of new aromatic plant fragments, as compared to moss addition. This response was especially observed during the first visit following the experimental change in nest plant composition. Nest composition treatment had no effect on the time spent in the nest. This study demonstrates that free-ranging blue tits detect changes in nest odour from outside the nest cavity.     

Intraspecific nest parasitism and anti-parasite behavior in the grey starling,Sturnus cineraceus

I studied intraspecific nest parasitism in the grey starlingSturnus cineraceus in 1992 and 1993. The population in this study consisted of 290 nests (157 nests in 1992 and 133 nests in 1993) in which the clutches were completed before May 10 in the year studied. Twenty-nine nests in 1992 and 32 nests in 1993 contained at least 1 parasitic egg. Hatching success per nest of parasitized nests was slightly higher than that of non-parasitized nests. However, fledging success per nest of parasitized nests was significantly lower than that of non-parasitized nests. Thus parasitism appeared to reduce the reproductive success of hosts. Hosts exhibited a few behaviors that minimized the potential cost of brood parasitism. These behaviors included throwing out the parasitic egg and nest guarding. Hosts threw out parasitic eggs before the onset of laying, but they never did so to parasitic eggs laid after that period. The nest guarding level was low during the hosts’ laying periods, and one observed nest was parasitized during this time. Thus, nest-guarding behavior was not effective as an anti-parasite behavior. Grey starlings do not appear to adopt strategies effective in reducing parasitism.     

Influence of radar radiation on breeding biology of tits (Parus sp.)

The aim of the present study is to observe the influence of long-term exposure to radar radiation on breeding biology of tits (Parus sp.), living and building nests around a military radar station, emitting pulse-modulated microwave radiation of 1,200-3,000 MHz. Two series of 36 nest-boxes each were located on the radar station area. Measurements of exposure were performed separately for each nest-box. Average power density (P(av), W/m(2)) and dose of exposure (W/m(2) x h) were recorded for each nest-box during 45 days. Control nest-boxes (N = 42) were located in other part of the same forests, free from radar radiation. The assessment of effects of radar exposure on breeding biology of tits included number of inhabited nest-boxes, number of eggs, and nestlings in the nest (Why not chick mortality?). Experimental nest-boxes were either exposed to relatively high levels of radiation (2.0-5.0 W/m(2), mean 3.41 +/- 1.38 W/m(2)) or an intermediate level of radiation that ranged from 0.1-2.0 W/m(2) (mean 1.12 +/- 0.84 W/m(2)). For control nest-boxes the exposure ranged from 0.001-0.01 W/m(2) (mean 0.0062 +/- 0.0007 W/m(2)). Only blue or great tits occupied all nest-boxes, used in the experiment. The number of nesting blue tits was higher in nest-boxes located on the radar station area than in the control boxes. In contrast, control nest-boxes were inhabited mainly by great tits. On the radar station area, blue tits nested in high exposed nest-boxes (67,0%) and great tit occupied mainly these boxes, which were exposed to low-level radiation (62,5%), the difference being statistically significant (p < 0.01). No statistically significant differences in other parameters of breeding biology (number of eggs per box, number of nestling per box) were observed between tits occupying exposed and control nest boxes. Results of the present study show that radar radiation generally does not lead to decrease of number of nesting tits, but may cause shifts in tits species living around the radar station. (But is the microhabitat, apart from the radiation level, around each nest box more likely to attract one species of tit or another?).     

Revisiting the influence of aggressive interactions on the survival of the first‐laid egg in crested penguins (genus Eudyptes)

Crested penguins Eudyptes spp. have evolved a unique form of breeding in which the first of two eggs laid is much smaller than the second and has a higher likelihood of being lost during egg laying and incubation. In this study, we quantified aggressive behaviour in nesting Snares penguins and undertook an egg survival analysis to examine which factors influence egg loss. During 120 h of observation of 50 nests, we recorded a total of 300 aggressive events in which females were repeatedly pecked, bitten and beaten. Aggressive events lasted from less than a minute to up to 55 min (mean 4.6 ± 7.4 min). Single males were the aggressor in 75% of aggressive events and in 50.7% of aggressive events the aggressor was identified as a neighbouring, breeding male. A greater percentage of the small first eggs (34%) were lost than the large second eggs (4%). We found that egg mortality was influenced by 1) whether the other egg within a nest had hatched, 2) who was present at the nest (father, mother or both) and 3) the average duration of aggressive events on the nest. When one egg within a nest had hatched, the other egg had a vastly increased mortality risk irrespective of aggression. However, long, aggressive events directed towards females after their partners had gone foraging, also increased the probability of egg loss. We suggest that the prolonged nest attendance by breeding males well beyond egg laying is in response to the high frequency of aggressive behaviour during this time. Our data show that A‐egg losses occur due to intraspecific aggression in this species. Further research is needed to clarify whether aggressive behaviour in breeding crested penguins is modulated by elevated testosterone levels in the males and whether any reproductive benefits accrue to the aggressors.     

Aromatic herbs in Corsican blue tit nests: The ‘Potpourri’ hypothesis

This study reports that Corsican blue tit (Parus caeruleus ogliastrae) nests contain between one to five aromatic herb species between the onset of egg laying till the chicks’ finished growth 13 d after hatching. An herb removal experiment during the chick stage shows that blue tits bring fresh aromatic material 1–5 d after herb removal. Nests with a series of distinct odour classes easily perceived by humans have never been reported in birds. A new ‘Potpourri’ hypothesis is proposed that may explain the functional significance of this behaviour.