Food diversity and niche-overlap of sympatric tits (Great Tit,Parus major,Blue Tit,Cyanistes caeruleus and Coal Tit Periparus ater) in the Hyrcanian Plain forests

Abstract

This study was carried out between early summer 1998 and late summer 1999 in the Hyrcanian Plain forests, the southern Caspian Sea woodlands. In total, the gizzard contents of 241 specimens of the Great Tit (Parus major Linnaeus, 1758), Blue Tit (Cyanistes caeruleus Linnaeus, 1758), and Coal Tit (Periparus ater (Linnaeus, 1758)) were collected so that the percentages of grit, plants and animal materials could be estimated and the plant and invertebrate species consumed identified. The data were tested by a series of Mann-Whitney U, ANOVA and Kruskal-Wallis tests. To survey niche overlap of food materials between species and season, the symmetrical overlap index developed by Pianka (1973) was used. The most important invertebrates (maximum of percentage per season) in the diet of P. major were Coleoptera (36.5%) and Lepidoptera (33.6%), for C. caeruleus Coleoptera (34.5%) and Araneae (31.2%), and for Pe. ater Coleoptera (38.2%) and Araneae (35.6%). The most important plants consumed by P. major were Sorghum halepense (68.7%) and Ficus carica (65.6%), by C. caeruleus were F. carica (67.5%) and Alnus glutinosa (39.4%), and by Pe. ater were Lonicera spp. (82.9%). There was a greater degree of overlap and competition for animal food, but plant feeding overlap among the three tit species was low. Two species, P. major and C. caeruleus, showed high feeding overlap for animal items (0.92), whereas Pe. ater and C. caeruleus had no significant feeding overlap. The Coal Tit had a greater tendency to forage for vegetable matter and P. major had a greater tendency to forage for animal matter. Analysis of gizzard contents of P. major and Pe. ater showed that, despite the high degree of feeding overlap, the extent of plant materials consumed indicated no significant competition between the two species. Because Pe. ater is present mostly in autumn and winter, this study cannot establish any degree of competition and food-niche overlap during the breeding season between this and other tit species.     

Habitat and laying date of Great and Blue Tit Parus major and P. caeruleus

Using 19 year's data from nine study areas at Ghent we investigated the effect of habitat on laying date, and found it to be quite different for Great and Blue Tits. In Great Tits we found a clear gradient in laying dates from urban over suburban to rural habitats. Blue Tits laid earliest in a suburban habitat and latest in a rural habitat. All other areas, including urban, suburban and rural ones, formed one intermediate group. The overall average laying date for the Great Tit, 18.9 April, was slightly earlier than that of the Blue Tit, 19.4 April. In urban areas Great Tits laid significantly earlier than Blue Tits, in optimal oak habitat and in one suburban area Blue Tits laid earlier, whereas in the five remaining areas no significant differences between the two species were found.
We argue that the differences in laying dates, although correlated to temperature, are probably caused by differences in the timing of food availability. We suggest that differences in laying dates of Great and Blue Tits are caused by a different response to environmental variations through differences in feeding ecology.     

Vigilance of a Nest-building Female Blue Tit (Parus caeruleus)

Observations of a single nest-building female blue tit (Parus caeruleus) supported the hypothesis that increasing proximity of domestic cats (Felis domesticus) to the nest box increases time spent scanning the nest site area. A hypothesised decrease in time in the nest box with increased cat proximity was not confirmed. A reason for this is forwarded, and differences in time spent in/and in the vicinity of the nest box reflecting position of the nest visit in a bout, and threat posed by individual cats are discussed.     

Paraphyly of the Blue Tit (Parus caeruleus) suggested from cytochrome b sequences

The phylogenetic relationships of the Blue Tit-Azure Tit assemblage (genus Parus; Aves: Passeriformes) were studied using mitochondrial DNA sequences of 24 specimens representing seven subspecies from Eurasia and North Africa. Previous work based on comparative morphological and acoustic data suggested a division of the Blue Tit (Parus caeruleus) into two species. Our analyses clearly indicate that the Blue Tit represents a paraphyletic assemblage, including a European/Middle Asian clade that is the sister group to the Azure Tit (Parus cyanus) and a North African clade. The North African clade (teneriffae subspecies group) is a sister group to the European Blue Tit/Azure Tit clade. We suggest a division of the Blue Tit into two separate species, Eurasian Blue Tit (Parus caeruleus s. str.) and African Blue Tit (Parus teneriffae). However, our data give no support for assigning species rank to Parus cyanus flavipectus, a subspecies of the Azure Tit, as suggested by several authors on morphological grounds.     

Pair bond and breeding success in Blue Tits Parus caeruleus and Great Tits Parus major

Data from 939 nests of the Blue Tit Parus caeruleus and 1008 nests of the Great Tit P. major from nestboxes provided in superabundance in mixed forest study sites between 1976 and 2001 were analysed to examine the effects of mate retention on breeding success and the relationship between mate fidelity and site fidelity. Most birds retained their former partner (76% in Great Tits and 65% in Blue Tits). The probability of a pair divorcing was affected by male age in Great Tits, divorce being more likely in pairs with first‐year males. Great Tit pairs breeding together for a second season bred earlier, but had no higher breeding success than pairs breeding together for the first time. In Blue Tits laying date and start of incubation tended to be earlier in pairs breeding together for a second season, but hatching and fledging dates were not earlier than in other pairs. Great Tit pairs breeding together for two consecutive seasons bred earlier in the second season than in the first, but breeding success did not differ significantly between years. In both species, breeding performance did not differ between pairs that divorced after a season and pairs that stayed together. Thus breeding success did not determine whether a pair divorced or bred together again. Neither Blue Tits nor Great Tits improved their breeding performance through divorce. Blue Tit females even had fewer fledglings in the year after divorce than in the year before. Mate retention affected breeding site fidelity. Blue Tit females had greater breeding dispersal distances between consecutive years when re‐mating than when breeding again with the same mate. In Great Tits both males and females dispersed more when re‐mating than when retaining the former partner, suggesting that mate retention increased the chance of retaining the breeding site. In both species, breeding dispersal distances did not differ between pairs that divorced and pairs in which one mate disappeared. Because no major advantage of mate retention was evident, we suggest that mate retention evolved under different conditions than those found in study sites with high breeding densities and a superabundance of artificial nesting sites.     

Reproduction and survival of polygynous and monogamous Blue Tit Parus caeruleus

In a seven-year study of Blue Tits in optimal habitat near Antwerp, Belgium, 45 polygynous broods involving 22 males out of 667 successful first broods were found. In another 43 nests no male was found, although a major effort was made to trap all adults. The estimated proportion of polygynous males is 3.4%, if only confirmed cases are considered, but 10.8% if all possible cases are included. One male was paired simultaneously to three females.
Primary females (laying earliest in a triangle) were as successful as monogamous ones. Secondary (laying later in a triangle) and deserted females (nests in which no male was trapped), although still quite successful, raised fewer young and in one plot had a lower probability of recruiting offspring.
Both in males and females, the frequency of polygyny was independent of age. Adult survival did not differ between monogamous and polygynous males. Among females no effect of pairing status on survival was found in one plot, but in a second plot monogamous females survived better than others. It is concluded that in any study of Blue Tits in optimal habitat one could expect to find polygyny.     

Personality and Nest Defence in the Great Tit (Parus major)

A growing number of studies have shown that individuals differ consistently in a suite of correlated behavioural traits across various contexts and situations. Yet, most work on animal personalities has been performed under laboratory conditions and still little is known about the ecological significance of differences in personality in the wild, and the behavioural mechanisms underlying possible fitness consequences. In this study, we investigated individual differences in personality in relation to nest defence behaviour in wild great tits. Nest defence is an important aspect of parental care and involves a trade‐off between two fitness components (i.e. survival and reproduction). As a measure of personality we used exploratory behaviour in a novel environment as this has been shown to be correlated with several other behavioural traits including risk‐taking and aggression, two important behavioural components of nest defence. We found that the intensity of alarm calling towards a human intruder was positively associated with exploratory behaviour, while there was a negative association between exploration score and number of movements during nest defence. Thus, fast explorers are shown to respond more boldly towards predators in the field. More generally we show that individuals with different personalities vary in their anti‐predator and reproductive investment strategies.     

Low Cross-Sex Genetic Correlation in Carotenoid-Based Plumage Traits in the Blue Tit Nestlings (Cyanistes caeruleus)

In some bird species, both adult and juvenile individuals are often brightly coloured. It has been commonly assumed that identical plumage colouration present in both sexes results from strong intersexual genetic correlations in colour-related traits. Here, we aimed at testing this hypothesis in juvenile individuals and looked at genetic parameters describing carotenoid-based colouration of blue tit nestlings in a wild population. To separate genetic and environmental sources of phenotypic variation we performed a cross-fostering experiment. Our analyses confirmed the existence of sexual dichromatism in blue tit nestlings and revealed a significant, although low, genetic component of carotenoid-based colouration. However, genetic effects are expressed differently across sexes as indicated by low cross-sex genetic correlations (r_mf). Thus our results do not support the prediction of generally high r_mf and suggest that intersexual constraints on the evolution of colouration traits may be weaker than expected. We hypothesise that observed patterns of genetic correlations result from sex-specific selective pressures acting on nestling plumage colouration.     

Similarity of growth among Great tits (Parus major) and Blue tits (P. caeruleus)

Differences in morphology among species are proximately caused by changes in the ontogeny of individuals. It is therefore of importance to analyse possible differences in growth parameters among closely related species in order to understand what parameters are most and least likely, respectively, to change in evolution. In this paper I analyse growth in two closely related sympatric species, namely Great tit, Parus major, and Blue tit, P. caeruleus. The former is considerably larger than the latter in all external traits. The growth rates of the two species were found to be very similar for all traits, thus excluding differences in growth rate as a potential cause of evolutionary size changes. Offset of growth occurred at relatively similar times in the two species, excluding this factor as a major cause of the final size differences. However, size differences at hatching were pronounced and remained so throughout ontogeny, pointing to initial size (egg size or hatching size) as the target of factors promoting change. Bivariate allometric relations of traits vs. body size (mass) were similar between the two species at all ontogenetic stages. There was a high correlation among traits especially at intermediate age stages (5 and 8 days), but these correlations became weaker at older age and approached the low pattern of integration found in adults. All this suggests the operation of a general growth factor affecting all parts of the phenotype simultaneously, which has its major influence at the time of maximal growth. If closely related species in general have highly similar growth patterns, strong evolutionary allometry as found in many avian taxa is to be expected.     

Carotenoid-based coloration correlates with the hatching date of Blue Tit Cyanistes caeruleus nestlings

Carotenoid-based coloration occurs predominantly in adult birds, yet in some species from the family Paridae, this trait is also present at the nestling stage. One of the factors proposed to affect the expression of this trait in immature birds is hatching date. Here, using the avian tetrahedral colour space model, we examined the influence of hatching date on the breast carotenoid-based plumage coloration of the Blue Tit Cyanistes caeruleus nestlings. Because Blue Tits are sexually dichromatic, we also investigated the potential interaction between hatching date and sex that could arise from differences in condition dependence of this trait between males and females. We found a positive relationship between UV chroma of breast feathers and hatching date. The amount of UV reflectance is thought to be negatively related to carotenoid content in feathers. The observed increase of UV chroma through the breeding season might therefore be caused by a seasonal decline in the availability and quality of Lepidoptera larvae – the main source of carotenoids in food of the Tits. We also observed a sex difference in the relationship between brightness of breast feathers (achromatic, structural component) and hatching date, which in males was negative and in females not significant. Our study provides further evidence that the timing of breeding is related to the expression of nestling carotenoid-based coloration, a potentially meaningful element of offspring–parent communication, and suggests a sex-specific effect of hatching date on its structural component.     

Competition and egg-weight in the Great Tit Parus major

The mean weight of eggs in clutches of the Great Tit Parus major varies markedly between clutches. Eggs are lighter in years of high density of breeding pairs than in years of low density, but egg weight is also inversely correlated with the density of breeding Blue Tits Parus caeruleus .     

Evidence for cuckoldry in the Great Tit Parus major

We estimated the frequency of cuckoldry (the proportion of offspring resulting from extra-pair copulations) in Great Tits Parus major using heritability estimates based on the resemblance of offspring tarsus-length to that of their parents. Our results suggest that cuckoldry occurs and may be relatively common in certain years.     

Characterizing the mammalian hair present in Great Tit (Parus major) nests

Capsule Many birds are known to incorporate mammal hair into their nest lining, but the frequency with which they use hair from different mammals is unknown. We performed one of the first detailed examinations of mammalian hair from 54 Great Tit nests. We identified 5317 hairs belonging to 21 mammal species. Almost all of the examined nests contained hair from Roe Deer (Capreolus capreolus), but hair from Wild Boars (Sus scrofa), European Hares (Lepus europaeus) and Red Foxes (Vulpes vulpes) was also common.     

Factors affecting divorce in the Great Tit Parus major

In a population of the Great Tit Parus major in Sagamihara, the divorce rate was 17.8% over 10 years from 1984/85 to 1993/94. Most of the divorces occurred in the year following pair formation. There were no differences in breeding success between faithful and divorced pairs. The birds were divided into three groups in relation to their home ranges: residents, emigrants and migrants. The residents established their home ranges within their flock ranges and remained there for life. Non-residents (of which two sorts were identified) left their flock ranges to breed, but returned to their original flock ranges after breeding. Therefore, their home ranges were separated into breeding and previous flock ranges. Most of the faithful pairs consisted of residents whose home ranges overlapped extensively. The divorce rate was higher in the pairs consisting of residents and non-residents and in those of non-residents. Since the non-breeding home ranges of emigrants and migrants did not overlap with their mates' non-breeding home ranges, an important factor affecting divorce seems to be the non-overlap of non-breeding home ranges between both mates of pairs after breeding.     

Sexual dimorphism and growth trade‐offs in Blue Tit Cyanistes caeruleus nestlings

The outcome of sibling competition for food is often determined by variation in body size within the brood and involves trade‐offs; traits that enhance competitive ability within the nest may be developed at the expense of traits that enable effective flight at fledging, or vice versa. We quantified growth of skeletal, body mass and feather traits in male and female Blue Tit Cyanistes caeruleus nestlings. Males were significantly heavier, had longer tarsi and tended to have greater head–bill lengths than females, whereas females were similar to males in wing flight feather growth. These differences in growth may result from sexual differences in selection of the traits. Females are likely to prioritize feather growth to facilitate synchronized fledging with the rest of the brood, and to enhance escape from predators. We suggest that males are heavier and develop longer tarsi because body size is an important determinant of male reproductive success.     

Clutch Defence in Great Tit (Parus major) Pairs and the Concorde Fallacy

We conducted an experiment to test whether great tits (Parus major) base their decisions of clutch defence on past investment or future benefits. Results were obtained by manipulation of future benefits. Great tit pairs of an experimental group with reduced clutch-size and thus diminished benefits defended their offspring against a live raptor significantly less than a comparable, non-manipulated control group with the same amount of past investment (clutch-size, incubation stage, time of year). While in the females the difference between the two groups was obvious from the beginning of a trial, the difference in the males developed only during the course of a trial, suggesting that the male bases its response on the female's response deficit. A more onerous explanation of the female signaling the male the egg loss by using a language-like symbol is not supported by the data. The experiment thereby permits the conclusion that at least the great tit female avoids committing the Concorde fallacy in the strict sense.     

Differences in reproductive success and parental qualities between habitats in the Great Tit Parus major

Reproductive success and parental attributes of Great Tits Parus major nesting in good and poor habitats were compared to investigate whether they differed and, if so, whether parental or environmental variation contributed most to the differences.
Monitored over 3 years, clutches in mature woodland were started earlier, were larger and produced larger broods of heavier chicks than those in marginal habitats such as gardens and hedgerows. Fledging success was significantly higher in woodland in 2 of the 3 years although egg weights were lower. Parents nesting in different habitats were very similar in body-size although those in woodland were in better condition. The prey delivered to nestlings in woodland was of significantly higher quality than that delivered in marginal habitats. Furthermore, the heritability of body-size (tarsus-length) was higher in woodland, suggesting that the environment limits nestling growth in poorer habitats. Reproductive success was considerably lower in poor habitats and appears to be constrained primarily by environmental quality, although this may operate partly through its affect on parental body condition.     

Parasites as mediators of heterozygosity-fitness correlations in the Great Tit (Parus major)

Positive correlations between heterozygosity and fitness traits are frequently observed, and it has been hypothesized, but rarely tested experimentally, that parasites play a key role in mediating the heterozygosity-fitness association. We evaluated this hypothesis in a wild great tit (Parus major) population by testing the prediction that the heterozygosity-fitness association would appear in broods experimentally infested with a common ectoparasite, but not in parasite-free broods. We simultaneously assessed the effects of parental and offspring heterozygosity on nestling growth and found that body mass of nestlings close to independence, which is a strong predictor of post-fledging survival, increased significantly with nestling levels of heterozygosity in experimentally infested nests, but not in parasite-free nests. Heterozygosity level of the fathers also showed a significant positive correlation with offspring body mass under an experimental parasite load, whereas there was no correlation with the mothers' level of heterozygosity. Thus, our results indicate a key role for parasites as mediators of the heterozygosity-fitness correlations.