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These studies were conducted in 1999–2010 on the territory of the Zvenigorod Biological Station of Moscow State University (western Moscow suburbs, 55°44′ N, 36°51′ E). Birds (Parus ater) were caught by mist-nets. All the birds were banded and weighed, and their fat reserves were determined; then, the birds were released. A total of 85 individuals were caught. The standard metabolic rate and respiratory quotient (by the method of indirect calorimetry) were measured in 46 experiments with 16 birds. Two peaks were distinguished in the daily locomotor activity: a strongly pronounced daily peak (from 6 a.m. to 4 p.m.) and a weak evening peak (from 6 to 10 p.m.). The body mass did not change during the day. However, some trend for an increase in the mean body mass toward the middle and end of the day was noted. The fat reserves drastically changed during the day. The metabolic rate and respiratory quotient had a well-pronounced diurnal rhythm with minimal values at night (from 12 p.m. to 4 a.m.) and maximum values in the afternoon (from 12 a.m. to 4 p.m.). The total energy budget of Parus ater in the autumn-winter period, energy balance, and the maintenance of constant flying weight along with the dynamics of fat reserves are discussed.

Parus ater parisi Jouard, 1928 1928 Revue française d’Ornithologie 12: 368 (material), 373 (description). Now Parus ater ater Linnaeus, 1758. See Greenway et al., 1967: 91. Syntypes: MHNG 930.013, adult male, collected in France (Montlouis, Pyrénées-Orientales) on 20 January 1928. MHNG 930.014, adult female, collected in France (Vernet-les-Bains, Pyrénées-Orientales) on 23 February 1928. Both collected by H. Jouard. Additional material examined (not types): Jouard (1928, p. 368) examined a total of nine specimens (including the two syntypes). Because he explicitly defined a type series, we treat the seven specimens as additional material and not as syntypes (Art. 72.4.6 of the Code). Two of them are in the collection: MHNG 930.015, adult female (?), collected in France (Font- Romeu, Pyrénées-Orientales) on 28 February 1928; MHNG 930.043, adult male, collected in France (Montlouis, Pyrénées-Orientales) on 6 February 1928. All collected by H. Jouard., Published as part of Cibois, Alice, Vallotton, Laurent, Othman, Nagwa, Weber, Claude & Ruedi, Manuel, 2016, Type specimens of birds in the collections of the Natural History Museum of Geneva, pp. 269-282 in Revue suisse de Zoologie 123 (2) on page 275, DOI: 10.5281/zenodo.155300, {"references":["Jouard H. 1928. De la variabilite geographique de Parus ater dans l'Europe Occidentale. Revue francaise d'Ornithologie 12: 355 - 374.","Linnaeus C. von 1758. Systema Naturae. Editio Duodecima, Reformata (Ed. 10). Impensis Direct Laurentii Salvii, Holmiae.","Greenway J. C., Mayr E., Moreau R. E., Rand A. L., Snow D. W., Paynter R. A. Jr. 1967. Check-list of birds of the world: a continuation of the work of James L. Peters. Vol. 12. Harvard University Press, Cambridge."]}

Since in oscine birds song divergence often leads to premating reproductive isolation, we asked which factors drive the evolution of song in a widespread species, the coal tit Parus ater. Based on an almost taxon-complete phylogeny we correlated song divergence with morphometric (ecology), genetic (time), and geographic distances (separation). We found eight well-supported mitochondrial lineages, one of which consists of melanolophus (usually treated as a separate species) and whose relationships remain generally unresolved. Due to intense song variability even sophisticated sonametric analyses failed to separate sub-specific units. We tested for the role of song in reproductive isolation by playing songs from China and the Himalayas to Central European males. They recognize them as conspecific, but react to their playbacks less aggressively than to local songs. While morphological divergence is correlated with neither time nor separation, song divergence coincides more with geographic than genetic distance. This is mainly driven by differences along longitude and is especially prominent in the southern part of the species’ range, where six lineages persisted during the last glaciation while the north was subsequently re-occupied by two lineages.

The motivation of this study was to investigate some hitherto unknown information on the breeding ecology of the Stripe-breasted Tit (Parus fasciiventer) in Bwindi Impenetrable National Park, south-western Uganda. Parus fasciiventer is one of the least studied and endemic bird species restricted to the montane forests of the Albertine Rift. Regionally, it is classified as near-threatened. The study was carried out around the Institute of Tropical Forest Conservation Ruhija camp and the period of study was from January to June 2003. Data were generated through direct observation at the nest box sites of three active nests. Each of the nest boxes was monitored from the time of nest building to the time the chicks fledged. Results and comparative assessments from this study demonstrate that P. fasciiventer, compared with its temperate congeners like Great Tits (Parus major), Marsh Tits (Parus palustris), Crested Tits (Parus cristatus), Coal Tits (Parus ater) and Blue Tits (Parus caeruleus), raised small broods and had longer nestling period. The findings further revealed that the species is capable of raising more than one brood in a single breeding season and provide further evidence that it is a cooperative breeder. Parents participated equally in raising the young, an indication of pure parenting in the species. Resume La raison de cette etude etait de rechercher certaines informations jusqu’alors inconnues sur l’ecologie de la reproduction de la mesange a ventre strie, Parus fasciiventer, dans le Parc National de la Foret impenetrable de Bwindi, dans le sud-ouest de l’Ouganda. Parus fasciiventer est une des especes d’oiseaux endemiques les moins etudiees; elle se limite aux forets de montagne du Rift Albertin. Au niveau regional, elle est classee comme quasi menacee. L’etude s’est realisee autour du camp de Ruhija de l’Institute of Tropical Forest Conservation de janvier a juin 2003. Les donnees furent obtenues par des observations directes sur les sites de trois nichoirs actifs. Chaque nid a ete suivi depuis sa construction jusqu’a la mue des oisillons. Les resultats et des evaluations comparatives de cette etude ont montre que P. fasciiventer, comparea ses congeneres des regions temperees comme la mesange charbonniere Parus major, la mesange nonnette Parus palustris, la mesange huppee Parus cristatus, la mesange noire Parus ater et la mesange bleue Parus caeruleus, elevait de plus petites nichees et avait une plus longue duree de nidification. Les resultats ont aussi revele que cette espece est capable d’elever plus d’une nichee au cours d’une meme saison de reproduction et apportent de nouvelles preuves du fait que c’est une espece qui pratique la reproduction cooperative. Les parents participaient de facon equitable a l’elevage des jeunes, un signe de parente directe chez cette espece.

Observations on the bionomy of Diprion pallidum during the gradation in Carinthia After the overstep of culmination-point of Diprion pallidum-gradation in the attack-district on 6.000 ha (reduced surface 5.000 ha) 1971/1972 stood on record: bare trees 86.95%, intact trees 13.04%. Spin in the 1. gradation on the bare lateral-shoots 76.19%, on intact lateral-shoots 21.64%, on intact leafs 2.19%. Cocon-quantity on intact terminal-shoots: with 1 cocon 69.56%, with 2 21.73%, with 3 or 4 along with 4.34%; on intact lateral-shoots: with 1 cocon = 81.01%, with 2 = 14.43%, with 3 = 3.03%, with 4 or 5 = 0.75%; on half-bare lateral-shoots: with 1 cocon = 88.88%, with 2 = 7.4%, with 3 = 3.7%; on bare lateral-shoots: with 1 cocon = 67.31–78.03%, with 2 = 14.11–17.15%, with 3 = 5.49–9.11%, with 4 = 1.56–4.05%, with 5 = 0.39–1.35%, with 6 = 0.60% and with 9 = 0.39–0.60%. The collective spin is not regular. Colour of cocons: faint dark greyish. The position of head in the cocon on shoots: upwards 93.70%, downwards 5.59%, lateral 0.71%. Parasitism in the cocons: 10.06%, predatory on the cocons 1.39%; the singbirds, especially great tit (Parus major) and coal tit (Parus ater) where feeding their nestlings in the spring with the hibernate cocons in the tree-crown. Beginning of flight: in the forest-country 1. April, in the laboratory 7. April. In one population slipped out the ♂, ♂ within 6, the ♀, ♀ with 25 days; the slipp: morning 88.88% (especially between 10 and 12 o'clock), afternoon 11.11%. Index of sexus: ♀, ♀ 71.43%, ♂, ♂ 28.57%. Longaevitas: ♀, ♀ 3–17 (maximum about 9 days: 26.66%), ♂, ♂ 3–9 days (maximum about 5 days: 44.40%). ♀, ♀ double long live (8.2 days) as ♂, ♂ (4.7 days). Zusammenfassung Nach der uberschreitung des Kulminationspunktes der Diprion pallidum-Gradation in eincm Befallsgebiet von 6000 ha (reduzierte Befallsflache 5000 ha) 1971/72 waren auf kahl-gefressenen Platzen 86,95% Baumchen kahlgefressen und 13,05% intakt. Das Einspinnen der 1. Generation erfolgte auf Baumchen mit intakten und kahlgefressenen Lateraltrieben: zu 76,19% auf intakten, zu 21,64% auf kahlgefressenen Lateral-trieben und zu 2,16% auf intakten Nadeln, Kokonanzahl auf intakten Terminaltrieben: ein Kokon 69,56%, zwei Kokons 21,73%, drei oder vier je 4,34%; auf intakten Lateral-trieben: ein Kokon 81,01%, zwei Kokons 14,43%, drei 3,03%, vier oder funf je 0,75%; auf halbgefressenen Lateraltrieben: ein Kokon 88,88%, zwei Kokons 7,4%, drei 3,7%; auf kahlgefressenen Lateraltrieben: ein Kokon 67,31–78,03%, zwei Kokons 14,11–17,15%, drei 5,49–9,11%, vier 1,56–4,05%, funf 0,39–1,35%, sechs 0,60% und neun 0,39–0,60%. Das kollektive Einspinnen ist nicht die Regel. Kokonfarbe: matt dunkelgrau. Kopflage beim Einspinnen: triebaufwarts 93,70%, triebabwarts 5,59%, lateral 0,71%. Parasitismus in den Kokons 10,06%, Raubertum auf Kokons 1,39%. Singvogel, vornehmlich Kohlmeise (Parus major) und Tannenmeise (Parus ater), futterten im Fruhjahr ihre Nestlinge mit den in den Baumkronen uberwinterten Kokons der 1. Generation. Flugzeitbeginn: im Freiland 1. April, im Laboratorium 7. April. In einer Population schlupfen die ♂, ♂ innerhalb von 6 Tagen, die ♀, ♀ innerhalb von 25 Tagen. Tageszeit des Ausschlupfens: vormittags 88,88% (vornehmlich zwischen 10 und 12 Uhr), nachmittags 11,11%. Sexualindex: ♀, ♀ 71,43%, ♂, ♂ 28,57%. Longaevitas: ♀, ♀ 3–17 Tage (hochste Individuenzahl bei 9 Tagen: 26,66%), ♂, ♂ 3–9 Tage (hochste Individuenzahl bei 5 Tagen: 44,40%); ♀, ♀ leben doppelt so lange wie ♂, ♂ (8,2 Tage zu 4,7 Tagen).

Literatur Verschiedene Insekten wurden auf ihre akute orale Toxizitat an Nestlingen der Meisenart Parus ater (Tannenmeise) und Adulten des Haussperlings Passer domesticus uberpruft. Phosphamidon war gegenuber Nestlingen und Adulten toxisch, die Wirkung auf letzte war starker. uberlebende Indi-viduen zeigten eine schnelle und vollstandige Erholung nach der Krisis. Dimethoate hat eine ahnliche Toxizitat, jedoch konnte keine Erholung nach starken Vergiftungssymptomen beobachtet werden. γ-BHC steht zwischen den beiden vorher erwahnten Verbindungen, und DDT wurde in den an-gewandten Dosen symptomlos ertragen.

Climate change has been shown to affect the phenology of many organisms, but interestingly these shifts are often unequal across trophic levels, causing a mismatch between the phenology of organisms and their food.We consider two alternative hypotheses: consumers are constrained to adjust sufficiently to the lower trophic level, or prey species react more strongly than their predators to reduce predation. We discuss both hypotheses with our analyses of changes in phenology across four trophic levels: tree budburst, peak biomass of herbivorous caterpillars, breeding phenology of four insectivorous bird species and an avian predator.In our long-term study, we show that between 1988 and 2005, budburst advanced (not significantly) with 0.17 d yr(-1), while between 1985 and 2005 both caterpillars (0.75 d year(-1)) and the hatching date of the passerine species (range for four species: 0.36-0.50 d year(-1)) have advanced, whereas raptor hatching dates showed no trend.The caterpillar peak date was closely correlated with budburst date, as were the passerine hatching dates with the peak caterpillar biomass date. In all these cases, however, the slopes were significantly less than unity, showing that the response of the consumers is weaker than that of their food. This was also true for the avian predator, for which hatching dates were not correlated with the peak availability of fledgling passerines. As a result, the match between food demand and availability deteriorated over time for both the passerines and the avian predators.These results could equally well be explained by consumers' insufficient responses as a consequence of constraints in adapting to climate change, or by them trying to escape predation from a higher trophic level, or both. Selection on phenology could thus be both from matches of phenology with higher and lower levels, and quantifying these can shed new light on why some organisms do adjust their phenology to climate change, while others do not.

The hypothesis that females of socially monogamous species obtain indirect benefits (good or compatible genes) from extra-pair mating behaviour has received enormous attention but much less generally accepted support. Here we ask whether selection for adult survival and fecundity or sexual selection contribute to indirect selection of the extra-pair mating behaviour in socially monogamous coal tits ( Periparus ater ). We tracked locally recruited individuals with known paternity status through their lives predicting that the extra-pair offspring (EPO) would outperform the within-pair offspring (WPO). No differences between the WPO and EPO recruits were detected in lifespan or age of first reproduction. However, the male WPO had a higher lifetime number of broods and higher lifetime number of social offspring compared with male EPO recruits, while no such differences were evident for female recruits. Male EPO recruits did not compensate for their lower social reproductive success by higher fertilization success within their social pair bonds. Thus, our results do not support the idea that enhanced adult survival, fecundity or within-pair fertilization success are manifestations of the genetic benefits of extra-pair matings. But we emphasize that a crucial fitness component, the extra-pair fertilization success of male recruits, has yet to be taken into account to fully appreciate the fitness consequences of extra-pair matings.

Many animals use hoarding as a long-term strategy to ensure a food supply at times of shortage. Hoarders employ strategies that enhance their ability to relocate caches such as remembering where caches are located. Long-term scatterhoarders, whose caches have potentially high pilferage rates, should also hoard in a way to reduce potential cache pilferers’ ability to find caches. Previous studies have demonstrated that this could be achieved by hyperdispersing caches to reduce the foraging efficiency of pilferers. This study investigates whether coal tits (Parusater) indeed place their caches away from existing ones. In our experiment, birds hoarded food in 3 conditions: when caches from a previous storage session were still present, when caches from a previous storage session were not present anymore because the bird had retrieved them, and when caches from a previous storage session had been removed by the experimenter. We show that coal tits hoard away from existing caches and that they do not use cues from extant caches to do this. This evidence is consistent with the use of memory for the locations of previous caches when deciding where to place new caches. This finding has important implications for our understanding of the selective pressures that have shaped spatial memory in food-hoarding birds. Key words: cache dispersal, coal tits, cognitive ecology, food-hoarding birds, Parus ater. [Behav Ecol]

The benefits of extrapair mating behaviour for females of socially monogamous bird species are unclear despite substantial research effort. The genetic compatibility hypothesis proposes genetic benefits such that females paired to social mates of low genetic compatibility avoid or diminish negative fitness consequences by mating with a more compatible extrapair mate, resulting in offspring of higher genetic quality. Furthermore, within the context of inbreeding depression observed in natural populations, a high degree of overall genetic similarity between social pair mates may be regarded as a special case of genetic incompatibility. We tested the hypothesis that female extrapair matings represent an adaptive behavioural response to avoid negative consequences of being paired to a genetically similar social pair male in the coal tit, Parus ater, a socially monogamous passerine with high rates of extrapair paternity. In contrast to what was predicted, we found no evidence for a positive association between the genetic similarity of social pair mates (measured as band-sharing coefficients from multilocus DNA fingerprints) and the occurrence of extrapair paternity. Furthermore, the genetic similarity of the cuckolding female with its social mate was not higher when compared pairwise to that of its extrapair mate in 63 uniquely composed triplets. Finally, three parameters of reproductive success were not related to the genetic similarity of social pair mates. We conclude that avoiding the potentially negative fitness consequences of being paired to a genetically similar social pair mate did not select for and thus cannot maintain female extrapair mating behaviour in our study population. (c) 2004 The Association for tire Study of Animal Behaviour Published by Elsevier Ltd. All rights reserved.

The costs and benefits of dominance status have been investigated in the past and it has generally been reported that subdominant birds are at a nutritional disadvantage owing to their low dominance status. The nutritional condition of birds during winter can be important in determining their likelihood of survival. This is particularly so in small passerines that are sensitive to severe weather conditions. Ptilochronology is an accurate method used to produce a long-term estimate of body condition spanning the number of days that it takes to grow a new feather. Ptilochronology was used during this study to estimate the nutritional condition of Coal Tits Pares ater during one winter and how condition was affected by dominance status. Dominant Coal Tits produced poorer quality feathers, which they grew at a slower rate, than did subdominant conspecifics. This study highlights a nutritional cost to high dominance status that could have long-term consequences because the induced tail feathers will not be replaced for at least 5 months.

Molecular analyses of genetic parentage in wild bird populations demonstrate that extra-pair paternity (EPP) is common, even in socially monogamous species. After having identified extra-pair offspring (EPO), an important step is to uncover the identity of extra-pair sires (EPS). Studies relying on multilocus DNA fingerprinting face the problem that simple between-gel comparisons of fingerprint banding patterns are imprecise in assigning parentage across gels. To tackle this problem, we developed a method to identify EPS by means of standardized across-gel comparisons of multilocus DNA fingerprints and applied it in a socially monogamous bird species with high levels of EPP, the coal tit (Parus ater). Scanned DNA fingerprint gels and standard image-editing software allowed to screen and reliably identify EPS. Overall, EPP was successfully assigned to 32 different EPS for 143 out of 210 (68.1%) EPO detected. This corresponds well to extra-pair parentage assignment rates revealed by microsatellite-based studies in congeneric species. Our approach proves especially useful whenever multilocus DNA fingerprints for parentage exclusion exist while microsatellite markers are not available.

We studied the role of early developed phenotypic characters associated to body size (i.e., tarsus, wing, and tail length) together with body mass in the determination of the winter residence and r...

Volumetric studies in a range of animals (London taxi-drivers, polygynous male voles, nest-parasitic female cowbirds, and a number of food-storing birds) have shown that the size of the hippocampus, a brain region essential to learning and memory, is correlated with tasks involving an extra demand for spatial learning and memory. In this paper, we report the quantitative advantage that food storers gain from such an enlargement. Coal tits ( Parus ater ) a food-storing species, performed better than great tits ( Parus major ), a nonstoring species, on a task that assessed memory persistence but not on a task that assessed memory resolution or on one that tested memory capacity. These results show that the advantage to the food-storing species associated with an enlarged hippocampus is one of memory persistence.

The probability of perception of prey may depend on the previous experience of the predator, its expectations and its search tactics. In this study we investigated the role of the search area. Based on previous results and on known facts about foraging ecology, we hypothesised that crested tits ( Parus cristatus ) would perform better when searching larger areas for cryptic prey compared with coal tits ( Parus ater ) that have been found to search small areas for cryptic prey more successfully. In our experiments these two tit species were required to search for cryptic prey in large and small dishes. Both species detected cryptic prey less frequently in the larger dishes, however, crested tits performed better initially. Coal tits improved over time. Both species needed more time for detecting prey in large dishes, and crested tits searched longer in small dishes than coal tits. Giving-up times, when no prey was found, also reflected these differences. The results suggest that the tits search with speciesspecifically set search areas and 'time windows'. It also appeared that coal tits are more flexible in adjusting search to a given situation.

Possible biases related to the use of recordings were assessed by comparing captures of Coal Tits Parus ater with mist nets in presence and absence of tape lures. Samples netted with and without tape lures differed in age composition but not in body size. Moreover, tape lured nets caught resident birds less frequently than expected. Group size of flocks captured with the use of recordings was larger than those captured without. Tape lured juveniles had higher levels of subcutaneous fat reserves than Coal Tits captured at random, whereas this difference was not detected in adults. Attraction of dispersing juveniles and habituation to the tape lure by more resident individuals seemed to explain the patterns observed. Caution and knowledge of the biases caused by decoys is recommended before designing bird population studies which rely on the capture of birds by this method.

Current theory predicts small birds should have a reduced body mass when daytime predation risk is high. However, the influence that nighttime predators might have on changes in body mass or daytime foraging behaviour has not been addressed. We therefore studied the effect of changes in perceived nocturnal predation risk on the body mass of captive coal tits. In a soundproof room, eight coal tits were housed in individual cages and an experiment was performed in which the birds were subjected to two treatments. First, morning and evening body mass was monitored following nights that were quiet. Second, these parameters were measured following nights when the call of a tawny owl had been played once per hour. Evening body mass was 3% greater on days following owl-disturbed nights, but morning body masses did not differ between treatments. To ensure this result was a response to the owl calls per se, and not a general response to increased disturbance, a second experiment was necessary. Here the coal tits were exposed hourly to the calls of a nightjar, a non-predatory nocturnal bird, but no increases in body mass were observed compared to quiet nights. We suggest the coal tits increased body mass in response to owl calls to offset increased nighttime energy expenditure in attentive behaviour., The study was funded by project PB95-0102-CO2-01. LMB was supported by a postdoctoral contract from Ministerio de Educación y Ciencia. S.J.L was supported by a Marie Curie Research Training Grant awarded under the Training and Mobility of Researchers Program by the European Commission Directorate General XII for Science, Research and Development.

Seven males and thirteen female Diplotriaena henryi Blanc 1919 were collected from body cavities (heart, sternum, and chest) of tits (one Parus major and three Parus ater) in plain woodland of Noor, in Northern Iran.

Sexual dimorphism of 2 plumage features and some morphological measurements was assessed in a sample of 140 museum skins and over 300 wild-trapped Coal Tits Parus ater britannicus. All the characters examined displayed highly significant sexual dimorphism, and have allowed the presentation of a recommended methodology which should allow over 90% of all British Coal Tits to be sexed correctly by fieldworkers. The validity of year-round sexing using characteristics of the throat feathering is proven by comparison with the results of a molecular DNA sexing technique.

Abstract. Two playback experiments were carried out to test the prediction on coal tits that singing rate could be used to assess the vigour of rival males, and that song length might be used in the same way, during singing interactions. Playbacks of songs with increasing rate inhibited the songs of responding birds, but induced more calls in response. The results suggested that a bout of singing with increasing rate is reacted to as a more serious threat than one with decreasing song output. The playback of long songs induced a greater number of calls in response, while fewer were produced during playback of shorter songs. The results suggest that a bout of long songs is responded to as a more serious threat than one of short songs.

Parus ater insularis Hellmayr Parus ater insularis Hellmayr, 1902: 36 (Surugano-kumi). Now Periparus ater insularis (Hellmayr, 1902). See Eck and Martens, 2006: 23–28, and Gosler and Clement, 2007: 722. LECTOTYPE: AMNH 681709, adult male, collected in Suruga Province (5 Suruga-nokumi, Morioka et al., 2005: 153), Japan, undated, by collectors for A. Owston (no. 215). From the Rothschild Collection. COMMENTS: In the original description, Hellmayr listed as types two specimens: a male in the Rothschild Collection from Suruga-no-kumi, no. 215, and one other specimen—a female, no. 2554 in the v. Tschusi Collection, from Musashi (Hondo) collected in November. Hartert (1920a: 444) listed the male adult from Suruga as the type, thereby designating it the lectotype. It bears a Rothschild type label. The second specimen listed by Hellmayr is a paralectotype in NMW (Morioka et al., 2005: 52). The only other Owston specimen in AMNH from Suruga is a female, also bearing the number 215, but not mentioned by Hellmayr. Both of these specimens bear an early Owston label without his name but a distinctive 3 inch by K inch cream cardboard rectangle with all four corners clipped off. One side bears what is probably a species number (215 in this case), the species name, and the sex of the specimen; the reverse has the locality., Published as part of Pachycephalidae, Aegithalidae, Remizidae, Paridae, Sittidae, Neosittidae, Certhiidae, Rhabdornithidae, Climacteridae, Dicaeidae, Pardalotidae, Nectariniidae, And & Lecroy, Mary, 2010, Type Specimens Of Birds In The American Museum Of Natural History Part 8. Passeriformes:, pp. 1-178 in Bulletin of the American Museum of Natural History 2010 (333) on page 71, {"references":["Hellmayr, C. E. 1902. Untersuchungen uber einige palaarctische Vogel. Ornithologisches Jahrbuch 13: 26 - 43.","Eck, S., and J. Martens. 2006. Systematic notes on Asian birds. 49. A preliminary review of the Aegithalidae, Remizidae and Paridae. Zoologische Mededelingen Leiden 80 - 5: 1 - 63, 3 pls.","Gosler, A. G., and P. Clement. 2007. Family Paridae (tits and chickadees). In J. del Hoyo, A. Elliott, and D. Christie (editors), Handbook of the birds of the world, volume 12: 662 - 750. Barcelona: Lynx Edicions, 816 pp, 56 pls., 436 photographs.","Morioka, H., E. C. Dickinson, T. Hiraoka, D. Allen, and T. Yamasaki. 2005. Types of Japanese birds. Tokyo: National Science Museum Monographs no. 28, 154 pp.","Hartert, E. 1920 a. Types of birds in the Tring Museum. B. Types in the general collection (contd.). Novitates Zoologicae 27: 425 - 505."]}

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Field experiments were designed to examine the effects of parental ability on clutch size, for the two sexes separately. To handicap parental ability we removed feathers from the wings and the tail of three tit species (Parus spp.) during the initial incubation period. We then removed their first clutches, and examined the effect of the handicap on the subsequent nesting. In Blue Tits (P. caeruleus), and Coal Tits (P. ater), handicapped females reduced the size of the repeat clutch more than control females. In Blue Tits handicapped females fed their young at a lower rate, lost more body mass during the nestling period, and at fledging time their broods were significantly smaller than those of controls. Their fledglings had a body mass, similar to the control fledglings. Handicapped female Great Tits (P. major) only reduced their clutch size a small amount; consequently, at fledging time, their body mass and those of their young were lower than for controls. The males of the three species of tits did not seem to increase parental investment when their mate was handicapped. When male Blue and Coal Tits were handicapped their mates did not reduce clutch size more than did those of the control group, despite a reduction in parental investment by handicapped males. However, breeding success was not significantly affected by handicapping males. We conclude that the clutch size determination of female tits is primarily based on their own ability to feed their young, and not on the quality of their mate. Male tits seem to have a lesser influence on breeding success than females, and males seem less willing or less able to increase their parental investment.

The performances of the coal tit (a food storer) and the great tit (a non-storer) were compared in two experiments in which the bird was rewarded in phase 2 of a trial for returning to sites where it had seen food in phase 1 (‘window-shopping’). In both experiments the differences in performance between the species were small. In the first experiment the storing species was better than the non-storer at returning to the site where it had seen a seed, and in the second experiment the storing species was better at discriminating in phase 2 between sites visited in phase 1 that contained a seed and sites visited in phase 1 that were empty. A third experiment demonstrated that the motivational variables of reward size and deprivation were unlikely to account for the difference between the species. The results might indicate that there are differences in the spatial memory of the storing and non-storing species, but a difference in perceptual discrimination cannot be excluded.

We compared food-storing tits' memory for the locations of items they had stored and of food they had only seen. The results are consistent with suggestions that memory for stored food is subserved by the same memory system as that for encountered food

In the majority of socially monogamous bird species, extra-pair paternity (EPP) has been shown to be common, while its frequency varies considerably among as well as within species. Our knowledge of factors affecting this variation, however, still remains comparatively poor. A thorough analysis of EPP patterns on an individual level may lead to a better understanding of the general diversity in EPP. Here, we analyse intra- and inter-annual as well as individual variation in the occurrence of EPP in first and second broods of coal tits (Parus ater) in three successive years. Based on a sample of 483 broods with 3559 offspring genotyped, we show that EPP was extraordinarily frequent in the study population, surpassing the values observed in other species of the genus Parus by far. On the population level, the rates of EPP were found to be remarkably similar between years, while there was pronounced intra-annual variation such that EPP rates increased significantly from first to second broods. Considering consecutive broods of individual females and males, the general occurrence of EPP was 'inconsistent' (i.e. EPP often affected only one of two broods), but the proportion of extra-pair young (EPY) per brood showed significant repeatability for both sexes in case of mate retention. When mate change occurred, repeatability collapsed, indicating that pair identity was more important in determining the repeatability of EPP than female or male identity alone. This was further supported by the fact that, besides female and male age, also their interaction (reflecting pair age combination) significantly predicted the proportion of EPY, though not in all breeding periods surveyed. Hence, the identity of the breeding pair (reflecting possible interactions of male and female characteristics) should be explicitly considered and accounted for in future studies investigating patterns of EPP on an individual level. Finally, we show that, depending on the boundary conditions, in principle, all coal tits experience EPP: When analysing three or more broods of the same individual, there were no completely 'faithful' females and also nearly no males, which were not cuckolded at least once.

Despite substantial research effort, the benefits of female extra-pair matings in socially monogamous bird species remain elusive. The “good genes” hypothesis assumes that females engage in extra-pair copulations with males of superior genetic quality compared to their respective social mate. Therefore, a negative association between the degree of cuckoldry and male survival is predicted, if genetic quality is phenotypically reflected by high viability. Furthermore, genetic sires of extra-pair offspring (EPO) should survive better than the social fathers they cuckolded. We tested these predictions in a nestbox population of the coal tit (Parus ater), a socially monogamous passerine with low breeding dispersal and high rates of extra-pair paternity (EPP). Based on 257 genotyped first broods of two consecutive years, we found no relationship between the incidence of EPP or the proportion of EPO within a given brood and male or female recapture probabilities. Furthermore, recapture rates did not differ between social and genetic fathers of EPO or males that did or did not appear as extra-pair sires in other broods. Our results were not affected by differential (short-range) breeding dispersal with respect to EPP or by other potentially confounding variables. Hence, they are not in accordance with the “good genes as viability genes” hypothesis.

Whether female birds choose extra-pair mating partners to obtain genetic fitness benefits is intensely debated. The most straightforward and crucial test of 'good genes' models of female extra-pair mating is the comparison of naturally 'cross-fostered' maternal half-siblings sharing the same rearing environment as any systematic differences in performance between the two categories of offspring phenotype can be attributed to differential paternal genetic contribution. We analysed local recruitment and first-year reproductive performance of maternal half-siblings in the coal tit (Parus ater), a passerine bird with high levels of extra-pair paternity. We provide a highly comprehensive measure of the long-term fitness consequences of female extra-pair matings based on a large sample of 736 within-pair offspring (WPO) and 368 extra-pair offspring (EPO) from 91 first and 55 second broods, from which 132 breeders recruited into the study population. In contrast to predictions derived from 'good genes' models, we found no differences in local recruitment and seven parameters of first-year reproductive performance when comparing WPO and EPO. These results question the universal validity of findings in other bird species supporting 'good genes' models, particularly as they are based on the best approximation to female fitness obtained so far.

In a dominance-structured flock, social status may determine priority of access to food. Birds of low social status may perceive present and future access to food as less predictable, and so have a higher risk of starvation, than birds of high social rank. Theoretical models predict that subordinate birds should carry larger fat reserves and incur higher mass-dependent costs than dominants. However, empirical tests of the assumptions of these models are still scarce and controversial. We investigated the effect of dominance rank on daily mass gain under conditions of fluctuating food availability in a laboratory experiment using four flocks of four coal tits (Parus ater) each. The same amount of food was delivered in two treatments, but in one treatment the food was offered at a constant rate between days (fixed treatment), while in the other treatment the daily food supply varied in an unpredictable sequence between days (variable treatment). All birds showed greater variance in body mass in the variable treatment than in the fixed treatment. Body mass within birds showed the same variability at dawn than at dusk in the fixed treatment, but less variability at dawn than at dusk in the variable treatment. This may be a mechanism to reduce the immediate risk of starvation at the beginning of the day, when fat reserves are at their lowest and the aggression between flock members when feeding highest. Subordinate birds were excluded from the feeders by dominants more often in the early morning than in the rest of the day, and they showed more variability in daily mass gain and body mass at dawn than dominant birds. These results support the hypothesis that subordinate birds have a reduced probability of surviving when food availability changes unexpectedly compared to dominants., This study was funded by DGCYT project PB95-0102-CO2-01 of Ministerio de Educación y Cultura (MEC, Spain)

During winter, small birds in temperate zones increase their internal fat reserves to combat harsh environmental conditions and the drastic reduction in predictability of resource access. High fat reserves provide some benefits related to starvation-risk reduction, but also imply some costs related to increased predation risk. In general, subordinate species should increase their level of fat reserves in response to the unpredictability of resource access when in competition with dominants. However, the magnitude of such reserve increases could differ between subordinate species if opportunity of access to resources varies interspecifically in relation to differences in niche breadth due to differences in morphology. We tested the hypothesis that subordinate species (Parus cristatus) with decreased access to food will accumulate more fat than subordinate species (Parus ater) with greater opportunities. Results supported the prediction. We found that crested tit, the species with lower ecological plasticity and thus less access to food under competition, stores more fat daily than the coal tit, the species with higher ecological plasticity owing to its ability to forage while upside down.

Current theory of risk-sensitive foraging predicts that foragers should choose feeding sites on the basis of variation in as well as mean reward rate when there is a shortfall in their food supply or a decrease in their energy budget. For a given mean reward delay, they should choose high variance feeding sites if they are running below energy requirement, but low variance sites if they are running above. It has been suggested that the smaller the animal size, the stronger the preference reversion between high and low variable feeding sites. Previous tests of the energy budget rule when there was time variability have used bird species heavier than 80 g. Hence we tested energy budget rule predictions with coal tits Parus ater, a bird of 9 g of body mass, foraging at two feeding sites with high or low variability in food delivering delay. We manipulated energy budgets by controlling air temperature in the laboratory. In one treatment (positive budget), individuals were allowed to eat at the level of their own ad-libitum daily consumption and the air temperature was set to 24°C, while for the other treatment (negative budget), temperature was set to 14°C, and food availability was limited to the maximum daily intake observed in the positive budget treatment. When air temperature was low, daily intake increased but body mass decreased. Birds were also less active in the low temperature treatment, hopping less times every day. Latency to peck decreased as well, pecking for food when it was available sooner than in the high temperature treatment. These results show that coal tits were living in a negative energy budget when air temperature was set to 14°C. Preference for the variable feeding site was greatest in the negative energy budget, as it was predicted by the energy budget rule. Contrary to the energy budget rule, coal tits consistently preferred the variable option also in the positive energy budget. Possible explanations for these results are explored, including alternative foraging models to the energy budget rule.

Several mechanisms can explain individual differences in foraging behaviour, such as variation in predation risk between patches, variation in the ability of individuals to detect or escape from predators, variation between individuals in their requirement for food, the quality and abundance of food in different patches, phenotypic variation giving rise to differences in resource use (exploitation hypothesis) and interference competition such as the exclusion of subordinate individuals by dominants. Subordinates can develop compensation mechanisms. One of these mechanisms is morphological differentiation. However not every change in morphology can account for the same variation in behaviour, since some morphologies can be ecologically more plastic than others (i.e. some morphs can exploit a broader niche than other morphs). Under controlled conditions in the Coal Tit Parus ater, we tested whether (1) differences in resource use were explained by either the exploitation hypothesis or by the interference hypothesis, and (2) the presumed costs of subordination can be reduced through different ecological plasticities associated with different morphologies. Our results support the interference hypothesis as there are no differences in hanging behaviour between dominants and subordinates when foraging solitarily; while in the presence of other individuals, we observed differences in foraging behaviour that varied with social status. Our results also show that body mass influenced foraging behaviour; lighter birds can exploit patches where hanging postures are needed more easily than heavier birds. Moreover, this relationship varied among individuals, as predicted by the ecological plasticity hypothesis. Lighter subordinate individuals used hanging postures more frequently than heavier ones, differentially reducing the costs of subordination. We propose that differences in the breadth of ecological niche due to differences in morphology can reduce the costs of subordination., This study was carried out with the logistic support of El Ventorrillo Field Station, and was funded by DGES project PB95-0103 to E.M. A.B. was supported by a CSIC-CAMMNCN postdoctoral grant. M.B. was supported by a PhD grant from El Ventorrillo. J.J. Sanz kindly provided us with the discriminant function for sexual dimorphism. A. Gosler, N. Metcalfe and an anonymous referee greatly improved an early draft of the manuscript.

Theoretical models predict that birds should decrease their body mass in response to increased predation risk because lighter birds take off faster and are more manoeuvrable. We studied the effect of predation risk by chasing coal tits in large outdoor aviaries thus simulating an attempt to capture them. With this increase in predation risk, both perceived and actual, coal tits lost significantly more weight than in a control situation when they were not pursued. This pattern was attributable to a smaller gain in weight only during the day; nocturnal weight did not change in relation to diurnal predation risk. The lower daily weight gain was not consistent with predictions from models of interrupted foraging, but was consistent with predictions from risk adjustment models. Moreover, there was no difference in weight gain over 2-h periods that included a 1-h fast and those in which feeding was ad libitum, suggesting that coal tits could easily regain their body mass after a predator had interrupted their feeding. Our results therefore suggest that pursuit by predators leads to a decrease in the body mass of small birds. Copyright 1999 The Association for the Study of Animal Behaviour.

We compared the ability of coal tits, Parus ater (a food-storing species), great tits, P. major, and blue tits, P. caeruleus (two nonstoring species) to remember spatial locations in a spatial delayed-matching-to-sample task. Presentation of a single sample image on a touch screen was followed by a choice phase containing two, three or four images, in which the bird had to choose the original image. Storers made more correct choices than did nonstorers. Performance was affected by the proximity of the distractors: both groups performed less well when distractors were close to the sample although storers were less affected by proximity of distractors than were nonstorers. Both groups made correct decisions sooner than errors. We conclude that the accuracy of spatial memory in food-storing birds is greater than that of nonstorers. Copyright 1999 The Association for the Study of Animal Behaviour.