Breeding behaviour of Nuthatch Sitta europaea in relation to natural hole attributes in a primeval forest

Capsule Nuthatches used holes with strong walls, typically in live trees with entrances reduced by plastering, and ‘oversized’ interiors filled with bark flakes.

Aims To describe patterns of nest-site utilization by Nuthatches in primeval conditions, to examine the influence of various hole attributes on nesting success and to consider the adaptive value of nest-site choice.

Methods Observations of birds living in undisturbed conditions in a strictly protected part of the Bia?owie?a National Park (Poland) during 27 breeding seasons coupled with measurements of hole attributes and observations of nests' fate.

Results Tree species used for breeding differed among habitats. Holes were on average 14.0 m above the ground, in trees with girth at breast height 206 cm; both parameters varied strongly among tree species. They were situated mainly in tree trunks (76%), in living trees (89%), in conical knotholes (51%); woodpecker-made holes constituted 32%. Nuthatches bred in very large (mean bottom area: 325 cm²) holes, the preferred tree species (Maple Acer platanoides, Ash Fraxinus excelsior) had larger holes than other tree species. Nuthatches reduced entrances by ‘plastering’ to a mean size of 2.9 × 3.3 cm. They made the holes substantially shallower by filling them with bark flakes (mean depth to nest level: 10 cm), eggs were laid in depressions among flakes, far from the entrance (mean distance: 21 cm), their ‘nests’ occupied only a fraction of the hole area. Broods in holes with smaller entrances were the most successful. Eggs and small young were covered with bark during the absence of the female. Larger nestlings, when endangered, moved to the rear wall of the hole, where they stayed tightly pressed. These behaviour patterns reduced the risk of predation.

Conclusion The features of holes used by Nuthatches (combination of strong walls, small entrances and large bottoms with copious filling) are probably an evolutionary solution to the need to evade predators while keeping contents of the nest dry.     

Phylogenetic relationships of Eurasian Nuthatches (Sitta europaea Linnaeus, 1758) from the Alborz and Zagros Mountains,Iran

The Eurasian Nuthatch (Sitta europaea Linnaeus, 1758) is a resident bird in the Alborz and Zagros deciduous forests. We investigated the phylogenetic relationships and the taxonomic position of the Eurasian Nuthatch among other separated lineages of Eurasia with the help of blood samples collected from 19 individuals belonging to four populations in the Eastern and Western Alborz, as well as in the Northern and Southern Zagros forests. Genetic variation was then analysed using complete ND2 gene sequence (1041?bp) and phylogenetic analysis was done using Bayesian and maximum likelihood inference. Additionally, a median-joining algorithm was used to reveal the relationships among haplotypes. The results of the phylogenetic and haplo-type network analyses indicated that Eurasian Nuthatch haplotypes from the Alborz and Zagros Mountains form lineages distinct from the Asian, Caucasian and European haplotypes. Furthermore, an analysis of molecular variance (AMOVA) detected significant (P<0.001) genetic structure among the lineages. The Asian, European, Caucasian and Alborz lineages diverged from one another by an uncorrected genetic distance ranging from 0.029 to 0.039, while the Zagros lineage showed a slightly lower genetic divergence from the Caucasian lineage (0.006), but it did not share any haplotype with the Caucasian lineage. Thus, we suggest considering five Conservation Significant Units (CSU) for the Eurasian Nuthatches as the result of used dataset.     

Individual contributions to territory defence in a cooperative breeder: weighing up the benefits and costs

While investment in territory defence is expected to be influenced by its benefits, the additional role that costs may play is rarely considered. Here, we quantify both benefits and costs of repelling prospecting males in cooperative meerkats, and demonstrate that both are required to explain the substantial variation in individual contributions to the defence observed. Males benefit more from repelling prospectors than females, as males may lose dominance and be expelled during intrusions. Accordingly, males invest the most in repelling prospectors. We also show that males experience an associated cost in the form of reduced weight gain and, as such, heavier males contribute more to chasing prospectors. Finally, we show evidence of a cost not restricted to individuals engaged in chasing: both males and females reduce their contributions to feeding dependent pups when prospectors are present, resulting in a reduction in pup weight gain in this context. Males appear to adjust their contributions to chasing in light of this cost, chasing at lower rates when their group contains dependent young. Our findings support the view that investment in cooperative behaviours can be attributed to benefits and costs, and highlight the additional importance of considering trade-offs in investment between cooperative behaviours.     

Nuthatches (Sitta europaea: Aves) in forest fragments: demography of a patchy population

Breeding density, local survival and summer recruitment of nuthatches were evaluated in a population scattered over many small (1–30 ha) forest fragments, and compared with study plots inside larger forests. Since most young birds settle outside the fragment in which they were born this population corresponds to the “patchy population” concept implying that patterns in abundance may be better explained by processes at the population level than by metapopulation processes. Mean breeding density was c. 50% lower in fragments and decreased with regional isolation (distance from larger forests) but not with local isolation (distance to nearby fragments). Local survival of adults and established (i.e. territorial) 1st-year birds was not related to forest size or isolation. However, fewer young birds settled in summer in the fragments compared with a large forest. This difference probably reflects high mortality during the sensitive dispersal phase. Moreover, the observed number of recruits and their estimated survival was insufficient to maintain the breeding population, suggesting significant net immigration from larger forests. This “rescue effect” explains why densities are affected by regional, but not local isolation. Received: 14 December 1998 / Accepted: 1 March 1999     

Selective neutrality of mitochondrial ND2 sequences, phylogeography and species limits in Sitta europaea

Variation and geographic differentiation in mitochondrial DNA (mtDNA) was studied in the widespread and phenotypically variable Eurasian nuthatch (Sitta europaea). To assess whether sequences were evolving in a selectively neutral fashion, we used McDonald–Kreitman [Nature 351 (1991) 652] tests and a tree-based method, which suggested that although ND2 sequences are affected by natural selection against slightly deleterious alleles, the effects do not compromise phylogeographic inferences. Three phylogenetic species-level clades of nuthatches were discovered, corresponding to the Caucasus, southern Europe, and northern Europe plus Asia. Unimodal mismatch distributions within each clade suggest that populations have undergone recent growth. A westward range expansion was inferred from the geographic pattern in nucleotide diversity. Although samples were insufficient, it is possible that nuthatches in England and Japan are recently differentiated. Two specimens of the subspecies S. e. arctica formed a sister group to all other S. europaea, differing by ca. 10% uncorrected sequence divergence, pointing the need for additional study of this phenotypically distinct taxon. As with other species, mtDNA data support major phenotypic distinctions, but not subspecies.     

A prudent hoarder: effects of long-term hoarding in the European nuthatch, Sitta europaea

Although hoarding has been studied intensively for many decades,few studies have attempted to measure its actual fitness consequences.To fill this gap, we used ptilochronology, the growth of replacementfeathers as a measure of nutritional status, and thus a reflectionof starvation risk, of individual European nuthatches (Sittaeuropaea) during winter. We found that nuthatches are long-termhoarders, retrieving stored food up to at least 98 days afterstoring it. Long-term hoarding enhanced the nutritional statusof individual birds significantly because those individualsexperimentally given an opportunity to store seeds during autumnregrew plucked rectrices faster and larger than did controlbirds. Nuthatches used their stored seeds prudently by adjustingthe amounts they ate to meet their requirements, as determinedby ambient temperatures. Nuthatches refrained from using storedfood during periods of relatively benign conditions so theycould use the food during periods of more severe conditions.     

Impact of shrub canopies on understorey vegetation in western Eurasian tundra

Question: How does the composition and species richness of understorey vegetation associate with changing abundance of deciduous shrub canopies? What are the species‐specific associations between shrubs and understorey plants? Location: Tundra habitats along an over 1000‐km long range, spanning from NW Fennoscandia to the Yamal Peninsula in northwest Russia. Methods: The data from 758 vegetation sample plots from 12 sites comprised cover estimates of all plant species, including bryophytes and lichens, and canopy height of deciduous shrubs. The relationships between shrub volume and cover of plant groups and species richness of vegetation were investigated. In addition, species‐specific associations between understorey species and shrub volume were analysed. Results: Shrub abundance was shown to be associated with the composition of understorey vegetation, and the association patterns were consistent across the study sites. Increased forb cover was positively associated with shrub volume, whereas bryophyte, lichen, dwarf shrub and graminoid cover decreased in association with increasing volume of deciduous shrubs. The total species richness of vegetation declined with increasing shrub volume. Conclusions: The results suggest that an increase of shrubs – due to climatic warming or a decrease in grazing pressure – is likely to have strong effects on plant–plant interactions and lead to a decrease in the diversity of understorey vegetation.     

The day after: effects of vocal interactions on territory defence in nightingales

1. Models on territory acquisition and tenure predict that territorial animals benefit by adjusting territorial defence behaviour to previous challenges they had experienced within the socially complex environment of communication networks. 2. Here, we addressed such issues of social cognition by investigating persisting effects of vocal contests on territory defence behaviour in nightingales Luscinia megarhynchos (Brehm). 3. Using interactive playback during nocturnal song of subjects, a rival was simulated to countersing either aggressively (by song overlapping) or moderately (by song alternating) from outside the subjects' territory. Thereby, the time-specific singing strategy provided an experimentally controlled source of information on the motivation of an unfamiliar rival. 4. Expecting that nightingales integrate information with time, the same rival was simulated to return as a moderately singing intruder on the following morning. 5. The results show that the vigour with which male nightingales responded to the simulated intrusion of an opponent during the day depended on the nature of the nocturnal vocal interaction experienced several hours before. 6. Males that had received the song overlapping playback the preceding night approached the simulated intruder more quickly and closer and sang more songs near the loudspeaker than did males that had received a song alternating playback. 7. This adjustment of territory defence strategies depending on information from prior signalling experience suggests that integrating information with time plays an important part in territory defence by affecting a male's decision making in a communication network.     

Foraging behaviour of Nuthatches(Sitta europaea) in relation to the presence of mates and mixed flocks

Summary Nuthatches (Sitta europaea) usually forage in pairs outside the breeding season. I investigated whether foraging site selection in winter (tree species, height, substrate size) differed between sexes and whether this difference was related to the presence of mixed-species flocks. Foraging sites of pair members foraging together were highly correlated. In the rare cases when each used different tree species, males foraged significantly more often on the preferred oak trees than females. Males foraged significantly lower than females in some territories, but not in others. The reasons for this significant variation between pairs remains unexplained. Males also foraged lower than females in the company of mixed flocks. These differences are probably better explained as avoidance of the dominant male by the subordinate female, rather than specialization in resource use to avoid competition.

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Nahrungssuche beim Kleiber (Sitta europaea) im Paarverbund und in gemischten Schwärmen

Zusammenfassung Außerhalb der Brutzeit fouragieren Kleiber gewöhnlich in Paaren. Ich untersuchte, ob sich die Geschlechter in der Wahl des winterlichen Nahrungsortes (Baumart, Höhe, Astdicke) unterscheiden und ob solche Unterschiede durch die Anwesenheit von gemischten Schwärmen beeinflußt ist. Die Ortswahl der gemeinschaftlich auf Nahrungssuche gehenden Paarpartner war hoch korreliert. In den seltenen Fällen, in denen sich die Partner an verschiedenen Bäumen aufhielten, fanden sich die Männchen signifikant mehr an den der Art bevorzugten Eichen als die Weibchen. In einigen Revieren fouragierten die Männchen an signifikant niedrigeren Standorten als die Weibchen, doch war dies nicht in allen Revieren der Fall. Die Ursachen für diese Unterschiede sind nicht bekannt. Auch in gemischten Schwärmen erfolgte die Nahrungssuche der Männchen niedriger als die der Weibchen. Möglicherweise vermeiden die Weibchen so die Konkurrenz durch die dominanten Männchen.

     

Seasonal acclimatization of metabolism in Eurasian tree sparrows (Passer montanus)

Acclimatization to winter conditions is an essential prerequisite for survival of small passerines of the northern temperate zone. Changes in photoperiod, ambient temperature and food availability trigger seasonal acclimatization in physiology and behavior of many birds. In the present study, seasonal adjustments in several physiological, hormonal, and biochemical markers were examined in wild-captured Eurasian tree sparrows (Passer montanus) from the Heilongjiang Province in China. In winter sparrows had higher body mass and basal metabolic rate (BMR). Consistently, the dry mass of liver, heart, gizzard, small intestine, large intestine and total digestive tract were higher in winter than in that in summer. The contents of mitochondrial protein in liver, and state-4 respiration and cytochrome c oxidase (COX) activity in liver and muscle increased significantly in winter. Circulating level of serum triiodothyronine (T₃) was significantly higher in winter than in summer. Together, these data suggest that tree sparrows mainly coped with cold by enhancing thermogenic capacities through increased organ masses and heightened activity of respiratory enzymes activities. The results support the view that prominent winter increases in BMR are manifestations of winter acclimatization in tree sparrows and that seasonal variation in metabolism in sparrows is similar to that in other small temperate-wintering birds.     

Structure of theory in vegetation science

In order to help evaluate progress in vegetation science, we present an outline of the components of theory. Theory has both empirically and conceptually based components, and is tied together by a hierarchical framework. Theory is related to observable phenomena to further the most general goal of science, understanding. More specific tactics to advance understanding include explanation, generalization, and testing. Different components of theory permit the operation of specific tactics. Most components of theory are exemplified by one or more aspects of vegetation science. However, there are some conceptual and empirical components in need of further development. We conclude that vegetation science does in fact conform to the contemporary view of a developing theory, and that this theory can serve as a fruitful guide to research and application of concepts to the field.     

Post-glacial vegetation in the South Patagonian territory of the giant ground sloth, Mylodon

By comparison with the major modern plant communities of southern Patagonia, the changing post-glacial vegetation of the region is reconstructed from macrofossils derived from deposits in the Cueva del Mylodon, Ultima Esperanza, S. Chile. The oldest deposits, carbon-dated as c. 12,400 B.P., comprise dung of the extinct ground sloth Mylodon darwinii and show the animal to have fed entirely on Cyperaceae, Gramineae and species associated with these in the modern cool, wet sedge-grasslands of western Patagonia, communities which would be expected after the retreat of the ice. Overlying deposits of well-preserved, wind-blown leaf-litter permitted some quantitative analyses which show a rise of evergreen forest dominated by Nothofagus betuloides that reached its maximum c. 7000 B.P. and then declined as it was replaced by deciduous N. pumilio forest. A break in the fossil plant record, covering the human occupation about 5643 B.P. and the subsequent final appearance of Mylodon remains, is followed by evidence of mixed evergreen/deciduous forest in which Nothofagus pumilio gradually increases in importance to give deciduous forest some 2500 years ago similar to that found in the environs of the cave in historical times. Comparison with pollen diagrams shows that the modern climatic and vegetation difference between E. Fudgia and Ultima Eeperanza has persisted throughout the post-glacial period.     

Growth and reproductive costs of larval defence in the aposematic lepidopteran Pieris brassicae

1.?Utilization of plant secondary compounds for antipredator defence is common in immature herbivorous insects. Such defences may incur a cost to the animal, either in terms of survival, growth rate or in the reproductive success. 2.?A common defence in lepidopterans is the regurgitation of semi-digested material containing the defensive compounds of the food plant, a defence which has led to gut specialization in this order. Regurgitation is often swift in response to cuticular stimulation and deters predators from consuming or parasitizing the larva. The loss of food and other gut material seems likely to impact on fitness, but evidence is lacking. 3.?Here, we raised larvae of the common crop pest Pieris brassicae on commercial cabbage leaves, simulated predator attacks throughout the larval period, and measured life-history responses. 4.?We found that the probability of survival to pupation decreased with increasing frequency of attacks, but this was because of regurgitation rather than the stimulation itself. There was a growth cost to the defence such that the more regurgitant that individuals produced over the growth period, the smaller they were at pupation. 5.?The number of mature eggs in adult females was positively related to pupal mass, but this relationship was only found when individuals were not subjected to a high frequency of predator simulation. This suggests that there might be cryptic fitness costs to common defensive responses that are paid despite apparent growth rate being maintained. 6.?Our results demonstrate a clear life-history cost of an antipredator defence in a model pest species and show that under certain conditions, such as high predation threat, the expected relationship between female body size and potential fecundity can be disrupted.     

Effects of resource distribution on activity and territory defence in house mice, Mus domesticus

How much an animal invests in defending a territory depends, in part, on the quality, quantity and distribution of resources, but do animals target their investment in defence within the territory according to the location of different resources? We examined whether wild-caught male and female house mice increased aggression towards intruders of the same or opposite sex when in areas containing their food/water or nest site. Pairs of resident mice were established in enclosures consisting of two areas. In split enclosures, food and water were in one area and a nest site in the other, while in clumped enclosures all resources were in one area with no resources in the other. Residents of both sexes were quicker to attack intruders, and were more aggressive, in areas containing resources than in an area containing no resources. There was no difference in aggression between areas containing food/water, a nest site or both. The effect of resources on territorial defence was due largely to changes in aggression against intruders of the opposite sex. Residents, especially males, attacked intruders of their own sex more than those of the opposite sex. This bias was much stronger when no resources were present, owing to low aggression against opposite-sex intruders in the absence of resources. There were no differences in intruder aggression, and the relative size of the intruder (heavier or lighter than the resident) had no effect on resident aggression. Residents also spent less time out of their nest in clumped enclosures owing to low activity in the nonresource area.     

Development of simple sequence repeats (SSRs) in olive tree (Olea europaea L.)

We report the development of microsatellites or simple sequence repeats (SSRs) in the olive tree (Olea europaea L.). Forty three positive clones obtained by the screening of a GA-enriched genomic library were sequenced and primers were designed for 13 microsatellite loci. Five primer pairs amplified polymorphic products of the expected size range. SSR polymorphism was explored in a set of 46 olive cultivars. A total of 26 alleles were detected for the five loci. Heterozygosity ranged from 0.46 to 0.71. Ninety one per cent of the cultivars had unique multilocus genotypes. Microsatellite segregation was studied in a complex population from a cross between the commercial cultivars ’Leccino’ and ’Dolce Agogia’. Received: 3 February 2000 / Accepted: 21 March 2000     

Patterns of territory size and defence level in rural and urban tawny owl (Strix aluco) populations

The effect of different variables on the territory size and defence level of tawny owl ( Strix aluco L.) populations was studied in two habitats: mixed farmland (1976–1978) and town (1986–1988). Territory size was determined by plotting the position of territorial challenges (hoots) of males which were individually recognizable in most cases. No differences were found in size, defence, number of competitors and biomass between territories in farmland and in town, but territories in town were more fragmented and showed a larger nearest neighbour distance. Size of rural territories was adequately predicted both by some structural habitat features and food supplies, while size of urban territories was affected only by habitat structure and by years of territory occupancy. No effect on territory size by competing neighbours was found in either habitat, thus confirming that behavioural interactions between the owners of territories and potential settlers prevent a decrease of territory size.
Defence level of urban territories was affected by 'social'variables (nearest neighbour distance and years of occupancy), structural variables (percentage of poplar grove and diversity), and food supply, while in rural territories only structural variables (fragmentation or natural vegetation) and food supplies affected defence level.
This was due to different settlement tactics in the two areas, depending on the extent and quality of suitable habitat.     

Activation of central opioid receptors may induce gastric mucosal defence in the rat.

The effect of different opioid peptides on acidified ethanol- and indomethacin-induced gastric mucosal lesions was studied following intracerebroventricular (i.c.v.) administration. It was found that both the selective delta opioid receptor agonists--deltorphin II, [D-Ala(2), D-Leu(5)]-enkephalin (DADLE), [D-Pen(2), D-Pen(5)]-enkephalin (DPDPE)-, mu-opioid receptor agonist--[D-Ala(2), Phe(4), GlyT-ol]-enkephalin (DAGO)--as well as beta-endorphin inhibited the mucosal damage induced by both ethanol and indomethacin in pmolar dose range. In contrast, the gastric acid secretion was not influenced by DADLE in the dose of 16 nmol/rat and only a slight reduction (40%) was induced by DAGO in the dose of 1.9 nmol/rat. The protective effect was abolished in both ulcer models by bilateral cervical vagotomy. N(G)-nitro-L-arginine, an inhibitor of NO synthase, reduced the protective action in ethanol-induced, but not in indomethacin-induced gastric damage. The results suggest that activation of supraspinal delta and mu-opioid receptors resulted in inhibition of gastric mucosal lesions elicited by ethanol or indomethacin. The gastroprotective action is independent from the effect of opioids on acid secretion. Vagal nerve is involved in conveying the central action to the periphery. The mechanism of the gastroprotective effect of opioids is different in ethanol- and indomethacin-ulcer models: prostaglandins and nitric oxide are likely to be involved in the protective action of opioid peptides in ethanol-, but not in the indomethacin-ulcer model.