Uncommon mechanism of egg incubation in the endemic Southern hermit crab Loxopagurus loxochelis: how is this phenomenon related to egg production?

The aim of the present work was to characterize the egg production of Loxopagurus loxochelis. A total of 71 ovigerous females were obtained, 28% of which were simultaneously incubating eggs at different developmental stages. This phenomenon can be the result of incomplete fertilization, or may represent a rapid gonadal cycle by this species in this area, which is, to our understanding, the best explanation of this phenomenon. Egg volume decreased 25.6% during the incubation period. The reproductive output based on dry and wet weight was 6.8 and 19.3%, respectively. Water was the prevailing component of the eggs, representing 86.0% of the total weight at initial stage, increasing to 95.1% at the final stage. Ash content increased at the same time as a decrease in the organic content occurred, indicating the consumption of yolk and absorption of salts from the water medium. In conclusion, we hypothesized that this population has the capacity to copulate and deposit another brood even before the release of the larvae from the previous one, intensifying the reproductive effort of these hermit crabs as a strategy of adaptation in a region considered the northern limit of the geographical distribution of this species.

Developmental characterization, function and regulation of a Laccase2 encoding gene in the honey bee, Apis mellifera (Hymenoptera, Apinae)

In insects, exoskeleton (cuticle) formation at each molt cycle includes complex biochemical pathways wherein the laccase enzymes (EC 1.10.3.2) may have a key role. We identified an Amlac2 gene that encodes a laccase2 in the honey bee, Apis mellifera, and investigated its function in exoskeleton differentiation. The Amlac2 gene consists of nine exons resulting in an ORE of 2193 nucleotides. The deduced translation product is a 731 amino acid protein of 81.5 kDa and a pl of 6.05. Amlac2 is highly expressed in the integument of pharate adults, and the expression precedes the onset of cuticle pigmentation and the intensification of sclerotization. In accordance with the temporal sequence of exoskeleton differentiation from anterior to posterior direction, the levels of Amlac2 transcript increase earlier in the thoracic than in the abdominal integument. The gene expression lasts even after the bees emerge from brood cells and begin activities in the nest, but declines after the transition to foraging stage, suggesting that maturation of the exoskeleton is completed at this stage. Post-transcriptional knockdown of Amlac2 gene expression resulted in structural abnormalities in the exoskeleton and drastically affected adult eclosion. By setting a ligature between the thorax and abdomen of early pupae we could delay the increase in hemolymph ecdysteroid levels in the abdomen. This severely impaired the increase in Amlac2 transcript levels and also the differentiation of the abdominal exoskeleton. Taken together, these results indicate that Amlac2 expression is controlled by ecdysteroids and has a critical role in the differentiation of the adult exoskeleton of honey bees. (C) 2010 Elsevier Ltd. All rights reserved.

Decision rules for egg recognition are related to functional roles and chemical cues in the queenless ant Dinoponera quadriceps

The capacity to distinguish colony members from strangers is a key component in social life. In social insects, this extends to the brood and involves discrimination of queen eggs. Chemical substances communicate colony affiliation for both adults and brood; thus, in theory, all colony members should be able to recognize fellow nestmates. In this study, we investigate the ability of Dinoponera quadriceps workers to discriminate nestmate and non-nestmate eggs based on cuticular hydrocarbon composition. We analyzed whether cuticular hydrocarbons present on the eggs provide cues of discrimination. The results show that egg recognition in D. quadriceps is related to both age and the functional role of workers. Brood care workers were able to distinguish nestmate from non-nestmate eggs, while callow and forager workers were unable to do so.

Nest site selection by a neotropical swarm-founding wasp: seasonal alternation of nest orientation

Nest orientation in social insects has been intensively studied in warmer and cooler climates, particularly in the northern hemisphere. Previous studies have consistently shown that species subjected to these climatic conditions prefer to select mostly southern locations where the nests can gain direct sunlight. However, very little is known on nest orientation in tropical and subtropical social insects. We studied nest orientations initiated by swarms throughout a year in a Brazilian swarm-founding wasp, Polybia paulista von Ihering (Hymenoptera: Polistinae). Swarms selected various orientations as nest sites, but there was a particular trend in that swarms in the winter period (May-August) preferred to build northward-facing nests. This preference is opposite from that of social wasps observed in the northern hemisphere. Colonies of this species can potentially last for many years with continuous nesting, but nesting activities of colonies during the winter are severely limited due to cool temperature and a shortened day length. Northward-facing nests are warmer through the gain of direct solar heat during the winter period; consequently, choosing northward-facing sites may be advantageous for swarms in terms of a shortened brood development and shortened time needed to increase metabolic rates during warm-up for flight.

Pollen substitutes increase honey bee haemolymph protein levels as much as or more than does pollen

Adequate substitutes for pollen are necessary for maintaining healthy bee colonies during periods of pollen dearth, but testing them objectively is both time consuming and expensive. We compared two commercial diets with bee collected pollen and acacia pod flour (used by beekeepers in some parts of Brazil) by measuring their effect on haemolymph protein contents of young bees exclusively fed on these diets, which is a fast and inexpensive assay. The commercial diets included a new, non-soy-based, pollen substitute diet (named Feed-Bee (R)) and a soy-based diet, named Bee-Pro (R). The diets were each given in patty form to groups of 100 Africanized honey bees in hoarding cages, maintained and fed from emergence until six days of age. Sucrose, in the form of sugar syrup, was used as a protein free control. Feed-Bee (R), Bee-Pro (R), pollen and acacia pod flour diets increased protein titers in the haemolymph by factors of 2.65, 2.51, 1.76 and 1.69, respectively, over protein titers in bees fed only sucrose solution. The bees fed Feed-Bee (R) and Bee-Pro (R) had their haemolymph significantly enriched in protein compared to the controls and those fed acacia pod flour and to titers slightly higher than those fed pollen. All four proteinaceous diets were significantly superior to sucrose alone.

Differential expression of hypoxia pathway genes in honey bee (Apis mellifera L.) caste development

Diphenism in social bees is essentially contingent on nutrient-induced cellular and systemic physiological responses resulting in divergent gene expression patterns. Analyses of juvenile hormone (JH) titers and functional genomics assays of the insulin-insulin-like signaling (IIS) pathway and its associated branch, target-of-rapamycin (TOR), revealed systemic responses underlying honey bee (Apis mellifera) caste development. Nevertheless, little attention has been paid to cellular metabolic responses. Following up earlier investigations showing major caste differences in oxidative metabolism and mitochondrial physiology, we herein identified honey bee homologs of hypoxia signaling factors, HIF alpha/Sima, HIF beta/Tango and PHD/Fatiga and we investigated their transcript levels throughout critical stages of larval development. Amsima, Amtango and Amfatiga showed correlated transcriptional activity, with two peaks of occurring in both queens and workers, the first one shortly after the last larval molt and the second during the cocoon-spinning phase. Transcript levels for the three genes were consistently higher in workers. As there is no evidence for major microenvironmental differences in oxygen levels within the brood nest area, this appears to be an inherent caste character. Quantitative PCR analyses on worker brain, ovary, and leg imaginal discs showed that these tissues differ in transcript levels. Being a highly conserved pathway and linked to IIS/TOR, the hypoxia gene expression pattern seen in honey bee larvae denotes that the hypoxia pathway has undergone a transformation, at least during larval development, from a response to environmental oxygen concentrations to an endogenous regulatory factor in the diphenic development of honey bee larvae. (C) 2010 Elsevier Ltd. All rights reserved.

Begging and food distribution in crimson rosella (Platycercus elegans) broods: why don't hungry chicks beg more?

Altricial nestlings solicit food by begging and engaging in scramble competition. Solicitation displays can thus signal both hunger and competitive ability. I examined nestling solicitation and parental responses in crimson rosellas (Platycercus elegans), a species in which parents engage in complex patterns of food allocation and appear to control the distribution of food. By manipulating the hunger of individual chicks and entire broods, I assessed how chick behaviours and parental food allocation varied with hatching rank, level of hunger, and intensity of nestling competition. Last-hatched chicks begged more than first-hatched chicks irrespective of individual hunger levels. The two parents combined fed individually hungry chicks more, but mothers and fathers varied in their responses to begging chicks: fathers fed last-hatched chicks in proportion to their begging intensity, whereas mothers fed chicks equally. Since fathers generally allocate more food to first-hatched chicks, fathers appear to use begging rates to adjust food allocation to non-preferred chicks within the brood. When I manipulated brood hunger levels, begging rates increased for first- and last-hatched chicks suggesting that chick begging rates are sensitive to the level of competition. This study shows that begging by rosella chicks does not correlate with hunger in a straightforward way and that the primary patterns of food allocation by parents art: not influenced by chick begging. Thus the benefits of increased begging may be limited for nestlings in this species.

Laying date and laying sequence influence the sex ratio of crimson rosella broods

We examine the patterns of sex allocation in crimson rosellas Platycercus elegans, a socially monogamous Australian parrot. Overall, 41.8% of nestlings were male, a significant female bias. However underlying this population-level bias were non-random patterns of sex allocation within broods. Broods produced early in the season were female-biased, but the proportion of males in a brood increased as the breeding season progressed. Female rosellas may obtain greater fitness benefits from early-fledging daughters than sons because daughters can breed as 1-year-olds whereas sons do not breed until they are at least 2 years old. Laying date and laying sequence also interacted to influence the sex ratio of eggs. The sex of early-laid eggs strongly followed the brood level pattern, whereas the sex of middle- and late-laid eggs did not change significantly as the season progressed. Nevertheless, late-laid eggs were very unlikely to be male at the end of the season. We argue these differing seasonal patterns reflect the relative costs and benefits to producing early-hatched males and females at different times of the season. Female rosellas appear to maximise the probability that daughters are able to breed early but to minimise competitive asymmetries within the brood. In particular, late-hatched male chicks are disadvantaged if their oldest sibling is male, explaining the dearth of broods containing late-hatched males at the end of the breeding season.

Extra-pair paternity and mate-guarding behaviour in the brown thornbill

We used multilocus DNA fingerprinting to assess parentage in the brown thornbill, Acanthiza pusilla, a socially monogamous Australian passerine. Extra-pair paternity was uncommon (6.2% of 178 offspring; 11.9% of 67 broods) and there was no evidence of intra-specific brood parasitism. Extra-pair paternity was limited because pairs spent more time together when females were fertile and males were able to evict intruding males before they could approach the female. Males were responsible for the close proximity of partners during the fertile period. Mate guarding therefore appears to be a male tactic aimed at preventing female infidelity rather than a cooperative behaviour of the pair aimed at preventing extra-pair copulations and/or female harassment. Females did not attempt to escape male guarding and were rarely observed to solicit copulations from intruding males. Nevertheless, females paired to smaller and younger males were more likely to cuckold their mates than females paired to larger and older males. This suggests that females may be more likely to seek or accept extra-pair matings when paired to small, young males or that old, large males are better at preventing their mates from engaging in extra-pair copulations. We found that male age but not male size influences mate-guarding behaviour. Older males tended to respond more aggressively to intruders. We therefore speculate that the relationship between male size/age and extra-pair paternity in brown thornbills may arise because female thornbills prefer large males as mates but are unable to express this preference as easily when paired to older males.

Sex allocation in the sexually monomorphic fairy martin

Offspring sex ratios were examined at the population and family level in the sexually monomorphic, socially monogamous fairy martin Petrochelidon ariel at five colony sites over a 4-year period (1993 1996). The sex of 465 nestlings from 169 broods % as determined using sex-specific PCR at the CHD locus. In accordance with predicted sex allocation patterns, population sex ratios at hatching and fledging did not differ from parity in an), year and the variance in brood sex ratios did not deviate from the binomial distribution, Further, brood sex ratio did not vary with hatching date during the season, brood number, brood size or colony size, The sex ratio or broods with extra-pair young did not differ from those without, while the sex ratio of broods fathered by males that gained extra-pair fertilizations did not differ from broods fathered by other males. Extra-pair chicks were as likely to be male as female. Neither the total number of feeding visits to the brood nor the relative feeding contribution by the sexes varied significantly with brood sex ratio. Brood sex ratios were also unrelated to paternal size, condition and breeding experience or maternal condition and breeding experience, However, contrary to our prediction, brood sex ratio was negatively correlated with maternal size. Generally, these results were consistent with our expectations that brood sex ratios would not vary with environmental factors or parental characteristics, and would not influence the level of parental provisioning. However, the finding that females with longer tarsi produced an excess of daughters is difficult to reconcile with our current understanding or fairy martin life history and breeding ecology.

Ascaris lumbricoides: reinfection in children bearing an established worm burden

To clarify the existance of reinfection in children bearing an established Ascaris lumbricoides infection, the authors evaluated the weight and the length of worms collected from ten cases of ascaridiasis. The worm burden was greater than 27 worms in nine cases. In seven cases the weight and the length of worms showed little variation, with unimodal distribution of values, suggesting that all the worms in each case belong to the same population, originated from a single brood infection or from successive infections over small time intervals. In three cases there was great variation in worm size indicated by the different values for the means and medians and by the high values for the standard deviation and coefficient of variation. In these three cases there was a bimodal distribution of worm's size suggesting the coexistance of two distinct populations: one, less numerous, composed of mature worms and the other, more numerous, composed ofimmature worms, in two cases, and two distinct populations of immature worms in one case. The existance of worms in different stages of maturation indicates that the less mature population was acquired when the mature worms were established in the gut. These results indicate that the reinfection with Ascaris in children bearing an established infection is not rare and resistance induced by a preexisting infection is not the rule.

Genetic breeding on the bee Melipona scutellaris (Apidae, Meliponinae)

A selection of queens of Melipona scutellaris through the most productive colonies were carried out during eight months in an orange honeyflow. Each of the colonies was evaluated by its production, that is, the gross weight production ( pollen, brood, geopropolis and wax of each hive). With this data a coefficient of repeatability was estimated by the intraclass correlation method, obtained r = 0.835 ± 0.071. The repeatibility is very high showing that the analysed data (production) is repeatable. Selection was then carried out using the regression coefficient of each colony and the respective production gain. Using these data the colonies were divided into three groups according to the method Vencovsky and Kerr (1982): a with the colonies of highest productivity, b of least productivity, and c of intermediary productivity. Colonies with the highest production (Group a) gave their queens to those of the lowest production (Group b) after their queens were taken out and killed; while those of intermediate (Group c) stayed with the same queens during the entire experiment both before and after the selection. The modifications in weight, that is, the genetic response was (R)= 7.98 gr per day which indicated a selection gain. The estimate of the realized herdability is twice the rate of the response to selection (R) by the selection differential (S2). That is then h²R=2(R/S2) then h²R= 0.166

Life history, biomass and production of Coronatella rectangula (Branchiopoda, Anomopoda, Chydoridae) from Minas Gerais

Species of Chydoridae provide the main diversity of the Cladocera. These organisms have been the subject of many studies; some dealing with their role in energy flow in aquatic ecosystems, since they inhabit the littoral region of water bodies which undergo the first impacts from anthropic activities. The aim of this study is to increase knowledge about the life cycle of Coronatella rectangula (Sars, 1861), a species found in several water bodies in the state of Minas Gerais, Brazil. The life cycle was determined by the culture of parthenogenetic females under controlled conditions in the laboratory. Experimental cultures were maintained in growth chambers at a constant temperature of 23.6(±0.5)ºC, through a 12 h light/12 h dark photoperiod. The organisms were fed on a suspension of Pseudokirchneriella subcapitata (Chlorophyceae) (10(5) cells.mL-1), and 0.02 mL of a mixed suspension of yeast and fish ration added per organism in equal proportions (1:1). Fifty parthenogenetic females with eggs were isolated and maintained until they produced neonates. Thirty of these neonates that had less than 24 hours were put in polypropylene bottles of 50 mL and kept in a germination chamber. These organisms were observed daily to obtain the parameters of the life cycle. Biomass and secondary production were also calculated. The embryonic development time of the specimens of C. rectangula was 1.68(±0.13) days and the time to reach primipara, was 2.48(±0.45) days. The mean fecundity of C. rectangula was two eggs/female/brood and the total number of eggs produced by the female during its life cycle was 27.8 eggs. During the whole life cycle, specimens of C. rectangula had a maximum of 14 seedlings, with two instars in the juvenile stage. Total biomass for C. rectangula was 36.66 µgDW.m-3(9.83 for the juvenile stage and 26.82 µgDW.m-3 for adults), and secondary production was 12.10 µgDW.m-3.day-1(8.34 µgDW.m-3.day-1 for egg production and 3.76 µgDW.m-3.day-1 for the juvenile stage).

Social chromosome variants differentially affect queen determination and the survival of workers in the fire ant Solenopsis invicta.

Intraspecific variation in social organization is common, yet the underlying causes are rarely known. An exception is the fire ant Solenopsis invicta in which the existence of two distinct forms of social colony organization is under the control of the two variants of a pair of social chromosomes, SB and Sb. Colonies containing exclusively SB/SB workers accept only one single queen and she must be SB/SB. By contrast, when colonies contain more than 10% of SB/Sb workers, they accept several queens but only SB/Sb queens. The variants of the social chromosome are associated with several additional important phenotypic differences, including the size, fecundity and dispersal strategies of queens, aggressiveness of workers, and sperm count in males. However, little is known about whether social chromosome variants affect fitness in other life stages. Here, we perform experiments to determine whether differential selection occurs during development and in adult workers. We find evidence that the Sb variant of the social chromosome increases the likelihood of female brood to develop into queens and that adult SB/Sb workers, the workers that cull SB/SB queens, are overrepresented in comparison to SB/SB workers. This demonstrates that supergenes such as the social chromosome can have complex effects on phenotypes at various stages of development.

Social evolution in the sweat bee Halictus scabiosae

The evolution of eusociality is one of the major evolutionary transitions of life on earth. For investigating the conditions and processes that are central to the origin of such integrated social organization, it is best to study organisms in which individuals have retained some flexibility in their reproductive strategies. Halictid bees are especially well suited as model organisms, because they show huge variation in social systems, both within and between species. In this thesis, I investigated female reproductive strategies in the primitively eusocial bee Halictus scabiosae, with a focus on the role of helpers, in order to get insight into the mechanisms governing the evolution and maintenance of eusociality. This species produces two broods per year. The females from the first brood can stay in the natal nest to help raise a second brood of males and gynes that become the next-generation foundresses in spring. We first compared the morphology of females from the two broods, as well as the nutrition they receive as larvae. Then we conducted a helper- removal experiment in the field to quantify the effects of the presence of helpers on colony survival and productivity. Finally, we reconstructed pedigree relationships of individuals using microsatellite markers in order to detect who reproduces in the nest and how much individuals drift between nests. We found that first brood females had a uniformly small size and low fat reserves, which may be caused by the restricted pollen and nectar provisions on which they develop. Colony survival and productivity was increased by the presence of a single helper, but the effect was small and mostly limited to small colonies. By inferring parentage within and across colonies, we could determine that females from the first brood rarely reproduce in their natal nests. However, foundresses are frequently replaced, and foundresses and females from the first brood occasionally move to and reproduce in foreign colonies. As a result, colonies often contain offspring from unrelated individuals, and the relatedness of females to the brood they rear is low. Overall, this thesis shows that the reproductive system of H. scabiosae is highly flexible. The production of helpers in the first brood is important for colony success and productivity, but there is a high colony failure rate and part of the first brood females drift and reproduce in foreign nests. Both foundresses and helpers appear to be constrained by harsh environmental conditions or social factors limiting reproduction and independent colony founding. - L'origine des insectes sociaux est un domaine fascinant pour la recherche. Pour comprendre les mécanismes et les conditions qui sont nécessaires pour l'évolution et le maintien de la vie en société, il est judicieux d'étudier des sociétés primitives d'insectes, où toutes les femelles ont conservé la capacité de se reproduire, même si leur rôle comportemental dans la colonie est d'aider sans se reproduire. Une des familles d'abeilles, les halictes, est idéale pour cette sorte de recherche, en raison de la grande variabilité dans leur comportement social. Dans cette thèse, j'ai étudié les stratégies reproductives des femelles de Halictus scabiosae pour mieux comprendre les mécanismes qui influencent l'évolution de la vie en société. Cette espèce produit deux cohortes de couvain par année. Les femelles du premier couvain restent souvent dans leur nid natal pour aider à élever le deuxième couvain, tandis que les femelles du deuxième couvain s'accouplent et hibernent pour devenir les nouvelles fondatrices au printemps suivant. Nous avons d'abord comparé la morphologie des femelles issues des deux couvains ainsi que leur nutrition au stade de larve. Puis, dans une expérience sur le terrain, nous avons quantifié l'apport d'une ouvrière pour la survie et la productivité de la colonie. Finalement, nous avons reconstruit des pedigrees en utilisant des marqueurs génétiques, pour savoir qui se reproduit dans la colonie et combien d'individus migrent entre colonies. Les résultats montrent que les femelles du premier couvain sont uniformément plus petites et plus maigres, ce qui indique que les fondatrices réduisent les provisions de nourriture pour leur premier couvain afin de les inciter à aider dans le nid au lieu de se reproduire indépendamment. Dans l'expérience sur le terrain, la survie et la productivité de la colonie augmentaient avec la présence d'une ouvrière additionnelle, mais l'effet était petit et limité aux petites colonies. Par la reconstruction de pedigrees, nous pouvions constater que les femelles du premier couvain pondent rarement dans leurs nids natals. Les fondatrices cependant sont souvent remplacées en cours de saison, et migrent fréquemment entre nids, tandis que les femelles du premier couvain pondent parfois des oeufs dans des nids étrangers. De ce fait, les colonies contiennent souvent des descendants d'individus étrangers, et la parenté génétique entre les femelles et le deuxième couvain est basse. Cette thèse démontre que le système reproductif de H. scabiosae est très flexible. La production d'ouvrières est importante pour la survie de la colonie et sa productivité, mais le taux d'échec est élevé et une partie des femelles du premier couvain migrent et pondent dans une colonie étrangère. Autant les fondatrices que les ouvrières semblent être contraintes par des conditions environnementales ou sociales qui limitent la reproduction et les nouvelles fondations de colonie. - Die Entstehung von sozialen Lebensformen ist eines der wichtigsten Entwicklungen in der Geschichte des Lebens. Um die Bedingungen oder Prozesse zu verstehen, welche bei der Entstehung und dem Erhalt von sozialen Merkmalen wichtig sind, sollte man Lebewesen untersuchen, welche je nach Umwelteinflüßen ihr soziales Verhalten flexibel ändern können. Furchenbienen (Halictidae) gehören dazu. Diese weisen nämlich ein breites Spektrum verschiedener sozialer Organisationsformen auf, oftmals sogar innerhalb der einzelnen Arten. In meiner Doktorarbeit befasste ich mich mit den Fortpflanzungsstrategien der Weibchen der Skabiosen-Furchenbiene Halictus scabiosae. Diese Art produziert zwei Brüten pro Jahr. Die Weibchen der ersten Brut bleiben dabei meist als Arbeiterinnen in ihrem Geburtsnest, wohingegen die Weibchen der zweiten Brut nach der Paarung überwintern, um im nächsten Frühling neue Kolonien zu gründen. In einem ersten Schritt verglichen wir die beiden Brüten bezüglich der Grösse und der Fettreserven der Weibchen sowie der Pollen-Nektar-Vorräte für die Larven. Dann bestimmten wir in einem Feldexperiment, wieviel eine zusätzliche Arbeiterin zum Überleben und zur Produktiviät der Kolonie beiträgt. Schliesslich ermittelten wir durch genetische Tests die Verwandtschaftsbeziehungen zwischen den Bienen, um herauszufinden, wer in den Kolonien tatsächlich die Eier legt und ob und wieviel die Bienen zwischen verschiedenen Nestern wandern. Wir stellten fest, dass die Weibchen von der ersten Brut einheitlich kleiner sind und weniger Fettreserven besitzen. Das weist daraufhin, dass die Nestgründerin die erste Brut unterernährt, um die Wahrscheinlichkeit zu erhöhen, dass diese Weibchen als Arbeiterinnen im Nest bleiben anstatt sich unabhängig fortzupflanzen. Schon eine einzelne zusätzliche Arbeiterin verbesserte die Überlebenschancen und Produktivität der Kolonie, der Effekt war allerdings klein und auf kleine Kolonien beschränkt. Die Verwandtschaftsanalysen zeigten, dass die Arbeiterinnen nur sehr selten ein Ei in ihr Geburtsnest legen. Erstaunlicherweise wanderten die Nestgründerinnen oft zwischen verschiedenen Nestern. Einige Weibchen der ersten Brut wanderten auch in ein fremdes Nest und produzierten dort Nachkommen. Diese Doktorarbeit zeigt, dass die Fortpflanzungsstrategien der Skabiosen-Furchenbiene tatsächlich sehr flexibel sind. Die Anwesenheit von Arbeiterinnen ist wichtig für das Überleben und die Produktivität der Kolonie. Die Misserfolgsraten bleiben jedoch hoch, und ein Teil der Weibchen der ersten Brut pflanzt sich in fremden Nestern fort. Sowohl die Nestgründerinnen als auch die Weibchen der ersten Brut scheinen durch Umweltsbedingungen oder durch soziale Faktoren in der Wahl ihrer Fortpflanzungs¬strategie eingeschränkt zu sein.