Managing the social environments of flamingos for reproductive success

Recent research has shown that large flamingo flocks are more likely to experience reproductive success than small flocks and that there is a positive relationship between behavioral stimulation from group displays and reproductive success. This study compares the group displays and reproductive success in a captive flock of Caribbean flamingos over 6 years at the National Zoological Park. Birds were added to the flock in two different years. In each case, the flock produced fertile eggs during the breeding season following the addition of new birds. Furthermore, the addition of new birds in 1988, and then again in 1991, significantly increased group display activity over the previous year when the flock size was lower. That the addition of birds stimulated group display activity suggests an alternative management strategy for promoting reproduction in captivity: separate birds in a flock and then reunite them prior to the breeding season. © 1994 Wiley-Liss, Inc.     

The timing of birds' breeding seasons: a review of experiments that manipulated timing of breeding

Reproductive success usually declines in the course of the season, which may be a direct effect of breeding time, an effect of quality (individuals with high phenotypic or environmental quality breeding early), or a combination of the two. Being able to distinguish between these possibilities is crucial when trying to understand individual variation in annual routines, for instance when to breed, moult and migrate. We review experiments with free-living birds performed to distinguish between the 'timing' and 'quality' hypothesis. 'Clean' manipulation of breeding time seems impossible, and we therefore discuss strong and weak points of different manipulation techniques. We find that the qualitative results were independent of manipulation technique (inducing replacement clutches versus cross-fostering early and late clutches). Given that the two techniques differ strongly in demands made on the birds, this suggests that potential experimental biases are limited. Overall, the evidence indicated that date and quality are both important, depending on fitness component and species, although evidence for the date hypothesis was found more frequently. We expected both effects to be prevalent, since only if date per se is important, does an incentive exist for high-quality birds to breed early. We discuss mechanisms mediating the seasonal decline in reproductive success, and distinguish between effects of absolute date and relative date, for instance timing relative to seasonal environmental fluctuations or conspecifics. The latter is important at least in some cases, suggesting that the optimal breeding time may be frequency dependent, but this has been little studied. A recurring pattern among cross-fostering studies was that delay experiments provided evidence for the quality hypothesis, while advance experiments provided evidence for the date hypothesis. This indicates that late pairs are constrained from producing a clutch earlier in the season, presumably by the fitness costs this would entail. This provides us with a paradox: evidence for the date hypothesis leads us to conclude that quality is important for the ability to breed early.     

Variable fledging age according to group size: trade-offs in a cooperatively breeding bird

Group living can provide individuals with several benefits, including cooperative vigilance and lower predation rates. Individuals in larger groups may be less vulnerable to predation due to dilution effects, efficient detection or greater ability to repel predators. Individuals in smaller groups may consequently employ alternative behavioural tactics to compensate for their greater vulnerability to predators. Here, we describe how pied babbler (Turdoides bicolor) fledging age varies with group size and the associated risk of nestling predation. Nestling predation is highest in smaller groups, but there is no effect of group size on fledgling predation. Consequently, small groups fledge young earlier, thereby reducing the risk of predation. However, there is a cost to this behaviour as younger fledglings are less mobile than older fledglings: they move shorter distances and are less likely to successfully reach the communal roost tree. The optimal age to fledge young appears to depend on the trade-off between reduced nestling predation and increased fledgling mobility. We suggest that such trade-offs may be common in species where group size critically affects individual survival and reproductive success.     

Age-dependent reproductive costs and the role of breeding skills in the Collared flycatcher

This study addressed whether there are any age‐related differences in reproductive costs. Of especial interest was whether young individuals increased their reproductive effort, and thereby their reproductive cost, as much as older birds when brood size was enlarged. To address these questions, a brood‐size manipulation experiment with reciprocal cross‐fostering of nestlings of young and middle‐aged female Collared flycatchers, Ficedula albicollis, was performed on the Swedish island of Gotland. Nestlings’ body mass, tarsus length and survival were recorded to estimate the parental ability and parental effort of the experimental female birds. Female survival and clutch size were recorded in the following years to estimate reproductive costs. We found that middle‐aged female flycatchers coped better with enlarged broods than younger females or invested more in reproduction. In the following year, young female birds that had raised enlarged broods laid smaller clutches than the females from all the other experimental groups. This result shows that the young female birds pay higher reproductive costs than the middle‐aged females. Both young and middle‐aged female flycatchers seemed to increase their reproductive effort when brood size was increased. However, such an increase resulted in higher reproductive costs for the young females. The difference in reproductive costs between birds of different ages is most likely a result of insufficient breeding skills of the young individuals.     

Costs and benefits of flocking in foraging white-fronted geese (Anser albifrons): effects of resource depletion

This study investigated the costs and benefits of flocking in white-fronted geese Anser albifrons foraging on rice grains in Japan. The time budgets of focal geese were recorded, and the effects of flock size on the proportions of time spent in vigilant and agonistic behaviour were tested. The results showed that the decline in vigilance level and consequent increase in foraging time were beneficial results of flocking whereas agonistic interactions, a potential cost of flocking, did not increase with increasing flock size. However, seasonal variation in flock size suggested that exploitative competition could be a cost of flocking; the sizes of flocks in spring, when resource depletion had progressed, were significantly reduced compared with those in autumn. An experimental increase in rice density resulted in a significant increase in flock size. We conclude that the flock size of foraging white-fronted geese is a result of compromise between a constant benefit of flocking (i.e. decline in vigilance level) and a cost of flocking varying with food abundance (i.e. exploitative competition).     

The role of group size and environmental factors on survival in a cooperatively breeding tropical passerine

1. Variation in survival, a major determinant of fitness, may be caused by individual or environmental characteristics. Furthermore, interactions between individuals may influence survival through the negative feedback effects of density dependence. Compared to species in temperate regions, we have little knowledge about population processes and variation in fitness in tropical bird species. 2. To investigate whether variation in survival could be explained by population size or climatic variables we used capture-recapture models in conjunction with a long-term data set from an island population of the territorial, cooperatively breeding Seychelles warbler (Acrocephalus sechellensis). The lack of migration out of the study population means that our results are not confounded by dispersal. 3. Annual survival was high, both for adults (84%) and juveniles (61%), and did not differ between the sexes. Although there was significant variation in survival between years, this variation could not be explained by overall population size or weather variables. 4. For territorial species, resource competition will work mainly on a local scale. The size of a territory and number of individuals living in it will therefore be a more appropriate measure of density than overall population density. Consequently, both an index of territory quality per individual (food availability) and local density, measured as group size, were included as individual covariates in our analyses. 5. Local density had a negative effect on survival; birds living in larger groups had lower survival probabilities than those living in small groups. Food availability did not affect survival. 6. Our study shows that, in a territorial species, although density-dependent effects might not be detectable at the population level they can be detected at the individual territory level - the scale at which individuals compete. These results will help to provide a better understanding of the small-scale processes involved in the dynamics of a population in general, but in particular in tropical species living in relatively stable environments.     

The evolution of cooperative breeding through group augmentation

Some individuals (helpers) in cooperatively breeding species provide alloparental care and often suppress their own reproduction. Kin selection is clearly an important explanation for such behaviour, but a possible alternative is group augmentation where individuals survive or reproduce better in large groups and where it therefore pays to recruit new members to the group. The evolutionary stability of group augmentation is currently disputed. We model evolutionarily stable helping strategies by following the dynamics of social groups with varying degrees of subordinate help. We also distinguish between passive augmentation, where a group member benefits from the mere presence of others, and active augmentation, where their presence as such is neutral or harmful, but where helping to recruit new group members may still be beneficial if they in turn actively provide help for the current reproductives ('delayed reciprocity'). The results show that group augmentation (either passive or active) can be evolutionarily stable and explain costly helping by non-reproductive subordinates, either alone or leading to elevated help levels when acting in concert with kin selection. Group augmentation can thus potentially explain the weak relationships between relatedness and helping behaviour that are observed in some cooperatively breeding species. In some cases, the superior mutualistic performance of cooperatively behaving groups can generate an incentive to stay and help which is strong enough to make ecological constraints unnecessary for explaining the stability of cooperatively breeding groups.     

Monitoring and managing genetic variation in group breeding populations without individual pedigrees

The genetic management of captive populations to conserve genetic variation is currently based on analyses of individual pedigrees to infer inbreeding and kinship coefficients and values of individuals as breeders. Such analyses require that individual pedigrees are known and individual pairing (mating) can be controlled. Many species in captivity, however, breed in groups due to various reasons, such as space constraints and fertility considerations for species living naturally in social groups, and thus have no pedigrees available for the traditional genetic analyses and management. In the absence of individual pedigree, such group breeding populations can still be genetically monitored, evaluated and managed by suitable population genetics models using population level information (such as census data). This article presents a simple genetic model of group breeding populations to demonstrate how to estimate the genetic variation maintained within and among populations and to optimise management based on these estimates. A numerical example is provided to illustrate the use of the proposed model. Some issues relevant to group breeding, such as the development and robustness evaluation of the population genetics model appropriate for a particular species under specific management and recording systems and the genetic monitoring with markers, are also briefly discussed.     

Variation in daily and seasonal foraging routines of non-breeding barnacle geese (Branta leucopsis): working harder does not overcome environmental constraints

This study describes the changes in stored body fat in Svalbard barnacle geese Branta leucopsis over the non-breeding period, and uses behavioural patterns to explain the variation in body fat stores. It was predicted that foraging effort would: (1) increase at low food densities; (2) increase when days were short; (3) decrease in smaller flocks; (4) be bimodally distributed throughout the day for long days, but constant for short days. Time constraints were found to be the major driving force behind foraging decisions during the shortest days of mid-winter, whereas food density was found to drive decisions during longer days. Field observations of fat stores showed that fat was rapidly accumulated at the start and end of the non-breeding period, but that fat stores remained constant during mid-winter. It is suggested that day length prevented a positive fat storage rate in mid-winter through the limitation of foraging time, even though foraging effort was high during this period. During a single day, evidence for a bimodal foraging routine was found, where feeding activity was concentrated in the early morning and late afternoon periods. This pattern was found in the full range of day lengths, which suggests that even for short days, feeding must be interrupted so that other essential activities can be conducted. It is concluded that the behavioural choices of barnacle geese were constrained by environmental conditions, and that these behavioural patterns allow the variation in fat stores to be explained.     

Classical heinrothian intimidation displays exhibited byGalago demidoff demidoff: A paradigm for pinpointing galagine species diversity

The existence of rhythmic micturation displays among bushbabies is well established but the function of such behavior has been a source of dispute, due largely to indiscriminate lumping of different kinds of phenomena. In this paper classical ethological evaluation of the rhythmic micturation intimidation displays regularly exhibited by dominant adult Galago demidoff demidoffallows comparison with other varieties of intra- /interspecific displaying and agression. These include (1) pure offensive behavior shared by both sexes; (2) defensively toned, antiintruder/antipredator displaying; (3) male- male rank demonstration confrontations; and (4) dominant- female, emotionally neutral self- advertising. The dominant- male, offensively toned displays couple the functions of rival intimidation and partner attraction; they vary between different species of bushbaby, and they represent an important taxonomic tool.     

Behavioral solutions to breeding cheetahs in captivity: Insights from the wild

Knowledge of cheetahs' behavior is increasingly seen as the key to solving the mystery of cheetahs' poor breeding performance in captivity. In the absence of zoos' maintaining systematic records of individuals' behavior during introductions, behavior of free-living animals can be informative. In the wild, most female cheetahs probably mate with males living in small groups or coalitions; thus, zoos may benefit from replicating these social conditions, provided injuries can be minimized. Relations between free-living coalition members are amicable, and escalated aggression was never witnessed during 4 years of observation. Some antagonism was seen in newly formed trios, although this had disappeared in longer established coalitions. Minor aggression occurred over carcasses, being greater at small and intermediate sized kills than at large ones. In the presence of females, mild intramale aggression was only seen within 1 of 7 coalitions. Free-living females showed similar rates of behavior in the presence of different numbers of males, aside from more frequent growling at large groups. These findings suggest that captive institutions should be less nervous about housing male cheetahs together and introducing females to groups of males for purposes of breeding. © 1993 Wiley-Liss, Inc.     

Japanese monkey group translocation: Effects on seasonal breeding

A 150-member troop of Japanese monkeys (Macaca fuscata)was translocated from its temperate native habitat, near Kyoto, Japan, to a 42-ha enclosure near Laredo, Texas, in February 1972. The seasonal timing and distribution of 430 births recorded over the period 1954–1971 were compared to those of 186 births recorded in Texas from 1973 to 1979. Despite striking climatological and environmental differences between the pre- and the postranslocation sites, the timing of the birth season remaines unchanged, although the distribution of births was altered. These findings, considered in light of other published data on the seasonality of mating in macaques, suggest that a phenomenon akin to social drift may account for much of the intertroop variation that occurs.     

Truppgr??en bei weidenden Bl??- und Saatg?nsen(Anser albifrons, A. fabalis) an der Unteren Mittelelbe und ihr Einflu? auf Fluchtdistanz und Zeitbudget

Zusammenfassung  In den Jahren 1994 bis 1998 wurden an der Unteren Mittelelbe Untersuchungen zum Aggregationsverhalten weidender Bläß- und Saatgänse durchgeführt. Mehr als 90% aller erfaßten Gänse hielten sich in Trupps auf, die aus mehr als 500 Vögeln bestanden. Eine verlängerte Freßzeit auf Kosten von Komfortverhalten und Ruhen belegte einen erhöhten Konkurrenzdruck mit zunehmender Truppgröße. Die Fluchtdistanzen kleiner Gänsegruppen war mit 60 bis 120 Metern geringer als bei größeren, doch nahmen sie bereits ab etwa 150 Vögeln bei einem Wert von ca. 200 Metern nicht weiter zu. Mit zunehmenden Bestandszahlen wuchs die Größe der Haupttrupps im Gebiet, zugleich erhöhte sich jedoch auch die Anzahl der Trupps.Eine mögliche Erklärung für die beobachtete Verteilung der Gänse auf die unterschiedlichen Truppgrößen besteht darin, daß ein großer Teil der Rastpopulation bei der Nahrungssuche opportunistisch vorgeht. Diese Gänse orientieren sich bei der Feldwahl an einigen wenigen, besonders erfolgreichen Art- bzw. Gattungsgenossen. Feldgröße und maximale Individuendichte waren wichtige Faktoren, welche die absoluten Truppgrößen auf den überwiegend aus Äckern bestehenden Nahrungsflächen begrenzten.

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Flock sizes in foraging White-fronted and Bean Geese in the Elbe valley and their effects on flight distance and time budget

Summary  Flocking behaviour of foraging Whitefronted and Bean Geese (Anser albifrons, A. fabalis) was studied in the valley of the lower River Elbe from 1994 to 1998. Geese were counted every forthnight in the winter season of 1994/95 in a study area of 170 km², and daily in 1995/96 and 1996/97 in an area of 40 km². In the winter of 1997/98, counts were conducted every second day. Feeding behaviour was sampled by scan sampling in 1995/96, and distances of flight reactions to an approaching car were estimated in 1996/97 and 1997/98.Usually, geese formed large flocks. More than 90% of individuals recorded stayed in groups of more than 500 birds. In large flocks (several thousand geese), prolonged feeding times at the cost of preening and resting behaviour indicated a severe competition between individuals. Flight distances were lower in small flocks, but did not increase further with flocks becoming larger than 150 birds. As shown in earlier studies, the benefit of flocking in terms of predator avoidance is unlikely to increase any further with groups exceeding a few hundred birds.One possible explanation of the observed flocking behaviour could be that most individuals in the population follow an opportunistic strategy when foraging. They join their foraging cnspecifics instead of looking for feeding sites on their own. Flock size was limited by population size on the one hand, and by field size on the other. Average maximum density of individuals in a flock was 0,1 birds per square meter. Thus, bird density together with field size are likely to be the main factors determining and constraining flock size on agricultural fields.

     

栖息地特征对褐马鸡种群密度和集群行为的影响

1998-2000年冬季在山西芦芽山自然保护区选择车道沟和梅洞 2个研究地,就栖息地特征对褐马鸡 (Cros soptilonmantchuricum)越冬种群密度和集群大小的影响进行了研究。经χ2 检验,针叶林是褐马鸡冬季经常利用的栖息地类型。对 2个研究地的栖息地结构比较发现,车道沟适宜栖息地的面积较梅洞大,连接性也较好,褐马鸡种群有更为广阔的取食空间和更多的隐蔽场所,而且活动的阻碍也较小;而梅洞栖息地的取食空间较小,隐蔽场所较少,其活动时受到的阻碍也较大。从 2个研究地微生境结构特征的差异,并结合褐马鸡越冬期栖息地选择来看,树高和高层盖度对褐马鸡在 2个栖息地的选择上起重要作用。车道沟乔木较粗大,不但有丰富的食物,而且可提供较好的夜宿条件;高层盖度越大,栖息地的隐蔽条件越好,能吸引更多的褐马鸡个体取食和越冬。研究结果表明,栖息地结构连接性、微生境结构特征的差异以及人为干扰是褐马鸡种群密度和集群行为差异的主要影响因素。     

Cooperative breeding in birds: the role of ecology

Theory predicts that cooperative breeding should only occurin species in which certain individuals are constrained frombreeding independently by some peculiarity of the species' ecology.Here, we use comparative methods to examine the role of variationin ecology in explaining differences between taxa in the frequencyof cooperative breeding. We address three questions. First,does the frequency of cooperative breeding vary at just one phylogeneticlevel, or across several levels? Second, are differences inthe frequency of cooperative breeding among closely-relatedspecies correlated with ecology? Last, are ecological differencesbetween ancient lineages important in predisposing certain lineagesto cooperative breeding? We find that variation in the frequencyof cooperative breeding occurs across all phylogenetic levels,with 40% among families and 60% within families. Also, variationin the frequency of cooperative breeding between closely related speciesis associated with ecological differences. However, differencesin the frequency of cooperative breeding among more ancientlineages are not correlated with differences in ecology. Together,our results suggest that cooperative breeding is not due toany single factor, but is a two step-process: life-history predispositionand ecological facilitation. Low annual mortality predisposescertain lineages to cooperative breeding. Subsequently, changesin ecology facilitate the evolution of cooperative breedingwithin these predisposed lineages. The key ecological changesappear to be sedentariness and living in a relatively invariableand warm climate. Thus, although ecological variation is notthe most important factor in predisposing lineages to cooperativebreeding, it is important in determining exactly which speciesor populations in a predisposed lineage will adopt cooperativebreeding.     

The endemic gastropod fauna of Lake Titicaca: correlation between molecular evolution and hydrographic history

Lake Titicaca, situated in the Altiplano high plateau, is the only ancient lake in South America. This 2- to 3-My-old (where My is million years) water body has had a complex history that included at least five major hydrological phases during the Pleistocene. It is generally assumed that these physical events helped shape the evolutionary history of the lake's biota. Herein, we study an endemic species assemblage in Lake Titicaca, composed of members of the microgastropod genus Heleobia, to determine whether the lake has functioned as a reservoir of relic species or the site of local diversification, to evaluate congruence of the regional paleohydrology and the evolutionary history of this assemblage, and to assess whether the geographic distributions of endemic lineages are hierarchical. Our phylogenetic analyses indicate that the Titicaca/Altiplano Heleobia fauna (together with few extralimital taxa) forms a species flock. A molecular clock analysis suggests that the most recent common ancestor (MRCAs) of the Altiplano taxa evolved 0.53 (0.28-0.80) My ago and the MRCAs of the Altiplano taxa and their extralimital sister group 0.92 (0.46-1.52) My ago. The endemic species of Lake Titicaca are younger than the lake itself, implying primarily intralacustrine speciation. Moreover, the timing of evolutionary branching events and the ages of two precursors of Lake Titicaca, lakes Cabana and Ballivián, is congruent. Although Lake Titicaca appears to have been the principal site of speciation for the regional Heleobia fauna, the contemporary spatial patterns of endemism have been masked by immigration and/or emigration events of local riverine taxa, which we attribute to the unstable hydrographic history of the Altiplano. Thus, a hierarchical distribution of endemism is not evident, but instead there is a single genetic break between two regional clades. We also discuss our findings in relation to studies of other regional biota and suggest that salinity tolerance was the most likely limiting factor in the evolution of Altiplano species flocks.     

The building-up of social relationships: behavioural types, social networks and cooperative breeding in a cichlid

Consistent individual differences in behavioural types may not only cause variation in life-history decisions, but may also affect the choice of social partners and sociality in general. Here, we tested whether and how behavioural type influences the establishment of social ties using the cooperatively breeding cichlid, Neolamprologus pulcher. In a habitat saturation experiment with individuals pre-tested for behavioural type, we first analysed whether behavioural type affected the likelihood of settlement (i.e. social status), group sizes, and the types of dominant and subordinate individuals accepted as group members. Corrected for effects of body size and sex, the behavioural type did not affect settlement. However, bold dominant males only accepted smaller females, and grouped with bold subordinates, while shy dominant males accepted larger females than themselves, and grouped with shy subordinates. Second, we analysed the relationships between behavioural type and the aggressiveness or affiliation social network. Behavioural type significantly affected the number and quality of connections within the two networks. We show that behavioural types affect group composition, social networks and status achieved, in interaction with body size. Thus, the interactions within groups may depend not only on age, size and sex, but also on the behavioural type of the individuals involved.     

The role of zoos and captive breeding in lemur conservation

All the lemurs of Madagascar are declining in the wild, but the species and numbers held in collections outside Madagascar are not sufficient to ensure the continuence of their genetic diversity in captivity. General recommendations as to how to rectify this situation are presented, including a strong plea for coordinated population management among collections holding lemurs.