Predation risk affects trade-off between nest guarding and foraging in Seychelles warblers

The fitness costs of egg loss for Seychelles warblers (Acrocephalus sechellensis)on Cousin Island are considerable because warblers have a single-eggclutch and no time to lay a successful replacement clutch. Onthe islands of Cousin and Cousine, with equal densities of Seychellesfodies (Foudia sechellarum), nearly 75% of artificial eggs placedin artificial nests were predated by fodies after 3 days. OnAride Island with no fodies present, loss of artificial eggswas not observed. Female warblers incubate the clutch, and malewarblers guard the clutch when females are absent. Deterrenceof fodies by male warblers is efficient: loss rate of eggs fromunattended warbler nests was seven times as high as from attendednests, and the more nest guarding, the lower the egg loss andthe higher the hatching success. Egg loss is independent ofthe amount of incubation by females. There is no trade-off betweenincubating and foraging by females. Nest guarding competes withforaging by males, and this trade-off has a more pronounced effecton egg loss when food availability is low. The transfer of breeding pairsfrom Cousin to either Cousine with egg-predating fodies or toAride without fodies allowed us to experimentally investigatethe presumed trade-off between nest guarding and foraging. OnCousine, individual males spent the same amount of time nestguarding and foraging as on Cousin, and egg loss was similarand inversely related to time spent nest guarding as on Cousin.Males that guarded their clutch on Cousin did not guard theclutch on Aride but allocated significantly more time to foragingand gained better body condition. Loss of warbler eggs on Aridewas not observed. Time allocation to incubating and foragingby individual females before and after both translocations remainedthe same.     

First record of the fledging period of Temminck's Courser,Cursorius temminckii

The Seychelles Fody, Foudia sechellarum, is a ploceid weaver occurring naturally on three islands in the Seychelles group in the Indian Ocean. The population on Cousine Island was studied between 30 June and 25 August 1997. The size of the population on the island was estimated at 458–614 individuals and densities varied in different habitat types. As Seychelles Fodies in non-breeding plumage are difficult to sex, we provide sexing criteria based on wing length. Breeding pairs form small, probably temporary, territories that are defended by both partners against other fodies, including the introduced Madagascar Fody, Foudia madagascariensis. The Seychelles Fody often breeds semi-colonially and we observed up to five nests close together. The birds are socially monogamous and both sexes share in nest building, nest defense and provisioning of the young. Only females incubate. Many nests were deserted before eggs were laid, including some that had been accepted by the female. Additional males and females were sometimes seen helping provision the young, but this cooperative breeding behaviour appeared to be uncommon. Non-breeding individuals congregate in large flocks, sometimes joined by breeding birds.     

The evolution of the mating system of the Carpenter bee Xylocopa varipuncta(Hymenoptera: Anthophoridae)

The Carpenter bee, Xylocopa varipuncta , appears to have evolved a dispersed lek mating system. Males leave the natal nest in the late afternoon in March and April to hover for periods of up to two hours in the crowns of non-flowering trees growing in desert washes. Females and other males occasionally visit a hovering male, probably drawn by a sex pheromone. Resident males repel intruders of the same sex and attempt to induce females to alight on foliage, where mating occurs. The hovering territories do not contain nests, or potential nest sites, or food resources for females. Lek territoriality may have evolved in this species because nests are difficult to locate and the food-plants are evenly dispersed over a large area, making economical defence of patches of this resource unfeasible. Males are left with the option of demonstrating social dominance to potential mates through control of a superior landmark territory.     

Don'T put all your eggs in one nest: spread them and cut time at risk

Abstract In many egg-laying animals, some females spread their clutch among several nests. The fitness effects of this reproductive tactic are obscure. Using mathematical modeling and field observations, we analyze an unexplored benefit of egg spreading in brood parasitic and other breeding systems: reduced time at risk for offspring. If a clutch takes many days to lay until incubation and embryo development starts after the last egg, by spreading her eggs a parasitic female can reduce offspring time in the vulnerable nest at risk of predation or other destruction. The model suggests that she can achieve much of this benefit by spreading her eggs among a few nests, even if her total clutch is large. Field data from goldeneye ducks Bucephala clangula show that egg spreading enables a fecund female to lay a clutch that is much larger than average without increasing offspring time at risk in a nest. This advantage increases with female condition (fecundity) and can markedly raise female reproductive success. These results help explain the puzzle of nesting parasites in some precocial birds, which lay eggs in the nests of other females before laying eggs in their own nest. Risk reduction by egg spreading may also play a role in the evolution of other breeding systems and taxa-for instance, polyandry with male parental care in some birds and fishes.     

Effects of nestboxes on the breeding biology of Southern House Wrens Troglodytes aedon bonariae in the southern temperate zone

Nestboxes are known to increase clutch size, enhance breeding success and affect the social mating system of several cavity nesters. Although in recent years various cavity nesters have been studied in nestboxes in South America, the effects of boxes on the biology of the study species are unknown. We evaluated the effects of nestboxes on the breeding biology and social mating system of Southern House Wrens Troglodytes aedon bonariae by comparing birds breeding in nestboxes and tree cavities in two cattle ranches in Buenos Aires Province, Argentina. Southern House Wrens nesting in boxes had higher breeding success but, contrary to studies on the temperate zone, we did not find differences in clutch size between Wrens breeding in nestboxes and tree cavities. The main causes of nest failure in tree cavities were nest predation and flooding of the cavity (70 and 23% of the failures, respectively) while in nestboxes predation and desertion were the most important causes of failure (38 and 34% of the failures, respectively). The social mating system of Southern House Wrens is monogamy with biparental care, and neither was affected by the boxes. Males did not attract secondary females to additional nestboxes; however, nestboxes are safer breeding sites than tree cavities, and females seemed to prefer males with nestboxes on their territory. These results suggest that nest quality alone might be not enough for secondary females to accept polygyny.     

INTRASPECIFIC NEST PARASITISM IN BIRDS

(1) The incidence of intraspecific nest parasitism in birds is reviewed. This phenomenon can be identified by various methods, ranging from biochemical examination of protein and enzyme polymorphisms to observations on the sequence of appearance of eggs in the nest as well as on the shape and colour of the eggs. The phenomenon has been reported for at least 53 species (Table I), mostly among precocial birds.
(2) The evidence suggests that the parasites are ( a ) young, unmated birds, ( b ) females which have lost their nests and (c) mated females which also lay in the nests of other females. High proportions of such females, and scarcity of suitable nest sites increase the rate of parasitism.
(3) Breeding success in parasitized nests is reduced by factors such as egg loss, desertion, inefficient incubation, reduced clutch size and late layings. The upper limit of the rate of intraspecific parasitism is discussed. It is argued that its incidence will be higher in the tropics than in extreme zones, for example, high latitudes and deserts, because there is higher synchronization of breeding in the latter, and increased predation in the former zones. Also, it will be more common in species which lay large clutches, such as precocial species.     

赛罕乌拉国家级自然保护区红角鸮(Otus scops)的繁殖生态观察

为了解红角鸮(Otus scops)的繁殖状态,2009年5～8月,在内蒙古赛罕乌拉国家级自然保护区,以直接观察法对红角鸮的繁殖生态进行了研究。结果表明,所观察的4个巢址全部是以柳树为主干的树洞巢,巢内铺垫物较少或无铺垫,巢距地面的平均高度为91.3cm,巢内径平均为13.5cm,巢外径平均为15.0cm,巢深平均32.0cm(n=4)。平均窝卵数为3.75,对其中一窝进行测量,4枚卵平均量度是28.9mm×24.6mm,重量9.5g。雏鸟孵出后,雌性亲鸟承担主要育雏任务,雄性亲鸟日间停歇于离巢不远的树上负责警戒工作,夜间与雌鸟共同哺育雏鸟。小型鼬科动物的捕食可能是干扰红角鸮繁殖成功率的主要因素。     

Nest location, clutch size and nest success in the Scarlet Ibis Eudocimus ruber

Breeding success and nest-site characteristics were studied during the 1996–1997 breeding season in a colony of Scarlet Ibises Eudocimus ruber in south-eastern Brazil to test the hypothesis that nest-site characteristics and clutch size affect nest success. Two nesting pulses produced young, the earlier being more successful. Predation accounted for most failures during the first pulse, wind destruction during the second. A third pulse with few nests produced no young. Adult Ibises abandoned nests when they lost sight of other incubating birds. Logistic regression analysis indicated that nest success during the first pulse was positively related to clutch size, number of nests in the nest tree and in the nearest tree, and negatively to the distance to the nearest neighbour. During the second pulse there were significant negative associations between success, nest height and distance to the fourth nearest nest, and a positive association between success and nest cover. The results agree with the 'selfish herd' hypothesis, indicating that nest aggregation may increase breeding success, but the nest-site characteristics affecting success can differ over the course of one breeding season.     

Is Carrying Feathers a Sexually Selected Trait in House Sparrows?

Males in several avian groups carry specialised materials as part of their courtship display. Females may vary their investment in reproduction in relation to their mates' attractiveness. The house sparrow (Passer domesticus) population on Dassen Island, South Africa, builds domed nests in the branches of trees. These nests are more or less spherical structures with a deep cup lined largely with feathers. When males collect feathers, they call to females and display the feather before it is added to the nest. We examined whether carrying feathers provides females with an index of male quality, which correlates with their subsequent investment in parental care. Additionally, we studied the potential importance of feathers in nest insulation. Feathers were collected mainly by males. Males also brought larger feathers, and deposited more feathers in nests, than females. Number of trips with feathers – which increased after feathers were experimentally removed from nests – and number of feathers brought varied among males. Volume of feathers influenced females' investment in reproduction and positively correlated with clutch size and chick feeding rates. We found more feathers during incubation and immediately after chicks hatched, when nest heat requirements peak. Furthermore, number of trips with feathers and number and size of feathers were greater during incubation. Our results suggest that this feather‐carrying behaviour by male house sparrows influences maternal reproduction investment and could be a kind of parental care per se by supplementing nest insulation.     

The functional significance of multiple nest-building in the Australian Reed Warbler Acrocephalus australis

The vast majority of bird species build a nest in which to breed. Some species build more than one nest, but the function of most multiple nest-building remains unclear. Here we describe the unusual nest-building behaviour of the Australian Reed Warbler Acrocephalus australis , and test experimentally the hypotheses that multiple nest-building is related to individual condition or territory quality, and plays a role in mate assessment. Australian Reed Warblers built two types of nest structures: 'type I' nests, which were used for eggs and nestlings, and 'type II' nests, which were structurally distinct from type I nests, did not support eggs, nestlings or adults and were not essential for successful breeding. The number of type II nests built in each territory varied. Type II nests were only built before breeding had commenced in a territory and females were not observed participating in their construction, supporting a role in female mate choice. Birds provided with supplementary food built significantly more type II nests than control birds. However, supplementary-fed birds did not have greater pairing success, and the addition of further type II nests to territories did not increase the pairing rate or type II nest construction in those territories. There was no relationship between the presence of type II nests and either reproductive success or likelihood of nest predation. We discuss the implications of these results in light of previous suggestions regarding the function of multiple nest-building in birds.     

Breeding of the Seychelles Magpie Robin Copsychus sechellarum and implications for its conservation

The total population of the Seychelles Magpie Robin Copsychus sechellarum declined from 38–41 birds in 12–13 territories in 1977–1978 to 17–21 birds in eight to nine territories in 1988–1990 and was entirely confined to Fregate Island (210 ha) in the Seychelles. After a successful cat eradication program in 1981–1982, recruitment improved, although the abandonment of agriculture had caused a reduction in the amount of feeding habitat and hence in the carrying capacity of the island. The population declined because of the failure of recruitment to compensate for the annual adult mortality. Foraging activity of the Magpie Robin was greater in high-quality territories (measured by soil invertebrates available), leading to increased reproductive success. Through supplementary feeding, five times as many recruits were produced. Of the 11.5 potential annual breeding recruits, 5.3 are required to compensate for adult mortality, and the other 6.2 recruits can be regarded as “surplus” contributing to an increase. Magpie Robins prefer to breed in rotten trees, which are a scarce resource. The greater the distance between the nest site and feeding areas, the less time was spent in incubation and nest guarding, resulting in greater egg loss. Because of lack of suitable areas for establishing territories, many young Magpie Robins became “floaters”. Nest disturbance, both by these floaters and by the introduced Indian Mynah Acridotheres tristis, had adverse effects on the breeding success of robins. A recovery plan has been designed to save the Magpie Robin. Territories have been improved for feeding (by tree planting) and for breeding (by providing nestboxes and reducing nest disturbances). Given the vulnerability of one small island, the presence of surplus birds (supported mainly by supplementary food) and the suitability of neighbouring Aride Island (68 ha), successful translocations to this island took place in 1992 and 1994. Given the presence now of 47 individuals on two islands, it is hoped that the species will pull back from the brink of extinction.     

Appearance and vulnerability of artificial duck nests to avian predators

Use of artificial nests in studies of nest success in birds has become increasingly popular, but this practice has been criticised because the appearance and success of artificial nests may differ from natural nests. Typically, artificial nests are presented with few visible eggs throughout the entire exposure period while natural nests during their life have different appearances, possibly producing stage-dependent predation rates. In ducks, nests may appear with openly visible eggs, eggs covered with nest material and covered by an incubating female. To test how nest appearance may affect nest survival, I simulated such duck nest appearances, and used direct observations to record responses of marsh harriers Circus aeruginosus , the dominant duck nest predator at the Lake Engure, Latvia, during the breeding seasons 2000–2002. I recorded 274 harrier flights over all types of artificial nests, 74 nest discoveries and 54 predation events. Logistic regression analyses showed that nest appearance was a significant predictor of the fate of the nest: nests with openly visible eggs were discovered more often than nests covered with nest material. Nests with dummy females had discovery rates similar to the average of all artificial nest types suggesting that they best reflect the predation risk of artificial nests and could be used in future studies. After discovering nests covered with dummy females, harriers never attacked but tried to scare them off to get access to the eggs. In 52% of cases of nest discovery, harriers gave up before touching the dummy. This shows that duck passive nest defence may prevent clutch predation.     

Dark-bellied brent geese Branta b. bernicla breeding near snowy owl Nyctea scandiaca nests lay more and larger eggs

Several studies have demonstrated that snowy owls Nyctea scandiaca defend an area around their nests against predators, hereby inadvertently creating safe havens for breeding dark-bellied brent geese Branta b. bernicla . However, studies investigating brent goose breeding ecology within the predator-exclusion zones of the snowy owls are absent. In 1999 and 2005, years of high lemming abundance Lemmus sibiricus and Dicrostonyx torquatus , brent geese were primarily breeding in association with snowy owls in the Medusa river catchment on western Taimyr, Russia. Goose nest failure, either as a result of nest abandonment by the adult birds or of nest depredation, increased with increasing distance from the owl nests. Within the brent goose colonies, clutch size as well as egg size increased with decreasing distance from the snowy owl nest, indicating an increasing adult quality closer to owl nests. However, as a result of the abandonment of eggs and goslings, the increasing clutch size did not result in a higher nest success during this study. Apparently brent geese compete for breeding sites close to owl nests, but details of this process remain unknown.     

Islands in a desert: breeding ecology of the African Reed Warbler Acrocephalus baeticatus in Namibia

The continental African Reed Warbler Acrocephalus baeticatus, like its relative the Seychelles Warbler Acrocephalus sechellensis, breeds in isolated patches. We studied the mating system of the African Reed Warbler to see whether this species, like the Seychelles Warbler, shows co-operative breeding. The African Reed Warbler is not polygynous. The majority breed monogamously (88%, n = 65), however in 12% of the territories three adult unrelated birds (mostly males) were observed participating in the brooding and feeding of nestlings, suggesting a polyandrous breeding system. Multilocus DNA fingerprinting revealed that the helping bird was unrelated to the pair birds. The percentage of nests with helpers was low compared to rates found in the Seychelles Warbler or Henderson Reed Warbler Acrocephalus vaughani taiti. This could be due to the scarcity of potential helpers or to the fact that, although limited, birds still had the opportunity to disperse within a meta-population structure in search of vacant territories. The presence of helpers was associated with increased hatching success due to lower predation rates, but not with increased fledging success. Another possible benefit of helping behaviour in this species could be improved predator detection and mobbing. Nest predation was high and warblers tended to build their nests in the highest, most dense reed patches available in their territory. There was no relation between habitat quality, measured as insect food availability, and the occurrence of helpers.     

Nesting success of understory forest birds in central Panama

Greater nest predation in tropical than temperate birds has been hypothesized to be a primary selective force generating latitudinal differences in avian life history traits. Few extensive data sets, however, have been available from tropical forests to compare with data from temperate forests. To increase the amount of empirical information available for addressing issues related to the evolution of life history traits of tropical birds, we measured the nesting success of understory birds in lowland forest of central Panama. We found and monitored the fates of 696 nests of 71 species over two breeding seasons. Daily nest predation rates for the ten species for which we obtained the largest samples ranged from 1.6 to 8.3%, equivalent to a loss of 43 to 92% of nests. These values overlapped extensively the range of daily predation rates experienced by ecologically similar species in North America. Proportion of nests fledging young, estimated with the Mayfield method, was significantly lower in tropical (range: 8 to 57%) than temperate (27 to 60%) species. Nesting success in Panama varied among years, however, being greater in 1996 than 1997. In 1996, nesting success was similar to that of species breeding in forest fragments of midwestern North America. When compared with success of nests in large, contiguous forest tracts of North America, however, tropical avian nesting success was consistently lower by approximately 23%. We conclude that nesting success in central Panama may be poor in most breeding seasons, but also may be punctuated by occasional years of relatively exceptional success, a possibility heretofore unappreciated because of a general paucity of data from the tropics. Furthermore, our results indicate substantial variation in levels of nesting success among species, and almost no variation in clutch size. Such large interspecific variation, as well as potentially large annual variation, in nesting success does not support the hypothesis that uniformly low levels of nesting success select for small tropical clutch sizes.     

Clutch size variation in passerine birds: The nest predation hypothesis

Summary The hypothesis that a negative relationship exists between clutch size and the probability that the nest will be robbed is tested, using data for passerine birds given in the literature. The data for four separate groups of species, viz. hole-nesters, semi hole-nesters and open-nesters nesting above and on the ground, respectively, were examined in relation to geographical gradients and seasonal and annual variation. In general, the data analysis results support the hypothesis, but cannot yet be considered as proven. More data on the riks of nest predation are needed. The most serious discrepancy is that for the Fieldfare, the protection from predation provided by nesting in colonies does not seem to be accompanied by a corresponding increase in clutch size. However, the clutch size of the Brambling, a species with seeks out such colonies for its breeding sites, does tend to increase in these hatitats.The pattern of clutch size variation was similar for the two groups of hole-nesting species, but differed significantly from that found for the two groups of open-nesters. The difference in the clutch size variation of the two groups of open-nesting species predicted from the hypothesis, viz. that, in northern regions, both latitudinal and altitudinal increases in clutch size should more commonly be found for those species which nest on the ground, compared to those nesting above ground level, was confirmed.A seasonal decrease in clutch size, in temperate regions, was found to be typical for species whose nests are subject to relatively little predation, particularly for those species which have a short breeding season. Advantages of laying small clutches, resulting in fewer nestlings than the number which would be possible for the parents to rear successfully, are discussed, and a simple model is presented which does not assume that nest predation is dependent on clutch size.     

Nestbox provisioning in a rural population of Eurasian Kestrels: breeding performance,nest predation and parasitism

The breeding biology of the Eurasian Kestrel Falco tinnunculusin nestboxes in farmland was studied to test for differences between artificial and natural sites. We report on the direct effect of nestbox provisioning on some life-history traits and how nestbox use affects nest predation and parasitism. Five types of nest-sites were available: nestboxes on poles and trees (artificial sites), stick nests on trees, stick nests on pylons and holes in buildings (‘natural’ sites). The Kestrel population increased from 23 pairs in 1993 (prior to nestbox installation) to 55 in 1998 as nestboxes were provided. In general, pairs breeding in trees started to lay later than those nesting in nestboxes on poles or in building holes, but this difference was probably associated with habitat quality rather than nest type. Differences in clutch size were found between nest-sites in some years, and were associated with laying date and, probably, with variation in territory quality. Using only data from successful nests, pairs breeding in nestboxes produce more fledglings than those in building holes or pylons. The frequency of nest predation was higher in natural sites than in nestboxes. The number of fledglings from pairs breeding in nestboxes was higher than from those breeding in old stick nests in trees when all nests were considered. Nestbox provisioning had no effect on the occurrence of the ectoparasite Carnus hemapterus, but chicks from nestboxes showed higher intensity of infection. Our results suggest that nestbox provisioning increases reproductive success and the frequency of nest predation or intensity of parasite infestation in Kestrels.     

Causes of reduced clutch size in a tidal marsh endemic

We tested three hypotheses of clutch size variation in two subspecies of the swamp sparrow (Melospiza georgiana georgiana and M. g. nigrescens). Swamp sparrows follow the pattern of other estuarine endemics, where clutch size is smaller among tidal salt marsh populations (M. g. nigrescens) than their closest inland relatives (M. g. georgiana). Our results support predation risk and temperature, but not adult survival, as explanations of this pattern in swamp sparrows. Coastal nests were twice as likely to fail as inland nests, and parental activity around the nest site was positively related to clutch size at both sites. When brood size was controlled for, coastal adults visited nests less often and females vocalized less frequently during visits than inland birds, which may decrease nest detectability to predators. Coastal parents waited longer than inland birds to feed offspring in the presence of a model nest predator, but there was no difference in their response to models of predators of adults, as would be expected if coastal birds possessed increased longevity. Additionally, coastal females laid more eggs than inland females over a single season, following a within-season bet-hedging strategy rather than reducing within-season investment. Coastal territories experienced ambient air temperatures above the physiological zero of egg development more often, and higher temperatures during laying correlated with smaller clutches and increased egg inviability among coastal birds. Similar effects were not seen among inland nests, where laying temperatures were generally below physiological zero. Both subspecies showed an increase in hatching asynchrony and a decrease in apparent incubation length under high temperatures. Coastal individuals, however, showed less hatching asynchrony overall despite higher temperatures. Both air temperatures during laying and predation risk could potentially explain reduced clutch size in not only coastal plain swamp sparrows, but also other tidal marsh endemics.     

Old nests are cues for suitable breeding sites in the Eurasian penduline tit Remiz pendulinus

Nest-sites often have a major influence on avian reproductive success. The use of reliable cues that assist nest-site selection should thus be favoured by natural selection. The old nests have been known to serve as a cue in nest-site selection in several species. To find out whether the old nests act as cue in nest-site selection in the Eurasian penduline tit Remiz pendulinus , we carried out two experiments in southern Hungary, where the penduline tits breed around fishponds and build sophisticated pendulous nests on tree branches that often hang over water. In April 2006, we choose 20 groups of two nearby trees, and hung an old nest on one of the trees in each group. The male penduline tits choose 12 of these groups to build a new nest, and every of the twelve nests were built on trees with an old nest. This suggests that the old nests serve as a cue in the selection of breeding sites for males when they enter a habitat. To find out whether the old nests are cues of plentiful nest building material, or to signal high quality breeding areas, we carried out a second experiment in 2007 by selecting 13 groups of three nearby trees. A "worn-out" old nest was hung on one of the trees, a "re-utilize" type of old nest on another tree, whereas the third tree was left without an old nest in each group. The rationale was that while the worn-out material of the old nests is of no use in building the new nests, material of the "re-utilize" nests is good enough to be used for building new nests. Males built a new nest in 10 of the 13 groups, and eight of the new nests were built on trees with an old nest. Of the eight new nests, five were built on trees with a "worn-out" old nest and three on trees with a "re-utilize" type of old nest. It appears that for the penduline tit males it is the presence of an existing old nest and not the quality of the old nest material that serves as cue during the selection of the suitable breeding sites.     

Plural Breeding in the Splendid Fairy-wren,Malurus splendens (Aves: Maluridae), a Cooperative Breeder

Groups of the cooperatively breeding splendid fairy-wren Malurus splendens may include more than one female. Previously this species has been described as singular breeding (only one female breeds). This paper describes the occurrence of plural breeding (PB) groups in 10% of group years, in which two females had separate nests. In all cases, the secondary female (Y) was related to the primary breeding female (X) and was generally a 2-year old female which had helped in the group during the previous breeding season. Plural breeding was correlated with an increase in population density and in the number of female helpers; PB groups were larger than singular-breeding groups. In most cases, the X female was occupied with her own nest or offspring when the Y female began to nest, and there was no aggression between them. Which birds helped the Y female to feed at her nest depended on the time between the hatching of the two nests. If the interval was small, some group members helped at each nest; with longer intervals, the group members began to feed at the earlier nest, and the other female was left to raise her brood alone. Female helpers were very active in feeding at single nestings, and the cost to an X female of a Y female breeding was mainly a loss of this assistance. The success of individual X nests was not affected. Effects on productivity were slight, but fewer X females in PB groups raised second broods than did experienced singular breeding females. Y females were less productive than X females, but no less productive than singular breeding novice females without helpers. It is not known whether Y females copulated with primary or secondary males within their group, or with males from outside the group. Certainly, they did not form an observable pairing with any male in the group. Plural breeding occurred in a minority of group years in response to extrinsic conditions and the current demographic situation, and shows the extreme plasticity of the mating system in M. splendens.