Artificial incubation of eggs of the green iguana (Iguana iguana)

Successful hatching of large numbers of artificially incubated eggs from the green iguana, Iguana iguana, are reported. Gravid females were captured at a nesting site in Summit Gardens, Soberania, Parque Nacional, Panama, and released into an enclosure with an artificial nesting area. Females dug their own tunnels and nest chambers or used artificial nest chambers for egg deposition. Eggs (n = 829) from 21 clutches were removed from the nests and artificially incubated. Average hatching success per clutch was 94.6% and the mean clutch incubation time was 92.1 days. A distinct odor was noticed in the incubation containers several days before the eggs began to hatch. At this time, the substrate layer was removed exposing the top surface of all eggs, and the eggs desiccated to some degree prior to hatching. High incubation temperature, hydric conditions, and egg arrangement are all implicated as contributing factors in low hatching success in previous studies using artificial incubation techniques.     

Patterns of hatching failure in the house sparrow Passer domesticus

Hatching failure is widespread in birds despite presumably strong selection against any wasted reproductive effort and yet there have been few attempts at uncovering sources of variation in failure within a species. Here we use 11 yr of data on the frequency of both undeveloped and unhatched eggs in a marked population of house sparrows Passer domesticus to examine the effects of multiple factors hypothesized to affect hatching success in birds including parental identity, phenotype and genotype, as well as several environmental variables. Unhatched eggs were present in 34.3% of 1523 clutches in which at least one egg hatched, and a total of 10.1% of eggs failed to hatch, of which 75.7% had no visible embryo. We found an effect of female identity on the proportion of eggs that developed and an effect of male identity on hatching success, as well as a positive effect of egg size on both the proportion of eggs that developed and hatching success between females. Neither the proportion of eggs that developed nor hatching success varied according to time of season or weather conditions, clutch size, the age of either parent, the size of the male's black bib, female size, female heterozygosity or the genetic similarity between members of a pair. There was also no positional bias in the occurrence of undeveloped eggs, although undeveloped eggs were more common among birds nesting at low density. Our results suggest that hatching failure in house sparrows has few environmental components and is primarily influenced by intrinsic differences among individuals.     

Developmental stage affects eggshell-breaking strength in two ground-nesting birds: the partridge (Alectoris rufa) and the quail (Coturnix japonica)

We examined the relationship between embryo development and egg hardness in two ground nesting bird species, the red-legged partridge (Alectoris rufa; n = 165 eggs) and the quail (Coturnix japonica; n = 148 eggs). For both species, we observed a strong effect of developmental stage on egg hardness. Eggs near hatching were significantly weaker than unincubated eggs (partridge: 18 and 23 N, respectively, and the quail 7 and 10 N, respectively). We additionally explored the effect of incubation on egg hardness in a control sample of non-fertilised quail eggs (i.e., without embryo development). The control eggs maintained in the incubator for the full incubation time (17 days) were significantly harder (7-9 N) than eggs containing fully developed chicks (5-7 N). Thus, the incubation conditions of high temperature and humidity alone seem not to have a significant effect on egg hardness, and support the important effect of calcium uptake.     

Costs of large communal clutches for male and female Greater Rheas Rhea americana

The breeding system of the Greater Rhea Rhea americana is almost unique among birds as it combines harem polygyny and sequential polyandry, with communal egg-laying and uniparental male care. In this species, large communal clutches (more than 30 eggs) are rare and have a lower hatching success than smaller clutches. Here we analyse the proximate causes of hatching failures and the costs of large communal clutches (and therefore the costs of extensive polygyny) for males and females. We evaluated if length of the nesting period, egg viability, egg losses during incubation and male parental activity at the nest were affected by clutch size. We also evaluated if chicks hatched from large clutches have a lower survival during the first 2 months after hatching. Large clutches had longer nesting period and lower hatching success, mainly as a result of bacterial contamination of the eggs and increased hatching asynchrony. In addition, large clutches tended to lose more eggs as a result of accidental breakage or predation. Male activity at the nest and chick survival were not related to clutch size. Low hatching success, nest predation risk and energetic costs associated with large clutches penalize females that join large harems and males that accept additional eggs into the nest.     

FREQUENCY-DEPENDENT FITNESS CONSEQUENCES OF INTRASPECIFIC NEST PARASITISM IN SNOW GEESE

The reproductive efficiency, defined as the number of breeding recruits produced per egg laid; of intraspecific nest parasites; of hosts in parasitized nests; and of unparasitized nesting females, was measured for 14 years for lesser snow geese Anser caerulescens caerulescens nesting near Churchill, Manitoba, Canada. Relative efficiencies were 0.71–0.88, 0.91, and 1.0 for eggs of parasites, hosts, and unparasitized birds, respectively. Differences in the hatching probabilities of the three classes of eggs produced the efficiency differences. Parasitic success was limited by the parasites' failure to place more eggs than expected by chance into nests at the appropriate time relative to host incubation. Host nesting success was lower when more than one parasitic egg was added to the clutch. No differences in gosling survival and breeding recruitment probabilities were detected among any categories of goslings. Thus, hatching parasitic young are at no disadvantage relative to parental young, and there is no support for the hypothesis that increased success of host young at later stages of reproduction might offset negative effects at the egg stage. The hatching efficiency of parasitic eggs declined more rapidly than that of parental eggs as the parasitism rate increased. Efficiencies were similar when 3–4% of the eggs laid per year were parasitic, but relative parasitic efficiency was significantly lower when the parasitism rate was 8–9% or more. Using ancillary information and assumptions about the fecundity, viability, and behavioral flexibility of parasitic and parental females, we conclude that intraspecific nest parasitism could compete with nesting as a reproductive strategy in this population. The conditional use of parasitism by a large component of the population in certain years, however, combined with negative-frequency dependent success, limits the potential spread of a purely parasitic strategy in this population.     

Observations of predation by corvids at a Marbled Murrelet nest

ABSTRACT.   Unlike other alcids, Marbled Murrelets ( Brachyramphus marmoratus ) typically nest solitarily on large platforms in the upper canopy of old-growth coniferous trees. Although Marbled Murrelets exhibit characteristics that minimize the risk of predation, habitat fragmentation has likely increased the risk of nest predation by corvids. Using a video camera at a nest in northern California, we observed nest fates for 4 yr (2002–2005). These recordings revealed two cases of egg predation by a Steller's Jay ( Cyanocitta stelleri ) and a Common Raven ( Corvus corax ), respectively. In both instances, the egg was carried from the nest site, leaving no evidence that nesting had occurred. Our observations revealed (1) the need to be conservative in estimates of nesting attempts and nesting success of Marbled Murrelets based only on the presence or absence of eggshell fragments or feces, (2) that not all predation attempts were successful, and (3) that murrelets may be more susceptible to predation early in the incubation period when adults may be less attentive to eggs and, after hatching, when young are less than 14 d old. We encourage efforts to minimize anthropogenic activities that may increase local corvid densities or disturb murrelets during the early incubation period.     

Incubation capacity contributes to constraints on maximal clutch size in Brent Geese Branta bernicla nigricans

Lack ( 1967 ) proposed that clutch size in species with precocial young was determined by nutrients available to females at the time of egg formation; since then others have suggested that regulation of clutch size in these species may be more complex. We tested whether incubation limitation contributes to ultimate constraints on maximal clutch size in Black Brent Geese (Black Brant) Branta bernicla nigricans. Specifically, we investigated the relationship between clutch size and duration of the nesting period (i.e. days between nest initiation and the first pipped egg) and the number of goslings leaving the nest. We used experimental clutch manipulations to assess these questions because they allowed us to create clutches that were larger than the typical maximum of five eggs in this species. We found that the per‐capita probability of egg success (i.e. the probability an egg hatched and the gosling left the nest) declined from 0.81 for two‐egg clutches to 0.50 for seven‐egg clutches. As a result of declining egg success, clutches containing more than five eggs produced, at best, only marginally more offspring. Manipulating clutch size at the beginning of incubation had no effect on the duration of the nesting period, but the nesting period increased with the number of eggs a female laid naturally prior to manipulation, from 25.4 days (95% CI 25.1–25.7) for three‐egg clutches to 27.7 days (95% CI 27.3–28.1) for six‐egg clutches. This delay in hatching may result in reduced gosling growth rates due to declining forage quality during the brood rearing period. Our results suggest that the strong right truncation of Brent clutches, which results in few clutches greater than five, is partially explained by the declining incubation capacity of females as clutch size increases and a delay in hatching with each additional egg laid. As a result, females laying clutches with more than five eggs would typically gain little fitness benefit above that associated with a five‐egg clutch.     

On a Poorly Known Rookery of Green Turtles (<Emphasis Type="Italic">Chelonia mydas</Emphasis>) Nesting at the Chabahar Beach,Northeastern Gulf of Oman

There is a poorly known rookery of green turtles (Chelonia mydas) nesting in sandy beaches of Chabahar town, northeastern Gulf of Oman, Iran.This study has been carried out to evaluate nesting activity of this small rookery in 2014 nesting season (June to October). In this study, total clutches were collected and transferred to an artificial hatchery. The peak of nesting occurred from the third parts of August to the end of September. Mean CCL was 106.3 ± 6 cm and mean CCW was 94.5 ± 5 cm. Females laid on average of 99.42 ± 47.8 eggs per clutch. The mean inter-nesting interval was 18.5 days. The observed clutch frequency was 3.4. Mean hatching success was 36.63 ± 4.1%. The incubation period was 61.07 ± 5.4 days. Nest status evaluation represented that major causes for the failure was unhatched egg with no obvious embryo followed by unhatched eggs with obvious but undeveloped embryos > unhatched eggs with developed embryos > hatched eggs but dead embryos. The results achieved in this study are a valuable contribution to cognition of the reproductive ecology of the green turtle population globally and regionally.     

Trans-shell infection by pathogenic micro-organisms reduces the shelf life of non-incubated bird's eggs: a constraint on the onset of incubation?

Many birds initiate incubation before clutch completion, which results in asynchronous hatching. The ensuing within-brood size disparity often places later-hatched nestlings at a developmental disadvantage, but the functional significance of the timing of the onset of incubation is poorly understood. Early incubation may serve to maintain the viability of early-laid eggs, which declines over time owing to the putative effects of ambient temperature. An unexplored risk to egg viability is trans-shell infection by micro-organisms. We experimentally investigated the rate and magnitude of microbial trans-shell infection of the egg, and the relative effects of ambient temperature and micro-organisms on hatching success. We show that infection of egg contents is prevalent and occurs within the time required to lay a clutch. The probability of infection depends on the climatic conditions, the exposure period and the phylogenetic composition of the eggshell microbiota. We also demonstrate that microbial infection and ambient temperature act independently to reduce egg viability considerably. Our results suggest that these two factors could affect the onset of avian incubation in a wide range of environments.     

Eggshell pigmentation pattern in relation to breeding performance of blue tits Cyanistes caeruleus

1. We test the consequences, in terms of breeding success and parental effort, of eggshell pigmentation pattern in a hole-nesting bird, the blue tit Cyanistes caeruleus that lays eggs asymmetrically speckled with reddish spots (maculated eggs). 2. We assess the effect of distribution of spots (pigment 'spread') and spot size and pigment intensity (pigment 'darkness') on eggshell physical properties and breeding parameters concerning nestling condition, investment of parents in offspring care and reproductive output in two different habitat types: a deciduous oakwoodland and an evergreen forest. 3. Blue tit clutches with more widely distributed spots showed a thicker eggshell, a shorter incubation period, a lesser amount of mass loss per day and a higher hatching probability than those with spots forming a 'corona' ring. While eggs with larger and darker (more pigment intensity) spots showed a thicker eggshell and a shorter incubation period. In the light of 'signal function hypothesis', these egg traits may reflect female health status and, consequently, this could affect male parental effort. 4. Here we show supports for some of the necessary assumptions of this hypothesis. We found a positive relationship between egg pigment 'spread' and male but not female provisioning rates per day. On the other hand, pigment 'darkness' of blue tits' clutches was positively related to female tarsus length, while pigment 'spread' was positively related to clutch size, male body mass and nestling tarsus length. Our study shows that eggshell pigment 'spread' can be used as an indicator of clutch quality. Further investigations are needed to understand the role of calcium availability as possible causal agent of deviant eggs and its relation to the maculation phenomenon.     

Seasonal variation in reproductive measures of tropical Roseate Terns Sterna dougallih previously undescribed breeding patterns in a seabird

Seasonal variation in egg-laying, egg size, hatching success, hatchling mass, fledging success and chick growth of Roseate Terms Sterna dougallii breeding on Aride Island (Seychelles), Indian Ocean, were studied in 1997 and 1998. I investigated to what extent two patterns, common in a range of species, were followed by tropical Roseate Terns: (a) seasonal decrease in clutch size, egg size and breeding success and (b) an increase in breeding success with increasing egg weight. In 1997 (a poor year), the earliest nesting birds laid significantly smaller eggs, and chicks were lighter at hatching than those of peak nesting birds. The mean clutch size, of 1.04 eggs, showed no seasonal variation and no 'b'-eggs hatched. In 1998 (a good year) the earliest nesting birds laid eggs of similar size and their chicks were of similar weight to those of peak nesting birds. Mean clutch size, of 1.25 eggs, increased significantly through the season and about 60% of the 'b'-eggs hatched. In 1997, hatching success was 57% whereas in 1998 it was 80%. In both years, breeding success declined significantly through the season. The fact that the earliest breeding birds laid smaller eggs in a poor year and smaller clutches in a good year is in marked contrast to a range of other species, and to temperate-nesting Roseate Terns. Egg volume explained about half of the variance in hatchling mass in both years, but only 15% of the variation in linear growth rate. Hatching date was the only variable with a significant effect on fledging success. Roseate Terns on Aride seemed to sacrifice egg size and clutch size for earliness of laying. Presumably it is a strategy of older birds to lay as early as possible and may be regarded as a response of tropical Roseate Terns to breeding under relatively poor, and seasonally declining, food conditions.     

人工巢箱条件下白眉姬鹟的繁殖参数

2005～2006年,在吉林省左家自然保护区的次生林中,对人工巢箱条件下白眉姬鹟(Ficedulazanthopygia)的繁殖参数开展了初步研究.结果表明,人工巢箱中自眉姬鹟的窝卵数为5～7枚,平均6.0枚;卵重平均为1.6 g.卵大小平均为17.0 mm×13.1 mm.孵化期平均为13.1 d,每巢平均出雏5.4只,育雏期平均为12.8 d,每巢平均出飞雏鸟5.3只.白眉姬鹟的营巢成功率为70.0%,繁殖成功率为81.3%.     

Determination of heavy metals in eggs of Little Grebe (Tachybaptus ruficollis) around the wastewater treatment ponds,Khon Kaen University

Bird eggs can often be used as a bioindicator of potential effects in environment pollutions. Little Grebe (Tachybaptus ruficollis) lays the eggs around wastewater treatment ponds, Khon Kaen University. The aim of this study is to determine the contamination level of Hg, Pb, Cd, Cr, and Cu in egg content, eggshell and to relate hatching success. Fifteen eggs, four sludge samples, and wastewater samples were collected during September 2016 to April 2017 and were digested and determined using ICP-MS. Clutch size was 1–6 eggs and the weight, width, and length of eggs (mean ± SD) were 11.64 ± 0.91 g., 24.96 ± 0.85 mm, 35.05 ± 1.32 mm, respectively. Hatching success (n = 15) was 96.7%. Cu and Pb were detected at low level in wastewater, and five heavy metals were contaminated in sludge samples. Cu> Cr> Pb> Hg were detected in egg samples (n = 15), and all concentrations in egg content were more than in eggshell. Cr, Cu, Pb, and Hg concentration were not correlated between eggshell and egg content. This study indicated that the heavy metals can be contaminated in sediment, wastewater and can be transferred to egg of Little Grebes. These suggest the ecological risk of heavy metals on other long-living vertebrates living in the area.     

Moisture during Egg Incubation Adversely Affects Hatching of the West Indian Sweetpotato Weevil,Euscepes postfasciatus (Coleoptera: Curculionidae), When the Eggs are Incubated En Masse

We compared the egg-hatching of Euscepes postfasciatus (Fairmaire) with and without moisture treatment to test whether moisture is necessary during egg incubation when a large number of eggs are gathered into a mass. Moisture treatment exhibited significant undesirable effects on hatching (reduction of hatchability, delay of hatching, and increase of the variance of hatching date) compared to no moisture treatment. In addition, moisture treatment significantly increased the incidence of fungus on the egg surface, which can subsequently contaminate the larval artificial diet. Based on these results, we concluded that moisture is not necessary for incubating E. postfasciatus eggs. Two possible explanations for the undesirable effects of moisture on hatching were discussed: a direct effect by preventing respiration of the eggs and an indirect effect through fungal infection of the eggs.     

Sources and timing of calcium intake during reproduction in flycatchers

Calcium availability may limit the reproductive output of birds and snail shells are considered to be the main source of calcium in many passerine species. This study of collared (Ficedula albicollis) and pied (F. hypoleuca) flycatchers evaluates calcium intake of a natural diet in Central Europe, and sex differences in the utilization of experimentally supplemented sources of calcium during the entire breeding period in aviary birds. The study provides the first evidence that successful reproduction of these species depends on the availability of woodlice (Isopoda) and millipedes (Diplopoda). Each of these two components provided about 3 times more calcium than the snail shells contained in a natural nestling diet. The breeding performance of aviary birds was poor when only snail shells and the fragments of eggshells were provided in food, i.e., irregular laying, smaller clutches, eggshell defects (25 of 53 eggs), and eggs dried-up during incubation. In contrast, no defective eggshell or dried-up eggs were found and the overall breeding performance increased 2–3 times when woodlice were added to the food. Females increased their intake of woodlice during both the pre-laying and laying periods, and both sexes did so during the nestling period. Both sexes took more woodlice in the evening than in the morning, independent of the nesting stage. Intake by females was low until 4 days before laying the first egg, then increased to the highest level, dropping immediately after laying the last egg. Intake of woodlice by both sexes increased steadily from hatching until the nestling age of about 10–12 days decreasing thereafter, which corresponds with the period of rapid skeletal growth. In contrast, the intake of mealworms increased until the nestling age of 13–14 days leveling off thereafter which corresponds with the growth curve of nestling body mass.Due to an error in the citation line, this revised PDF (published in December 2003) deviates from the printed version, and is the correct and authoritative version of the paper.     

Ecology and Physiology of Nesting and Early Development of the American Alligator

Ambient temperatures were significantly correlated with alligator(Alligator mississippiensis) nesting activity. Nesting occurredearliest when March-April-May ambient temperatures were highest.Rainfall had no significant relationship with time of nestingactivity although water levels did affect the degree of nesting.The bulk of the nesting took place within a 2-week period eachyear. Nesting chronology and factors affecting nesting were investigatedat Rockefeller Refuge from 1964 through 1987. Courtship activitiesgenerally began in early April and progressively intensifieduntil early June. From late May through the first week of Junecourtship and copulation were intense, the females ovulated,and the high point of spermatogenesis occurred. Factors whichseriously affected nesting success were floods, droughts, andpredation. Salinity levels affected nesting densities alongthe coast. Nesting success (% of nests which produced live hatchlings)averaged 68.3% for 266 nests followed during a 4-yr study. Hatchingsuccess (% of eggs which hatched) averaged 58.2% for 154 nestsfollowed during two nesting seasons. Habitat selection, home ranges, and daily movements were delineatedfor adult males and females. Courtship occurs in deep open waterduring springtime. Adult females selectively seek out densevegetation adjacent to isolated ponds for nesting during thesummer. Adult and sub-adult males tend to prefer open waterall year round and only occasionally venture into the more secludedand heavily vegetated areas used by the females. Home rangesizes and daily movements were much larger for adult males thanfor adult females. Laboratory studies evaluated the effectsof four different egg incubation temperatures on hatching success,sex determination, size at hatching, and growth rates to 18mo of age. The results indicate that incubation temperaturecan have a profound effect on sex ratios, hatchling size, andpost-hatching growth. No significant differences were observedin hatching success at the four different temperatures.     

Egg temperature and initial brood patch area determine hatching asynchrony in Magellanic penguin Spheniscus magellanicus

In birds, the adaptive significance of hatching asynchrony has been under debate for many years and the parental effects on hatching asynchrony have been largely assumed but not often tested. Some authors suggest that hatching asynchrony depends on the incubation onset and many factors have been shown to influence hatching asynchrony in different species. Our objective was to analyze the exact timing of the onset of incubation and if this affects hatching asynchrony; and, in addition, which other factors (brood patch development, incubation position, adult body condition, intra‐clutch egg dimorphism, laying date and year) affect hatching asynchrony in Magellanic penguins Spheniscus magellanicus. We first estimated the eggshell temperature at which embryo development starts, with a non‐destructive and novel method. We then recorded individual egg temperatures in 61 nests during incubation, and related them, and other breeding parameters, to hatching asynchrony. We also observed incubation positions in 307 nests. We found a significant positive relationship between hatching asynchrony and the temperature that the first‐laid egg experienced during egg laying and between hatching asynchrony and the initial brood patch area. We also found a negative relationship between hatching asynchrony and the difference in temperature between second and first‐laid eggs within a clutch, measured after the egg‐laying period was finished. We ruled out position of the eggs during incubation, adult body condition, egg volume, laying date, and study year as factors influencing hatching asynchrony. The egg temperature during laying and the difference in temperature between eggs of a clutch are determinants of hatching asynchrony in Magellanic penguins.     

Some analyses of artificially incubated eggs and hatchlings of Green and Loggerhead sea turtles

Eggs of the Green sea turtle ( Chelonia mydas ) and the Loggerhead sea turtle ( Caretta caretta ) were incubated in coral sand or silica sand moistened with distilled water. The eggs and hatchlings were analysed for calcium, magnesium and phosphorus. There was less magnesium but more calcium in the hatchlings than in the original egg contents but this was not influenced by the type of sand used during incubation. The results are interpreted as indicating that the eggshell is the major source of calcium for the developing embryos. The Logger-head sea turtle shows a greater efficiency in the incorporation of yolk magnesium and phosphorus into the embryo than does the Green sea turtle.     

Embryonic death is linked to maternal identity in the leatherback turtle (Dermochelys coriacea)

Leatherback turtles have an average global hatching success rate of ~50%, lower than other marine turtle species. Embryonic death has been linked to environmental factors such as precipitation and temperature, although, there is still a lot of variability that remains to be explained. We examined how nesting season, the time of nesting each season, the relative position of each clutch laid by each female each season, maternal identity and associated factors such as reproductive experience of the female (new nester versus remigrant) and period of egg retention between clutches (interclutch interval) affected hatching success and stage of embryonic death in failed eggs of leatherback turtles nesting at Playa Grande, Costa Rica. Data were collected during five nesting seasons from 2004/05 to 2008/09. Mean hatching success was 50.4%. Nesting season significantly influenced hatching success in addition to early and late stage embryonic death. Neither clutch position nor nesting time during the season had a significant affect on hatching success or the stage of embryonic death. Some leatherback females consistently produced nests with higher hatching success rates than others. Remigrant females arrived earlier to nest, produced more clutches and had higher rates of hatching success than new nesters. Reproductive experience did not affect stage of death or the duration of the interclutch interval. The length of interclutch interval had a significant affect on the proportion of eggs that failed in each clutch and the developmental stage they died at. Intrinsic factors such as maternal identity are playing a role in affecting embryonic death in the leatherback turtle.     

Effects of Shiny Cowbird Molothrus bonariensis parasitism on different components of House Wren Troglodytes aedon reproductive success

Avian brood parasites, including cuckoos and cowbirds, have multiple negative effects on their hosts. We analysed the effects of Shiny Cowbird Molothrus bonariensis parasitism on different components (e.g. egg losses, hatching success, chick survival and nest abandonment) of House Wren Troglodytes aedon reproductive success. We also conducted an experiment to discriminate between two mechanisms that may reduce hatching success in parasitized clutches: lower efficiency of incubation due to the increase in clutch volume and disruption of host incubation by the early hatching of Cowbirds. Egg puncturing by Shiny Cowbirds reduced host clutch size at hatching by 10–20%, and parasitized nests had a decrease in hatching success of 40–80%. Egg losses and hatching failures were positively associated with the intensity of parasitism. Brood reduction was greater in parasitized nests, but the growth rate of the chicks that fledged was similar to that in unparasitized nests. The combined effects of egg losses, hatching failures and brood reduction decreased the number of fledged chicks by 80%. In addition, egg puncturing increased the likelihood of nest abandonment by Wrens. Experimental data showed that hatching failures occurred when there was a combination of: (1) an increase in the volume of the clutch by the addition of the Cowbird egg without removal of host eggs, and (2) the addition of the Cowbird egg before the onset of incubation. This was relatively common in House Wren nests, as Cowbirds generally parasitize before the onset of incubation. Our results indicate that Shiny Cowbird parasitism imposes a major impact on House Wrens, as it affects all components of the Wren's reproductive success.