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Nest productivity in male-only cared and female-only cared Eurasian Penduline Tit nests from Hungary as offspring development advances . Box plots indicate the same nests in four subsequent reproductive stages (filled boxes: n = 7 male-only cared nests; open boxes: n = 40 female-only cared nests). " Eggs " represent clutch size, and " Nestlings d10 " indicates the number of 10-day-old nestlings. Box plots show the median and 25th and 75th percentiles, whiskers indicate data points within 1.5 interquartile range from the first and third quartiles, and open circles represent outliers that do not fall within this range.
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Diverse patterns of parental care, including uniparental care by either the male or the female, provide excellent opportunities to investigate how variation in social traits is maintained in wild populations. Coexistence of different parental strategies within the same population is expected when they exhibit similar cost-benefit ratios. We investi...
Contexts in source publication
Context 1
... Two generalized linear models (GdLM; with binomial error) of offspring survival are given; GdLM 1 included all 142 nests for which we had data (33 male-only cared [MC] auk, vol. 129 stage (i.e., eggs, hatchlings, 10-day-old nestlings, and fledglings) than MC nests (GLMM, F = 19.75, df = 1 and 42, P < 0.001, n = 47 nests; Table 3 and Fig. 5). The difference in nest productivity be- tween MC and FC nests decreased from clutch size to fledglings as indicated by a significant care-giving sex * reproductive stage in- teraction (F = 6.65, df = 3 and 135, P < 0.001; Table 3). Therefore, FC nests that had twice as many median eggs (Wilcoxon rank sum test, W = 272.5, Z = 4.09, n ...
Context 2
... teraction (F = 6.65, df = 3 and 135, P < 0.001; Table 3). Therefore, FC nests that had twice as many median eggs (Wilcoxon rank sum test, W = 272.5, Z = 4.09, n FC = 40, n MC = 7, P < 0.001, effect size: r = 0.60) yielded the same median number of fledglings as MC nests (W = 174.0, Z = 1.04, n FC = 40, n MC = 7, P = 0.304, effect size: r = 0.15; Fig. 5). There was no difference in nestling size (mean body mass or mean tarsus length) between MC and FC nests (GLM, body mass: F = 0.63, df = 1 and 82, P = 0.428, n = 87 nests; tarsus length: F < 0.01, df = 1 and 82, P = 0.991; Table 4). Nestling mass decreased with egg-laying date and with the number of hatchlings (Table 4), whereas both ...
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Capsule
The experimental removal of either the male or female from breeding pairs of Cinereous Tits Parus cinereus revealed that both sexes could increase their nestling provisioning rate to compensate fully for the loss. However, single females but not males delivered lower quality food.
Aims
Parents would benefit from transferring the burden of...
Citations
... Caring males, however, may enhance the reproductive success of a brood they provide parental care to by producing higher quality offspring. Most studies conducted on this species have already shown the initial difference in clutch size between female-only and male-only cared clutches (Persson and Öhrström 1989;Czyż 2005;Pogány et al. 2012). This initial difference resulted in different productivity of female-only and male-only clutches in some populations being studied (Persson and Öhrström 1989;Czyż 2008), whilst in others, the productivity of nests did not depend on the type of parental care (Pogány et al. 2012). ...
... Most studies conducted on this species have already shown the initial difference in clutch size between female-only and male-only cared clutches (Persson and Öhrström 1989;Czyż 2005;Pogány et al. 2012). This initial difference resulted in different productivity of female-only and male-only clutches in some populations being studied (Persson and Öhrström 1989;Czyż 2008), whilst in others, the productivity of nests did not depend on the type of parental care (Pogány et al. 2012). That means higher brood reduction of female-only compared to male-only cared broods in the latter population, resulting in no difference between the number of fledglings reared in female-only and male-only cared broods. ...
... That means higher brood reduction of female-only compared to male-only cared broods in the latter population, resulting in no difference between the number of fledglings reared in female-only and male-only cared broods. In our study population, brood reduction in female-only cared clutches was lower than in the Hungarian population (medians were 6/5/4 in our population vs. 6/4/3 eggs/nestlings/fledglings, respectively, for the Hungarian population, Pogány et al. 2012) whilst these parameters for male-only cared clutches were similar. These differences in populations could result from many factors, e.g. ...
Conditions encountered during early development affect future survival and reproduction in many bird species. For parents, it is important what body condition the nestlings will achieve at fledging because the condition affects the offspring's chances to survive and reproduce in the future. However, there is a trade-off between the number of nestlings and their condition. We studied parental behaviour and nestling body condition in uniparental Penduline Tits. In this small passerine, the parental care (incubation and food provisioning) is provided by the female only (49% of clutches in the study population) or the male only (15%). In addition, over a third of clutches are deserted by both parents before the start of incubation. We found that female-only cared clutches had more eggs and nestlings and produced more fledglings than male-only cared clutches. The incubation behaviour and incubation temperature in both types of clutches were similar. The provisioning rate per brood was positively, and the provisioning rate per nestling was negatively, correlated with brood size. Although males cared for smaller clutches, parent sex was not significant in both models of provisioning rates (per brood and per nestling). Moreover, the provisioning rate did not predict the brood's average nestling condition. However, nestlings reared in broods with male care were in better condition than those reared by females. At the age of 13 days, they had a higher scaled mass index (describes the relative size of energy reserves) and higher haemoglobin levels. The results suggest that the lower productivity of male-only cared clutches, compared to those cared for only by females, may be compensated by the higher condition of nestlings. Information about the recruitment success of broods cared for by males and females will be necessary to test this prediction.
... After clutch desertion, both males and females may find another partner and rebreed during a single breeding season (Zheng et al. 2018). This breeding system is reminiscent of the one exhibited by Eurasian penduline tits, Remiz pendulinus (Persson and Öhrström 1989;Pogány et al. 2012;van Dijk et al. 2012), although the latter exhibits strictly uniparental care and biparental brood desertion rendering the nest a failure. The behavioral flexibility of Chinese penduline tits provides excellent opportunities to investigate parental decisions of males and females. ...
... The "nest initiation day" was estimated from the building stage when the nest was found (stages A-C, see Zheng et al. 2018), since the number of days spent on different nest-building stages (A, B, or C) were relatively similar for all males (for details on the method of calculating the start day and stages of nest building, see Zheng et al. 2018). On nest initiation day, males start displaying and singing to attract females (Hoi et al. 1996;Pogány et al. 2012;Zheng et al. 2018). We observed each nest with binoculars from a distance of~15 m every 2 days for 20 min as mentioned in Smith (1995) and Bleeker et al. (2005). ...
Parents are expected to make fine-tuned decisions by weighing the benefits of providing care to increase offspring survival against that of deserting to pursue future mating opportunities. A higher incentive for the rarer sex in the population indicates an impact of mating opportunities on parental care decisions. However, in a dynamic breeding system, deserting the offspring and searching for a new mate would influence mating opportunities for both sexes. Sex-specific costs and benefits are expected to influence males’ and females’ parenting strategies in different ways. Here, we investigated Chinese penduline tits, Remiz consobrinus, which exhibit flexible parental care strategies: uniparental care by the male or female, biparental care, and biparental desertion occur in the same population. We show that male penduline tits change their parental behavior over the breeding season; they desert clutches produced early in the season but care for the late season clutches. The change in male parenting behavior is consistent with the seasonal decline in mating opportunities. In contrast, parenting by females did not change over the breeding season, nor was it associated with seasonal variation in mate availability. Taken together, mating opportunities have different associations with parental behavior of male and female Chinese penduline tits. We recommend an inclusion of mating opportunities for both sexes simultaneously in order to understand one of the fundamental decisions in parental care evolution—care or desert.
Significance statement
Divorce is a common feature of both human and nonhuman animal societies. Theoretical studies suggest that one of the drivers of divorce is enhanced mating opportunity, i.e., parents with higher mating opportunities are more likely to abandon their family than those with low mating opportunities. Here, we investigate the dynamics of parental behavior and mating opportunities in a wild population of a small songbird, the Chinese penduline tit Remiz consobrinus. This species exhibits one of the most diverse avian breeding systems wherein both uniparental (male or female) and biparental rearing can be seen in a single population. We show that male penduline tits abandon their offspring in response to enhanced mating opportunities while the female parental behavior remains unaffected. This implies the relationship between mating opportunities and parental care is more complex than currently acknowledged and requires further investigation.
... In Kentish plovers (Charadrius alexandrinus), a precocial bird species, removal experiments also indicated better remating opportunities for females than males (Székely, Cuthill, & Kis, 1999). In contrast, males are more likely to desert in another altricial species, the penduline tit (Remiz pendulinus; Pogány, Dijk, Horváth, & Székely, 2012;van Dijk et al., 2012). ...
Comparative studies have established the necessity for biparental care as an important factor for monogamy in freshwater fish and birds. However, whether two parents are really needed for offspring care remains an open question in many cases. I experimentally studied female and male contributions to offspring care in the white‐browed coucal (Centropus superciliosus), a monogamous and biparental cuckoo with a balanced adult sex ratio, and contrasted it with the sympatric black coucal (C. grillii), a classically polyandrous species with a male‐biased adult sex ratio and male‐only care. To study the necessity for biparental care, I temporarily removed one partner for 2 days to see whether the remaining parent compensated for the absence of its partner. Both female and male white‐browed coucals approximately doubled their feeding rates when their partner was absent, thus fully compensating the number of feeding visits to the nest. However, nestlings maintained their growth only, when males were present and females were removed. When males were removed and only females were present, nestling growth declined. Hence, only male white‐browed coucals fully compensated for the temporary loss of the partner, suggesting that females could benefit most from nesting with additional males—if these should become available. Removing female black coucals had no consequence for nestling feeding rates of male black coucals. But male black coucals had to be returned to their territories within a few hours to avoid harming the brood because female black coucals typically would not commence feeding their offspring. In conclusion, the breeding system of white‐browed coucals seems quite flexible and the relatively balanced adult sex ratio may stabilize monogamy in this species. Should ecological factors ever favour a stronger bias in the adult sex ratio towards males, female white‐browed coucals may easily become polyandrous and relinquish parental care entirely to males.
... In Chinese Penduline Tits, the resolution of sexual conflict may differ from that of the Eurasian Penduline Tit. Clutch desertion occurred at the egg-laying stage in Eurasian Penduline Tits, which directly induced nearly twice the clutch size in femaleonly care nests than that in male-only care (Persson and Öhrström 1989;Pogány et al. 2012). It has been argued that by deserting the smaller clutches and leaving them for Schleicher et al. 1993;Szentirmai et al. 2007; males to care, females can relatively reduce her partner's reproductive success ). ...
The Chinese Penduline Tit (Remiz consobrinus) is a small passerine that breeds in Northeast Asia. Despite its common appearance and its high encounter rate, the breeding biology of this species has remained largely unknown. In this paper, we describe for the first time the breeding biology and parental care system of the Chinese Penduline Tit in the coastal area of Northeast China. As a migratory species of birds, it arrives at the Liaohe Delta in late April, the males establish territories in early May, and breeding pairs build their nests from May to July. Most nest building occurs from mid-May to early June. The females produce small white eggs (egg mass, 1.0 ± 0.1 g) and a clutch size of 6.8 ± 0.6 eggs. The parental care system is complicated in Chinese Penduline Tits: uniparental care is dominant (80%) in this population, with a majority (71%) of female-only care and 9% male care for all brooding nests, together with 4% biparental care and 16% biparental desertion. Chicks hatch after 13.9 ± 1.0 days of incubation and fledge after 22.1 ± 1.0 days. The hatching success and nestling survival are 86.7 and 80.6%, respectively. Furthermore, breeding failure is 23.4%, which is commonly caused by predation and nests blown away by strong winds. Compared to other species of the Remiz genus, the Chinese Penduline Tit shows much similarity to the Eurasian Penduline Tit (Remiz pendulinus) but large differences compared to the White-crowned Penduline Tit (Remiz coronatus) with respect to breeding habitat, breeding density, nest site selection and parental care system. Moreover, the good nest attributes, relatively high nestling survival and low risk of nest failure may contribute to the prevalence of the uniparental care system in this Chinese Penduline Tit population.
... Most of the evidence for social environmental effects comes from negotiation models of parental care which suggest that the amount of care of the focal individual is not independent of other family members (McNamara et al. 1999;Hinde and Kilner 2007). Empirical studies report various aspects of parenting to vary in line with these negotiation models, including incubation and brooding time (AlRashidi et al. 2010;Pogány et al. 2012;Delia et al. 2013), frequency and length of food provisioning (Amat 1995;Lycett et al. 1998;Dawson and Bortolotti 2002;Eldegard and Sonerud 2010), facultative male parental care (Whittingham and Robertson 1994), length of the post-fledging dependence period and provisioning (Ferrer 1992;Zarybnicka 2009) and energy expenditure (Jodice et al. 2002). ...
In sexually reproducing species, parents and offspring have different optima in terms of the amount of parental investment. For offspring, higher investment than the parental optimum generally increases their fitness. However, such higher investment will, in theory, result in net parental fitness loss because increased benefits from current offspring will be more than offset by a potential decrease in parental survival or future reproduction. Whether and how parents respond to a shortfall in resources might therefore have important fitness consequences. Here, we manipulate the nutritional condition of captive zebra finch families to investigate how variation in environmental conditions influences parental resource allocation between themselves and their offspring. By allowing the same zebra finch pairs to raise one brood under high and another brood under low nutritional conditions, we found that parents lost more body mass and their offspring grew at slower rates under low nutritional conditions. Therefore, both parents and their offspring were affected by the nutritional treatments. Offspring incurred greater costs, because slower growth rates under low nutritional conditions resulted in an overall lower body mass of nestlings after reaching independence. Nutritional treatments had sex-specific effects on parental body mass, because, in contrast to fathers, only mothers recouped their losses under high nutritional conditions. Furthermore, both parents spent less time brooding their nestlings under low nutritional conditions. Parental strategies hence vary with the nutritional quality of their food, even in captivity, supporting the idea that parental resource allocation may be driven by the expected costs and benefits from current offspring.
Significance statement
Parental care is among the most influential traits affecting fitness in many sexually reproducing animals. The more care offspring receive, the higher their chances of survival and reproduction. However, caring for their young does not only provide benefits for parents: all costs of parental care are paid by them. Parental resource allocation is therefore expected to be optimised. But how should parents respond to changing environmental conditions when they are already breeding? By manipulating nutritional conditions in a captive population of zebra finches Taeniopygia guttata, we show that both parents and their offspring lose body mass under low nutritional conditions. The effect on offspring was more pronounced and long lasting, and parents also decreased their parental effort. Our results show plasticity in parental investment of zebra finches in response to nutritional conditions and reproductive value of their offspring.
... Previous studies frequently monitored time spent in the nest and number of nest visits during incubation and/or nestling provisioning (e.g. [8,[9][10][11][12]). In the majority of these studies the nest is either directly observed, or more frequently, video-recorded for behavioural coding from the inside or outside for a pre-defined period at a standardized time (i.e. starting at a given time of a given day of the reproductive stage). ...
... Finally, proportion of male feedings were calculated as the ratio of the number of feedings by the male to the sum of feedings by the male and female, to control for between-nest differences in number of feeding visits arising from differences in brood size (cf. [11]). Zebra finches feed their nestlings by regurgitating food a number of times during each visit to the nest, and we quantified these from the recordings taken from the inside of the nest box. ...
Parental care plays a key role in ontogeny, life-history trade-offs, sexual selection and intra-familial conflict. Studies focusing on understanding causes and consequences of variation in parental effort need to quantify parental behaviour accurately. The applied methods are, however, diverse even for a given species and type of parental effort, and rarely validated for accuracy. Here we focus on variability of parental behaviour from a methodological perspective to investigate the effect of different samplings on various estimates of parental effort. We used nest box cameras in a captive breeding population of zebra finches, Taeniopygia guttata, a widely used model system of sexual selection, intra-familial dynamics and parental care. We investigated diurnal and reproductive stage-dependent variation in parental effort (including incubation, brooding, nest attendance and number of feedings) based on 12h and 3h continuous video-recordings taken at various reproductive stages. We then investigated whether shorter (1h) sampling periods provided comparable estimates of overall parental effort and division of labour to those of longer (3h) sampling periods. Our study confirmed female-biased division of labour during incubation, and showed that the difference between female and male effort diminishes with advancing reproductive stage. We found individually consistent parental behaviours within given days of incubation and nestling provisioning. Furthermore, parental behaviour was consistent over the different stages of incubation, however, only female brooding was consistent over nestling provisioning. Parental effort during incubation did not predict parental effort during nestling provisioning. Our analyses revealed that 1h sampling may be influenced heavily by stochastic and diurnal variation. We suggest using a single longer sampling period (3h) may provide a consistent and accurate estimate for overall parental effort during incubation in zebra finches. Due to the large within-individual variation, we suggest repeated longer sampling over the reproductive stage may be necessary for accurate estimates of parental effort post-hatching.
... The higher adult mortality and larger clutch size of the Cape Penduline Tit does not fit the pattern of relatively low adult mortality and small clutch sizes typical of many southern hemisphere birds (Martin 1996;Martin et al. 2000Martin et al. , 2006 and of the rest of the breeding bird community in the study area (Lloyd et al. 2014). These trait patterns are inconsistent with the seasonality and food limitation hypothesis, but are consistent with the adult mortality hypothesis, which predicts that species with higher adult mortality should have larger clutches Staav (1998), Cramp et al. (1993 and Persson and Ö hrström (1989) b Pogány et al. (2012) and Szentirmai et al. (2007) c Taylor (1971), Austin (1977) and Klimkiewicz et al. (1983) d George (1987) e This study, Lloyd et al. (2014) and Lloyd and Martin (2016) f Climate data sourced from http://www.worldweatheronline.com g Excludes nests deserted by both parents prior to incubation h Measured at age 10 days in EPT (n = 29) and at age 8-12 days in CPT (n = 13) (Williams 1966;Martin 1996Martin , 2015Martin et al. 2000;Ghalambor and Martin 2001). Given the higher adult mortality of the Cape Penduline Tit compared with that of the Eurasian Penduline Tit based on maximum longevities, a key prediction of the adult mortality hypothesis is that the Cape Penduline Tit should exhibit higher nestling feeding rates, but lower parental investment per offspring. ...
... While both sexes contribute to nest building in all species, gender roles diverge thereafter. The Eurasian Penduline Tit pair members compete to desert the nest first towards the end of egg-laying, leaving the other member of the pair to incubate and raise the young alone while the deserting individual seeks additional breeding opportunities (Pogány et al. 2012), a strategy that may allow a greater number of nesting attempts within the short breeding season (Persson and Ö hrström 1989). Verdin females incubate and tend the young in their first week alone, whereafter the male contributes equally to nestling feeding (Taylor 1971). ...
We studied the breeding biology of the south temperate Cape Penduline Tit (Anthoscopus minutus) in order to compare its life history traits with those of related north temperate members of the family Remizidae, namely the Eurasian Penduline Tit (Remiz pendulinus) and the Verdin (Auriparus flaviceps). We used this comparison to test key predictions of three hypotheses thought to explain latitudinal variation in life histories among bird species—the seasonality and food limitation hypothesis, nest predation hypothesis and adult mortality hypothesis. Contrary to the general pattern of smaller clutch size and lower adult mortality among south-temperate birds living in less seasonal environments, the Cape Penduline Tit has a clutch size larger than that of the Verdin and similar to that of the Eurasian Penduline Tit, and higher adult mortality than both of the other two species. The most notable difference between the Cape Penduline Tit and the two other species is in parental behavioural strategy, with the former exhibiting bi-parental care at all stages of nesting together with facultative cooperative breeding, whereas the Eurasian Penduline Tit has uni-parental care and the Verdin has a combination of female-only incubation but bi-parental nestling care. Consequently, in comparison to the other two species, the Cape Penduline Tit exhibits greater nest attentiveness during incubation, a similar per-nestling feeding rate and greater post-fledging survival. Its relatively large clutch size, high parental investment and associated high adult mortality in a less seasonal environment are consistent with key predictions of the adult mortality hypothesis but not with key predictions of the seasonality and food limitation hypothesis in explaining life history variation among Remizidae species. These results add to a growing body of evidence of the importance of age-specific mortality in shaping life history evolution.
... Care type and reproductive stage were established by daily observations of attending parent(s), and recording clutch size daily during egg-laying (van Dijk et al., 2007a). Since male-only care is less frequent than female-only care (14% vs. 47%; Pogány et al., 2012;Persson and Öhrström, 1989), we compared biparentally deserted with female-only cared for eggs in this experiment. We collected 78 eggs from 37 nests (18 biparentally deserted and 19 female-only nests). ...
... Deserted eggs were collected from inactive nests where no bird was seen at the nest for at least 20 min on two consecutive days (van Dijk et al., 2007a). 2 × 20 min total observation time is sufficient to assign desertion by the male, the female or both parents in this species, as males and females appear within 3.7 ± 5.2 min (mean ± SD) and 6.8 ± 7.5 min, (respectively) from start of observation at active nests (van Dijk et al., 2007a). In addition, none of the birds classified as 'deserted' based on this method returned to their nests afterwards during any of our field studies to date including this study (e.g., Pogány et al., 2012;van Dijk et al., 2007avan Dijk et al., , 2010a. Cared for eggs were collected during the course of egg-laying (on day four or five) after male desertion was confirmed (i.e., only the female was present at the nest during two consecutive checks). ...
... Because of the suboptimal conditions for hatching in the incubator the hatching success was lower than in nature (33% as opposed to 73% in nature, the latter based on data collected for Pogány et al. (2012)). Only 8 eggs hatched in the deserted, and 12 in the cared for experimental groups, therefore hatching success was not analysed further. ...
... Using data from a Hungarian population of penduline tits, van Dijk et al. (2012) estimated the average seasonal reproductive success of males and females given their decision in their first breeding attempt. These population averages suggest that there are two alternative ESSs, female-only and male-only care. ...
In his paper 'Parental investment: a prospective analysis', Maynard Smith (1977, Animal Behaviour, 25,1-9) introduced a game-theoretic approach to understanding the evolution of parental behaviour and addressed the broad issue of which sex should provide care for the young. This paperwas important in that it introduced the use of game theory to the analysis of parental care. It also stimulated empirical work on care. We identify progress that has been made since the publication of the paper. In particular, although Model 2 of Maynard Smith (1977) has been used in several textbooks to explain the evolution of care, subsequent work has shown that this model is not built on a consistent view of how parental care influences future reproductive success through its effect on the sex ratio. Several models incorporate a consistent account in which opportunities to remate after desertion emerge from the analysis, rather than being specified in advance. More generally, it is not possible to consider parental care in isolation from factors such as paternity, mating preferences and mate choice behaviour. We identify various theoretical and empirical issues in the area of parental care research that we believe deserve further study if our understanding of care decisions is to advance. Taken together, the landmark paper of Maynard Smith (1977) stimulated new theoretical and empirical studies in parental care research and led to new insights into the behavioural interactions between males and females.
... In many bids species like, for instance, kentish plovers (Charadrius alexandrinus) or Eurasian penduline tits (Remiz pendulinus), flexible mating strategies exist, and plasticity in mating behaviours is viewed as an adaptive response to varying environmental and social contexts (Valera et al., 1997;Amat et al., 1999;Székely et al., 1999;van Dijk et al., 2007van Dijk et al., , 2010Pogány et al., 2008Pogány et al., , 2012. Rock sparrows exhibit a wide array of mating patterns including monogamy and multiple forms of polygamy such as sequential or simultaneous polyandry (Pilastro et al., 2001;Tavecchia et al., 2002;Griggio et al., 2003;. ...
Relatively few bird species show complex social mating systems whose preponderance in a population is likely to affect the patterns of parental care observed there. In turn, parental investment is likely to be related to the expression of certain ornaments, which may reveal information on the bearer's individual quality. Here we address both issues in a species characterized by several forms of parental care (both biparental and uniparental care) and in which both sexes possess a yellow breast patch, the rock sparrow (Petronia petronia). In our colony, males contributed more to the care of the young in comparison with other populations. Social monogamy was the most frequent mating option and the percentage of cases of female (or male) brood desertion was lower with respect to that reported in previous studies, suggesting a
flexible behaviour of this species to deal with different social environments. Birds did not pair assortatively with respect to the size of the yellow breast patch and we found no significant relationship between this secondary sexual trait and the frequency with which parents provisioned their chicks. However, we observed a positive relationship between male yellow patch size and nestling tarsus length, which suggests that more
ornamented males are better parents. Males, but not females, differentially allocated parental investment in response to female ornamentation, although the benefits that males may gain from choosing more attractive females remain unidentified. Our results on paternal care investment along with previous studies on this species, reinforcing the view that the rock sparrow constitutes a good model to study sexual conflict over parental care under different social environments.
