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1.
Why are birds’ eggs speckled?   总被引:8,自引:0,他引:8  
Birds are unique in laying eggs with pigmented shells, but for most species (e.g. most passerines, which lay white eggs speckled with reddish spots of protoporphyrin) the pigments’ function is unknown. We studied a bird population at a geologically variable site, and considered a hitherto untested hypothesis: that protoporphyrin pigments might compensate for reduced eggshell‐thickness (caused partly by calcium deficiency), which is known to reduce eggshell‐strength and increase eggshell‐permeability. We found that pigment spots specifically demarcated thinner areas of shell, with darker spots marking yet thinner shell than paler spots. Variation in pigmentation was thus associated with variation in shell thickness both within and between clutches, so accounting for the eggshell's characteristic spot patterns. Geological variability at this site has resulted in a great range of calcium availability and, as predicted by the hypothesis, variation in calcium availability was found to affect between‐clutch variation in both eggshell‐mass (+) and pigmentation characteristics (?). We suggest a physiological mechanism and some important implications of these findings.  相似文献   

2.
Higham JP  Gosler AG 《Oecologia》2006,149(4):561-570
Many small passerine birds worldwide lay white eggs speckled with red, brown and black protoporphyrin pigment spots (maculation). Unlike some patterns of avian eggshell pigmentation which clearly serve a crypsis or signalling function, the ubiquity of maculation among passerines suggests that its origins lie in another function, not specific to any particular ecological or behavioural group. Elsewhere, we have presented evidence that protoporphyrin pigments serve a structural function related to eggshell thickness and calcium availability: eggshell maculation in the great tit Parus major increases with decreasing soil calcium levels, pigments demarcate thinner areas of shell, and both the pigment intensity and distribution are related to shell thickness. Here we show that maculation also affects the rate of water loss from the egg during incubation (≈ Mass Loss per Day or MLD, which is critical to egg viability), but not that of unincubated eggs. We also demonstrate, both by observation and experiment, that the effect of female incubation behaviour on MLD compensates in some way for variation in egg characteristics, and that differences between females in the degree of such compensation are related to differences in clutch maculation. Our results suggest that, while a principal function of maculation in this species may be to strengthen the eggshell, it may also reduce eggshell permeability when large amounts of pigment are used, and that this necessitates a behavioural adjustment from the female during incubation. We discuss these findings and make further testable predictions from our model.  相似文献   

3.
Inter‐ and intraspecific variation in eggshell colouration has long fascinated evolutionary biologists. Among species, such variation may accomplish different functions, the most obvious of which is camouflage and background matching. Within species, it has been proposed that inter‐female variation in eggshell pigmentation patterns can reflect egg, maternal or paternal traits and hence may provide cues to conspecifics about egg, maternal or paternal phenotypic quality. However, the relationship between protoporphyrin‐based eggshell pigmentation and egg or maternal/paternal traits appears to be highly variable among species. We investigated patterns of intraspecific variation in Eurasian barn swallow Hirundo r. rustica protoporphyrin‐based eggshell pigmentation, and analysed its association with egg and clutch characteristics, maternal/paternal phenotypic traits and parental feeding effort. Eggshell pigmentation pattern significantly varied between breeding colonies, was significantly repeatable in first clutches laid by the same females in different years (intraclass correlation coefficient ranging between 0.56 and 0.63), but it was not significantly associated with egg traits, such as position in the laying sequence, egg mass, yolk testosterone concentration and antioxidant capacity. It was weakly or non‐significantly associated with female and male traits (sexual ornaments), but females laying darker (higher pigment intensity) first clutches had higher hatching success, suggesting that eggshell pigment intensity may predict fitness. Male nestling feeding effort was not predicted by eggshell pigmentation. In addition, females with darker breast plumage colouration (a melanin‐based trait related to fitness) laid highly protoporphyrin‐covered eggs, suggesting the presence of a previously unappreciated link between protoporphyrin biosynthesis and plumage melanisation. Moreover, the proportion of male offspring increased in clutches originating from highly protoporphyrin‐covered eggs, suggesting that parents could acquire visual cues about their future brood sex composition before egg hatching. Our results support the idea that intraspecific signalling via eggshell pigmentation is a species‐specific rather than a general feature of avian taxa.  相似文献   

4.
One of the hypotheses to explain egg colour variation in birds lays in the context of sexual selection, where egg colour may signal the female’s physical condition and antioxidant capacity. We tested one of the assumptions following from this hypothesis, that eggshell pigment deposition should be limited for females. The study was conducted in a captive house sparrow (Passer domesticus) population over several years under constant environmental conditions. This multi-brooded species lays eggs which vary in ground colour (biliverdin pigment) and in the intensity and distribution of brownish-red spots (protoporphyrin pigment). Spot darkness, spread and ground colour diminished along the laying sequence, suggesting that the deposition of both pigments was limiting for females over the short term. Also, the proportion of eggs with biliverdin diminished in consecutive clutches laid by the same female over the breeding season, suggesting a long-term cost of biliverdin deposition. On the other hand, spots were darker at the end of the breeding season, indicating that protoporphyrin deposition was probably not limited over the long term. This result could indicate a lower capacity for calcium deposition over the long term, which was compensated for by darker spots. Female age also significantly affected the proportion of bluish eggs and spot patterns. Egg pigmentation decreased with age, indicating that senescing passerine females lay less pigmented eggs. Clutch size was positively related to the proportion of bluish eggs and to spot patterns (darker and more evenly spotted). These results are in accordance with assumptions for the sexual selection hypothesis.  相似文献   

5.
According to the structural-function hypothesis, the eggshell pigment protoporphyrin, deposited at weak spots, can strengthen the shell structure when calcium is lacking in avian species. However, this hypothesis has not been tested in species that produce pigmented eggs of uniform ground colour without spotting patterns. We tested the structural-function hypothesis using 435 eggs produced on seven calcium diets (0.2–4.5%) given to Common Pheasants Phasianus colchicus, a species that produces a large number of eggs on a low-calcium diet with unspotted eggshells composed of a uniform ground colour of mainly protoporphyrin. We found that pheasants on the lowest calcium diet (0.2%), which had thinner eggshells, produced eggs containing more protoporphyrin-based coloration than four of six other diets, suggesting this species employs pigmentation as ground colour to strengthen eggshells when available calcium is low. Our results provide the first, at least partial support for a structural function for eggshell pigments producing ground colour without spotting in a species that is often calcium-limited. This pattern may be more widespread in other ground-nesting taxa that also produce large numbers of eggs with protoporphyrin-based ground colour and are potentially limited by calcium during breeding.  相似文献   

6.
S. JAMES REYNOLDS 《Ibis》2001,143(2):205-215
Many small passerines forage intensively for calcium-rich foods during laying. Increased incidences of shell defects in eggs of small passerines have been reported, particularly in western Europe, and these have been explained in terms of declining calcium availability in soils, resulting from prolonged anthropogenic acid deposition. Studies in the field have provided laying birds, nesting in areas of low calcium availability, with calcium supplements. An alternative approach was adopted in this study by allowing captive Zebra Finches Taeniopygia guttata to lay first clutches on ad libitum calcium, switching them to a low calcium diet for 72 hours for the formation of all but the first egg of the second clutch and reinstating ad libitum calcium for the final clutch. Control females had access to ad libitum calcium for all three clutches. Clutch sizes did not vary significantly between birds on low calcium and controls. The former took over three days longer to lay clutch 3 than did controls but the difference was not statistically significant. Birds on low calcium laid eggs that declined in shell ash mass with laying sequence, indicating that birds may have been calcium-limited. Although not statistically significant, eggshell thickness also declined with laying sequence in clutches laid by females on low calcium. The remaining egg measurements (shell mass, shell surface area and volume] of clutches laid by birds on low calcium did not differ significantly from those of controls. Furthermore, females on low calcium did not resort to skeletal reserves to provide sufficient calcium for egg formation. Dietary calcium appears to be of paramount importance in providing sufficient calcium for clutch formation.  相似文献   

7.
Many passerine birds lay white eggs with reddish brown speckles produced by protoporphyrin pigment. However, the function of these spots is contested. Recently, the sexually selected eggshell coloration (SSEC) hypothesis proposed that eggshell color is a sexually selected signal through which a female advertises her quality (and hence the potential quality of her future young) to her male partner, thereby encouraging him to contribute more to breeding attempts. We performed a test of the SSEC hypothesis in a common passerine, the great tit Parus major. We used a double cross-fostering design to determine whether males change their provisioning behavior based on eggshell patterns they observe at the nest. We also tested the assumption that egg patterning reflects female and/or offspring quality. Because birds differ from humans in their color and pattern perception, we used digital photography and models of bird vision to quantify egg patterns objectively. Neither male provisioning nor chick growth was related to the pattern of eggs males observed during incubation. Although heavy females laid paler, less speckled eggs, these eggs did not produce chicks that grew faster. Therefore, we conclude that the SSEC hypothesis is an unlikely explanation for the evolution of egg speckling in great tits.  相似文献   

8.
The expression and impact of maternal effects may vary greatly between populations and environments. However, little is known about large‐scale geographical patterns of variation in maternal deposition to eggs. In birds, as in other oviparous animals, the outermost maternal component of an egg is the shell, which protects the embryo, provides essential mineral resources and allows its interaction with the environment in the form of gas exchange. In this study, we explored variation of eggshell traits (mass, thickness, pore density and pigmentation) across 15 pied flycatcher populations at a large geographic scale. We found significant between‐population variation in all eggshell traits, except in pore density, suggesting spatial variation in their adaptive benefits or in the females’ physiological limitations during egg laying. Between‐ population variation in shell structure was not due to geographic location (latitude and longitude) or habitat type. However, eggshells were thicker in populations that experienced higher ambient temperature during egg laying. This could be a result of maternal resource allocation to the shell being constrained under low temperatures or of an adaptation to reduce egg water loss under high temperatures. We also found that eggshell colour intensity was positively associated with biliverdin pigment concentration, shell thickness and pore density. To conclude, our findings reveal large‐ scale between‐population variation of eggshell traits, although we found little environmental dependency in their expression. Our findings call for further studies that explore other environmental factors (e.g. calcium availability and pollution levels) and social factors like sexual selection intensity that may account for differences in shell structure between populations.  相似文献   

9.
Egg characteristics have been used to determine egg maternity for birds in situations where two or more females lay eggs in a single nest (as in communal breeders and intraspecific parasites). We assessed the applicability of egg morphometry and eggshell appearance in ascribing egg ownership in communal clutches of guira cuckoos Guira guira , a species where up to seven females may lay eggs in a joint nest. We used both combined variables (including egg mass, length, width, shape and two eggshell variables), and a shape index to test whether eggs laid by each female were similar but different from eggs laid by other females. Also, we conducted discriminant function analyses to verify if eggs could be correctly classified to their mothers based on their characteristics. The correct maternity was determined by yolk protein electrophoresis of freshly-laid eggs. Individual female chutches were separated through egg characteristics or shape alone in 29% and 41% of the groups tested, respectively. Differences were mostly due to a single female that differed from her nest-mates in a unique egg variable. On average, 55% of the eggs analyzed were not assigned to the correct mother using egg dimensions and eggshell speckling pattern. In conclusion, the egg characteristics used do not reliably indicate maternity in guira cuckoo communal clutches. We therefore recommend a protein-based verification of egg appearance characteristics for assigning maternity in other species in which multiple female laying occurs.  相似文献   

10.
The striking variation in colour and maculation of bird eggs has fascinated biologists since centuries, and many hypotheses based on mechanical, physiological or signalling functions have been proposed for its evolution by natural and sexual selection. Protoporphyrin is the main eggshell pigment found in brown maculated eggs, and is assumed to function as a mediator of these selection processes. It is a precursor of heme with pro‐oxidant properties, and hence a link between brown maculation and female condition has been proposed and tested in a number of studies, albeit with contrasting results. A variety of different visual methods have been used to quantify outer eggshell pigmentation, which has been assumed to correspond to overall quantity of protoporphyrin in the shell. Yet, this relationship has rarely been tested. The aim of this study was to apply four commonly used methods to assess pigmentation in great tit eggs with protoporphyrin as the predominant eggshell pigment, and to compare the results of these methods. Specifically, we 1) ranked eggshell pigmentation by human naked eye, 2) applied a granularity approach and 3) measured spectrophotometric reflectance of eggshell pigments. Second, we estimated the relationship between outer eggshell pigmentation (i.e. estimated by three different methods above) and true protoporphyrin concentration deposited in the entire shell measured by HPLC. Among‐method estimates were significantly correlated for the traits describing pigment ‘darkness’ only. While the model including scores based on human naked eyes explained 16% of the variance of pigment concentration in the entire shell, spectrometry explained 27%, and the granularity approach explained 40%. Thus, the estimation of true pigment concentration in the entire shell from the visible outer side of the shell is most reliable with the granularity approach. It is relevant for studies where the maintenance of the integrity of the eggs is essential.  相似文献   

11.
De Coster G  De Neve L  Lens L 《Oecologia》2012,170(2):297-304
Many passerine species lay eggs that are speckled with dark protoporphyrin pigmentation. Because protoporphyrin is mainly derived from the blood, we here formulate and test a new hypothesis that links an increase in anaemia along the laying sequence to within-clutch variation in egg pigmentation. More intense pigmentation is expected if pigments accumulate during enhanced red blood cell production in response to anaemia. Reduced pigmentation is expected if pigments are derived from the degradation of red blood cells that circulate in smaller numbers due to blood loss. To test this hypothesis, we manipulated anaemia in great tit (Parus major) females by infesting the nests with hen fleas (Ceratophyllus gallinae) prior to egg laying. Polychromatophil (i.e., immature red blood cells) percentage, as a measure of blood cell production, was positively correlated with parasite load confirming that female great tits experienced stronger anaemia when infested with haematophagous parasites during egg laying. We found a positive relationship between spot darkness and laying order that weakened under high parasite load. This result suggests that anaemia in females due to blood-sucking parasites led to diminished protoporphyrin from disintegrated red blood cells and hence a decreased deposition of protoporphyrin. However, the overall increase in pigment darkness along the laying sequence suggests that pigments also accumulate by enhanced red blood cell production caused by anaemia due to egg production itself.  相似文献   

12.
Avian eggshell color seems to fulfill multiple functions, some of them being structural and others signaling. In this study, we tested whether or not eggshell coloration may play a role in sexual selection of Tree Sparrows (Passer montanus). According to the “Sexually selected eggshell coloration” hypothesis, eggshell coloration signals female, egg or chick quality and males adjust parental investment according to this signal. Eggs of this species are covered with brown spots and patches, and variation between clutches is high. We found that eggshell coloration correlates with both protoporphyrin and biliverdin, but protoporphyrin concentrations are ten times higher. Eggshell coloration reflects egg and offspring quality, but not female quality. Thus, eggshell coloration may signal female postmating investment in offspring rather than female quality. Furthermore, differential allocation in terms of maternal investment is supported by the fact that females lay more pigmented clutches when mated to males with bigger melanin‐based ornaments relative to their own. Moreover, males invested proportionally more to chicks that hatched from more pigmented clutches. Our correlative results thus seem to support a role of sexual selection in the evolution of eggshell coloration in birds laying brown eggs, pigmented mainly by protoporphyrin.  相似文献   

13.
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.  相似文献   

14.
During egg laying, females face a trade‐off between self‐maintenance and investment into current reproduction, since providing eggs with resources is energetically demanding, in particular if females lay one egg per day. However, the costs of egg laying not only relate to energetic requirements, but also depend on the availability of specific resources that are vital for egg production and embryonic development. One of these compounds are carotenoids, pigments with immuno‐stimulatory properties, which are crucial during embryonic development. In this study, we explore how carotenoid availability alleviates this trade‐off and facilitates egg laying in a small bird species, the blue tit (Cyanistes caeruleus). Blue tits have among the largest clutch size of all European passerines and they usually lay one egg per day, although laying interruptions are frequent. We performed a lutein supplementation experiment and measured potential consequences for egg laying capacity and egg quality. We found that lutein‐supplemented females had less laying interruptions and thus completed their clutch faster than control females. No effects of treatment were found on the onset of egg laying or clutch size. Experimentally enhanced carotenoid availability did not elevate yolk carotenoid levels or egg mass, but negatively affected eggshell thickness. Our results provide hence evidence on the limiting role of carotenoids during egg laying. However, the benefits of laying faster following lutein supplementation were counterbalanced by a lower accumulation of calcium in the eggshell. Thus, even though single components may constrain egg laying, it is the combined availability of a range of different resources which ultimately determines egg quality and thus embryonic development.  相似文献   

15.
The blue-green eggshell pigment biliverdin is an efficient antioxidant, the deposition of which may be costly and act as a handicap to the female's antioxidant capacity. Thus, it is hypothesized that there exists a positive correlation between the degree of eggshell coloration and the condition and health state of the mother. In this study, we measured the blue-green chroma of collared flycatcher Ficedula albicollis eggs, and investigated the associations between egg colour and parental characteristics, laying date, clutch size and laying order. We also analysed the possible correlations between egg size, yolk carotenoid and antibody levels and eggshell pigmentation. Our results revealed no significant relationships between egg colour and female age, morphological and serological variables. Blue-green colour intensity of eggs increased throughout the breeding season, which could be the result of either a proximate constraint or an adaptive strategy. We found a positive association between egg colour saturation and yolk lutein concentration. This relationship could be the result of the seasonal improvement of caterpillar supply, which could influence the deposition of both components into the egg. Furthermore, we found that females mated to lower-quality males produced eggs with higher biliverdin pigmentation. It is possible that deposition of biliverdin is correlated to some unmeasured aspect of egg quality that could enhance the viability of offspring of lower-quality fathers. We found that intensity of blue-green coloration increased with laying order between the first and fifth eggs.  相似文献   

16.
Current research into the evolution and adaptive function of avian eggshell pigmentation, including maculation, has focused mostly on signalling‐based and structural function hypotheses but ignored the potential consequences of shell pigmentation for the developing avian embryo, especially in moderating the embryo's interaction with its light environment. The exposure of the eggs to sunlight that frequently accompanies avian incubation behaviour is one of the major evolutionary steps setting apart birds and reptiles, and coincides with the appearance of eggshell pigmentation. This suggests that shell pigments could play a major role in ensuring the successful development of the avian embryo. We propose that the effects of shell pigments on the egg contents should be considered in addition to established hypotheses of shell pigmentation such as crypsis, egg recognition or a possible structural function. This approach has the potential to identify trade‐offs between different pigment functions and to resolve some of the long standing paradoxa in the evolution of eggshell colour, such as the occurrence of conspicuous blue eggs in passerines or the secondary evolution of white eggshells in cavity nesters. In particular, we identify seven hypotheses, which address how the interaction of eggshell pigments and the light environment may influence embryonic development. These hypotheses are the: thermo‐regulation; UV‐B protection; photo‐acceleration; lateralization; circadian rhythm; photo‐reactivation; and antimicrobial defence. We believe that the understanding of eggshell pigmentation will greatly benefit from taking these hypotheses into consideration when studying the functional significance of eggshell pigmentation and suggest a number of promising directions for future experimental and comparative research.  相似文献   

17.
Reproduction is a critical period for birds as they have to cope with many stressful events. One consequence of an acute exposure to stress is the release of corticosterone, the avian stress hormone. Prolonged stress can have negative impacts on the immune system, resulting in, for example, increased oxidative stress. Through maternal effects, females are known to modulate their investment in eggs content according to their own physiological condition. Less is known about maternal investment in eggshells, especially in pigments. The two main eggshell pigments may possess opposite antioxidant properties: protoporphyrin (brown) is a pro-oxidant, whereas biliverdin (blue-green) is an antioxidant. In Japanese quail, we know that the deposition of both pigments is related to female body condition. Thus, a chronic stress response may be reflected in eggshell coloration. Using female Japanese quails that lay brown-spotted eggs, we explored whether physiological exposure to corticosterone induces a change in female basal stress and antioxidant factors, and eggshell pigment concentration, spectrophotometric reflectance, and maculation coverage. We supplemented adult females over a 2 week period with either peanut oil (control) or corticosterone (treatment). We collected pre- and post-supplementation eggs and analysed the effect of corticosterone treatment on female physiology and eggshell appearance parameters. Except for corticosterone-fed birds which laid eggs with brighter spots, supplementation had no significant effect on female physiology or eggshell pigment concentration, reflectance and maculation. The change in eggshell spot brightness was not detected by a photoreceptor noise-limited color opponent model of avian visual perception. Our data confirms that eggshell reflectance in spotted eggs varies over the laying sequence, and spot reflectance may be a key factor that is affected by females CORT exposure, even if the changes are not detected by an avian visual model.  相似文献   

18.
Most penguin species lay two eggs 3–4 days apart that hatch 1–2 days apart. Hypotheses to explain the shorter incubation period for the second egg include differences in porosity of the eggshells. Eggs with more or larger pores and/or thinner shells have higher gas exchange rates and faster embryonic development. We used eggshells from hatchling Magellanic Penguins Spheniscus magellanicus to test whether eggshell thickness and pore density affect incubation period. We expected: (i) second eggs to have thinner shells and/or more pores than first eggs; (ii) eggs to have thinner shells and/or more pores if they were laid by older females, later in the season, or in burrow nests (more humid than open nests under bushes). We found no support for these hypotheses. Egg pores and shell thickness were not related to incubation period. Eggs from the same clutch were similar in eggshell thickness and pore density. Female age, laying date and nest type were poor predictors of eggshell thickness and pore density. Egg pores, we conclude, have little explanatory power for differences in incubation time of eggs in Magellanic Penguins, and suggest that synchronous onset of incubation matters.  相似文献   

19.
We explore the relationship between general eggshell colour (background and spots) and proportion of surface covered by spots on the eggs of blue tits Cyanistes caeruleus and several variables reflecting female health and condition. Females laying more spotted eggs showed a poorer body condition, higher cellular concentration of the stress protein HSP70 and marginally lower total immunoglobulin levels in blood. In addition, these females were paired with males with higher levels of HSP70 and lower concentrations of immunoglobulins. We interpret these results based on the properties of the eggshell pigment protoporphyrin, which is known as an inductor of oxidative stress in females. Although observational, this study presents, for the first time, clear evidence of eggshell spottiness and general colour as indicators of general condition and stress in avian populations.  相似文献   

20.
Variation in blue‐green and brown coloration of avian eggshells could be affected by several factors, including environmental nutritional constraints. Better availability of nutrients could influence the synthesis and deposition of pigments into the eggshell, so we may expect a link between the food availability during egg formation, the body condition of the female and intensity of eggshell coloration. This hypothesis has received mixed support so far: in some bird species a positive correlation between female body condition and eggshell blue‐green coloration could be demonstrated, but other studies failed to find a significant link. In this study, we experimentally limited the food availability for domestic canaries Serinus canaria prior to and during egg laying, and examined its effects on the biliverdin‐ and protoporphyrin‐based eggshell coloration. Treatment had no significant effect on eggshell blue‐green chroma and biliverdin concentration. However, we found a significant positive relationship between female body condition and eggshell background blue‐green chroma in the control group, but not in the food restricted group. Females possibly experiencing a decline in antioxidant capacity due to food limitation may not be able to produce a blue‐green eggshell colour intensity reliably indicating their body condition. Furthermore, food‐restricted canary females laid eggs with significantly higher eggshell brown chroma, spot intensity, and protoporphyrin concentration. Therefore, our results suggest that limitation in actual nutrient availability increases deposition of protoporphyrin into the eggshell, and it may also modify the association between female body condition and intensity of blue‐green eggshell coloration.  相似文献   

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