Body size development of captive and free‐ranging Leopard tortoises (Geochelone pardalis)

The growth and weight development of Leopard tortoise hatchings (Geochelone pardalis) kept at the Al Wabra Wildlife Preservation (AWWP), Qatar, was observed for more than four years, and compared to data in literature for free‐ranging animals on body weight or carapace measurements. The results document a distinctively faster growth in the captive animals. Indications for the same phenomenon in other tortoise species (Galapagos giant tortoises, G. nigra; Spur‐thighed tortoises, Testudo graeca; Desert tortoises, Gopherus agassizi) were found in the literature. The cause of the high growth rate most likely is the constant provision with highly digestible food of low fiber content. Increased growth rates are suspected to have negative consequences such as obesity, high mortality, gastrointestinal illnesses, renal diseases, “pyramiding,” fibrous osteodystrophy or metabolic bone disease. The apparently widespread occurrence of high growth rates in intensively managed tortoises underlines how easily ectothermic animals can be oversupplemented with nutrients. Zoo Biol 29:517–525, 2010. © 2009 Wiley‐Liss, Inc.     

Intraspecific nest parasitism in the grey starling (Sturnus cineraceus)

We studied intraspecific nest parasitism in the grey starling (Sturnus cineraceus) in 1992 and 1993. We used three criteria to detect nest parasitism: (i) the appearance of more than one egg per day while the host was laying; (ii) the appearance of extra eggs after the host completed its clutch; and (iii) the appearance of eggs which were of a different shape, size and color to other eggs in the clutch. There were 290 nests (157 nests in 1992; 133 nests in 1993) in which the clutch was completed early (clutches initiated before May 10). Twenty-nine (1992) and 32 (1993) nests contained at least one parasitic egg. Parasitic eggs hatched if they were laid during the laying period and early in the incubation period of their host, and a few of them fledged. Fledging success of parasitic eggs was not different from that of eggs in non-parasitized nests if parasitic eggs were laid during the host's laying period. However, fledging success of all parasitic eggs was fewer than that of eggs in non-parasitized nests. By comparison, fledging success of parasitized nests was not a great as that of non-parasitized nests.     

Intraspecific nest parasitism in the Spotless Starling Sturnus unicolor

Intraspecific nest parasitism in two colonies of Spotless Starling Sturnus unicolor breeding in nestboxes was studied in central Spain from 1991 to 1994. Nests were monitored regularly and three criteria were used to detect nest parasitism: the appearance of more than one egg per day during the laying period of the host; the appearance of an egg after the start of incubation; eggs with unusual shape or pigmentation. The proportion of parasitized nests in first clutches (37%) was twice that of intermediate (19%) or second (20%) clutches in colony B, whereas parasitism occurred in first (35%) and intermediate (12%) but not in second clutches in colony A. Most clutches (52–70%) were parasitized during the host's laying period and received one parasitic egg. In 10% of the parasitized clutches in colony B, one of the host's eggs disappeared on the day the parasitic egg was added, suggesting that the parasitic female removed this egg. Although parasitism increased clutch size significantly, it led to a decrease in host breeding success, mainly through the removal of eggs and the loss of host nestlings and the survival of parasitic chicks. Observations suggested that parasitic females were young individuals without their own nests and/or those whose breeding attempt had been disrupted while laying in their own nest.     

Laying‐sequence variation in yolk carotenoids and egg characteristics in the red‐winged blackbird Agelaius phoeniceus

In many bird species with asynchronous hatching, smaller, later‐hatched nestlings are out‐competed for food by their larger, earlier‐hatched siblings and therefore suffer increased mortality via starvation. It is thought that female birds can either maintain or reduce the survival disadvantage of later‐hatched nestlings by differentially allocating maternal resources across the eggs of a clutch. Carotenoid pigments are an example of resources that female birds allocate differentially when producing a clutch, but laying sequence patterns for these pigments remain poorly studied in North American songbirds. We examined intraclutch variation in yolk carotenoids and egg metrics in 27 full clutches of red‐winged blackbird Agelaius phoeniceus eggs collected from eight wetlands in central Alberta, Canada. We predicted that carotenoids would decrease across the laying sequence, as in this species, later‐hatched, marginal nestlings suffer greater mortality than earlier‐hatched, core nestlings. We found nine carotenoid pigments in red‐winged blackbird egg yolks, including two that have never been described from avian yolks: α‐doradexanthin and adonirubin. As predicted, concentrations and amounts of most carotenoids decreased across the laying sequence, suggesting that female red‐winged blackbirds depleted their carotenoid resources as they laid more eggs. However, egg mass and yolk mass both increased across the laying sequence, suggesting that female red‐winged blackbirds may use other maternal resources to compensate for the size and survival disadvantage experienced by later‐hatched, marginal nestlings.     

Egg clutch patterning in Lestes virens (Odonata,Lestidae) with evolutionary emphasis on endophytic oviposition in lestid dragonflies

Egg deposition within plants is one of the most widely distributed and ancient behaviors in Odonata. The resulting clutch consists of eggs placed in peculiar pattern that can be a characteristic for certain groups of Odonata. Despite their importance for paleontological and evolutionary research, data on egg‐clutch positioning are missing or insufficient for most species. Here, patterning of egg clutches in Lestes virens was measured and described in detail for the first time. The female usually produces a linear row of single eggs directed at an angle rightward or leftward to the longitudinal axis of plant substrate. Less often eggs are arranged in egg‐sets consisting of up to 4 eggs. Apparently, the female insect follows the rigid behavior stereotypes during oviposition and is unable to easily switch to the alternate stereotypical behavior of single egg deposition or production of multiegg sets. Based on a literature review and original data, egg clutch patterning of European Lestidae is overlaid on preexisting phylogenies. The resulting evolutionary scenario of egg‐clutch patterning can be considered in the framework of egg‐laying behavior in Lestidae.     

Seed Dispersal by Geochelone carbonaria and Geochelone denticulata in Northwestern Brazil1

The role of red‐footed tortoises (Geochelone carbonaria) and yellow‐footed tortoises (G. denticulata) as seed dispersal agents was investigated in northwestern Brazil from 5 to 26 January 2002 by analyzing fecal samples for frequency and viability of seed species and estimating daily displacement of tortoises from recaptured and thread‐trailed individuals. Fourteen of 19 fecal samples contained a total of 646 seeds represented by 11 plant species. The most abundant species was Ficus sp. (N= 400) with 100 percent of seeds viable, followed by Aechmea sp. (N = 88) with 93 percent of seeds viable, and Genipa americana (N= 59) with 91 percent of seeds viable. Mean minimum retention time of seeds was 1.6 d and mean daily displacement of tortoises based on recaptured (N= 7) and thread‐trailed tortoises (N= 2) was 57 m. Thus, the diversity and proportion of viable seeds consumed by tortoises, combined with the seed retention times and daily movements, suggest they may be effective dispersal agents. These preliminary findings warrant further investigation into the ecological role of these tortoises in Neotropical ecosystems and their contribution to the maintenance of species diversity and forest structure.     

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.     

Reproductive assessment of the great hornbill (Buceros bicornis) by fecal hormone analysis

The population of great hornbills (Buceros bicornis) in the United States is rapidly aging, and captive breeding efforts have not met population managers' expectations for a sustainable captive group. Little is known about the reproductive physiology of these birds. This study reports the first data on the re‐productive endocrinology of the great hornbill. The hormone profiles of the only pair of these birds that hatched a chick in the 1999–2000 breeding season are compared to the profiles of six other pairs of hornbills, from different institutions in the United States, that did not reproduce successfully that season. The study investigates the estradiol, corticosterone, and testosterone profiles of these seven pairs of birds, establishing a base of knowledge from which endocrine data may be used to improve the success of captive breeding programs. The estradiol profiles from this study indicate a difference in hormonal patterns between laying and non‐laying female great hornbills. Egg‐laying females had significantly higher estradiol concentrations during the breeding season than the non‐laying females (P<0.003). Testosterone concentrations of the males were not significantly different between the mates of egg‐laying and non‐egg‐laying females. The corticosterone concentrations tended to be lower in the females that laid eggs vs. the non‐egg‐laying group. The males of the egg‐laying pairs showed a significantly lower (P<0.036) corticosterone concentration than the non‐egg‐laying male pairs. This, combined with the extremely low corticosterone levels (compared to the other birds in the study) of the pair of hornbills that hatched a chick in the 1999–2000 breeding season, suggests that adrenal activity may play a role in the reproductive failure of some captive great hornbills. Zoo Biol 22:135–145, 2003. © 2003 Wiley‐Liss, Inc.     

Intraspecific variation in reproductive traits of burrowing owls

Reviews of hatching asynchrony in birds recommended more studies on intraspecific variation in the extent of hatching asynchrony. We examined intraspecific variation in clutch size, laying chronology, onset of incubation, incubation period, and hatching asynchrony in burrowing owls (Athene cunicularia) in the Imperial Valley of California. Mean clutch size was 7.4 eggs and owls averaged 0.5 eggs laid per day. Females varied considerably in laying interval and onset of incubation (range?=?1st to 9th egg in the clutch). The mean incubation period was 21.9?days. Hatching interval also varied greatly among females ( $ \overline{x} $ ?=?0.8, range 0.1–2.0?days between successively hatched eggs). Past burrowing owl studies have largely overlooked the substantial intraspecific variation in these traits or have reported estimates that differ from ours. Future studies designed to identify the environmental factors that explain the large intraspecific variation in these traits will likely provide insights into the constraints on local abundance.     

Incubation prior to clutch completion accelerates embryonic development and so hatch date for eggs laid earlier in a clutch in the great tit Parus major

The ability of parents to respond to changes in food supply within a season will have a large effect on fitness through the number and quality of chicks fledged. Great tits, Parus major, attempt to synchronise their production of chicks with a seasonal food peak, but when food supply fails, hatching asynchrony of chicks provides a mechanism by which some young can be fledged because more developed chicks outcompete their less developed siblings for the reduced parental food supply. We tested whether female great tits can potentially control the degree of hatching asynchrony by using incubation before clutch completion, so that early laid eggs develop faster and hatch sooner. The temperature of an artificial egg placed in 29 nests during the laying period was measured with data loggers, and nocturnal incubation of eggs similar to incubation post clutch completion was recorded in all nests. We then demonstrated that eggs removed from the nest for 72 hour periods prior to clutch completion hatched later than eggs remaining in the nest for the entirety of the laying period. Our results show that variable pre clutch completion incubation (which was mostly nocturnal) can lead to faster embryo development and earlier hatching, so potentially providing a mechanism for adaptive female control of degree of hatching asynchrony.     

BREEDING SUCCESS IN SOME AFRICAN EAGLES RELATED TO THEORIES ABOUT SIBLING AGGRESSION AND ITS EFFECTS

Brown, L. H., Gargett, V. & Steyn, P. 1977. Breeding success in some African Eagles related to theories about sibling aggression and its effects. Ostrich 48:65-71. Previous explanations for fatal inter-sibling strife in eagles (lack of food, extra aggressiveness which enhances survival, and an expression of the innate aggressiveness or territoriality of raptors) can not be supported by recent evidence. The latest theory, that the second egg acts as a “reserve”, is examined. If so, eagles normally laying two eggs should have better reproductive success than eagles laying a single egg. However there was no significant difference between the mean reproductive success of five African eagle species that lay two eggs and that of five African eagle species laying one egg. Even excluding inequalities due to sample size, and other factors, the overall finding is the same. In Verreaux's Eagle Aquila verreauxi for which the largest number of records is available there was a significantly higher total loss in clutches recorded as c/l than in c/2 clutches, but the reasons for the laying of a one-egg clutch, and its success or failure, have not been investigated. In three out of 120 c/2 the second-laid egg acted as a “reserve”, but in 110 cases the second-laid egg was “wasted”. Direct evidence and a comparison of reproductive success between two-egg and one-egg African eagles does not support the latest theory, therefore inter-sibling strife which effectively reduces the clutch of two to one, is still inexplicable.     

Partial incubation during egg laying reduces eggshell microbial loads in a temperate‐breeding passerine

Incubation prior to clutch completion may be adaptive if it maintains egg viability by inhibiting eggshell microbial growth, thus reducing the likelihood that the embryo becomes infected. To test this hypothesis, we examined the effect of partial incubation during egg laying on eggshell microbial loads in eastern bluebirds Sialia sialis breeding at a temperate‐zone site. We sampled eggshell microbes prior to and following four days of exposure to either partial incubation during the laying period or ambient environmental conditions without incubation (experimental eggs). Microbial colony counts declined significantly for eggs left in the nest during the laying period but did not vary significantly for eggs exposed to ambient conditions. Initial microbial loads were more similar to those previously reported from tropical than temperate environments, and microbes from potentially pathogenic groups were detected on 88% of first‐laid eggs on the day of laying. Egg viability was maintained when eggs were held indoors for four days without incubation but declined sharply thereafter. Our results suggest that partial incubation during egg laying may enhance egg viability in eastern bluebirds by reducing eggshell microbial loads; these effects appear stronger than those usually reported from the temperate zone.     

Reproductive biology and ecology of white‐winged trumpeters (Psophia leucoptera) and recommendations for the breeding of captive trumpeters

The reproductive biology and ecology of a wild population of white‐winged trumpeters (Psophia leucoptera) were studied in southeastern Peru from 1983 to 1987. Because little information is available about any of the trumpeter species and because trumpeters have proven difficult to breed in captivity, information relevant to breeding and management of captive trumpeters is reported in this paper. White‐winged trumpeters lived in territorial social groups that ranged in size from four to 13 individuals. A typical territorial group contained three adult males, two adult females, and several sexually immature offspring, but smaller temporary groups sometimes formed for the duration of the breeding season. Only the dominant female contributed eggs to the clutch, and all adult males in the group competed to obtain copulations with her. Eggs were laid in elevated nesting cavities and no nest was constructed. The average clutch size was three eggs and incubation was not begun until the final egg was laid. The dominant male and female shared most of the incubation duties, but subordinate males covered approximately 15% of the incubation shifts. Eggs hatched approximately 27 days after incubation was begun and chicks left the nesting cavity the day after they hatched. Chicks were completely dependent on older birds to feed them for their first 3 weeks and then gradually began to feed themselves more and more food. The subordinate adult males fed chicks the most food, the dominant male and female and older offspring fed chicks an intermediate amount, and the subordinate adult female fed chicks the least. Young chicks behaved aggressively toward each other but were separated by adults before they injured each other. If at least one chick from the clutch survived, trumpeters did not breed again until the beginning of the next breeding season the following year. Zoo Biol 19:65–84, 2000. © 2000 Wiley‐Liss, Inc.     

Egg cannibalism in Ascia monuste in the field; opportunistic, preferential and very frequent

Newly hatched caterpillars of the specialist herbivorous species Ascia monuste (Lepidoptera, Pieridae) feed on egg chorion before ingesting plant material. They also ingest their exoskeletons and perform egg cannibalism in both the field and the laboratory. Females lay eggs on plants which already have conspecifics (eggs and/or caterpillars) leading to the observation of cannibalism in the field. Two favorable conditions are required for such an occurrence: (1) eggs laid close together on the same leaf, and (2) the finding by older caterpillars of eggs on the same plant. Oviposition preference experiments in cages showed that females do not avoid laying on plants with eggs. However, they do avoid laying eggs on plants predated by caterpillars. All instars perform cannibalism in the field and, in addition to this, the ingested eggs are healthy and cannibalism occurs in the presence of abundant food. The ingestion of eggs from their own clutch was more intense when the clutch was large. Contact with eggs is also important for cannibalism. Caterpillars ingested healthy eggs during three successive days. According to the results, it is possible to state that egg cannibalism in A. monuste is opportunistic, preferential and highly frequent and that there is a tolerance to a diet based on animal protein in this species. Since A. monuste obtains nutrients from both animal and vegetable resources, it is probably better to consider it as opportunistic omnivorous.     

1. A CENSUS AND STUDY OF BIRDS IN ALBANY OPEN THORNBUSHVELD

Williams, A. J. &; Cooper, J. 1983. The Crowned Cormorant: breeding biology, diet, and offspring reduction strategy. Ostrich 54:213-219.

Crowned Cormorants Phalacrocorax coronatus were studied at Dassen and Marcus Islands. The most frequent clutch was three eggs. Egg size varied within clutches with first-laid eggs being largest and heaviest and subsequent eggs progressively smaller and lighter, The mean laying interval was 2,2 days, the mean laying-to-hatching interval was 23,0 days, and the hatching interval was one day. The normal incubation period was 22.4 days. The weight of hatchings was related to the position of the originating egg in the laying sequence. Chicks were fed within 24 h of hatching. Chick development is described over the first 35 days. One chick could fly at 35 days. Hatching success was 48,2%. Hatching success was greatest in second-laid eggs, least in last-laid eggs. The mean number of chicks hatched at a nest was two. Mean diving time was 23,5 s. Most food was fish, particularly klipfish Clinidae and pipefish Syngnathus, 60–160 mm long. The number of offspring produced can be related to food availability by interaction of difference in egg size, hatching asynchrony, and the preferential feeding by adults of the strongest-begging chick. There is a trend towards producing two chicks, normally those from the first two eggs to be laid.     

Breeding ecology of the African Spoonbill Platalea alba in the Free State,South Africa

Kopij, G. 1997. Breeding ecology of the African Spoonbill Platalea alba in the Free State, South Africa. Ostrich 68 (2–4): 77–79.

The breeding ecology of the African Spoonbill is surprisingly little known. In the Free State, the number of nesting African Spoonbills appears to have declined; 120 nests in 11 colonies were located in 1972–1973, compared to 32 nests in four colonies in 1993–1996 (though this later survey may have missed a few small colonies). In 1976–1977, a colony with 15 active spoonbill nests was monitored for an entire breeding season. Mean clutch size was 2.6 (n = 15); 56% of eggs hatched; 91% of hatched eggs resulted in fledglings; and chicks fledged from 53% of nests. The main food of spoonbill chicks was frogs, mainly Rana angolensis and Xenopus laevis, and aquatic invertebrates.     

Oviposition behavior and progeny allocation of the polyembryonic waspCopidosoma floridanum (Hymenoptera: Encyrtidae)

Oviposition behavior was used to determine the primary clutch size and sex ratio of the polyembryonic wasp Copidosoma floridanumAshmead (Hymenoptera: Encyrtidae) parasitizing Pseudoplusia includens(Walker) (Lepidoptera: Noctuidae). The laying of a female egg was associated with a pause in abdominal contractions during oviposition, while the laying of a male egg was associated with uninterrupted abdominal contractions. Although unmated females produced only male broods, they also displayed male and female egg oviposition movements. Wasps always laid a primary clutch of one or two eggs. For mated females if only one egg was laid, the emerging secondary clutch was all male or female, but if two eggs were laid a mixed brood of males and females was almost always produced. The secondary clutch of single sex broods was usually between 1000 and 1200 individuals, but the secondary clutch of mixed broods averaged 1143 females and 49 males. Thus, the primary sex ratio for mixed broods was 0.5 (frequency males), but the secondary sex ratio was 0.042. Manipulation of the sequence of male and female egg oviposition or of the primary clutch did not produce major alterations in the secondary clutch size or sex ratio.     

Egg development time in the Zebra Finch Taeniopygia guttata varies with laying order and clutch size

As female birds are able to lay no more than a single egg each day, in those species producing larger clutches the first laid eggs may get a developmental head‐start over later eggs in the clutch. All other things being equal, the differential pattern of development across the clutch may contribute to hatching asynchrony and subsequent inequity in the competition between brood mates, and ultimately increase variance in the quality and fitness of first‐ and last‐laid offspring. It has been suggested that females might allocate resources differently across the laying sequence to moderate the developmental rate and hatching time of different embryos. We tested this theory in the Zebra Finch Taeniopygia guttata, a common model species for investigating maternal effects in birds. We removed 758 eggs from 160 nests shortly after they were laid and used artificial incubators to control for parental effects and monitor hatching times. Eggs from larger clutches consistently hatched sooner than those from average‐sized clutches, demonstrating that the intrinsic properties of an egg can alter the developmental time of embryos. There were also differences in the development time of eggs across the laying sequence, but these patterns were weaker, inconsistent and unrelated to sequential investment across the laying sequence in a straightforward way. This study indicates that maternal resource allocation to eggs across the laying sequence and across clutch sizes can influence development times and play a potentially important role in determining the competitive dynamics of broods.     

Breeding ecology of Red‐faced Cormorants in the Pribilof Islands,Alaska

ABSTRACT Red‐faced Cormorants (Phalacrocorax urile) are North Pacific endemics recognized as a vulnerable species, but little is known about their breeding ecology. We studied Red‐faced Cormorants on St. Paul Island, Alaska, from 1975 to 2009, with more detailed data collected in 2004 and 2005. Mean clutch sizes in 2004 (3.2 ± 0.8 [SD] eggs) and 2005 (3.1 ± 0.8 eggs) were similar to the long‐term average (2.9 ± 0.3 eggs from 1976 to 2009). The mean laying interval in 2004 and 2005 was 2.15 ± 0.80 d (N= 407), and the mean egg period (number of days between laying of an egg and hatching) was 31.1 ± 1.4 d (N= 158). Approximately 64 ± 17% of eggs hatched during the period from 1975 to 2009. The mean number of chicks per nest in 2004 and 2005 was 2.8 ± 0.8 (N= 232), and the mean number of fledglings per initiated nest in all years was 1.22 ± 0.52. Chicks fledged 46 to 66 d posthatching. In 2004 and 2005, the primary causes of egg loss were predation by Arctic foxes (Vulpes lagopus) and destruction of eggs and abandonment of nests due to storms. Starvation was the primary cause of nestling mortality in both years. Because chicks are dependent on parents to provide food for over 45 d, consistent near‐shore foraging opportunities must be available. From 1975 to 2009, Red‐faced Cormorants experienced only 1 yr of complete reproductive failure (1984). The consistent reproductive success of Red‐faced Cormorants suggests that conditions may be relatively stable for this species on St. Paul Island, or that the variability in their breeding ecology (e.g., phenology, clutch sizes, and incubation strategies) provides the flexibility needed to successfully fledge some chicks nearly every year.     

The role of partial incubation and egg repositioning within the clutch in hatching asynchrony and subsequent effects on breeding success

The main mechanism to achieve hatching asynchrony (HA) for incubating birds is to start heating the eggs before clutch completion. This might be achieved through partial incubation and/or early incubation. Even in the absence of incubation behaviour during the laying phase, clutches still experience a certain degree of asynchrony. Recent studies have shown that eggs located in the centre of the nest receive more heat than peripheral ones during incubation. As eggs receiving more heat would develop faster, we hypothesized that HA should be shorter in nests where eggs were moved homogeneously along the centre–periphery space during incubation than in those nests where eggs repeatedly remained in the same locations, either centrally or peripherally. We explored the relative roles of egg repositioning and partial incubation in determining HA in wild birds by (1) removing eggs from 20 Great Tit Parus major nests on the day of laying and replacing them with fake eggs to avoid partial incubation, and returning them when full incubation began; (2) monitoring twice a day the position of each individually marked egg relative to the clutch centre during incubation, and estimating the coefficient of variation of the distances; and (3) determining HA in each nest. Preventing partial incubation reduced HA by 51% days in experimental nests. It also caused negative effects for the incubating females (lengthening the full incubation period) and positive effects for the brood (increasing fledging success). However, our hypothesis about the role of egg repositioning on HA was not supported: all the females moved the eggs with remarkable consistency, generally attaining a coefficient of variation of the distances around 33%, and it was not related to the HA experienced. We therefore conclude that partial incubation is an important factor regulating HA, and females compensate for the potential effects of differential heating by moving the eggs homogeneously within the clutch.