Lifetime egg production and egg mortality in the damselflyPyrrhosoma nymphula (Sulzer) (Zygoptera: Coenagrionidae)

Artificial oviposition sites were used to estimate egg deposition rates in the field. Females laid an average of 10.76 eggs/minute with a mean duration of 22.81 minutes, giving an average clutch size of 245 eggs. Since one mating corresponded to one clutch of eggs, lifetime mating success was used as a measure of the number of clutches produced. Mean lifetime clutch production was 5.91 clutches per female, equating to 1447 eggs per female per lifetime. Eggs were hatched in the laboratory at temperatures comparable with those in the field. Hatching was highly synchronised and the overall hatching success was 75.1%. Causes of egg mortality in the laboratory were limited to infertility and unhatchability. Since no other sources of egg mortality could be found at the study site, this value was a good reflection of hatching success in the field. Lifetime egg production and hatching success were used to estimate the number of viable offspring produced per female, giving a higher order estimate of reproductive success than has previously been published for a zygopteran.     

Patterns of variation in size of Boat-tailed Grackle Quiscalus major eggs

Boat-tailed Grackle Quiscalus major eggs averaged 8^.09 g wet weight. Mean egg weight represented 8 05 % of female body weight. Based on egg weight, Grackle eggs have a shorter incubation period than the general passerine pattern.
Egg weight varied significantly with sequence of laying and between clutches in both two-egg and three-egg clutches. Last laid eggs weighed less than first laid eggs. The mean weight of the first two eggs laid in three-egg clutches did not differ from the mean egg weight of two-egg clutches. The average clutch of two and three eggs weighed 16.33 g and 24.16 g, respectively.
Mean egg weight varied between study colonies and with season. Grackles at East Lake laid eggs that weighed less than grackles at either Alligator Lake or North Lake. Eggs laid during March averaged less than eggs laid later in the season. The locality variation reflects the different timing of nesting between sites rather than food abundance. As nutrient provisioning increases with an increase in egg weight, the seasonal change in egg weight probably reflects improved feeding conditions for the female. This suggests that selection favours beginning laying before reserves of the female are sufficient to lay the largest egg possible.
The size of the young at hatching was correlated with egg size. Newly hate had young averaged 79.6% of fresh egg weight. Egg weight did not determine the probability of hatching or starvation of the young. Females do not appear to adjust egg size facultatively depending on the sex of the young. Eggs producing males and females did not vary significantly in weight, whether comparisons were made within or between clutches.     

Effects of embryonic responses to clutch mates on egg hatching patterns of Pentatomidae (Heteroptera)

Stink bugs and shield bugs of the family Pentatomidae (Heteroptera) generally produce a clutch of densely deposited eggs. In a few species of this family, embryos hatch in response to some form of cues associated with the preceding hatching to synchronize egg hatching with clutch mates. The aim of the present study is to obtain a family‐wide understanding of the extent to which the hatching response to clutch mates accelerates hatching within egg clutches. Accordingly, the hatching patterns in intact egg clutches and eggs individually detached from egg clutches are compared in eight species among different genera. In Halyomorpha halys, hatching is significantly and highly synchronized by the effect of the hatching response: when eggs are not attached to each other, the hatching rate is only 3.8% at 15 min and exceeds 95% at 200 min. By contrast, when eggs are attached to each other, the hatching rate reaches more than 95% at 15 min. Hatching is also significantly synchronized by the hatching response in Nezara viridula (which shows relatively high hatching synchronization) and in Piezodorus hybneri and Plautia stali (both of which show milder hatching synchronization). Synchronization of hatching is not found to be promoted by a hatching response in Aelia fieberi, Dolycoris baccarum, Eurydema rugosum or Palomena angulosa. These findings reveal that the hatching response varies depending on the species in Pentatomidae, with a wide spectrum of effects on the hatching patterns of the egg clutches.     

Plasticity in incubation behavior and shading by king rails Rallus elegans in response to temperature

King rails experience a wide range of temperatures during the course of the breeding season throughout their rapidly contracting geographic range. Incubating parent birds are adapted to keep their eggs within a temperature range appropriate for embryo development, but king rail clutches are at risk of exceeding lethal temperatures in the latter half of the nesting season. We investigated whether behavioral plasticity during incubation enables parents to maintain clutch temperature within tolerable limits for embryo development. Video revealed that king rail parents interrupted incubation to stand above and shade their eggs. We tested the hypothesis that the onset of shading was a direct response to ambient temperature (adaptive plasticity). We monitored clutch temperature directly by experimentally adding into clutches a model egg embedded with a programmable iButton. We measured ambient temperature at the nest site simultaneously. Parents spent proportionately more time shading and less time incubating their eggs at higher ambient temperatures. Shading may primarily function in cooling the parent. The frequency and duration of shading bouts were significantly greater at higher ambient temperatures. Parents also took more frequent but shorter recesses in hotter conditions. Diurnal recesses exposed eggs to direct sunlight, and the highest clutch temperatures were recorded under these conditions. Complete hatching failure in at least one nest was attributable to high clutch temperature for an extended period. Because mean ambient temperature increases throughout the breeding season, we investigated seasonal patterns in onset of incubation and its effect on hatching rate. Later in the season, parents tended to initiate incubation earlier, and hatching asynchrony increased significantly. Together these results suggest that breeding king rails may be constrained in their ability to cope with sustained high temperatures should seasonal averages continue to rise as predicted.     

CLUTCH-SIZE, INCUBATION AND HATCHING SUCCESS IN THE HOUSE SPARROW AND TREE SPARROW PASSER SPP. AT OXFORD

Clutch-size, incubation and hatching success were studied in P. domesticus and P. montanus in 1961 and 1963–64 at Oxford. The most frequent clutch-size was four eggs in P. domesticus and five eggs in P. montanus . With one exception, colonies of P. domesticus showed no significant annual or local variations in its mean clutch-size; in P. montanus , however, there were significant annual variations in the mean clutch-size. Both species showed a seasonal increase followed by a decrease in their mean clutch-sizes.
Partial incubation occurred during the laying period of the clutch; sufficient incubation for continuous development of the embryo was apparently achieved when the last egg had been laid in clutches of two and three eggs in P. domesticus and in clutches of four eggs in P. montanus , but when the penultimate egg had been laid in larger clutches of both species. On average, hatching in P. domesticus occurred more or less synchronously in all eggs in clutches of two and three eggs, and in all eggs except the last one laid in larger clutches; the last egg in the larger clutches hatched up to a day after the others. It is suggested that this pattern of hatching was brought about by the pattern of incubation during the laying period.
P. domesticus had a lower hatching success than P. montanus , probably because fewer of its eggs were fertile.     

Effects of clutch size and egg-laying order on the breeding success in the Little Tern Sterna albifrons on the Nakdong Estuary, Republic of Korea

Survivorship of Little Tern Sterna albifrons eggs and chicks was followed on an islet in the Nakdong Estuary, Republic of Korea, in 1995 and 1996. Mean egg size and incubation period were significantly different between the 2 years. The maximum clutch size was three eggs, and the second egg in the clutch often hatched earlier than the first, while most of the third eggs hatched last. In 1996, when the fate of 249 eggs from 106 nests was followed for 40 days, hatching success, fledging success and breeding success were 77%, 40% and 31%, respectively. High mortality occurred in the early chick stage, mostly because of rain and predation by Weasels Mustela sibirica. The breeding success per egg was 14% in one-egg clutches, 28% in two-egg clutches and 34% in three-egg clutches. This difference was mainly attributed to the lower hatching success in the smaller clutches. In three-egg clutches, the third egg showed significantly lower breeding success than siblings. The main foods of the Little Tern were Tridentiger obscurus, Engraulis japonicus, Hyporhamphus intermedius, Acanthogobius flavimanus (all fish), Palaemon sp. and Crangon affinis (shrimps). The feeding frequency was, apparently, not affected by time of day and age of chicks but was probably influenced by weather conditions. Newly hatched chicks failed to eat 25% of the prey brought to them, although this decreased with the age of the chicks.     

Corticosterone stimulates hatching of late-term tree lizard embryos

The regulation of hatching in oviparous animals is important for successful reproduction and survival, but is poorly understood. We unexpectedly found that RU-486, a progesterone and glucocorticoid antagonist, interferes with hatching of viable tree lizard (Urosaurus ornatus) embryos in a dose-dependent manner and hypothesized that embryonic glucocorticoids regulate hatching. To test this hypothesis, we treated eggs with corticosterone (CORT) or vehicle on Day 30 (85%) of incubation, left other eggs untreated, and observed relative hatch order and hatch time. In one study, the CORT egg hatched first in 9 of 11 clutches. In a second study, the CORT egg hatched first in 9 of 12 clutches, before vehicle-treated eggs in 10 of 12 clutches, and before untreated eggs in 7 of 9 clutches. On average, CORT eggs hatched 18.2 h before vehicle-treated eggs and 11.6 h before untreated eggs. Thus, CORT accelerates hatching of near-term embryos and RU-486 appears to block this effect. CORT may mobilize energy substrates that fuel hatching and/or accelerate lung development, and may provide a mechanism by which stressed embryos escape environmental stressors.     

Incubation capacity limits maximum clutch size in black-tailed gulls Larus crassirostris

Factors determining clutch size of birds have long been the central issue in studies in life histories. It is assumed that the configuration of brood patches could limit the maximum clutch size. To test this hypothesis we manipulated clutch sizes and measured egg temperature as well as reproductive consequences in black-tailed gulls Larus crassirostris , which usually lay two egg clutches and have three brood patches. Mean egg temperature in 4-egg clutches (32.6±1.0°C) was significantly lower than in 2-egg (34.6±0.4°C) and 3-egg clutches (34.1±0.4°C), because egg temperature of the coolest egg within a 4-egg clutch was often substantially lower than the other three eggs. The proportion of eggs hatching from 4-egg clutches (11.6%) was lower than those of 2-egg (49.1%) and 3-egg clutches (52.0%). Four-egg clutches had longer incubation periods (29.6±1.3 day) than 2-egg (28.1±1.7 day) and 3-egg clutches (28.0 ±1.3 day). The results indicate that incubation capacity, which may be determined by the configuration of brood patches, limits the maximum clutch size in black tailed gulls, but not the actual clutch size typically laid.     

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

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

Natural growth and reproduction in Triops granarius (Lucas) (Crustacea: Notostraca)

Field sampling during successive inundations was followed by laboratory studies on growth, egg production and hatching of Triops granarius from temporary pans in southern Africa.The species was common where mean inundation was less than one month. Sexual maturity was reached in as little as six days at a carapace length of 6.5 mm. A biomass of 2.0 g, at carapace length of 25 mm was achieved in two months. Males and females occurred in similar numbers and mating was a prerequisite for sustained fertile egg production. Eggs were in similar numbers in both pouches, while successive clutches were larger. Hatching was light-dependent. Desiccation of eggs was not necessary but a period of up to 20 days was required before undesiccated eggs would hatch. In the event of desiccation, no minimum period thereof was necessary and hatching occurred within as little as a day of rehydration. Hatching success seldom exceeded 50 percent.As a candidate for aquaculture, the species has failings such as low hatching rate and cannibalism, but it grows rapidly and produces eggs over an extended period.     

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

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

Multiple maternity in black-headed gull Larus ridibundus clutches as revealed by protein fingerprinting

Social monogamy with biparental care is the norm in gulls Laridae , but egg colour variation suggests that some nests may contain mixed clutches laid by more than one female. Here we use protein fingerprinting of egg albumen to assess the occurrence of mixed maternity clutches in three colonies of black-headed gulls. Among 160 analysed clutches with >1 egg, 34% contained eggs from more than one female, and 15% of the eggs in clutches >1 came from other females than the major female (laying most eggs in nest). Among clutches with 2–3 eggs 28% were mixed, and among clutches with 4 or more eggs 89% contained eggs from two or more females. There were significantly fewer eggs from the major female in mixed nests (mean=2.06±0.63 SD) than in non-mixed nests (mean=2.82±0.43 SD). In nests without evidence of female conflict, hatching success of minority eggs was similar to that of eggs from the major female (12.5 and 8.4%, respectively). In 21% of mixed maternity nests, one or more minority eggs was buried or punctured, and 25% of eggs from major females were also found evicted, suggesting conflict between females and rejection of eggs. Intra-specific nest parasitism seems the most likely cause of mixed clutches, but there are also other possible causes.     

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

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

CLUTCH SIZE, EGG SIZE, HATCH WEIGHT AND LAYING DATE IN RELATION TO EARLY MORTALITY IN RED GROUSE LAGOPUS LAGOPUS SCOTICUS CHICKS

Clutches of Red Grouse eggs were collected from the wild and subsequent hatching and rearing done in standard conditions in captivity. Variations in chick survival from one clutch to another in the same year were related to differences in hatch weight. Hatch weight was determined only partly by egg size. Weight loss between laying and hatching was related to survival independently of egg size. Variation in this weight loss obscured any simple relationship between egg size and survival, except in eggs laid by captive hens. Intrinsic differences amongst hens caused some variations in laying date, egg size, hatch weight and chick survival. Variations in egg size and hatch weight accounted for less than half the variation in survival; other unmeasured intrinsic factors were also important. Big clutches hatched earlier than small ones. The commonest clutches were of seven and eight eggs, with six and nine frequent. Very big clutches of ten or more eggs were infrequent and chicks from them sometimes survived worse than from smaller clutches. As in other species, the commonest clutch sizes were not the most productive. There was no simple relationship between egg size and clutch size.     

Egg size, offspring sex and hatching asynchrony in zebra finches Taeniopygia guttata

Experimental synchronization of onset of incubation was employed in laboratory held zebra finches Taeniopygia guttata to study whether differential resource allocation and possible bias of offspring sex in subsequent eggs in the laying order could mitigate the effects of hatching asynchrony. We found that egg mass increased with laying order, but offspring sex was not related to laying order. Among synchronized clutches, eggs hatched more synchronously than eggs from control nests. Survival probability was related to egg mass, and as expected, this effect differed between experimental groups: it was positive among synchronized broods and not significantly related among asynchronous broods. This suggests that increase in egg mass with the laying order might reduce disparities between early and late hatching chicks. Female nestlings survived better than male nestlings. However their growth was impaired in synchronized broods, whilst growth of males was not affected by hatching synchronization.     

Development and the influence of nurse egg allotment on hatching size in Searlesia dira (Reeve, 1846) (Prosobranchia : Buccinidae)

The reproductive biology of the intertidal prosobranch Searlesia dira (Reeve, 1846) was examined with special attention given to variability in the nurse egg to embryo ratio among capsules, among clutches and among geographically isolated populations. Embryos and nurse eggs were distributed among the capsules in a manner consistent with the hypothesis that nurse eggs were genetically predetermined, that each female had a genetically defined nurse egg to embryo ratio, and that each capsule represented a random sample of that ratio. The binomial distribution of embryos and nurse eggs among the capsules resulted in some capsules receiving many more embryos per nurse egg than others. The number of nurse eggs an embryo succeeded in eating was proportional to the number of capsule-mates sharing a capsule. Embryos eating more nurse eggs hatched out at a larger size. Differences in the nurse egg to embryo ratios among capsules in the same clutch were much larger than that of the mean ratios among clutches. Among-site differences in the mean nurse egg to embryo ratios suggest that selection pressure for different mean hatching sizes may have acted on the mean nurse egg to embryo ratios.In contrast to the predictions of optimal hatching size theory, hatching size varied widely within clutches as a consequence of differences in nurse egg to embryo ratios among capsules. This variance may be adaptive for species that lay their eggs months before juveniles emerge into an unpredictable environment, or simply be a consequence of an imperfect mechanism for increasing hatching size.     

Different Temperature and Cooling Patterns at the Blunt and Sharp Egg Poles Reflect the Arrangement of Eggs in an Avian Clutch

Incubation is an energetically demanding process during which birds apply heat to their eggs to ensure embryonic development. Parent behaviours such as egg turning and exchanging the outer and central eggs in the nest cup affect the amount of heat lost to the environment from individual eggs. Little is known, however, about whether and how egg surface temperature and cooling rates vary among the different areas of an egg and how the arrangement of eggs within the clutch influences heat loss. We performed laboratory (using Japanese quail eggs) and field (with northern lapwing eggs) experiments using infrared imaging to assess the temperature and cooling patterns of heated eggs and clutches. We found that (i) the sharp poles of individual quail eggs warmed to a higher egg surface temperature than did the blunt poles, resulting in faster cooling at the sharp poles compared to the blunt poles; (ii) both quail and lapwing clutches with the sharp poles oriented towards the clutch centre (arranged clutches) maintained higher temperatures over the central part of the clutch than occurred in those clutches where most of the sharp egg poles were oriented towards the exterior (scattered clutches); and (iii) the arranged clutches of both quail and lapwing showed slower cooling rates at both the inner and outer clutch positions than did the respective parts of scattered clutches. Our results demonstrate that egg surface temperature and cooling rates differ between the sharp and blunt poles of the egg and that the orientation of individual eggs within the nest cup can significantly affect cooling of the clutch as a whole. We suggest that birds can arrange their eggs within the nest cup to optimise thermoregulation of the clutch.     

Effect of brood size and hatching sequence on prefledging mortality of Sandwich terns: why lay two eggs?

The mortality of Sandwich tern Sterna sandvicensis chicks held in enclosures was studied in colonies on Griend, in the Dutch Wadden Sea, from 1992 to 1999, and on Hirsholm, in the Danish Kattegat, in 1997. Survival of chicks until fledging was 73% for chicks hatching from first-laid eggs or single-egg clutches and 59–64% for partially hatched two-egg clutches, whereas 6% of second hatchlings survived until fledging. Less than 2% of all two-chick broods actually fledged two chicks. Because 18% of the two-egg clutches only hatched one egg, 7% of fledglings of two-egg clutches originated from a second-laid egg. In nests where both eggs hatched, the number of chicks was usually reduced soon after hatching. Within five days of hatching more than 50% of the second hatchlings died of starvation or were preyed upon. It seems that overproduction commonly occurs in Sandwich terns and that investment in a surplus egg mainly serves as an insurance mechanism. On Griend and Hirsholm, chick productivity of two-egg clutches was somewhat higher than for one-egg clutches. Undernourishment was an important cause of death, either directly by starvation or by selective predation of chicks in poor condition. This, in combination with earlier, studies suggests that Sandwich tern parents on Griend are exposed to severe food stress.     

Brood Pumping Modulation and the Benefits of Paternal Care in Abedus breviceps (Hemiptera: Belostomatidae)

Males of the subfamily Belostomatinae, within the giant waterbugs (Hemiptera: Belostomatidae), brood eggs on their backs from oviposition until egg-hatching. The eggs in the clutch must be kept moistened and exposed to enough oxygen flow for them to develop properly and maximize hatching success. When submerged, males regularly perform different behavioral patterns that promote oxygen flow to the eggs. One of these patterns, brood pumping, consists of 'push-up' like movements using their hind legs. In this study, we assessed if temperature, oxygen dissolved in the water and the developmental stage of the clutch influence brood pumping rate in Abedus breviceps . In the wild, we found that males covered with bigger eggs showed a higher brood pumping rate than that of males with smaller eggs. An analysis of egg growth curves under laboratory conditions demonstrated that males that did not perform brood pumping were carrying broods that were in the first third of their developmental period. We also found that non-brooded clutches had a much lower hatching success than those brooded by males, even when exposed to controlled and favorable environmental conditions. We discuss the persistence and modulation of paternal care intensity in belostomatines as an aptative character conditioned by environmental and anatomical features.     

Incubation of eggs of tuatara, Sphenodon punctatus

Eggs of the tuatara, Sphenodon punctatus , were incubated either buried or half buried in vermiculite at constant temperatures of 15, 18, 20, 22 and 25 °C and constant water potentials between —90 and —400 kPa. Many clutches failed completely, possibly because they had been taken from females prior to proper shell development. Failed eggs were significantly smaller than successful eggs. Incubation is unsuccessful at 15 °C. Hatching success is high between 18 and 22 °C but low at 25 °C, but equally successful between 18 and 22°C. Incubation is strongly influenced by temperature, with mean incubation periods of 328 days at 18 °C, 259 days at 20 °C, 169 days at 22 °C and 150 days at 25 °C. Water potential generally has little influence on incubation time at a given temperature. Buried eggs hatch sooner than partially buried eggs at 20 °C but the large range makes significance dubious.
Eggs on the driest substrata at 18 and 20 °C lose water initially but then gain water through the rest of incubation. Eggs in all other conditions gain water throughout incubation, with the rate of i water absorption being maintained or increasing late in incubation. The suggestion that increasing rate of water absorption late in incubation facilitates explosive hatching is not supported. Egg mass at the time of hatching varies from 132 to 398% of initial values, depending on incubation conditions. Final egg mass is not affected significantly by incubation temperature. Hence, rates of absorption increase with temperature.
Water potential has no influence on hatchling size. However, hatchlings from buried eggs generally are significantly larger than those from partially buried eggs.