The point of a Guillemot's egg

The adaptive significance of avian egg shape in birds is poorly understood. The pyriform (pear‐like) shape of the Common Guillemot's Uria aalge egg has long been considered to be an adaptation to prevent eggs rolling off the bare cliff ledges on which this species breeds. Rolling was thought to be prevented either by the egg spinning like a top, which is not the case, or by rolling in an arc, which it does but with little influence on whether the egg will fall from a ledge. We therefore sought alternative explanations for the pyriform shape of the Common Guillemot's egg. This species breeds in extremely dense colonies, which makes their eggs vulnerable to mechanical damage from conspecifics, and to contamination by debris such as faeces and soil. We present evidence consistent with both these possible explanations. First, the pyriform shape of Common Guillemot eggs means that a higher proportion of the eggshell lies in contact with the substrate and this may minimize the effect of impacts. Resistance to impacts may be further enhanced because their eggshells are especially thick where they are in contact with the substrate. Secondly, Common Guillemot eggs are often heavily contaminated with faecal material and other debris during incubation. Most contamination is on the pointed end of the egg where it is in contact with the substrate; the pyriform shape thus keeps the blunt end of the egg, which has the highest porosity, relatively free of contamination, which in turn may facilitate both gas exchange during incubation and the hatching process, because the chick emerges from the blunt end of the egg.     

Counterstrategy to Egg Dumping in a Coreid Bug: Recipient Individuals Discard Eggs from Their Backs

The golden egg bug, Phyllomorpha laciniata Vill. (Heteroptera: Coreidae), is the only terrestrial insect in which females oviposit on the backs of female and male conspecifics. Eggs do not survive unless carried by a bug. Herein, I report laboratory observations that egg-carrying individuals actively brush their backs against the host plant seemingly in an effort to rub off eggs. Egg scraping is more common among individuals carrying many eggs than among those carrying only a few eggs. The most recently received eggs were rubbed off first. Females did not avoid laying eggs on the backs of egg-loaded individuals, nor did bugs carrying several eggs resist oviposition attempts more often than unloaded ones. Some males were likely to have fertilized the eggs they scraped off their backs. Laboratory results of active egg removal correspond with egg loss in the field, suggesting that egg scraping may explain egg loss in nature. The data indicate a cost of egg carrying to an individual and an evolutionary arms race between oviposition and discarding behavior.     

Egg size, composition and offspring quality in some Alcidae (Aves: Charadriiformes)

The composition of fresh and fully developed (pipping) eggs of four alcids, Razorbill, Common guillemot, Brünnich's guillemot and Atlantic puffin was examined. There were no differences in egg composition between the semi-precocial Atlantic puffin and the three species with "intermediate" developmental patterns. The absolute amount of yolk increased with egg size in fresh eggs, and the relative amount remained constant with egg size for Common guillemot and Razorbill, but decreased in the Atlantic puffin. In fully developed eggs chick weight and egg weight were closely correlated, and this was due mainly to larger eggs producing chicks with larger yolk reserves. Under some conditions chicks from larger eggs do better than those from smaller eggs. Several factors influence egg size; a comparison of first and replacement eggs laid by the same females showed that a maternal effect accounted for 60–90% of the variance in egg-size and that laying date accounted for most of the remaining variance.     

Social interactions among breeding Brünnich's Guillemots Uria lomvia suggest constraints in relation to offspring vulnerability

Brünnich's Guillemots Uria lomvia are adapted to high-density breeding in large colonies on steep cliffs. Because they breed on narrow ledges, egg loss through dislodgement is an important cause of breeding failure. Fighting among breeders presumably raises the risk of accidental dislodgement. In this study, we investigated whether social behaviour among Brünnich's Guillemots shows any adaptations to reduce accidental egg loss by modifying behaviour during incubation. We found that the amount of aggression increased significantly at the time of hatching, perhaps in response to the reduced risk of breeding failure through dislodgement of a chick, compared with an egg. Allopreening followed an inverse trend, falling significantly after the day of hatch. This supports the hypothesis that allopreening is used to reduce aggressive interactions. At the same time, the frequency of allopreening was greatly increased on days when mosquitoes affected the birds, consistent with the hypothesis that allopreening is part of a defence strategy against ectoparasites.     

Morphometric study of embryonic development of Macrobrachium borellii (Arthropoda: Crustacea)

Summary

Embryo morphometry and developing time from just-laid eggs until hatching were described in the palaemonid prawn, Macrobrachium borellii, using a rapid non-invasive staging method. Embryos were kept in the laboratory under controlled conditions and development divided into seven stages according to major morphological characteristics. This lecithotrophic, freshwater shrimp has a highly abbreviated type of hatching development after 39 ± 2 days as postlarvae at 24°C. Morphometry was recorded using a stereoscopic microscope with an image analyzer. Area, perimeter, maximum and minimum diameters and shape were measured in yolk sac, egg-coat and eye, respectively, and they were statistically selected as the best to define the stages. Eggs are ovoid with a maximum diameter that varies from the moment of oviposition to the time of hatching from 1.5 to 2.0 mm, respectively. Water content and egg size increase along with development, whereas egg shape only varies just before hatching when the egg becomes strongly ovoid. Egg coat and eye variables significantly increase as the embryo develops while all yolk variables decrease as the embryo consumes the vitellus. Yolk represents more than 95% of the egg at the time of oviposition, falling to 22% by the time of hatching. The major yolk area decrease is observed between stages 4 and 5, which is coincident with a marked increase in the catabolism. Using only egg coat, yolk and eye shape and maximum diameter, a researcher can straightforwardly identify a developing stage with an accuracy ranging from 70 to 100%. This tool may be employed in other species provided they have transparent chorion.     

Comparison of egg period, hatching rate, and first-instar nymphs among three species of stonefly

Egg period was compared among several temperature conditions (11°C, 16°C, 20°C, 23°C) in Sweltsa sp., Stavsolus japonicus, and Isoperla aizuana (Plecoptera). The shortest mean egg incubation period was 27.8 days at 20°C in Sweltsa sp., 118.1 days at 16°C in Stavsolus japonicus, and 162.0 days at 20°C in Isoperla aizuana on average. Egg hatching rate was also the highest at the water temperature that provided the shortest egg incubation period. Based on laboratory data, eggs of Sweltsa sp. were considered to be deposited in May and hatched in June in the field. Thus, they must have spent the summer as nymphs in the field. Eggs of Stavsolus japonicus and Isoperla aizuana were considered to be deposited in April to May and hatched in September to October in the field. Visible eyes of Stavsolus japonicus and Isoperla aizuana appeared in August. It is likely that the long egg period of Stavsolus japonicus and Isoperla aizuana reflects that these two species spend the summer as dormant eggs in the field.     

Sex-biased aggression and male-only care at sea in Brünnich's Guillemots Uria lomvia and Razorbills Alca torda

In several groups in the order Charadriformes, biparental care is followed by a period of male‐only care. Several hypotheses attempting to explain extended male parental care in shorebirds do not fit the Alcini. In a previous study of Brünnich’s Guillemots Uria lomvia and Razorbills Alca torda, we did not find support for female‐biased parental effort at the breeding site that would lead to males being in better condition to care for chicks at sea. However, in both species, males spent more off‐duty time at the breeding site than females, suggesting greater involvement in the defence of egg or chick, breeding site and mate. We predicted that there would be a male bias in size and aggressive behaviour associated with parental roles. To test this, body size and aggression of attending male and female Brünnich’s Guillemots and Razorbills were measured during incubation and brooding on the Gannet Islands, Labrador. Parental aggression was measured using natural observations of all agonistic interactions and, in Razorbills only, in situ responses to presentations of a predator model. In both species, males were significantly larger than females in culmen and gape length. Guillemot males initiated agonistic interactions more frequently than females during incubation. In contrast, female Guillemots were subjected to aggression more frequently than males and as a result were involved in more fights. In addition, the few chicks that were seen to die were being attended by single females. During the brooding period, Razorbill males responded aggressively to intruders more frequently than females, made more aggressive responses than females, and responded aggressively more frequently and more intensely than females to a predator model. In both species there was a similar male bias in morphology and behaviour that is consistent with male parents being more capable of protecting their chick, a probable advantage to chick survival during the uniparental care phase of some Charadriformes.     

New insights from old eggs – the shape and thickness of Great Auk Pinguinus impennis eggs

We compared the shape and eggshell thickness of Great Auk Pinguinus impennis eggs with those of its closest relatives, the Razorbill Alca torda, Common Guillemot Uria aalge and Brünnich's Guillemot Uria lomvia, in order to gain additional insights into the breeding biology of the extinct Great Auk. The egg of the Great Auk was most similar in shape to that of Brünnich's Guillemot. The absolute thickness of the Great Auk eggshell was greater than that of the Common Guillemot and Razorbill egg, which is as expected given its greater size, but the relative shell thickness at the equator and pointed end (compared with the blunt end) was more similar to that of the Common Guillemot. On the basis of these and other results we suggest that Great Auk incubated in an upright posture in open habitat with little or no nest, where its pyriform egg shape provided stability and allowed safe manoeuvrability during incubation. On the basis of a recent phylogeny of the Alcidae, we speculate that a single brood patch, a pyriform egg and upright incubation posture, as in the Great Auk and the two Uria guillemots, is the ancestral state, and that the Razorbill – the Great Auk's closest relative – secondarily evolved two brood patches and an elliptical egg as adaptations for horizontal incubation, which provides flexibility in incubation site selection, allowing breeding in enclosed spaces such as crevices, burrows or under boulders, as well as on open ledges.     

Variation in butterfly egg adhesion: adaptation to local host plant senescence characteristics?

Most butterflies that overwinter as diapausing eggs have evolved oviposition behaviours where egg placement is near or on perennating portions of their host plant. We describe an alternative strategy used by alpine populations of a Lycaenid butterfly species complex where eggs are not attached strongly to the leaf substrate and subsequently fall off the plant. Other populations occurring at lower elevations attach their eggs strongly to the plant. In the laboratory, we measured the amount of force required to detach eggs from the leaf substrate. Eggs of the non‐alpine populations required more than five times the force required to detach eggs of the alpine population. Field surveys of the host plant characteristics used by various populations suggest that easy detachment may be adaptive for alpine populations because, unlike the host plants used by other populations, nearly all of the preceding year's vegetative growth is blown away by strong winds during the winter months.     

FACTORS AFFECTING BREEDING OF RAZORBILLS ALCA TORDA ON SKOKHOLM

A study of the breeding biology of the Razorbill was carried out on Skokholm (South Wales) during 1971-73. Birds ringed or colour ringed before the study began provided additional information upon the effects of age on breeding. Mean laying date was delayed in 1972, compared with 1971; the effect is attributed chiefly to stormy weather which upset colony attendance. Eggs were also smaller in 1972. A seasonal decline in egg size (volume) was noted in all three years, attributed mainly to the later laying of young birds. Egg size increased with age, at least up to the fifteenth year. Eggs lost totalled 30% of those laid; 73% of this total was due to predation by Herring Gulls and of Jackdaws. Most losses (45%) occurred during the first 10 days after laying. Of lost eggs, 25% were replaced, usually 14 days after the loss of the original; only eggs laid and lost early in the season could be replaced. Only 7% of the chicks which hatched failed to fledge. Most (62.5%) chick losses occurred in the first week of nestling life, when chick weight was related to egg size. Afterwards, both growth rate and fledging weight were independent of egg size. The chicks fledging early in the season were heavier than later chicks. Failure to fledge was mainly due to a breakdown in behaviour between parent and young, rather than to predation. Breeding success was highest for birds breeding early in the season, most of which were older, more experienced breeders. These laid early enough to replace an egg if it was lost; they produced large eggs, and their chicks were therefore both heavier than average during the critical first 7–10 days of life, and fledged at a high weight. Thus experience accumulated with age, and the ability to lay early in the season are important for successful breeding in the Razorbill.     

A review of the evidence for the translocation of eggs and young by nightjars (Caprimulgidae)

I undertook a review of evidence for the translocation of eggs and young by nightjars through a comprehensive search of the literature. I found that most of the evidence was based on hearsay, supposition, a misunderstanding of nightjar behaviour, or on the repetition of a story going back 200 years to Le Vaillant, via Audubon. There is no satisfactory direct evidence of any nightjar deliberately airlifting its eggs or young away from a disturbance. This conclusion is based on 10 studies by ornithologists across five continents, involving over 522 nests of 13 species. Accidental airlifting occurs occasionally when an egg or young chick gets stuck to the ventral plumage of the sitting adult. This is most likely to occur near the time of hatching. A nightjar can move an egg along the ground for a short distance, either by placing its lower mandible over the egg and then walking backwards while rolling it, or by rolling it forwards with its feet. Nightjars regularly move their chicks, especially after a disturbance. They do so by first moving away themselves and then calling. The chicks, who are highly mobile within hours of hatching, respond immediately by running to the parent birds.     

The early embryonic mitosis in normal and cooled eggs of the silkworm,Bombyx mori

Early embryonic mitosis of the silkworm, Bombyx mori, was morphologically studied in the normal eggs and in the eggs treated by low temperature (?10°C). The first embryonic mitosis is observed in the eggs at 120 to 150 minutes after deposition at 26°C. After egg and sperm pronuclei unite, a spindle is formed in each of the pronuclei independently. At metaphase and anaphase paternal and maternal chromosomes are in separate groups on a spindle (gonomeric) and karyogamy takes place at telophase when they reach the poles. The second embryonic mitosis is shown in the eggs at 180 to 210 minutes after deposition. The division of two nuclei is not synchronous in the silkworm, and the mitosis is not gonomeric. In the eggs treated by low temperature, spindle fibers are not observed at all at ?10°C, and chromosomes, which form two deeply stained masses of irregular shape, are seen in the less stained area of spindle shape. When the eggs are returned to 26°C, some eggs go into normal gonomeric division, while some form two small and compact spindles, which seem to be derived from each of the pronuclei. It was observed that these compact spindles are able to continue mitosis.     

Reproductive performance of the hemlock looper,Lambdina fiscellaria,as a function of temperature and population origin

In a recent study of the hemlock looper (HL), Lambdina fiscellaria (Guenée) (Lepidoptera: Geometridae), long exposure of early‐diapausing eggs to high temperatures considerably reduced their ability to hatch. This finding raised the possibility that adults could also be negatively affected by increasing temperatures if they reproduced too early in the season in response to global warming. To investigate this hypothesis, newly formed HL pupae from three populations of eastern Canada ‐ Quebec (QC), Newfoundland (NL), and Labrador (LB) ‐ were submitted to four constant temperatures (10, 15, 20, or 25 °C) during pupal and adult development. The effect of population origin on HL reproduction was generally negligible. Mating probability was high at 15 and 20 °C (0.86 and 0.83, respectively), quite low at 10 °C (0.53), and even lower at 25 °C (0.38). Mating started earlier in the night and lasted longer as temperature decreased. Both productivity and absolute fecundity increased when temperature increased from 10 to 15 °C and then decreased slowly as temperature increased further. Over populations and temperatures, relative fecundity averaged 0.95, indicating that females had enough time to lay most of their eggs before they died. High temperatures had a deleterious effect on egg fertility: between 10 and 20 °C, relative fertility was about 0.90, but it dropped to 0.51 at 25 °C. The average proportion of fertile eggs declined from 0.88 in the first quarter of the egg‐laying period to 0.57 in the last quarter, suggesting lower sperm count or viability, or deterioration of the oocytes as the egg‐laying period progresses. Based on these findings, we argue that the production of an additional fifth instar among HL populations of southern origin can be viewed as an adaptive mechanism allowing adults to postpone reproduction or the egg‐laying period in order to mitigate the detrimental effect of high temperatures on their probability of mating successfully or that of laying fertile eggs.     

A brood parasite selects for its own egg traits

Many brood parasitic birds lay eggs that mimic their hosts'' eggs in appearance. This typically arises from selection from discriminating hosts that reject eggs which differ from their own. However, selection on parasitic eggs may also arise from parasites themselves, because it should pay a laying parasitic female to detect and destroy another parasitic egg previously laid in the same host nest by a different female. In this study, I experimentally test the source of selection on greater honeyguide (Indicator indicator) egg size and shape, which is correlated with that of its several host species, all of which breed in dark holes. Its commonest host species did not discriminate against experimental eggs that differed from their own in size and shape, but laying female honeyguides preferentially punctured experimental eggs more than host or control eggs. This should improve offspring survival given that multiple parasitism by this species is common, and that honeyguide chicks kill all other nest occupants. Hence, selection on egg size in greater honeyguides parasitizing bee-eaters appears to be imposed not by host defences but by interference competition among parasites themselves.     

Egg characteristics are unreliable in determining maternity in communal clutches of guira cuckoos Guira guira

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.     

Biology and behaviour of the neotropical ant-mimicking spider Aphantochilus rogersi (Araneae: Aphantochilidae): nesting, maternal care and ontogeny of ant-hunting techniques

Aphantochilus rogersi is an ant-mimicking spider that preys exclusively on cephalotine ants. The spiders oviposit in close proximity to nests of the model ant Zacryptocerus pusillus , and emergent spiderlings tend to remain in the vicinity of natal egg sacs. Females of A. rogersi actively defend their egg sacs against approaching workers of Z. pusillus , but the latter may sometimes destroy the eggs. Feeding specialization on these ants is confirmed by more than 300 observation of young and adult A. rogersi carrying ant corpses in the field. Although A. rogersi possesses several behavioural traits which may reduce the risk of being injured by ants during subjugation, field and laboratory observations showed that social defence by Z. pusillus may cause mutilation to the spiders. Tests in captivity revealed an ontogenetic change in the prey-capture techniques employed by A. rogersi. Early-instar spiderlings can apparently only seize the ant's petiole tightly if they are able to approach the ant from the front. As the ant is paralysed, the spiderling positions itself vertically in relation to the substratum. Larger spiders, on the other hand, attack ants most frequently from behind, and seem better equipped to seize the ant's petiole firmly with their larger chelicerae. Owing to their greater strength, late-instar spiders are able to lift the struggling ant aloft. The selection of a suitable oviposition site, the mother's ability to defend herself and the eggs from nearby ants, and the capacity to capture and subdue ants safely from emergence to maturity, are regarded as crucial traits inherent in the mimetic and feeding specialization by A. rogersi.     

THERMAL EVOLUTION OF EGG SIZE IN DROSOPHILA MELANOGASTER

We measured the size of eggs produced by populations of Drosophila melanogaster that had been collected along latitudinal gradients in different continents or that had undergone several years of culture at different temperatures in the laboratory. Australian and South American populations from higher latitudes produced larger eggs when all were compared at a standard temperature. Laboratory populations that had been evolving at 16.5°C produced larger eggs than populations that had evolved at 25°C or 29°C, suggesting that temperature may be an important selective agent in producing the latitudinal clines. Flies from laboratory populations produced larger eggs at an experimental temperature of 16.5°C than at 25°C, and there was no indication of genotype-environment interaction for egg size. Evolution of egg size in response to temperature cannot be accounted for by differences in adult body size between populations. It is not clear which life-history traits are direct targets of thermal selection and which are showing correlated responses, and disentangling these is a task for the future.     

Mothers modify eggs into shields to protect offspring from parasitism

Eggs are an immobile, vulnerable stage of development and their success often depends on the oviposition decisions of the mother. Studies show that female animals, and sometimes males, may invest parental resources in order to increase the survival of their offspring. Here, we describe a unique form of parental investment in offspring survival. The seed beetle Mimosestes amicus may lay eggs singly, or may cover eggs with additional egg(s). This egg stacking serves to significantly reduce the mortality of the protected egg from parasitism by the parasitic wasp, Uscana semifumipennis. The smaller top eggs serve only as protective shields; they are inviable, and wasps that develop in them suffer negative fitness consequences. Further, we found egg stacking to be inducible; M. amicus increase the number of stacks they lay when parasitoids are present. However, stacking invokes a cost. When wasps are absent, beetles lay more single eggs, and produce more offspring, highlighting the adaptive value of this extraordinary example of behavioural plasticity in parental investment.     

Prospects of herbivore egg‐killing plant defenses for sustainable crop protection

Due to a growing demand of food production worldwide, new strategies are suggested to allow for sustainable production of food with minimal effects on natural resources. A promising alternative to the application of chemical pesticides is the implementation of crops resistant to insect pests. Plants produce compounds that are harmful to a wide range of attackers, including insect pests; thus, exploitation of their natural defense system can be the key for the development of pest‐resistant crops. Interestingly, some plants possess a unique first line of defense that eliminates the enemy before it becomes destructive: egg‐killing. Insect eggs can trigger (1) direct defenses, mostly including plant cell tissue growth or cell death that lead to eggs desiccating, being crushed or falling off the plant or (2) indirect defenses, plant chemical cues recruiting natural enemies that kill the egg or hatching larvae (parasitoids). The consequences of plant responses to eggs are that insect larvae do not hatch or that they are impeded in development, and damage to the plant is reduced. Here, we provide an overview on the ubiquity and evolutionary history of egg‐killing traits within the plant kingdom including crops. Up to now, little is known on the mechanisms and on the genetic basis of egg‐killing traits. Making use of egg‐killing defense traits in crops is a promising new way to sustainably reduce losses of crop yield. We provide suggestions for new breeding strategies to grow egg‐killing crops and improve biological control.     

How avian incubation behaviour influences egg surface temperatures: relationships with egg position,development and clutch size

While understanding heat exchange between incubating adults and their eggs is central to the study of avian incubation energetics, current theory based on thermal measurements from dummy eggs reveals little about the mechanisms of this heat exchange or behavioural implications for the incubating bird. For example, we know little about how birds distribute their eggs based on temperature differences among egg positions within the nest cup. We studied the great tit Parus major, a species with a large clutch size, to investigate surface cooling rates of individual eggs within the nest cup across a range of ambient temperatures in a field situation. Using state‐of‐the‐art portable infrared imaging and digital photography we tested for associations between egg surface temperature (and rate of cooling) and a combination of egg specific (mass, shape, laying order, position within clutch) and incubation specific (clutch size, ambient temperature, day of incubation) variables. Egg surface temperature and cooling rates were related to the position of the eggs within the nest cup, with outer eggs being initially colder and cooling quicker than central eggs. Between foraging bouts, females moved outer eggs significantly more than centrally positioned eggs. Our results demonstrate that females are capable of responding to individual egg temperature by moving eggs around the nest cup, and that the energy cost to the female may increase as incubation proceeds. In addition, our results showing that smaller clutches experience lower initial incubation temperatures and cool quicker than larger clutches warrant further attention for optimal clutch size theory and studies of energetic constraints during incubation. Finally, researchers using dummy eggs to record egg temperature have ignored important elements of contact‐incubation, namely the complexity of how eggs cool and how females respond to these changes.