Genetic diversity and population history of two related seabird species based on mitochondrial DNA control region sequences

Geographical variation in two related seabird species, the razorbill (Alca torda) and common guillemot (Uria aalge), was investigated using sequence analysis of mitochondrial DNA (mtDNA) control regions. We determined the nucleotide sequence of the variable 5' segment of the control region in razorbills and common guillemots from breeding colonies across the Atlantic Ocean. The ecology and life history characteristics of razorbill and common guillemot are in many respects similar. They are both considered highly philopatric and have largely overlapping distributions in temperate and subarctic regions of the North Atlantic, yet the species were found to differ widely in the extent and spatial distribution of mtDNA variation. Moreover, the differences in genetic differentiation and diversity were in the opposite direction to that expected from a consideration of traditional classifications and current population sizes. Indices of genetic diversity were highest in razorbill and varied among colonies, as did genotype frequencies, suggestive of restrictions to gene flow. The distribution of genetic variation suggests that razorbills originated from a refugial population in the south-western Atlantic Ocean through sequential founder events and subsequent expansion in the east and north. In common guillemots, genetic diversity was low and there was a lack of geographical structure, consistent with a recent population bottleneck, expansion and gene flow. We suggest that the reduced level of genetic diversity and differentiation in the common guillemot is caused by an inherent propensity for repeated population bottlenecks and concomitantly unstable population structure related to their specialized feeding ecology.     

Effects of body size,sex, parental care and moult strategies on auk diving behaviour outside the breeding season

Information on seabird foraging behaviour outside the breeding season is currently limited. This knowledge gap is critical as this period is energetically demanding due to post‐fledging parental care, feather moult and changing environmental conditions. Based on species’ body size, post‐fledging parental strategy and primary moult schedule we tested predictions for key aspects of foraging behaviour (maximum dive depth (MDD), daily time submerged (DTS) and diurnal dive activity (DDA)) using dive depth data collected from three seabird species (common guillemot Uria aalge, razorbill Alca torda and Atlantic puffin Fratercula arctica) from the end of the breeding season (July) to mid‐winter (January). We found partial support for predictions associated with body size; guillemots had greater MDD than razorbills but MDD did not differ between razorbills and puffins, despite the former being 35% heavier. In accordance with sexual monomorphism in all three species, MDD did not differ overall between the sexes. However, in guillemots and razorbills there were sex‐specific differences, such that male guillemots made deeper dives than females, and males of both species had higher DTS. In contrast, there were no marked sex differences in dive behaviour of puffins in July and August in accordance with their lack of post‐fledging parental care and variable moult schedule. We found support for the prediction that diving effort would be greater in mid‐winter compared to the period after the breeding season. Despite reduced daylight in mid‐winter, this increase in DTS occurred predominantly during the day and only guillemots appeared to dive nocturnally to any great extent. In comparison to diving behaviour of these species recorded during the breeding season, MDD was shallower and DTS was greater during the non‐breeding period. Such differences in diving behaviour during the post‐breeding period are relevant when identifying potential energetic bottlenecks, known to be key drivers of seabird population dynamics.     

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.     

Male chicks are more costly to rear than females in a monogamous seabird, the Common Murre

Feeding rates and mass loss during chick rearing were comparedfor individually marked parents of male and female Common Murre(Uria aalge) chicks at Great Island, Newfoundland, Canada, from1997–2001. Both parents in this socially monogamous seabirdspecies share parental care duties until colony departure, afterwhich the single chick is fed only by its father. Because murresprovision their single chicks with one clearly visible fishper trip, it is possible to accurately determine whether parentsdifferentially feed male and female chicks. Based on slightlygreater mass of males in adulthood, possibly favored by sexdifferences in breeding roles, we predicted that male nestlingswould be fed more than females. Fathers' feeding rate to sons,but not daughters, increased with chick age, whereas maternalfeeding rate increased with chick age for both sexes. When year-correctedfeeding rates of pairs rearing both sexes were compared, bothmothers and fathers fed their sons significantly more than theirdaughters in the later part of the chick-rearing period. Moreover,parents rearing male chicks lost mass at a significantly higherrate than those rearing females. There was no difference infledging age for sons and daughters. These results indicatethat differential parental allocation occurs and has measurablecosts even in a species with only slight adult sexual dimorphism.     

Development and Validation of a Wing Key to Improve Harvest Management of Alcids in the Northwest Atlantic

Abstract: Murres (thick-billed [Uria lomvia] and common [U. aalge]) are legally hunted along the coast of Newfoundland and Labrador, Canada. Razorbills (Alca torda) are also incidentally taken. Only irregular estimates of the total murre harvest are available, so a tool to derive estimates of age- and species-specific harvest is required to effectively monitor the hunt and manage a sustainable harvest. We collected 293 murre and razorbill wings from hunters between 1999–2004, with the goal of identifying wing characteristics that could be used to discriminate age and species. We found that murres and razorbills could be reliably aged (first-yr vs. older) on the basis of molt limits of greater wing coverts. Using a discriminant function (DF) incorporating length of the first primary and second secondary feather, we classified 95–96% of common murres and 99–100% of thick-billed murres correctly to species. First-year thick-billed and common murres also differed in number of pale secondary coverts (median = 12 and 3, respectively), providing another species-specific trait. We developed a key to age and assign species based on these results. We assessed applicability and accuracy of the wing-key with novice observers, who differentiated between murre and razorbill wings using feather-pattern coloration with high accuracy (95 ± 9%) and were able to differentiate between the 2 murres species using 3 techniques: visual assessment of wing shape (83 ± 14% accuracy), the DF (94 ± 6%), and number of worn secondary coverts for first-year birds only (83 ± 5%). Experience increased success rates of aging and species classification using wing shape and number of worn secondary coverts but not using the DF. Despite differences in measurement accuracy and repeatability among observers, the DF proved to be robust. Our results will facilitate implementation of a species composition survey for the murre hunt and will improve identification rates of carcasses found during beached bird surveys in the Northwest Atlantic, aiding in monitoring of alcid populations vulnerable to anthropogenic activities.     

Mechanisms of egg contamination by Salmonella Enteritidis

Salmonella Enteritidis (SE) has been the major cause of the food-borne salmonellosis pandemic in humans over the last 20 years, during which contaminated hen's eggs were the most important vehicle of the infection. Eggs can be contaminated on the outer shell surface and internally. Internal contamination can be the result of penetration through the eggshell or by direct contamination of egg contents before oviposition, originating from infection of the reproductive organs. Once inside the egg, the bacteria need to cope with antimicrobial factors in the albumen and vitelline membrane before migration to the yolk can occur. It would seem that serotype Enteritidis has intrinsic characteristics that allow an epidemiological association with hen eggs that are still undefined. There are indications that SE survives the attacks with the help of antimicrobial molecules during the formation of the egg in the hen's oviduct and inside the egg. This appears to require a unique combination of genes encoding for improved cell wall protection and repairing cellular and molecular damage, among others.     

Effects of experimental calcium availability,egg parameters and laying order on Great Tit Parus major eggshell pigmentation patterns

Many bird species lay eggs speckled with protoporphyrin‐based spots, however, for most of them the function of eggshell spotting is unknown. A plausible hypothesis is that protoporphyrin might have a structural function in strengthening the eggshell and is therefore deposited when calcium is scarce. In this study, we experimentally provided Great Tit Parus major females with supplemental calcium to examine its effect on the protoporphyrin‐based maculation of their eggs. In addition, we studied variation in eggshell pigmentation patterns in relation to other egg parameters and laying order. Calcium‐supplemented females laid larger eggs but shell thickness was not significantly affected by the treatment. Calcium supplementation may reduce the time and energy females devote to searching for calcium‐rich material, so that they can collect more nutrients and so lay larger eggs. Furthermore, pigment darkness was associated with egg volume and shape, which suggests that female quality and environmental food availability may also influence the shell pigmentation pattern. Within clutches, later‐laid eggs had larger and darker spots that were distributed more unevenly on the shell surface. This within‐clutch pattern could be explained by the increase in egg volume and egg shape and a decline in shell thickness with egg‐laying order, which characteristics were all related to shell‐spotting pattern. Eggs with a coronal ring had thinner shells, but pigment intensity and spot size were not related to shell thickness. Thus, our results suggest that concentrated spotting distribution may have a mechanical function, supporting the structural‐function hypothesis.     

Effects of back-mounted biologgers on condition,diving and flight performance in a breeding seabird

Biologging devices are providing detailed insights into the behaviour and movement of animals in their natural environments. It is usually assumed that this method of gathering data does not impact on the behaviour observed. However, potential negative effects on birds have rarely been investigated before field-based studies are initiated. Seabirds which both fly and use pursuit diving may be particularly sensitive to increases in drag and load resulting from carrying biologging devices. We studied chick-rearing adult common guillemots Uria aalge equipped with and without back-mounted GPS tags over short deployments of a few days. Concurrently guillemots carried small leg-mounted TDR devices (time-depth recorders) providing activity data throughout. Changes in body mass and breeding success were followed for device equipped and control guillemots. At the colony level guillemots lost body mass throughout the chick-rearing period. When-equipped with the additional GPS tag, the guillemots lost mass at close to twice the rate they did when equipped with only the smaller leg-mounted TDR device. The elevated mass loss suggests an impact on energy expenditure or foraging performance. When equipped with GPS tags diving performance, time-activity budgets and daily patterns of activity were unchanged, yet dive depth distributions differed. We review studies of tag-effects in guillemots Uria sp. finding elevated mass loss and reduced chick-provisioning to be the most commonly observed effects. Less information is available for behavioural measures, and results vary between studies. In general, small tags deployed over several days appear to have small or no measurable effect on the behavioural variables commonly observed in most guillemot tagging studies. However, there may still be impacts on fitness via physiological effects and/or reduced chick-provisioning, while more detailed measures of behaviour (e.g. using accelerometery) may reveal effects on diving and flight performance.     

Seasonal changes of impala (Aepyceros melampus, Lichtenstein, 1812) faecal helminth egg counts through a one-year period

The study was done in Lake Mburo area, Uganda, consisting of Lake Mburo National Park and surrounding ranches. The study involved monitoring changes of output of helminth eggs per gram (e.p.g.) by impala for 12 months. Fresh faecal samples, 178 in total, were collected from the rangeland and subjected to floatation and culture tests. There was a peak faecal e.p.g. output from March to April and from October to November. This period coincided with time of high rainfall and calving peak of impala. There was no faecal e.p.g. observed during the dry spell from June to July.     

The modelling of avian visual perception predicts behavioural rejection responses to foreign egg colours

How do birds tell the colours of their own and foreign eggs apart? We demonstrate that perceptual modelling of avian visual discrimination can predict behavioural rejection responses to foreign eggs in the nest of wild birds. We use a photoreceptor noise-limited colour opponent model of visual perception to evaluate its accuracy as a predictor of behavioural rates of experimental egg discrimination in the song thrush Turdus philomelos. The visual modelling of experimental and natural eggshell colours suggests that photon capture from the ultraviolet and short wavelength-sensitive cones elicits egg rejection decisions in song thrushes, while inter-clutch variation of egg coloration provides sufficient contrasts for detecting conspecific parasitism in this species. Biologically realistic sensory models provide an important tool for relating variability of behavioural responses to perceived phenotypic variation.     

Effect of population bottlenecks on the egg morphology of introduced birds in New Zealand

Hatching failure is inversely correlated with population bottleneck size among exotic birds introduced to New Zealand, but the mechanism for this is unclear. We assess whether the bottlenecks these species experienced during their introduction have affected hatching failure through deleterious changes in egg morphology. We measured egg size and shape of 13 species that passed through bottlenecks of 11–800 individuals during their establishment in the 19th century. Eggs were also measured in the source populations ('pre-bottleneck') of each species to compare pre- and post-bottleneck egg morphology directly. Significant changes in egg volume were found in six of 13 species, with most laying smaller eggs in New Zealand. Egg shape changed in four of 13 species but there was no directional bias; two species developed more elongated eggs and two species broader eggs. There was no relationship between bottleneck size and change in egg volume, but species passing through severe bottlenecks had greater variability in egg volume and were more likely to have eggs that deviated in shape from their source populations. There was no relationship between changes in either egg volume or shape and rates of hatching failure. Further work is needed to assess whether changes in egg morphology have negative consequences on offspring fitness and whether the observed changes are the result of differing environmental conditions in the introduced range.     

A new egg with avian egg shape from the Upper Cretaceous of Zhejiang Province,China

A new egg is described from the Upper Cretaceous lacustrine deposit of the Chichengshan Formation in the Tiantai Basin, Zhejiang Province, southeast China. The new specimen shares eggshell micro-features with members of the oofamily Stalicoolithidae, Paraspheroolithus of the oofamily Spheroolithidae and Mosaicoolithus (oofamily indet.), with barrel-shaped cones, prolatocanaliculate pore system, horizontal accretion lines and light stripes throughout the eggshell. However, the new egg differs from the aforementioned ootaxa by its small size and asymmetrical shape, revealing new morphological variation among eggs with microstructure similar to that of Paraspheroolithus, Mosaicoolithus and Stalicoolithidae. We refer the new egg to a new oogenus and oospecies, thus increasing the diversity of the Tiantai Basin oofauna. The Tiantai Basin has yielded a variety of dinosaur eggs and one turtle clutch. Comparatively, the new egg is surprisingly small and ovoid, a morphology usually associated with avian eggs, although the absence of a squamatic layer excludes the egg from being referred to this group.     

Quantifying the costs and benefits of protective egg coating in a Chrysomelid beetle

Abstract.  1. The costs and benefits of behavioural care of offspring can often be easily quantified through observations and experiments. Other forms of parental investment, on the other hand, are usually less amenable to cost–benefit analysis.
2. Here, the costs and benefits are estimated for protective egg coating by a chrysomelid beetle, Cryptocephalus hypochaeridis , where the female spends a considerable amount of time adding extra structural components to each of the eggs after laying them.
3. Adding this protective coating was very costly, both in terms of material and energy used: the mass of the extrachorion is equivalent to half the mass of the egg, and water loss and energy expenditure while coating the egg is equivalent to half what would be lost while laying a further egg.
4. Choice tests with egg predators demonstrated that these high costs are offset by benefits in terms of protection against predation: whereas uncoated eggs are readily eaten by predators, coated eggs are always rejected.     

Allometry of egg and hatchling mass in birds and reptiles: roles of developmental maturity, eggshell structure and phylogeny

The factors determining hatchling mass (HM) are investigated in a wide range of birds and reptiles using regression analysis, analysis of covariance and comparative analysis by independent contrasts. In birds, initial egg mass (IEM) at laying is the most important factor affecting HM and phylogenetic relatedness has no significant effects on HM. Developmental maturity of the avian neonates did not affect the proportion of IEM converted into HM. For all reptile species, IEM also significantly affected HM but phylogenetic relatedness did not. By contrast, allometric relationships between IEM and HM in the different orders of reptiles were affected by shell type. The robustness of allometric relationships across taxa in birds and reptiles suggests that there is a physiological link between IEM and HM, which contrasts with that observed for the relationship between egg mass and incubation period. This result has significant implications for the inter-relationships between IEM and embryonic growth, which are discussed for birds and reptiles.     

Energetics of reproduction: consequences of divergent selection on egg size, food limitation, and female age for egg composition and reproductive effort in a butterfly

Using lines artificially selected on egg size and being subjected to a restricted and an unrestricted feeding treatment, we examined the relationships between egg size, egg number, egg composition, and reproductive investment in the butterfly Bicyclus anynana . Despite a successful manipulation of egg size, correlated responses to selection in larval time, pupal mass, pupal time, longevity, fecundity, or the amount of energy allocated to reproduction were virtually absent. Thus, there was no indication for an evolutionary link between offspring size and reproductive investment. Egg composition, in contrast, was affected by selection, with larger eggs containing relatively more lipid and water, but less protein and energy compared to smaller eggs. Hence, females producing large eggs did not have to sacrifice fecundity due to adjustments in egg composition. Food limitation per se caused only minor changes in egg composition, and there was no general reduction in egg provisioning with female age. The latter was restricted to food-limited females, whereas egg quality remained remarkably similar throughout the females' life in control groups. We conclude that neglecting changes in biochemical egg composition, depending on genetic background, food availability, and female age, may introduce substantial error when estimating reproductive effort, and may ultimately lead to invalid conclusions.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 403–418.     

The evolution of egg shape in birds: selection during the incubation period

A recent broad comparative study suggested that factors during egg formation – in particular ‘flight efficiency’, which explained only 4% of the interspecific variation – are the main forces of selection on the evolution of egg shape in birds. As an alternative, we tested whether selection during the incubation period might also influence egg shape in two taxa with a wide range of egg shapes, the alcids (Alcidae) and the penguins (Spheniscidae). To do this, we analysed data from 30 species of these two distantly related but ecologically similar bird families with egg shapes ranging from nearly spherical to the most pyriform eggs found in birds. The shape of pyriform eggs, in particular, has previously proven difficult to quantify. Using three egg‐shape indices – pointedness, polar‐asymmetry and elongation – that accurately describe the shapes of all birds’ eggs, we examined the effects of egg size, chick developmental mode, clutch size and incubation site on egg shape. Linear models that include only these factors explained 70–85% of the variation in these egg‐shape indices, with incubation site consistently explaining > 60% of the variation in shape. The five species of alcids and penguins that produce the most pyriform eggs all incubate in an upright posture on flat or sloping substrates, whereas species that incubate in a cup nest have more spherical eggs. We suggest that breeding sites and incubation posture influence the ability of parents to manipulate egg position, and thus selection acting during incubation may influence egg‐shape variation across birds as a whole.     

The capitula of phasmid eggs: an update with a review of the current state of phasmid ootaxonomy

The relationship between egg shape and presence or absence of a capitulum is considered. The distribution of capitula of different types within the order Phasmida is reviewed. These tend to confirm tribal and subfamily groupings but suggest that the suborder Anareolatae is polyphyletic and some suggestions are made for its regrouping. The species of Phasmida which have been recently studied from an ootaxonomic viewpoint are surveyed.     

Detection of the nifH gene of Methanobrevibacter smithii: a potential tool to identify sewage pollution in recreational waters

AIMS: The goal of this study was to develop and test the efficacy of a PCR assay for the environmental detection of the nifH gene of Methanobrevibacter smithii, a methanogen found in human faeces and sewage. METHODS AND RESULTS: PCR primers for the nifH gene of M. smithii were designed, tested and used to detect the presence or absence of this organism in faecal and environmental samples. Specificity analysis showed that the Mnif primers amplified products only in M. smithii pure culture strains (100%), human faeces (29%), human sewage samples (93%) and sewage-contaminated water samples (100%). No amplification was observed when primers were tested against 43 bacterial stock cultures, 204 animal faecal samples, 548 environmental bacterial isolates and water samples from a bovine waste lagoon and adjacent polluted creek. Sequencing of PCR products from sewers demonstrated that a 222-bp product was the nifH gene of M. smithii. The minimal amount of total DNA required for the detection of M. smithii was 10 ng for human faeces, 10 ng for faecally contaminated water and 5 ng for sewage. Recreational water seeded with M. smithii established a lower detection limit of 13 cells ml(-1). CONCLUSIONS: The Mnif assay developed during this investigation showed successful detection of M. smithii in individual human faecal samples, sewage and sewage-contaminated water but not in uncontaminated marine water or bovine-contaminated waters. The Mnif assay appears to be a potentially useful method to detect sewage-polluted coastal waters. SIGNIFICANCE AND IMPACT OF THE STUDY: This study was the first to utilize methanogens as an indicator of sewage pollution. Mnif PCR detection of M. smithii was shown to be a rapid, inexpensive and reliable test for determining the presence or absence of sewage pollution in coastal recreational waters.     

Effects of dietary combination of chromium and biotin on egg production, serum metabolites, and egg yolk mineral and cholesterol concentrations in heat-distressed laying quails

Chromium picolinate is used in the poultry diet because of its antistress effects in addition to the fact that the requirement for it is increased during stress. This study was conducted to determine if the negative effects of high ambient temperature (34°C) on egg production, egg quality, antioxidant status, and cholesterol and mineral content of egg yolk could be alleviated by combination of chromium picolinate and biotin (0.6/2.0; Diachrome^™, as formulated by Nutrition 21 Inc.), in laying Japanese quails (Coturnix coturnix japanica), Quails (n=240; 50 d old) were divided into 8 groups, 30 birds per group. The quails were fed either a basal diet or the basal diet supplemented with 2, 4, or 8 mg of Diachrome/kg diet. Birds were kept at 22°C and 53% relative humidity (RH). At 14 wk of age, the thermoneutral (TN) group remained in the same temperature as at the beginning of experiment, whereas the heat stress (HS) group was kept in an environment-controlled room (34°C and 41% RH) for 3 wk. Heat exposure decreased performance when the basal diet was fed (p=0.001). Diachrome supplementation at 4 and 8 mg/kg diet, increased feed intake (p=0.05), egg production (p=0.05), feed efficiency (p=0.01), egg weight (p=0.05), and Haugh unit (p=0.01) in quails reared under heat stress conditions. Heat exposure increased concentrations of serum malondialdehyde (MDA) (p=0.001), glucose, and cholesterol (p=0.01), which were elevated by supplemental Diachrome (p≤0.05). Egg yolk Cr, Zn, and Fe (p=0.01) concentrations increased linearly, whereas MDA and cholesterol concentrations decreased (p=0.05) as dietary Diachrome supplementation increased in HS groups. Similar effects of supplementation on serum levels of glucose and cholesterol (p=0.05) and egg yolk concentrations of cholesterol (p=0.05) and Cr (p=0.01) were observed in TN groups. No significant differences in other values were observed in the TN groups. Results of the present study suggest that supplementation with Diachrome protects the quail by reducing the negative effects of heat stress.     

Two forms of egg defence in a chrysomelid beetle: egg clumping and excrement cover

Abstract.

• 1 Field experiments demonstrated that the faecal covering that female Microrhopala vittata (Fabr.) (Coleoptera: Chrysomelidae) place over their egg masses significantly reduces egg mortality.
• 2 The bottom egg in the egg mass, which lies against the leaf of the host plant, suffers significantly higher mortality than the other eggs in the egg mass.
• 3 The parasite Chrysonotomyia sp. (Hymenoptera: Eulophidae) generally parasitizes only the bottom egg in the egg mass, while predators rarely penetrate the faecal covering.
• 4 No significant relationship was found between mortality and the number of egg masses per leaf.
• 5 By placing eggs in masses, females may ensure that at least the upper eggs in any egg mass are effectively protected against virtually all natural enemies likely to pose a threat.