Egg-size variation and reproductive success in the Herring Gull Lams argentatus: adaptive or constrained size of the last egg?

Herring Gull Larus argentatus eggs from a study colony in the Baltic showed a slight but significant variation in egg size within the laying sequence. Last-laid eggs were only about 5% smaller by volume than first eggs. There was no significant difference in dry yolk weight or dry albumen weight, although possible differences were evident. The chicks had nearly equal hatching weights and equally long tarsi. There was no differential mortality in the third chick in the study colony, and there were no indications of egg-size-mediated mortality. The birds in the colony produced an average of 1.45 fledglings per pair. Compared with several other studied colonies, the difference in egg size within a clutch was low, and a comparison of colonies from northwestern Europe suggests that variance within the clutch is negatively correlated with fledging success, so that a large difference in size between first and last eggs is associated with low fledging success. We suggest that the size of the last egg in the clutch reflects the feeding potential in the environment and is mainly a nonadaptive response to poor feeding conditions during laying.     

Seasonal variation in reproductive measures of tropical Roseate Terns Sterna dougallih previously undescribed breeding patterns in a seabird

Seasonal variation in egg-laying, egg size, hatching success, hatchling mass, fledging success and chick growth of Roseate Terms Sterna dougallii breeding on Aride Island (Seychelles), Indian Ocean, were studied in 1997 and 1998. I investigated to what extent two patterns, common in a range of species, were followed by tropical Roseate Terns: (a) seasonal decrease in clutch size, egg size and breeding success and (b) an increase in breeding success with increasing egg weight. In 1997 (a poor year), the earliest nesting birds laid significantly smaller eggs, and chicks were lighter at hatching than those of peak nesting birds. The mean clutch size, of 1.04 eggs, showed no seasonal variation and no 'b'-eggs hatched. In 1998 (a good year) the earliest nesting birds laid eggs of similar size and their chicks were of similar weight to those of peak nesting birds. Mean clutch size, of 1.25 eggs, increased significantly through the season and about 60% of the 'b'-eggs hatched. In 1997, hatching success was 57% whereas in 1998 it was 80%. In both years, breeding success declined significantly through the season. The fact that the earliest breeding birds laid smaller eggs in a poor year and smaller clutches in a good year is in marked contrast to a range of other species, and to temperate-nesting Roseate Terns. Egg volume explained about half of the variance in hatchling mass in both years, but only 15% of the variation in linear growth rate. Hatching date was the only variable with a significant effect on fledging success. Roseate Terns on Aride seemed to sacrifice egg size and clutch size for earliness of laying. Presumably it is a strategy of older birds to lay as early as possible and may be regarded as a response of tropical Roseate Terns to breeding under relatively poor, and seasonally declining, food conditions.     

Fitness consequences of egg-size variation in the lesser snow goose

We investigated the relationship between eggsize variation and (a) egg hatching success, (b) chick survival to fledging and recruitment, and (c) adult female survival, over 12 years in the lesser snow goose (Anser caerulescens caerulescens). By comparing the means and variances of egg size for successful and unsuccessful eggs, our aim was to assess the relative fitness of eggs of different sizes and to determine the type of selection operating on egg size in this species. As both egg size and reproductive success vary with age in the lesser snow goose we controlled for the effects of female age. Egg-size variation is very marked in this population, varying by up to 52% for eggs hatching successfully. However, there was no relationship between egg size and post-hatching survival of goslings to fledging or recruitment, either within or between broods, pooling across years. Egg size varied significantly between successful and unsuccessful clutches in only 2 of 33 individual year comparisons. First-laid eggs surviving to onset of incubation, and eggs hatching successfully, were on average larger than unsuccessful eggs, but this was probably due to the confounding effects of female age-specific and sequence-specific egg survival. Variance of egg size differed significantly between successful and unsuccessful eggs in only 3 of 24, and 0 of 21, individual year comparisons for pre- and post-hatching survival respectively. We therefore found little evidence for a relationship between egg-size variation and offspring fitness, or for strong directional, normalising or diversifying selection operating on egg size, in the lesser snow goose. In addition, there was only weak support for the hypothesis that egg-size variation is maintained by temporal variation in selection pressure (sensu Ankney and Bisset 1973). It is likely that egg-size variation represents the pleiotropic expression of alleles affecting more general physiological or metabolic processes. While this does not rule out the existence of alleles with more direct effects on egg size we suggest that their contribution to heritable egg size is small.     

Breeding biology of the Barn Owl Tyto alba in central Mali

Data were obtained on 178 clutches of African Barn Owls in central Mali from four breeding seasons during 1979–1983. Significantly more clutches were laid in 1979–1980 and significantly fewer in 1980– 1981 than the average for the 4 years and there were significantly more clutches laid in the middle period of the annual breeding season. The egg volume was significantly smaller at the beginning of the breeding season and significantly larger in the middle than the overall mean with eggs of second clutches being larger than those of first clutches. The clutch size was 605 eggs of which 479 hatched. The number of young fledged per successful nest was 319 and was 1 83 for all nesting attempts. The month was the only variable shown to affect significantly the clutch size, eggs hatched and fledging rate, the highest success rates being associated with the middle of the breeding period. The average interval between the hatching of eggs was 2–31 days. Survival rates (47'1%) to fledging were significantly affected by year (1981–1982 being the least) and month (mid-season birds the best). The order of hatching significantly affected age at death or disappearance, the first-hatched birds surviving the longest. The year significantly affected age at fledging, the young from the year in which most clutches were laid leaving the nest at the youngest age and those associated with the year having the least number of clutches remaining in the nest the longest. The month of hatching also affected fledging age, birds at the extremes of the breeding season fledging at older ages. The discussion compares these data with those from elsewhere.     

Egg size and clutch size in the reproductive investment of American Kestrels

We studied causes and consequences of egg-size variation among clutches of American kestrels ( Falco sparverius ). Egg. from 275 clutches were measured 1990 to 1992. To test the hypothesis that the size of eggs was contrained by food availability in the pre-laying period, we censused small mammal populations in the three years and performed a food supplementation experiment in 1990 and 1991. Kestrels did not advance the date they laid their first egg but did lay significantly larger eggs in response to extra food. The size of eggs was correlated with small mammal abundance on the territoty, and females in good body condition tended to lay large eggs. Body size did not affect egg size, and there were no relationship between agg size and laying date except in 1900, the poorest food year. Clutches with a large mean egg volume had better hatching success than clutches containing small eggs. We argue that there is a phonetypic component to egg size in kestrels, and that kestrels use egg size to fine-tune reproductive investiment to available resources.     

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.     

Population quality,dispersal and numerical change in the gypsy moth,Lymantria dispar (L.)

Summary First instars from small and large gypsy moth eggs differ significantly in their head capsule width, weight, hatching time and the length of thoracic setae. Pupal weight and the developmental period of immature stages of the gypsy moth originating from small or large eggs do not differ significantly. The mean number of eggs per mass produced by females originating from small eggs is greater than that of females from large eggs although not statistically significant. Highly significant differences in mean egg size of egg masses of each type of female were also observed. The relationship between egg size and dispersal strategies are discussed.Paper No. 2229 Massachusetts Agricultural Experiment Station. University of Massachusetts at Amherst. This research supported (in part) from Experiment Station Project No. 355     

Feeding behaviour of cod (Gadus morhua) in relation to spawning

The food consumption and egg production of 26 adult (13 female and 13 male) Atlantic cod ( Gadus morhua ) were monitored during prespawning, spawning and postspawning periods. Females spawned from late January to mid-April. Feeding activity occurred from December to early January and ceased for females, on average, 36 days (15–54 days) before the onset of spawning. The duration of spawning by females was, on average, 42 days (10–61 days) and feeding was suppressed by both sexes during the first three-quarters of each female's spawning period. Mature females went, on average, 70 days or 19% of the year without eating. An abrupt increase in feeding activity, particularly by females, occurred during the last quarter of spawning or shortly after the release of the last egg batch (on average, feeding started again after 91% of a female's eggs had been released or 82% of egg batches). Females consumed greater quantities of food than males during both winter and postspawning feeding periods. During spawning, females lost, on average, 29% of their body weight and males 14%. Fecundity ranged from 0.75 to 3.97 million eggs per female. The volume of eggs produced by four individual females (range = 1285–5995 ml in four to 11 batches) ranged from 99 to 195% (mean 150%) of a female's postspawning body volume. Six immature cod fed throughout the experimental period and gained, on average, 8% of initial body weight. Laboratory results were supported by stomach fullness index values of Georges Bank cod exhibiting different maturity states.     

High variability in egg size and energetic content among intertidal mussels

Maternal investment is a fundamentally important parameter in life-history theory and models, yet the scales at which it varies (among individuals vs. among populations) is rarely reported. In this study, variability in attributes of eggs and early larvae of Mytilus californianus was examined from four sites spanning Point Conception, California, in June and September 2001. The effects of female, site, and month were examined for the following variables: egg volume (microl), egg energy content (microg carbon per egg), and initial larval size (microm). The only significant effect on both egg traits was that of female. Females differed by up to 57% in mean egg volume and 116% in mean egg energetic content. Although there were significant effects of rearing environment, female, site, and month on initial larval size, variability in larval length was small compared to the egg traits. Mean larval length was maximally 11% different among females. Neither female body weight nor length was correlated to mean offspring traits, and there were also no significant relationships between egg traits and initial larval size. The primary source of variation in maternal investment in this system appears to be among individual females rather than over space or time.     

Reproductive ecology of the lacertid lizard Podarcis bocagei

This paper presents data on aspects of the reproductive ecology of a population of Podarcis bocagei in northwestern Spain, as monitored over a two-year period (1990–1991) Data were obtained principally on the basis of mark-recapture experiments, but also from laboratory hatching studies Mating took place between the end of March and July During the laying period, from May to July, 8 5% of reproductive females produced three clutches, 52 1% two clutches, and 39 4% one clutch In general, single clutches were produced by small females Only a small proportion of large females produced three clutches Mean clutch size was 4 8 eggs (range 4–7) in May, 4 3 (2–6) in June and 3 9 (2–4) in July There was sigmficant variation in the mean snout-to-vent length (SVL) of females laying in each month of the season Both clutch size and mean single-egg volume increased with mother's SVL There was a significant partial correlation between egg volume and clutch size when both mother's SVL and month of laying were held constant There was no significant between-year variation in clutch size, breeding females' SVL, egg weight or relative clutch mass A delay in the timing of reproductive events in one year (1991) is attributable to adverse weather conditions during early spring Hatching occurred between July and September Hatch success (as estimated in 1989, 1990 and 1991 from natural nests at the study site) was high, ranging from 83% in 1991 to 91% in 1989 The mean SVL of female hatchlings was greater than that of male hatchlings By contrast, adult females had lower mean SVL than adult males     

Experimental manipulation of female reproduction reveals an intraspecific egg size-clutch size trade-off

A negative relationship, or trade-off, between egg size and clutch size is a central and long-standing component of life-history theory, yet there is little empirical evidence for such a trade-off, especially at the intraspecific level. Here, I show that female zebra finches (Taeniopygia guttata) treated chronically during egg formation with the anti-oestrogen tamoxifen lay smaller eggs (by 8%) but produce larger clutches (on average two eggs more) than controls. Decreased egg mass in tamoxifen-treated females was associated with a 50% decrease in plasma levels of the two yolk precursors, vitellogenin and very-low-density lipoprotein. Although tamoxifen-treated females laid more, smaller eggs (and had a higher total expenditure in their clutch), they did not differ from controls in the number of chicks fledged, the mass or size of these chicks at fledging, or the chicks' egg-production performance at three months of age. However, tamoxifen-treated females had lower relative hatching success: they laid more eggs but hatched the same number of chicks. Among individual tamoxifen-treated females, birds that laid the smallest eggs early in their laying sequence laid the largest number of additional eggs, that is, there was a negative correlation, or trade-off, between egg size and clutch size.     

Heritability, plasticity and canalization of Ural owl egg size in a cyclic environment

Avian egg size is highly variable on the population level, but is considered inflexible on the individual level. On the basis of 2969 measurements of individual eggs collected during 1981-2005, we analysed heritability, plasticity and selection on egg size in the Ural owl, a long-lived bird that preys on voles. Vole abundance varied in a 3-year cycle, creating varying food supply across the cycle's phases. Ural owl egg size is heritable (h(2) = 60%). Ural owls lay larger eggs in improved food conditions. On the basis of repeated breeding records of 59 females that bred in all vole cycle phases, we show that intra-individual adjustment (plasticity) explained 22.4% of the variation in egg size across phases. Egg size was under stabilizing selection. Extremely small and extremely large eggs had reduced hatchability, and individuals who laid either large or small eggs had lower lifetime fledgling production than the ones laying intermediately sized eggs. Our findings illustrate how maternal investment in egg size can both be heritable and highly responsive to variable environmental conditions, and suggest that variation in the investment in egg size across individuals is canalized.     

The importance of energetic versus pelvic constraints on reproductive allocation by the eastern fence lizard (Sceloporus undulatus)

In ectothermic species, females often produce larger eggs in colder environments. Models based on energetic constraints suggest that this pattern is an adaptation to compensate for the slower growth of offspring in the cold. Yet, females in cold environments also tend to be larger than females in warm environments. Consequently, thermal clines in egg size could be caused by pelvic constraints, which stem from the inability of large eggs to pass through a small pelvic aperture. Models based on energetic constraints and models based on pelvic constraints predict similar relationships between maternal size and egg size. However, pelvic constraints should produce these relationships both within and among populations, whereas energetic constraints would not necessarily do so. If pelvic constraints are important, we might also expect small females to compensate by producing eggs that are relatively rich in lipids (i.e. high energy density). The present study aimed to assess whether energetic or pelvic constraints generate geographical variation in egg size of the lizard Sceloporus undulatus . Pelvic width is very highly correlated with body length in S. undulatus , making maternal size a suitable measure of pelvic constraint. Although maternal size and egg mass (dry and wet) covaried among populations, these variables were generally not related within populations. Energetic density of eggs tended to increase with decreasing egg mass (dry and wet), but this relationship was strongest in populations where no relationship between maternal size and egg mass was observed. Our results do not support the pelvic constraint model and thus indicate energetic constraints play a greater role in generating geographical variation in egg size.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 513–521.     

Patterns of life history among cyclopoid copepods of central Europe

• 1 Life history characters (body size of adults, egg diameter, egg sac length and breadth) of nineteen species of central European cyclopoid copepods were measured and sexual size dimorphism (adult female length x adult male length^?1), relative egg size (egg weight X body weight^?1), weight of adult females and of eggs, egg sac shape (egg sac length x egg sac breadth^?1), and reproductive effort (clutch weight produced per female weight per day) were calculated to detect trends in life history strategies.
• 2 Typical planktonic species exhibited the lowest reproductive effort. Among planktonic species, the value for egg sac shape increased with clutch size.
• 3 Large species and small species exhibited different trends in life history characters. Large species had larger clutches, larger eggs, and a greater sex size dimorphism than small species. However, small species had a greater relative egg size.
• 4 Large species live in cold water and reproduce during the spring bloom of phytoplankton where the production of large clutches with relatively small eggs is advantageous. Reserves are unnecessary for juveniles because food is abundant. Small species generally are most abundant during the warm season, when conditions are less predictable, and relatively large eggs, possibly provided with reserves, are advantageous.

     

Predicting fluctuations in the size of newly emerged sea-trout fry in a Lake District stream

The objective was to predict interannual fluctuations in the size of sea-trout fry when they emerged from the redd, using models developed from field data for 70 excavated redds (≥three per year), and from experimental data on egg and alevin development at 30 constant temperatures in the laboratory (range 1·5—10·5) C with 100 naturally fertilized eggs at each temperature). Egg weight increased with female length and also with the number of eggs laid in a redd, both relationships being well described by a power function. Early spawners were the largest females laying the largest and most numerous eggs, whilst late spawners were the smallest females laying the smallest and least numerous eggs, with middle spawners being intermediate between these two extremes. Mean values for egg weight and number of eggs per redd were obtained for these three groups. The numbers of early, middle and late spawners for each year of a 30-year study and the mean values from the excavated redds were used to estimate weighted means for the number of eggs per unit area and egg weight. Mean values varied considerably between years (30-year ranges: 518–7964 eggs per 60 m2; 112–138 mg wet weight). In the laboratory, mean weights of newly hatched alevins and newly emerged fry were both related positively to mean egg weights. Alevin and fry mean weights were independent of the number of days required for 50% of the eggs to hatch or fry to emerge. Models described in a previous paper formed the basis of those used to predict fry weights over the emergence period. Model predictions were validated by field data for the whole emergence period in 8 years (1967–1971, 1974, 1975, 1980), and by pre-fry weights on single dates in 21 years (1967–1987). As pre-fry densities on these single dates were very similar to egg densities for the same year class, mortality in the egg and alevin stages was very low. The chief objective was therefore fulfilled, and the extent of interannual fluctuations for the 30-year study showed some variation in mean fry weight (30-year ranges: 153–193 mg for both the whole emergence period and the date on which 50% of fry emerged) but a progressive decrease in fry weight through the emergence period. Possible reasons for this variation are discussed, and it is concluded that the size of the female spawners is the dominant factor.     

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.     

BREEDING OF THE GREY WARBLER GERYGONE IGATA AT KAIKOURA, NEW ZEALAND

I studied the breeding of Grey Warblers Gerygone igata (Muscicapidae: Acanthizinae) in forest near Kaikoura, New Zealand, between 1976 and 1979. Only males sang and singing occurred all year. From late July to January pairs defended self-contained territories of 0·25–1·73 ha but they occupied larger home ranges when not breeding. Territorial adults were strictly sedentary all year. The average annual mortality of breeding adults was 18·5% and the predicted life-expectancy 4·9 years, which is remarkable in a bird weighing 6–7 g. The breeding season from first building to last fledging was six months long and it began early. Exceptionally, Grey Warblers may build and lay before the shortest day. As the season progressed warblers nested lower on average, both in absolute terms and relative to the tree nested in and canopy at the site. Warblers built in 7–27 days then delayed up to eight days before laying. Only females built and at no stage of breeding did males feed their mates. Both sexes fed the young. Grey Warblers laid for 15–16 weeks of the year and first clutches were laid asynchronously during 5–6 weeks. Eggs of a clutch appeared at two-day intervals and each egg weighed 1·5 g when fresh (23% of mean adult weight). Clutch size was nearly constant (mean 3·9, mode 4, range 3–5). The incubation period was 17–21 days (mean 19·5 days) and the nestling period 15–19 days (mean 17·2 days). On average the clutch hatched over 1·4 days, even though incubation commenced with the laying of the last egg. Nestlings reached maximum weight on Days 13–14 on average and then receded in weight by 4%, apparently through loss of water. All healthy nestlings exceeded mean adult weight during development by up to 39%. Nestlings from broods of two were at first lighter on average than those from larger broods, but in the second half of the nestling period twins were significantly the heaviest. Grey Warblers were fed for 28–35 days after fledging and they survived well while dependent on parents. Fledglings dispersed up to 3 km or more at independence and only 5% per annum joined the breeding population. Of nests that received eggs, 42% produced at least one fledgling. On average each breeding adult raised 2·0 fledglings per season. Of 265 eggs in 73 nests 70% hatched and 38% produced fledglings. Of 185 nestlings 54% fledged. Probably the main cause of mortality of eggs and nestlings was predation by introduced rodents and mustelids. Grey Warblers raise two small broods slowly during a long breeding season, rather than investing in one large quickly-reared brood. In New Zealand's mild climate the warbler's food supply may not decline severely in winter, and the population of warblers may remain so close to the limit set by food that extra for breeding is hard to obtain. Thus the breeding strategy may be adapted to a restricted food supply.     

THE BREEDING BIOLOGY OF THE GANNET SULA BASSANA ON THE BASS ROCK, SCOTLAND

The Bass colony is increasing—in 1962 there were 5,350–5,700 breeding pairs; 1,340–1,430 pairs of non-breeders with nests or sites (mainly pairs in their season before first breeding) and 2,000–2,500 “club” birds without nest or site. 15% of nests were occupied by both birds of a pair at the time of the count. Oldest males return to the colony in January, followed by experienced females, considerably later by young adult-plumaged birds, immature birds later still, and the few one year-olds that return usually not until May and June. Mid-cliff sites are the first to be re-colonized each year. Gannets usually breed in their fifth year and there is some evidence that females breed earlier than males. The characteristics of Gannet nests and sites are described. Nests function in raising the egg and young above the morass of the breeding colony, and reach a density of about 2.3 per square metre. Nests are demolished and their positions changed more often than might be suspected. The extremely strong social tendency which causes Gannets to establish their sites amongst or very close to existing breeders probably is the factor ensuring high density and this contributes to synchronization of laying. Egg laying is analysed. Experienced pairs forming an isolated group of 20 nests began laying later and showed less synchronization than two other groups of the same size but from the middle of a dense mass, probably due to the greater social stimulation experienced by the non-isolated groups. The date for first and median eggs was also earlier in larger than smaller groups in the same year. The effect of density as distinct from group size is also discussed. Early eggs are mainly laid on cliff or cliff-edge sites and in large nests. Different groups within the colony produced the median egg within 2–3 days of the end of April each year. In the fullest documented group the mean date was also constant from year to year and closely approached the median, implying a considerable degree of synchronization within the gannetry as a whole. Laying in the observation colony became progressively earlier in successive years, probably due to recovery from previous disturbance. Nevertheless, individual females tended to lay in a fixed position each year with relation to the mean for the group. Increasing age of the female causes earlier egg laying and heavier eggs for up to at least five years. It is suggested that the survival value of seasonally synchronized laying in the Gannet is maximum utilization of a seasonally dependable and abundant food supply for the production of young with the optimal chance of post-fledging survival. The spread of laying acts as an insurance against possible adverse conditions. There is a considerable reserve of unutilized breeding capability within the colony (adult non-breeders, a pre-maturity period longer than physiologically necessary for egg production, and a one-egg clutch when in fact two young can easily be reared). The mean of 393 Gannet eggs was 104.5 gm. (range 81–130). Eggs constitute about 3.4% of adult female weight and lose 9–13% in weight during incubation. Replacement laying of the invariable one-egg clutch takes 6–32 days. The mean incubation period was 43.6 days. Male incubation spells averaged 35.6 hours; female's 32.0 hours. Copulation ceases immediately after egg laying. During three seasons, 82% of eggs laid in the observation colony hatched. Inexperienced pairs hatched 62.6% of eggs laid; experienced pairs hatched 86%. Some of the processes of incubation and chick rearing depend on the maturation of innate abilities and not on experience Inexperienced breeders do not seem inferior to experienced ones in finding enough food for their young. Parental care of the new chick is described; the pterylosis of the chick is figured. A summarized account of plumage development is given. Food fish and chick feeding are described. The average frequency of feeds throughout a continuous two-day watch was 2.7 feeding bouts per chick per day. Adult fishing trips usually took 7–13 hours and the estimated fishing range is over 100 miles, and possibly up to 400 miles, from the breeding colony. Despite this, 15% of daylight hours are spent by the pair together at the nest in addition to the constant guarding of the chick by one or other. Gannet young have a very compressed growth period compared with boobies and fledge at 3,100–4,100 gm., after a steady growth uninterrupted by periods of starvation or arrested development after an average 90 days. 92.3% of all eggs which hatched in 500 nests in the observation colony during three years of the study gave fledged young. Excluding inexperienced birds, there was no difference in the fledging success of eggs laid at different dates in the breeding season, in accordance with the proved abund- ance of food. However, post-fledging survival is probably higher among young fledging at the peak period (first half of September) than later and the few relevant ringing returns tend to support this. Breeding success at the small colony at Bempton was less compared with groups from the Bass for the years 1961–3. Causes of chick loss before and during fledging are discussed. They are unimportant compared with the great mortality in the first year of life after fledging. The adaptive significance of black plumage in the juvenile Gannet probably lies in reduced attack-releasing qualities of such plumage on the male parent. The Gannet alone in the Sulidae produces young which leave the nest with large fat deposits, and which are not fed at all by the parents after fledging. This is possibly another result of adaptation to using a seasonally abundant food supply to the maximum. The present Gannet population increase is discussed in relation to cessation of human predation, the possible impetus given by a temporary but large increase in pelagic fish during the war, but also the overall steady downward trend in fishing returns since the early part of this century. One cannot explain the steady and considerable rise in Gannet numbers only in terms of increased food supply. The fact that, at a time of population expansion and obviously favourable conditions, Gannets are still far from utilizing their full recruiting powers, needs investigating further. It may be partly due to the relative slowness of evolutionary change in a long-lived species with slow population turn-over, if the Gannet has evolved its characteristics in response to an environment different from the present one.     

Annual variation in the seasonal shift in egg size and clutch size in Sceloporus woodi

Summary Early and late season clutch parameters were examined over a three year period in the Florida scrub lizard, Sceloporus woodi. Precipitation levels were monitored throughout the study. In the early and late season of 1984 and the early season of 1986 precipitation levels approximated long-term mean levels of precipitation. In 1985 a severe winter drought occurred. Clutch size was positively related to body size in all samples in every year. In 1984 and 1986, egg size was not related to clutch size, whereas, in 1985 egg size was negatively related to clutch size. In 1985, females produced large clutches of small eggs early in the season and small clutches of large eggs late in the season. In 1984, no seasonal changes in egg or clutch size occurred. In the late season of 1986, females produced the largest clutches and the smallest eggs of all the samples, but egg and clutch size were not statistically different from the early season egg and clutch size of 1986. Total clutch dry weight, an estimate of total clutch energy, was not different in any of the six sampling periods. These data do not support current adaptationist models that attempt to explain the control of clutch and egg size in lizards. It is argued in this paper that egg and clutch size may vary in response to past environments that affect a female's physical condition, as well as, current resources that may be important for maintenance and reproduction. Egg and clutch size appear to be plastic traits selected to respond to proximal environmental variation, whereas, the investment of total dry matter/clutch has been optimized.     

Maternal and interannual comparison of the ovulatory periodicity, egg production and egg quality of the batch-spawning yellowtail flounder

Estimates of the ovulatory periodicity of yellowtail flounder indicate that a 1-day interval, which predominated over all other intervals, may characterize regular ovulation patterns. Females produced a mean number of 14–22 batches in 1994 and 1995, respectively. Batch fecundities usually remained within a range of 10 000–60 000 eggs. Mean egg production increased from 549 756 eggs per female in 1994 to 1 186 881 eggs in 1995. Mean fertilization rates rose interannually from 38 to 57%, while hatching rates, tested in 1994, had a mean of 63%. Maternal variation in egg production and egg quality was large and independent of size differences among females. Some females had disrupted ovulation patterns which affected the realization of potential fecundity contained within the prespawning ovary. High interbatch variation in egg quality was not related to progressive decreases in egg diameter and dry weight over time. Batches with high survival rates appeared at random within a female's duration of ovulation.