FACTORS AFFECTING THE EGG SIZE OF RED-BILLED GULLS LARUS NOVAEHOLLANDIAE SCOPULINUS

The factors influencing the egg size of the Red-billed Gull Larus novaehollandiae scopulinus were studied at Kaikoura, New Zealand, between 1964 and 1972. In two- and three-egg clutches there was a trend for the eggs to become smaller in the sequence of laying. Length, breadth and volume of eggs of one-, two- and three-egg clutches declined significantly as the season progressed. The size of eggs from single-egg clutches tended to be smaller than eggs from two-egg clutches laid at the same time. There were correlations between the proportions of one-egg and of three-egg clutches being laid at a given period and the mean egg volume of two-egg clutches. When the mean egg volume of two-egg clutches increased there was a corresponding increase in the proportion of two- and three-egg clutches laid. When the mean egg volume of two-egg clutches decreased there was an increase in the proportion of single-egg clutches laid. The egg size of the Red-billed Gull showed no direct correlation with the abundance or availability of food; the largest eggs were produced early in the season when food was in short supply. In spite of an increase in the food supply in the middle of the breeding season, birds laying at this time produced smaller eggs than birds which laid earlier in the season. However, early breeders which relayed at the peak in food abundance on average produced a larger replacement clutch than originals laid early in the season. It is suggested that the birds nesting early in the season are able to produce the largest eggs because they are the most efficient foragers for food, and those which nest later in the season produce smaller eggs, even at peak food abundance, because of their inefficiency or inexperience. Early breeders laying replacement clutches tended to lay larger eggs and larger clutches than birds which are producing their first clutches at the same time. Two-year-old females laid eggs which were significantly shorter than older aged birds while the breadth and volume of the egg increased with the age of the female up to the fifth year. There was a trend for females to lay larger eggs when mated with older rather than younger males. No statistical differences in egg size were detected between females changing or retaining the partner of the previous season. Female body weight and egg volume were positively correlated in females weighing less than 275 g but not for heavier females. It is suggested that the seasonal decline in egg size and clutch size results from a decrease in the availability of food and the ability of the individual to exploit the resource.     

ADAPTATIONS TO CLIFF-NESTING IN SOME ARCTIC GULLS (LARUS)

Various aspects of the breeding of three gulls, Larus thayeri, glaucoides and hyperboreus, which nested in colonies both on cliff ledges and on level ground in the eastern Canadian Arctic, were compared with those of the ground-nesting L. argentatus and the cliff-nesting Rissa tridactyla. As the result of adaptation to cliff ledge nesting, many aspects of the breeding biology of R. tridactyla were strikingly different from those of the ground-nesting European L. argentatus, but the behaviour of L. thayeri, glaucoides and hyperboreus clearly spanned these differences. Cliff-nesting individuals of thayeri and glaucoides were most like Rissa; ground-nesting individuals of these species were most like argentatus. L. thayeri was more like Rissa than was glaucoides. With but few exceptions, both cliff- and ground-nesting individuals of hyperboreus were most like argentatus. The factors responsible for the intra-specific differences between cliff- and ground-nesters of thayeri and glaucoides are not clear. Limited gene exchange between cliff and ground colonies occurs. Because of physical features of the nest, first-laid eggs were more liable to fall from ledges than second or third eggs. L. thayeri and glaucoides have evolved separate mechanisms to cope with this problem. Egg shape was multimodal in thayeri and glaucoides. Long pyriform eggs were less liable to fall from ledges than eggs of other shapes. L. thayeri laid more long pyriform eggs as first eggs than did glaucoides. L. thayeri lost fewer eggs than did glaucoides, but glaucoides replaced all lost eggs while thayeri did not. Delayed follicular atresia provided glaucoides with insurance of egg replacement. In thayeri, accessory follicles were reabsorbed after the first egg was laid; in argentatus, after the second egg, and in glaucoides after the third egg. At the approach of a predator, it was advantageous for cliff-dwelling chicks to remain motionless but for ground-dwelling chicks to flee their nests and to hide. Among the cliff-nesters, the “freezing in place” reaction of chicks was best developed in thayeri, to a lesser extent in glaucoides, and least in hyperboreus. Among the ground-nesters, chicks of glaucoides and hyperboreus behaved like those of argentatus and fled their nests when disturbed, but chicks of thayeri froze like their cliff-dwelling siblings. Reciprocal transfers of eggs and chicks between cliff and ground colonies indicated that in argentatus, glaucoides and hyperboreus, the factors determining a chick's reaction to disturbance came into play between hatching and the eighth day. In thayeri, the reaction appeared to be effectively innate. Chicks of glaucoides showed a greater predisposition to this behaviour than chicks of argentatus after both had received identical experience on cliff ledges. In thayeri, stereotypy of the freezing reaction has probably been a factor limiting the colonisation of areas where cliffs are scarce but predators present. In argentatus, lack of perfection of this behaviour (compared to glaucoides and thayeri) has probably been a factor preventing argentatus from attaining cliff ledges. L. hyperboreus, although nowhere abundant, is a widespread species nesting on level ground and on cliff ledges but lacking the modifications observed in glaucoides and thayeri; this is due to its size and aggressiveness, the fact that it picks nest-sites before glaucoides and thayeri arrive in the colonies, and that on cliffs it chooses the largest and most level ledges.     

DISEASE IN A COLONY OF BLACK-HEADED GULLS LARUS RIDIBUNDUS

An account is given of the disease factors encountered in a colony of Black-headed Gulls in the breeding season 1956.     

CLUTCH SIZE AND EGG SIZE IN FREE-LIVING AND PARASITIC COPEPODS: A COMPARATIVE ANALYSIS

The evolution of reproductive strategies and the trade-off between number and size of eggs were investigated in a comparative analysis of free-living and parasitic copepods. Data from 1038 copepod species were used to obtain family averages for 105 families; the phylogenetic relationships among these families include 94 branching events or 94 independent contrasts on which the analysis was based. Transition from a free-living existence to parasitism on invertebrates resulted in small increases in body size. Transition from parasitism on invertebrates to parasitism on fish was associated with greater increases in body size. After controlling for body size, a switch to fish hosts resulted in an increase in the number of eggs produced and a reduction in egg size. Among all contrasts, there was a negative relationship between changes in relative clutch size and changes in relative egg size, suggesting the existence of a trade-off between egg size and numbers. However, opposite changes in these measures of clutch size and egg size were not quite more frequent than expected by chance, therefore indicating that investments into egg numbers are not necessarily made at the expense of egg size, and vice versa. Latitude affected copepod body size, clutch size, and egg size, whereas the effects of freshwater colonization or size of the fish host were not significant. Comparative analyses at either the genus or species levels within given taxa of copepods parasitic on fish provided limited support for a trade-off between clutch size and egg size, but were hampered by the small number of independent phylogenetic contrasts available. From the family-level comparative analysis, it appears that the evolutionary transition from a free life to parasitism on invertebrates, and the transition from parasitism on invertebrates to parasitism on fish, have led to changes in life-history traits in response to the different selective pressures associated with the different modes of life.     

FACTORS DETERMINING VARIOUS FORMS IN CLADOSIPHON ZOSTERAE (PHAEOPHYCEAE)

The brown alga Cladosiphon zosterae (J. Ag.) Kylin (Chordariaceae) is found to exist in culture in a variety of forms, including filaments, discoid or crustose forms, and two types of erect cylindrical thalli. Asexual spores from plurilocular sporangia of each form can produce any and all forms. Three factors controlling morphological expression in C. zosterae have been identified. These factors include: (1) heteroblasty (formation of filamentous and discoid germlings as a consequence of alternate modes of spore germination), which was independent of all environmental factors tested including temperature, photoperiod, nitrogen source, and general fertility of the growth medium; (2) influence of certain nitrogen sources, notably ammonium, which induce a compact discoid or crustose form; and (3) a morphogenetic factor presumed to be bacterial which determines mode of development and subsequent morphology of the erect Cladosiphon thallus.     

HERITABILITY AND SELECTION ON CLUTCH SIZE IN DARWIN'S MEDIUM GROUND FINCHES (GEOSPIZA FORTIS)

I studied the causes of variation and selection on clutch size in a population of Darwin's Medium Ground Finches (Geospiza fortis) on Isla Daphne Major, using data collected over a nine-year period (1976–1984). Quantitative-genetic analyses were carried out using the first clutch laid by a female in a given year. I used both unadjusted clutch-size values and values adjusted for between-year differences in mean clutch size for repeatability and regression analyses. Repeatability of clutch size was small (≤8%) and nonsignificant in all cases. Sib-sib analyses and parent-offspring regressions gave no evidence of a significant additive genetic component to clutch-size variation. Slopes of mother-daughter regressions were actually negative, suggesting possible maternal effects of mother's clutch size on daughter's clutch size. There was a small positive relationship between female age and clutch size but no effect of male or female body size or of large-scale differences in habitat quality on clutch size. Selection on clutch size was generally directional and positive: in almost all years in which successful breeding occurred, large clutches tended to fledge more chicks and produce more young surviving to the following year, possibly because there was no trade-off between clutch size and the weights of individual chicks at fledging. Thus, sustained directional selection for large clutch size may have reduced additive genetic variation in clutch size to low levels in this population. The size of a female's clutch may be primarily determined by unidentified proximate environmental factors which vary from year to year, rather than by any long-term optimization of clutch size with respect to adult survival.     

THE EFFECTS OF DIURNAL AND TIDAL PERIODICITIES IN THE NUMBERS AND ACTIVITIES OF HERRING GULLS LARUS ARGENTATUS IN A COLONY

The average numbers of Herring Gulls Larus argentatus present in a breeding colony on Walney Island, Cumbria, were found to vary with the tidal cycle but to remain effectively constant with time of day through the breeding season. An activity survey, based on 50 Herring Gulls observed at half-hourly intervals during March and April 1973, showed that sleep and rest varied inversely with each other with sleep increasing to 50 per cent at midday. After a peak in the proportion of gulls asleep four hours before low tide, sleeping progressively decreased until low tide; seemingly a result of resident gulls waking and remaining more alert as others left the colony in search of food. Preening was constant throughout the day and tide cycle. Other behaviours (mostly courtship and agonistic behaviour associated with territory defence) increased slightly during low tide and were more common early and late in the day. Night observations of the gulls' activities showed that there was a peak of sleeping between midnight and 02.00 hours. It is suggested that Herring Gulls have a bimodal diel sleep pattern.     

FOOD SIZE AND AGGRESSIVE INTERACTIONS BETWEEN TWO SPECIES OF GULLS: AN EXPERIMENTAL APPROACH TO RESOURCE PARTITIONING

Duffy, D. C., Heseltine, S. & La Cock, G. D. 1987. Food size and aggressive interactions between two species of gulls: an experimental approach to resource partitioning. Ostrich 58: 164–167.

In feeding experiments with different sizes offish, Hartlaub's Gulls Larus hartlaubi reached prey first, but lost larger items to the larger, dominant Kelp Gull L. dominicanus. Hartlaub's Gulls took longer than Kelp Gulls to consume prey of the same size.     

ECOLOGICAL FACTORS IN WOOD EVOLUTION: A FLORISTIC APPROACH

Wood florulas from southwestern Australia were analyzed to determine whether wood anatomy is sufficiently correlated with ecology so that vessel element features can be said to have a predictive value. Indices for vulnerability (vessel diam: vessels per sq. mm) and mesomorphy (vulnerability × vessel element length) were calculated for each species in the following florulas: karri forest understory, coastal granitic slopes, bogs, sand heaths, and desert. Wood indices for the species studied and for each florula show that these florulas form a sequence in increasing xeromorphy in the order listed. Genera represented in more than one florula validate the trends. Data for Gyrostemonaceae, Loranthaceae, and Cupressaceae are calculated separately because these are succulents, epiparasites, and conifers, respectively. Comparison with categories from floras elsewhere in the world shows the flora of Western Australia as a whole to be relatively xeromorphic. The indices devised show promise of great reliability because correlations with rainfall, temperature, and other factors are very close. Functional nature of the vessel element is thereby believed to be clarified.     

BREEDING SUCCESS AND POPULATION GROWTH IN A COLONY OF HERRING AND LESSER BLACK-BACKED GULLS LARUS ARGENTATUS AND L. FUSCUS

This paper describes the results of investigations into the factors affecting breeding success of the Herring and Lesser Black-backed Gulls Larus argentatus and L. fuscus, in the large colony on Walney Island, northwest Lancashire, between 1962 and 1965. These investigations were concerned with the incubation period, and the first ten days after hatching. The survival of chicks to ten days is 67% in Herring Gulls, and 56% in Lesser Black-backs. Most of these losses occur in the period just after hatching and are due to “cannibalism” by other gulls. This form of predation does not appear to be masking any effects from starvation or disease. The following factors contribute to egg or chick mortality:breeding too late (and, to a much smaller extent, too early); breeding in the open, as opposed to amongst cover; the facts that eggs in small clutches have a lower hatching success than those in large ones and that Herring Gull (but not Lesser Black-back) chicks in small broods are less likely to survive to ten days than are those in large broods. Chick mortality after the first ten days is not certainly known. About 30% of the eggs laid gave rise to fledged young— or about one fledged chick per pair. In the Herring Gull, the average clutch size (2.56) is lower than that of the Lesser Black-back (2.76). Both species show a seasonal decline in clutch size—this occurs earlier in the Herring Gulls than in the Lesser Black-backs. The Walney population, which stood at about 700 pairs in 1950, had reached 12,000 in 1957, and is at present about 18–19,000 pairs. It is suggested that this increase may be linked to the greater availability, or exploitation, of human garbage in the Morecambe Bay area. The population explosion between 1950 and 1957 must have been partly due to massive immigration and could not have come about through natural increase alone. The possible influences of the gulls' behaviour on the population growth are discussed. There is no evidence of any “shock disease”, although the Walney colony is very crowded. “Cannibalism” is regarded, not as evidence of a failing food supply, but as an extension of the normal hunting behaviour of these omnivorous gulls; it will be an economical means of obtaining food only in a large, dense colony, such as Walney. It may be offset by increased breeding efficiency due to social factors.     

SEX DIFFERENCES IN BODY SIZE AND BODY CONDITION OF CALIFORNIA SEA LION (ZALOPHUS CALIFORNIANUS) PUPS FROM THE GULF OF CALIFORNIA

The Gulf of California harbors about 15% of the total California sea lion ( Zalophus californianus californianus ) population. We studied pup morphometrics from eight Gulf of California rookeries during the 1996 and 1997 reproductive seasons to describe sex differences in body size and body condition indices. Newborn pup body size was not different from previous reports. Male pups were heavier and larger than female pups in terms of all linear dimensions. Morphometric relationships, however, showed that males were 3%-4% denser, and that after removing the effects of length, they were about 2% heavier than females. Sculp depth adjusted for length was 12% larger in female than in male pups. Our data provide further evidence that male otariid pups may allocate a larger fraction of milk energy to muscular and skeletal growth compared to female pups.     

INCREASED HARASSMENT OF RIGHT WHALES (EUBALAENA AUSTRALIS) BY KELP GULLS (LARUS DOMINICANUS) AT PENÍNSULA VALDÉS, ARGENTINA

Kelp gulls at Península Valdés, Argentina, have recently developed the habit of feeding on pieces of skin and blubber that they gouge from the backs of southern right whales. In response, the whales flinch violently, submerge, and swim rapidly away underwater. The level of harassment in 1995 was almost five times higher than when first studied in 1984 by Thomas (1988). In 1995, 67% of attacks were aimed at large white lesions on the whales' backs. The proportion of whales with lesions increased from 0.01 in 1974 to 0.32 in 1990. Mother-calf pairs that were attacked traveled at medium and fast speeds for 3.1 h per day, compared to 0.8 h for undisturbed pairs. Mother-calf pairs are estimated to spend approximately 24% of their daylight hours in states of gullinduced disturbance. Little food is available at Península Valdés, so mothers must rely on blubber reserves to support their calves' growth, behavioral development, and migration to the feeding grounds. Even when undisturbed by gulls, mothers often curtail their calves' play and nursing bouts, suggesting that their energy reserves are limited. Increasingly intense harassment by gulls may therefore compromise calf development and might even induce right whales to abandon Península Valdés for other calving grounds.     

THE INFLUENCE OF THE CALIFORNIA MARINE LAYER ON BILL SIZE IN A GENERALIST SONGBIRD

The hypothesis is tested that birds in hotter and drier environments may have larger bills to increase the surface area for heat dissipation. California provides a climatic gradient to test the influence of climate on bill size. Much of California experiences dry warm/hot summers and coastal areas experience cooler summers than interior localities. Based on measurements from 1488 museum skins, song sparrows showed increasing body‐size‐corrected bill surface area from the coast to the interior and declining in the far eastern desert. As predicted by Newton's convective heat transfer equation, relative bill size increased monotonically with temperature, and then decreased where average high temperatures exceed body temperature. Of the variables considered, distance from coast, average high summer temperature, and potential evapotranspiration showed a strong quadratic association with bill size and rainfall had a weaker negative relationship. Song sparrows on larger, warmer islands also had larger bills. A subsample of radiographed specimens showed that skeletal bill size is also correlated with temperature, demonstrating that bill size differences are not a result of variation in growth and wear of keratin. Combined with recent thermographic studies of heat loss in song sparrow bills, these results support the hypothesis that bill size in California song sparrows is selected for heat dissipation.