Postnatal development of northern fulmar chicks, Fulmarus glacialis

The slow growth and large fat stores characteristic of many pelagic seabird chicks were generally assumed to reflect infrequent and unpredictable food provisioning by parents. Much less attention has been focused on the importance of intrinsic physiological processes in shaping patterns of development. In this study, we examined postnatal growth and changes in water content of different organs in fulmar chicks, Fulmarus glacialis, from Fair Isle, United Kingdom. After correcting for body size, mass growth rate was as high as in inshore-feeding species, which did not support the notion of an external constraint on growth imposed by the unpredictability of pelagic prey. Pectoral muscles and plumage grew more rapidly than other tissues. Pectorals also had a high water index, probably indicating slower maturation compared with leg muscles, which need to generate heat earlier on to free adults from brooding requirements. Lean dry mass of liver, kidney, and gut decreased markedly toward fledging, presumably because of high energetic costs of maintaining large metabolic machinery in older chicks and analogous to the situation in adult waders before migration. These results suggest that the general pattern of development of fulmars may be linked to changes in resource allocation as chicks grow and possibly a compromise at the tissue level between cell division and the attainment of mature function.     

Unravelling dispersal patterns in an expanding population of a highly mobile seabird,the northern fulmar (Fulmarus glacialis)

The northern fulmar (Fulmarus glacialis) is an abundant seabird whose Northeast Atlantic population has expanded dramatically over the past 100 years. Archaeological evidence suggests that Iceland and St Kilda were the ancestral populations from which essentially all other colonies in the region were derived. We collected samples from seven breeding colonies around the North Atlantic and used mitochondrial DNA analysis to ask whether population structure was present and, if so, where there was evidence about which colony was the dominant source population. Our data reveal a pattern consistent with isolation by distance, suggesting that, even though capable of flying great distances, most birds return to breed either at their own or neighbouring colonies. Interestingly, although most colonizers appear to have come originally from Iceland, our analysis also identifies St Kilda as a possible source. However, this secondary pattern appears to be largely an artefact, and can be attributed to the low haplotype diversity on St Kilda which yields a much clearer isolation by distance signal than that generated by birds dispersing from Iceland, where haplotype diversity is extremely high. Consequently, we urge caution when interpreting patterns in which populations vary greatly in the genetic diversity they harbour.     

Competition between two closely related Tetrabothrius cestodes of the fulmar (Fulmarus glacialis L.).

The two congeneic Tetrabothrius cestodes of the fulmar are so closely related that they can only be reliably distinguished by differences in the number of testes. The two species show a considerable distributional overlap along the intestine. Statistically it can be shown that high numbers of the common, anteriorly situated species (T. minor) affect the distribution of the rarer, more posteriorly situated species (T. procerus), causing the latter to occupy a more posterior site in the intestine. Several possibilities are discussed concerning the mechanism by which T. procerus is affected, and how this mildly competitive situation is maintained. It is probable that niche segregation between the two species is more or less complete, but that this is reinforced in concurrent infections by interactive site segregation.     

Ectoparasites of northern fulmars Fulmarus glacialis (Procellariiformes: Procellariidae) from the Canadian Arctic

We studied the prevalence and intensity of infestation of ectoparasites on northern fulmars (Fulmarus glacialis L.) from a breeding colony in Arctic Canada in June–August 2003. No fleas or ticks were found on any fulmars, but three species of chewing lice (Phthiraptera) were recorded: Ischnocera: Perineus nigrolimbatus (Giebel 1874), Ischnocera: Saemundssonia occidentalis (Kellogg 1896), and Amblycera: Ancistrona vagelli (Fabricius 1787). Non-breeding birds had a higher prevalence of lice than breeding birds, and prevalence varied markedly among louse species. Our study is an important baseline for the occurrence of ectoparasites on northern fulmars in the high Arctic, a region undergoing extensive environmental change due to global warming, and an area where parasites are expected to extend ranges or increase in prevalence under changing annual temperature regimes.     

Context-dependent effects of ranaviral infection on northern leopard frog life history traits

Pathogens have important effects on host life-history traits, but the magnitude of these effects is often strongly context-dependent. The outcome of an interaction between a host and an infectious agent is often associated with the level of stress experienced by the host. Ranavirus causes disease and mortality in amphibian populations in various locations around the world, but most known cases of ranaviral infection have occurred in North America and the United Kingdom. While Ranavirus virulence has been investigated, the outcome of Ranavirus infection has seldom been related to the host environment. In a factorial experiment, we exposed Northern leopard frog (Lithobates pipiens, formerly Rana pipiens) tadpoles to different concentrations of Ranavirus and investigated the effect of host density on certain life-history traits, namely survival, growth rate, developmental stage and number of days from virus exposure to death. Our results suggest a prominent role of density in driving the direction of the interaction between L. pipiens tadpoles and Ranavirus. We showed that increasing animal holding density is detrimental for host fitness as mortality rate is higher, day of death earlier, development longer and growth rate significantly lower in high-density tanks. We observed a linear increase of detrimental effects when Ranavirus doses increased in low-density conditions, with control tadpoles having a significantly higher overall relative fitness. However, this pattern was no longer observed in high-density conditions, where the effects of increasing Ranavirus dose were limited. Infected and control animals fitness were consequently similar. We speculate that the host may eventually diverts the energy required for a metabolic/immune response triggered by the infection (i.e., direct costs of the infection) to better cope with the increase in environmental "stress" associated with high density (i.e., indirect benefits of the infection). Our results illustrate how the net fitness of organisms may be shaped by ecological context and emphasize the necessity of examining the direct/indirect costs and benefits balance to fully understand host-pathogen interactions.     

The reproductive cycle and life history of the Arctic copepod Calanus glacialis in the White Sea

Seasonal variations in the gonad development and sex ratio of copepodite stage V (CV) and adults were examined from February to November in order to understand the reproductive cycle and the life history of Calanus glacialis in the White Sea. Gonad maturation, sexual differentiation and moulting to adults take place during the 2nd year of development. Energy accumulation takes place in the spring and summer of the 2nd year. The following autumn/winter is the major period of CV maturation, which occurs independent of food supply. Maturation of males precedes that of females by 2–3 months. The maximum proportions of CV and adult males are found in the population in October and November. The onset of female maturation is observed in February and March, ca. 2 months prior to the spring phytoplankton bloom. Reproduction takes place between April and June. Its termination in the second half of June coincides with the warming of the surface water layer where egg laying takes place. Variations in the gonad morphology throughout the year suggest long life spans and iteroparity of females of C. glacialis in the White Sea. Many of them survive for several months after reproduction and are able to overwinter again. Therefore, females with different life histories co-occur in the population in winter: “young” females recently moulted from the overwintering CVs, and “old” females which have spawned at least once in their life, after which they return to overwintering conditions. In contrast, males have shorter life spans of 3–4 months resulting in a sex ratio skewed toward females at all seasons. Accepted: 27 April 1999     

Diets of northern fulmar (<Emphasis Type="Italic">Fulmarus glacialis</Emphasis>) chicks in the northwest Atlantic Ocean

Northern fulmars (Fulmarus glacialis) have recently expanded their breeding range in the northwest Atlantic Ocean. We studied their diet in their largest colony in the northwest Atlantic on Funk Island, eastern Canada, by collecting dietary samples from chicks during 1999 and 2000. Fish, primarily capelin (Mallotus villosus), and offal from commercial fisheries were the most common foods in the diets of fulmar chicks. Crustaceans were also common prey fed to the chicks. Squids (Gonatus fabricii) were an important food in 2000. Chick diets varied considerably between the 2 years of the study. The diets of northern fulmar chicks on Funk Island were opportunistic and similar to those of chicks at other colonies in the eastern and northern North Atlantic Ocean. Compared to other regions in the North Atlantic Ocean, the diet of birds on Funk Island seems to be most similar to those from Iceland, and least resemble those from Shetland. Long-term studies of the feeding ecology on northern fulmars may be helpful in discerning factors influencing changes in the species distribution and abundance.     

Breeding ecology of the Fulmar Fulmarus glacialis and the Kittiwake Rissa tridactyla in high-arctic northeastern Greenland, 1993

The breeding ecology of the Fulmar Fulmarus glacialis and the Kittiwake Rissa tridactyla in the high Arctic was studied in relation to the occurrence of the northeast water polynya in northeasternmost Greenland (80̀N). Mean laying dates were 31 May in the Fulmar and 18 June in the Kittiwake; the total nesting season for the Fulmar just matched the time window of the polynya opening period. Fulmar colony attendance fluctuated within a period of 11.6 days because of variation in nonbreeding prospectors but showed no clear diurnal variation. Fulmar incubation shifts, on average, lasted 6.1 days (range 1–13 days), which is significantly longer than elsewhere, and the average chick-guard period of 10.9 days (range 1–17 days) was significantly shorter than in other studies. Egg neglect occurred in 18% of Fulmar nests or 0.7% of nests per day. Overall breeding success (chicks fledged per egg laid) was 0.56 in the Fulmar and 0.67 in the Kittiwake; the latter produced 1.4 young per active nest or 1.2 per completed nest. Mean Kittiwake clutch size was 2.03; larger clutches were laid early. Nest site characteristics (presumably reflecting nest predation risk) and breeding behaviour affected breeding success. in the Fulmar, hatching success was negatively correlated with laying date and the proportion of egg neglect, while overall breeding success was correlated negatively with distance to nearest neighbouring site and positively with the length of the chick-guard period. Kittiwake breeding success was negatively correlated with laying date. Using seabirds as indicators of marine food supply, breeding success in both species suggested moderate to good food supply in the northeast water polynya in 1993, although at least in the Fulmar the high reproductive output appeared partly maintained by behavioural buffering; long incubation shifts, egg neglect and short chick-guard periods were symptoms of foraging constraints.     

Perinatal life history traits in New World monkeys

Gestation length, neonatal and maternal body weight, and neonatal and adult brain weight data were collected for New World monkeys in an attempt to establish typical patterns of perinatal life history. This study attempts to illuminate the most accurate values from the available data, which suggest that gestation length and prenatal growth rate are broadly conserved in relation to maternal size in New World monkeys. Exceptions to the patterns evident in the data point to derivations in life history strategies. In particular, this study suggests that the extended gestation length of callitrichines is a function of minimum viable neonate size and not exclusively energy minimization associated with simultaneous lactation. Cebus is shown to undergo more postnatal brain growth relative to other New World monkeys, but not as much as previously believed. Alouatta is shown to be relatively small brained at birth as well as in adulthood. Saimiri is shown to present the most unusual package of perinatal life history traits, in which precocial neonates are gestated for a relatively long time and at a slightly faster growth rate than is typical for New World monkeys. © 1996 Wiley-Liss, Inc.     

Correlational selection on life history traits in the pitcher-plant mosquito

We examined the short-term effects of genetic architecture on the response to correlational selection in the pitcher-plant mosquito, Wyeomyia smithii. Previously, a positive phenotypic and genetic correlation between development time and propensity to diapause had been identified. Correlational selection consisted of all four combinations of (1) fast and slow development time and (2) propensity to diapause and not diapause, 3 replicates per selection regime, for 1 to 3 generations. Response to selection in the direction of the correlation was rapid and generally consistent with the direction of selection while response orthogonal to the correlation was inconsistent and frequently in a direction opposite to the direction of selection. We conclude that genetic architecture can have a significant, and quite asymmetrical and non-directional, effect on the response to correlational selection.     

Genetic architecture of life history traits and environment-specific trade-offs

Life history theory predicts the evolution of trait combinations that enhance fitness, and the occurrence of trade-offs depends in part on the magnitude of variation in growth rate or acquisition. Using recombinant inbred lines, we examined the genetic architecture of age and size at reproduction across abiotic conditions encountered by cultivars and naturalized populations of Brassica rapa. We found that genotypes are plastic to seasonal setting, such that reproduction was accelerated under conditions encountered by summer annual populations and genetic variances for age at reproduction varied across simulated seasonal settings. Using an acquisition-allocation model, we predicted the likelihood of trade-offs. Consistent with predicted relationships, we observed a trade-off where early maturity is associated with small size at maturity under simulated summer and fall annual conditions but not under winter annual conditions. The trade-off in the summer annual setting was observed despite significant genotypic variation in growth rate, which is often expected to decouple age and size at reproduction because rapidly growing genotypes could mature early and attain a larger size relative to slowly growing genotypes that mature later. The absence of a trade-off in the winter setting is presumably attributable to the absence of genotypic differences in age at reproduction. We observed QTL for age at reproduction that jointly regulated size at reproduction in both the summer and fall annual settings, but these QTL were environment-specific (i.e. different QTL contributed to the trade-off in the fall vs. summer annual settings). Thus, at least some of the genetic mechanisms underlying observed trade-offs differed across environments.     

The life history of Ranunculus glacialis, an arctic-alpine perennial herb, in Finnish Lapland

Dynamics and performance of established individuals of Ranunculus glacialis at the time of peak flowering were monitored in a mountain area in Finnish Lapland for five years (1982–1986). The plants reproduced only sexually, and their spatial distribution in 40 1-m² quadrats was clumped. During the study period the mean density rose from 8.5 to 10.3 individuals m⁻². The mean annual growth rate and birth rate of the population was 4% and 8.5%, respectively. The mean annual death rate of established individuals was 3.7%. The estimated mean life expectancy of an established individual was 27 yr. Death rate per quadrat was positively related to R. glacialis density, but birth rate per quadrat was negatively related to the percentage cover of stones per quadrat. High R. glacialis density per quadrat had negative effects, but high nitrogen availability had positive effects on the number of leaves per individual. An important factor reducing flowering was small rodent herbivory. In the same genets, the correlation between the number of leaves per individual in year t and the number of flowers per individual in year t+4 was significant. The below-ground biomass of R. glacialis comprised about 70% of the total biomass, and there was a close correlation between the above- and the below-ground biomass of an individual. The individuals flowered abundantly each year and produced a great number of small-sized achenes.     

Cost of reproduction and covariation of life history traits in birds

Evolution of life history traits can be studied at two different levels: (1) current selection processes, including trade-offs in life history traits in natural populations as revealed by observations or, preferably, exieriments; and (2) patterns of variation in life history traits with each other and with ecology among extant species. Selection is not evolution, but selection pressures must have caused evolutionary change and led to current patterns of life history traits. These problems are exemplified by recent research on clutch size in birds.     

Multiple growth-correlated life history traits estimated simultaneously in individuals

We present a new methodology to estimate rates of energy acquisition, maintenance, reproductive investment and the onset of maturation (four-trait estimation) by fitting an energy allocation model to individual growth trajectories. The accuracy and precision of the method is evaluated on simulated growth trajectories. In the deterministic case, all life history parameters are well estimated with negligible bias over realistic parameter ranges. Adding environmental variability reduces precision, causes the maintenance and reproductive investment to be confounded with a negative error correlation, and tends, if strong, to result in an underestimation of the energy acquisition and maintenance and an overestimation of the age and size at the onset of maturation. Assuming a priori incorrect allometric scaling exponents also leads to a general but fairly predictable bias. To avoid confounding in applications we propose to assume a constant maintenance (three-trait estimation), which can be obtained by fitting reproductive investment simultaneously to size at age on population data. The results become qualitatively more robust but the improvement of the estimate of the onset of maturation is not significant. When applied to growth curves back-calculated from otoliths of female North Sea plaice Pleuronectes platessa , the four-trait and three-trait estimation produced estimates for the onset of maturation very similar to those obtained by direct observation. The correlations between life-history traits match expectations. We discuss the potential of the methodology in studies of the ecology and evolution of life history parameters in wild populations.     

Coevolution of life history traits and morphology in female subterranean amphipods

Subterranean species show a distinct morphology, yet the adaptive significance of some traits, like body size and shape, is poorly understood and cannot be explained solely by distinct environmental conditions (darkness, less food). We predicted that in females some morphological changes may have co‐evolved with life history traits, and that co‐evolving life history traits provide at least part of the explanation for evolutionary changes of morphology. Using museum material we tested this prediction on the subterranean amphipod genus Niphargus. We studied six species found in springs and eight species found in cave lakes. We treated them as two ecologically distinct groups, and the major ecological differences between them were the availability of nutrients and the water currents. Cave species were found to be larger and stouter (as inferred from the shape of coxal plates, which are part of the marsupium), they had larger eggs and lower reproductive effort per brood, whereas the egg number and brood volume if corrected for the body size were not different. Using phylogenetic independent contrasts, we found a positive correlation between body shape and egg volume, a positive correlation between body size and egg volume, and a negative correlation between body size and reproductive effort per brood. We tentatively conclude that evolutions of morphology and life histories are functionally connected and that co‐evolving traits contribute to overall selective regime.     

Variation in life history and genetic traits of Hawaiian mosquitofish populations

Mosquitofish (Gambusia affinis) were collected from 17 reservoirs on three islands in Hawaii, USA. Genetic and life history traits for adult females from these populations were used to evaluate hypotheses concerning short-term evolutionary divergence of populations recently established from a common ancestral source. The effects of founder events and drift on genetic variability and population differentiation were also examined. Significant differences in life history characteristics, allele frequencies, and multi-locus heterozygosities (H) were found among fish populations collected from different reservoirs and between reservoirs classified as stable or fluctuating on the basis of temporal fluctuation in water level. Females from stable reservoirs exhibited greater standard length (35.1 vs 32.8 mm), lower fecundity (11.9 vs 15.2 embryos), lower reproductive allocation (18.2% vs 22.8%), but larger mean embryo size (1.95 vs 1.67 mg) than females from fluctuating reservoirs. Consistency in means among replicates of each reservoir class and concordance in direction and magnitude of differences reported here and results of sampling conducted from these same locations 10 years previously (Stearns, 1983a) suggest that ecological factors intrinsic to these two environments are important in determing population life history traits. Females from stable reservoirs exhibited lower heterozygosity than females from fluctuating reservoirs (0.134 vs 0.158, respectively). Levels and direction od differences in heterozygosity, the high proportion of polymorphic loci and lack of fixation of alternative alleles argue against a purely stochastic explanation for genetic and life history variation among reservoir populations. Levels of genetic variability and interpopulation differentiation were similar to those observed in mainland populations of this species. A high proportion of the genetic diversity was apportioned between populations and within populations due to differences between juveniles and adults. Significant genotypic differences between adult and juvenile age classes suggest that the genetic divergence of local populations may occur over short periods of time.