BODY SIZE AND SEXUAL SIZE DIMORPHISM IN MARINE IGUANAS FLUCTUATE AS A RESULT OF OPPOSING NATURAL AND SEXUAL SELECTION: AN ISLAND COMPARISON

Body size is often assumed to represent the outcome of conflicting selection pressures of natural and sexual selection. Marine iguana (Amblyrhynchus cristatus) populations in the Galápagos exhibit 10-fold differences in body mass between island populations. There is also strong sexual size dimorphism, with males being about twice as heavy as females. To understand the evolutionary processes shaping body size in marine iguanas, we analyzed the selection differentials on body size in two island populations (max. male mass 900 g in Genovesa, 3500 g in Santa Fé). Factors that usually confound any evolutionary analysis of body sizes—predation, interspecific food competition, reproductive role division—are ruled out for marine iguanas. We show that, above hatchlings, mortality rates increased with body size in both sexes to the same extent. This effect was independent of individual age. The largest animals (males) of each island were the first to die once environmental conditions deteriorated (e.g., during El Niños). This sex-biased mortality was the result of sexual size dimorphism, but at the same time caused sexual size dimorphism to fluctuate. Mortality differed between seasons (selection differentials as low as –1.4) and acted on different absolute body sizes between islands. Both males and females did not cease growth when an optimal body size for survival was reached, as demonstrated by the fact that individual adult body size phenotypically increased in each population under favorable environmental conditions beyond naturally selected limits. But why did marine iguanas grow “too large” for survival? Due to lek mating, sexual selection constantly favored large body size in males (selection differentials up to +0.77). Females only need to reach a body size sufficient to produce surviving offspring. Thereafter, large body size of females was less favored by fertility selection than large size in males. Resulting from these different selection pressures on male and female size, sexual size dimorphism was mechanistically caused by the fact that females matured at an earlier age and size than males, whereafter they constantly allocated resources into eggs, which slowed growth. The observed allometric increase in sexual size dimorphism is explained by the fact that the difference between these selective processes becomes larger as energy abundance in the environment increases. Because body size is generally highly heritable, these selective processes are expected to lead to genetic differences in body size between islands. We propose a common-garden experiment to determine the influence of genetic factors and phenotypic reaction norms of final body size.     

Wingbeat frequency and flap-pause ratio during natural migratory flight in thrushes

Powered flapping flight has evolved independently in many differenttaxa. For flapping fliers, wingbeat parameters such as frequencyand amplitude are the primary determinants of these animals’energetic expenditure during flight. Here we present data onwingbeat frequency and amplitude for three New World thrushspecies during 15 entire nocturnal migratory flights over theMidwestern United States. Using continuous (non-pulsing) radiotransmitters, we were able to measure wingbeat frequency andrelative amplitude of wingbeats as well as the characteristicsof flap-pauses. Contrary to previous telemetric findings, allof the individuals we followed used both flapping-only and flap-pauseflight. During migratory flights, wingbeat frequency, effectivewingbeat frequency, and amplitude were highest during initialascent. Effective wingbeat frequency and amplitude were lowestduring final descent. We show that identification of speciesbased solely on characteristics of the wingbeat e.g., duringradar studies, can be difficult because variables such as wingbeatfrequency and amplitude, wingbeat pausing, and pattern of beatsand pauses vary between individuals of the same species andeven within individual flights. We also show that observed wingbeatfrequencies were lower than those predicted by theoretical models.We speculate that this may be because theoretical predictionsare generally based on (1) data from larger birds and (2) datafrom diurnal flights. We found that diurnal wingbeat frequenciesof thrushes were generally higher than were those during nocturnalmigratory flight. Finally, we suggest that rather than remainingat a single altitude during flight or climbing slightly as theoreticalmodels predict, thrushes often moved up and down in the aircolumn, perhaps searching for favorable atmospheric conditions.     

A theoretical investigation of the effect of latitude on avian life histories

Tropical birds lay smaller clutches than birds breeding in temperate regions and care for their young for longer. We develop a model in which birds choose when and how often to breed and their clutch size, depending on their foraging ability and the food availability. The food supply is density dependent. Seasonal environments necessarily have a high food peak in summer; in winter, food levels drop below those characteristic of constant environments. A bird that cannot balance its energy needs during a week dies of starvation. If adult predation is negligible, birds in low seasonal environments are constrained by low food during breeding seasons, whereas birds in high seasonal environments die during the winter. Low food seasonality selects for small clutch sizes, long parental care times, greater age at first breeding, and high juvenile survival. The inclusion of adult predation has no major effect on any life-history variables. However, increased nest predation reduces clutch size. The same trends with seasonality are also found in a version of the model that includes a condition variable. Our results show that seasonal changes in food supply are sufficient to explain the observed trends in clutch size, care times, and age at first breeding.     

Responding to inflammatory challenges is less costly for a successful avian invader,the house sparrow (<Emphasis Type="Italic">Passer domesticus</Emphasis>), than its less-invasive congener

When introduced into new regions, invading organisms leave many native pathogens behind and also encounter evolutionarily novel disease threats. In the presence of predominantly novel pathogens that have not co-evolved to avoid inducing a strong host immune response, costly and potentially dangerous defenses such as the systemic inflammatory response could become more harmful than protective to the host. We therefore hypothesized that introduced populations exhibiting dampened inflammatory responses will tend to be more invasive. To provide initial data to assess this hypothesis, we measured metabolic, locomotor, and reproductive responses to inflammatory challenges in North American populations of the highly invasive house sparrow (Passer domesticus) and its less-invasive relative, the tree sparrow (Passer montanus). In the house sparrow, there was no effect of phytohemagglutinin (PHA) challenge on metabolic rate, and there were no detectable differences in locomotor activity between lipopolysaccharide (LPS)-injected birds and saline-injected controls. In contrast, tree sparrows injected with PHA had metabolic rates 20–25% lower than controls, and LPS injection resulted in a 35% drop in locomotor activity. In a common garden captive breeding experiment, there was no effect of killed-bacteria injections on reproduction in the house sparrow, while tree sparrows challenged with bacteria decreased egg production by 40% compared to saline-injected controls. These results provide some of the first data correlating variation in immune defenses with invasion success in introduced-vertebrate populations.     

Hormonal correlates of siblicide in Nazca boobies: support for the Challenge Hypothesis

The androgen hormone testosterone (T) mediates vertebrate aggression in many contexts and according to the Challenge Hypothesis is up-regulated during social challenges. While originally applied to challenges experienced by breeding adults, we show for the first time that T is similarly up-regulated during deadly sibling aggression in young birds. When two nestling Nazca boobies hatch, one--usually the older chick--virtually always kills the other chick by pushing it from the nest. We compared concentrations of T, dehydroepiandrosterone (DHEA; a precursor of T), and corticosterone (Cort; a stress hormone) of chicks at various stages. T was elevated during fights in both chicks in two-chick broods, but not before and after fights, and not in chicks lacking a nest mate. DHEA was elevated 1 day after hatching and declined with age but appeared not to vary in concert with aggression. Cort did not vary across fighting and nonfighting periods. In conjunction with an earlier study [Tarlow, E.M., Wikelski, M., Anderson, D.J., 2001. Hormonal correlates of siblicide in Galapagos Nazca boobies. Horm. Behav. 40:14-20], these results indicate that T is temporarily up-regulated around the time of fights, as predicted by the Challenge Hypothesis. Our data suggest a general role for T during challenges at any time in life, not just during breeding.     

Great flexibility in autumn movement patterns of European gadwalls Anas strepera

The annual migration cycle of waterbirds often involves several distinct movement stages, for example within‐winter movements or the moult migration during summer, which require a high degree of individual flexibility in migration direction. Here, we investigate whether such flexibility is a common characteristic of waterbird migration by analysing movement behaviour of a dabbling duck, the gadwall Anas strepera, during the little studied, intermediate autumn period. The tracking of individuals via satellite transmitters (n = 7) as well as the ring re‐encounter analysis of three European gadwall populations (Germany, England, Russia) revealed that autumn movements were multidirectional. Furthermore, the comparison with winter re‐encounters suggested that autumn movements were partly independent of the movements towards subsequently used south to southwestern wintering areas. Some individuals even travelled long distances north‐ or eastwards. Accordingly, some autumn locations were characterized by a harsh climate, thus serving as temporary staging sites but necessitating further movements when wetlands freeze during winter. The occurrence of such detours or reversals of migration was confirmed by the transmitter data. Inter‐individual variability in distance and direction of autumn movements was found for both sexes and age‐classes indicating that gadwalls, in general, followed flexible movement strategies. Based on the extent of multidirectional autumn movements, we hypothesize important benefits of such flights and suggest that the analysis of year‐round movement patterns of individual animals during their distinct life‐history stages is essential to understand how they can successfully reproduce and survive.     

Spatial and Temporal Patterns of Frugivorous Hornbill Movements in Central Africa and their Implications for Rain Forest Conservation

Tropical forest conservation and restoration require an understanding of the movements and habitat preferences of important seed dispersers. With forests now being altered at an unprecedented rate, avian frugivores are becoming increasingly vital for forest regeneration. Seed movement, however, is highly dependent on the behavioral characteristics of their dispersers. Here, we examined the movements, habitat preferences, and range sizes of two African frugivores: the Black‐casqued (Ceratogymna atrata) and the White‐thighed (Bycanistes albotibialis) Hornbill, in the lowland rain forests of southern Cameroon. Using satellite transmitters, we tracked eight hornbills for 3 yr to characterize their movements and relate them to environmental landscape features. Hornbill movements differed significantly, with B. albotibialis ranging over larger areas (mean = 20,274 ha) than C. atrata (mean = 5604 ha), and females of both species covering over 15 times the area of males. Evidence suggests that movements are irruptive during particular periods, perhaps driven by low resource availability. In addition, hornbills often returned to the same localities within a year, although movements were not characterized as migratory. Both species displayed significant differences in habitat preference, with B. albotibialis utilizing disturbed habitat more frequently than C. atrata (t = ?22.04, P = 2.2 × 10^?16). Major roads were found to act as barriers for C. atrata, but not for B. albotibialis. The ability of both hornbill species to move large distances suggests hornbills will play a vital role in the maintenance and regeneration of rain forests in Central Africa as forest fragmentation increases and terrestrial vertebrates decline in numbers.     

Simple rules guide dragonfly migration

Every year billions of butterflies, dragonflies, moths and other insects migrate across continents, and considerable progress has been made in understanding population-level migratory phenomena. However, little is known about destinations and strategies of individual insects. We attached miniaturized radio transmitters (ca 300 mg) to the thoraxes of 14 individual dragonflies (common green darners, Anax junius) and followed them during their autumn migration for up to 12 days, using receiver-equipped Cessna airplanes and ground teams. Green darners exhibited distinct stopover and migration days. On average, they migrated every 2.9+/-0.3 days, and their average net advance was 58+/-11 km in 6.1+/-0.9 days (11.9+/-2.8 km d-1) in a generally southward direction (186+/-52 degrees). They migrated exclusively during the daytime, when wind speeds were less than 25 km h-1, regardless of wind direction, but only after two nights of successively lower temperatures (decrease of 2.1+/-0.6 degrees C in minimum temperature). The migratory patterns and apparent decision rules of green darners are strikingly similar to those proposed for songbirds, and may represent a general migration strategy for long-distance migration of organisms with high self-propelled flight speeds.     

Corticosterone,testosterone and life-history strategies of birds

Steroid hormones have similar functions across vertebrates, but circulating concentrations can vary dramatically among species. We examined the hypothesis that variation in titres of corticosterone (Cort) and testosterone (T) is related to life-history traits of avian species. We predicted that Cort would reach higher levels under stress in species with higher annual adult survival rates since Cort is thought to promote physiological and behavioural responses that reduce risk to the individual. Conversely, we predicted that peak T during the breeding season would be higher in short-lived species with high mating effort as this hormone is known to promote male fecundity traits. We quantified circulating hormone concentrations and key life-history traits (annual adult survival rate, breeding season length, body mass) in males of free-living bird species during the breeding season at a temperate site (northern USA) and a tropical site (central Panama). We analysed our original data by themselves, and also combined with published data on passerine birds to enhance sample size. In both approaches, variation in baseline Cort (Cort0) among species was inversely related to breeding season length and body mass. Stress-induced corticosterone (MaxCort) also varied inversely with body mass and, as predicted, also varied positively with annual adult survival rates. Furthermore, species from drier and colder environments exhibited lower MaxCort than mesic and tropical species; T was lowest in species from tropical environments. These findings suggest that Cort0, MaxCort and T modulate key vertebrate life-history responses to the environment, with Cort0 supporting energetically demanding processes, MaxCort promoting survival and T being related to mating success.