Phylogenetic relationships within the Alcidae (Charadriiformes: Aves) inferred from total molecular evidence

The Alcidae is a unique assemblage of Northern Hemisphere seabirds that forage by "flying" underwater. Despite obvious affinities among the species, their evolutionary relationships are unclear. We analyzed nucleotide sequences of 1,045 base pairs of the mitochondrial cytochrome b gene and allelic profiles for 37 allozyme loci in all 22 extant species. Trees were constructed on independent and combined data sets using maximum parsimony and distance methods that correct for superimposed changes. Alternative methods of analysis produced only minor differences in relationships that were supported strongly by bootstrapping or standard error tests. Combining sequence and allozyme data into a single analysis provided the greatest number of relationships receiving strong support. Addition of published morphological and ecological data did not improve support for any additional relationship. All analyses grouped species into six distinct lineages: (1) the dovekie (Alle alle) and auks, (2) guillemots, (3) brachyramphine murrelets, (4) synthliboramphine murrelets, (5) true auklets, and (6) the rhinoceros auklet (Cerorhinca monocerata) and puffins. The two murres (genus Uria) were sister taxa, and the black guillemot (Cepphus grylle) was basal to the other guillemots. The Asian subspecies of the marbled murrelet (Brachyramphus marmoratus perdix) was the most divergent brachyramphine murrelet, and two distinct lineages occurred within the synthliboramphine murrelets. Cassin's auklet (Ptychoramphus aleuticus) and the rhinoceros auklet were basal to the other auklets and puffins, respectively, and the Atlantic (Fratercula arctica) and horned (Fratercula corniculata) puffins were sister taxa. Several relationships among tribes, among the dovekie and auks, and among the auklets could not be resolved but resembled "star" phylogenies indicative of adaptive radiations at different depths within the trees.      

Yolk carotenoids and stable isotopes reveal links among environment, foraging behavior and seabird breeding success

Nutrients that are limited in availability, such as carotenoids, are potentially involved in trade-offs between homeostasis and reproduction. Despite their importance, factors that affect the capacity of female birds to meet their carotenoid requirements are poorly understood. We used δ¹⁵N stable isotope analysis to relate foraging behavior to yolk carotenoid deposition in two seabirds, Cassin’s auklet (Ptychoramphus aleuticus) and rhinoceros auklet (Cerorhinca monocerata), during each of five years. As expected from their narrower trophic range, Cassin’s auklets produced yolks with fewer carotenoid types than did rhinoceros auklets (one vs. three). Cassin’s auklets also fed on a lower trophic level diet richer in carotenoids, yet had lower total yolk carotenoid levels, which suggests a role for species-specific adaptations for carotenoid uptake and utilization. Within both species, lower trophic-level feeding was linked to higher yolk carotenoid levels, but through different mechanisms. In Cassin’s auklets, it was due to a population-wide response to environmental variation: in warm-water years, all females fed at a low trophic level and produced carotenoid-rich yolks. In rhinoceros auklets, it was due to individual differences similarly expressed in all years: females fed across a wide trophic range, and those that fed at a low trophic level produced carotenoid-rich yolks. Rhinoceros auklets bred more successfully in years when their yolks were rich in carotenoids, probably due to a correlated response to stronger marine primary production. Our results are novel because they link avian yolk carotenoid deposition to behavioral and environmental variations.     

Population differentiation in whiskered auklets Aethia pygmaea: do diurnal and nocturnal colonies differ in genetics,morphometry and acoustics?

Despite a great number of empirical studies, the mechanisms of population differentiation and the factors that influence this process, particularly in seabirds, remain insufficiently understood. Here we analyzed population structure in the whiskered auklet Aethia pygmaea, a previously poorly studied alcid species with unusual differentiation in colony attendance rhythms (i.e. diurnal in the Sea of Okhotsk vs nocturnal in the Bering Sea), and examined the influence of it on intraspecific differentiation. For this study, we analyzed morphometric measurements, acoustic variables, mitochondrial control region fragment and five microsatellite loci from nine whiskered auklet colonies spanning the breeding range. Previous research has shown a clinal morphometric variation in this species. We build on this analysis by adding auklets from more colonies, for the first time analyzing vocalizations from different colonies and genetic structure of this species. Our data supports a clinal variation in morphometric and acoustic characters with the largest size and the lowest call frequency in western birds, and the smallest size and highest call frequency in the east. We also found two mitochondrial lineages – whiskered auklets from colonies in the Sea of Okhotsk and the Commander Is. (Bering Sea) and from the Aleutian Is. (Bering Sea), that were presumably formed during Sangamonian interglacial period (115 000–130 000 years ago). Genetic clusters found did not reflect differences in colony attendance rhythms, suggesting that they were shaped by other factors (e.g. differences in predator pressure) and are unlikely to have participated in the formation of population structure. Colony fidelity, mobility of birds, proximity of foraging grounds and location of colonies in relation to seasonal ice pack, seem to be more likely determinants of population differentiation.     

Differential use of zooplankton prey by Ancient murrelets and Cassin's auklets in the Queen Charlotte Islands

The distribution at sea and the food of two similar sized plankton-feedingalcids were examined during the 1981 breeding seasons in thenorthwestern Queen Charlotte Islands, British Columbia. Thetwo alcids, the Ancient murrelet (Synthliboramphus antiquus)and the Cassin's auklet (Prychoramphus aleuticus) have differentchick-rearing strategies. Both species fed predominantly atthe shelf break, although the Cassin's auklet also foraged overseamounts. The feeding distributions of the species appear tobe related to those of their prey. Zooplankton sampling indicatedthat each alcid selects a small and different portion of thezooplankton available in surface waters. The Ancient murrelet'smain foods were euphausiids (Thysanoessa spinifera and Euphausiapacifica) and larval and juvenile fishes. The Cassin's aukletchicks fed chiefly on calanoid copepods (Neocalanus cristatus).euphausiids (mostly Thysanoessa longipes in 1981, but in otheryears also Thysanoessa spinifera), and larval and juvenile fishes.The Cassin's auklets took smaller prey than the Ancient murrelet.Differences in the diets of the two alcid species were associatedwith differences in morphology and chick-rearing strategies.     

Rates and consequences of relaying in Cassin's auklets Ptychoramphus aleuticus and rhinoceros auklets Cerorhinca monocerata breeding in a seasonal environment

We removed first eggs from early‐laying females to measure rates and consequences of relaying in Cassin's auklets Ptychoramphus aleuticus and rhinoceros auklets Cerorhinca monocerata at Triangle Island, British Columbia, Canada. Based on egg size and composition, the investment that Cassin's auklets made in first eggs was very close to that predicted from adult body mass, whereas rhinoceros auklets invested more. In both species, a high percentage of females relaid (90% of Cassin's and 87% of rhinoceros auklets). Breeding success declined weakly with later laying among control Cassin's auklet pairs, but pairs that we induced to relay bred more successfully than naturally late pairs, and similar to values predicted from laying dates of their first eggs. Their chicks also fledged heavier and younger than late control chicks, and similar to values in early control chicks, but followed the population‐wide seasonal decline in wing length at fledging. Nestling diets were dominated by Neocalanus copepods until late in the season, a sign that feeding conditions remained favourable until late. In contrast, rhinoceros auklet pairs induced to relay followed the population‐wide seasonal decline in breeding success, which was driven by a decline in hatching success. Pacific sandlance Ammodytes hexapterus, thought to be a preferred prey species, virtually disappeared from nestling diets in mid‐to‐late season, yet there was no seasonal decline in fledging mass. However, chicks from replacement eggs followed the declines among control chicks in both age and wing length at fledging. Despite the female having produced a replacement egg, and despite delayed breeding, there appeared to be little immediate consequence associated with relaying for Cassin's auklets, except for a tendency for their chicks to fledge with short wings. Consequences were more marked in rhinoceros auklets (greatly reduced hatching success, and having their chicks fledge with short wings), and this may have been due to the large investment made in eggs, and/or to delayed breeding. Results of this study show that attributes of Cassin's and rhinoceros auklets that lay at different times in the season can be important in driving seasonal declines in breeding performance, as found in studies on other Alcidae. They also show how decisions taken during the egg stage can have variable yet potentially important implications for fitness, even in relatively long‐lived species that lay single‐egg clutches.     

Individuality in trumpet calls of the crested auklet (<Emphasis Type="Italic">Aethia cristatella</Emphasis>), a highly social species

Crested auklet is a unique representative of the family Alcidae with a complex social behavior nesting in dense colonies with a size of up to one million individuals. Here we study specific individual features in their trumpet calls, loud vocalizations used by auklets as an advertising display. In 2008 and 2009, we recorded 231 calls of 24 individually marked male crested auklets from the Talan Island (Sea of Okhotsk). According to a stepwise discriminant function analysis based on seven variables in the call introduction, the rate of correct call assignment to an individual was 84.8% and that in the case of seven variables of the main part of the call was 82.3%, which considerably exceeds a random classification. Thus, a single introduction or a single main part of the call is sufficient for a reliable individual identification of a calling bird. We assume that such redundancy of information for individual recognition can enhance identification of calls of a large number of auklet conspecifics in dense colonies with a nesting density of several tens of pairs per 1 m².     

Heterospecific mating preferences for a feather ornament in least auklets

Auklets (Alddae, Aethiini) include five species of small, sociallymonogamous, sexually monomorphic seabirds that display a varietyof feather and bare-part ornaments during the breeding season.Previous experimental work on two auklet species has demonstratedthat some ornaments are likely to be favored by sexual selectionbecause mutual male and female mating preferences benefit individualswith the most elaborate expression of these traits. In thisstudy we experimentally investigated whether naturally crestlessleast auklets Aethia pusilla have a maring preference for foreheadcrests similar to the most prominent ornament of two other species,crested A. cristatella and whiskered auklets A. pygmaea. Ourobjective was to investigate the function of this ornament asa species-recognition mechanism or as a product of one or moreof three proposed sexual selection models that address the originof elaborate traits and preferences. During the experiment,least auklets reacted to realistic models equipped with artificialforehead crests with approximately an order of magnitude morefrequent sexual displays and greater interest, consistent withthe idea that they have a mating preference for crests, eventhough they do not naturally express this ornament This heterospecificpreference also favored large crest size. These results refutethe possibility that least auklet forehead ornamentation alonedetermines species recognition at present Among models of sexualselection considered, the results are consistent with the sensoryexploitation model, although this could not be established unequivocallybecause a viability indicator or Fisherian mechanism could havebeen involved if least auklets had an ancestor with a foreheadcrest.     

Limited consequences of infestation with a blood‐feeding ectoparasite for the nestlings of two North Pacific seabirds

The seabird tick Ixodes uriae parasitizes over 60 host species in the circumpolar regions of both hemispheres. To assess the impacts of these ticks on the growth and development of nestling seabirds, we used a logistic growth model to interpolate between successive measures of mass (g) and wing chord (mm) for 558 Cassin's auklet Ptychoramphus aleuticus and 344 rhinoceros auklet Cerorhinca monocerata chicks over 11 years (1997–2008, less 2003) on Triangle Island, British Columbia, Canada. From the model, we estimated the asymptotic measure and the age at inflection point for each chick's growth trajectory, and assessed their relationships with tick load relative to other sources of annual and seasonal variation in growth. Most chicks (72.4% of Cassin's auklets, 62.2% of rhinoceros auklets) hosted ≥ 1 ticks, and the median tick load on infested chicks was two in both species. Infestation rates varied by a factor of about two among years (0.42 to 0.87 overall), but were uncorrelated between species and with air temperatures over the preceding winter. The probability of hosting a tick declined strongly with chick age, mainly in the first 20 days after hatching, and to near zero by fledging. Asymptotic weights and/or wing lengths declined with tick load in both species, but at normal loads the reductions were minor relative to those imposed by other factors; only at very high loads, which were rare, were effects likely to be biologically relevant. Tick load and survival to fledging were unrelated in both species. While our study found little influence of ticks, we believe there is need for further study of the relationships between parasites and seabird demography, especially in light of ongoing environmental change.     

Varying foraging patterns in response to competition? A multicolony approach in a generalist seabird

Reducing resource competition is a crucial requirement for colonial seabirds to ensure adequate self‐ and chick‐provisioning during breeding season. Spatial segregation is a common avoidance strategy among and within species from neighboring breeding colonies. We determined whether the foraging behaviors of incubating lesser black‐backed gulls (Larus fuscus) differed between six colonies varying in size and distance to mainland, and whether any differences could be related to the foraging habitats visited. Seventy‐nine incubating individuals from six study colonies along the German North Sea coast were equipped with GPS data loggers in multiple years. Dietary information was gained by sampling food pellets, and blood samples were taken for stable isotope analyses. Foraging patterns clearly differed among and within colonies. Foraging range increased with increasing colony size and decreased with increasing colony distance from the mainland, although the latter might be due to the inclusion of the only offshore colony. Gulls from larger colonies with consequently greater density‐dependent competition were more likely to forage at land instead of at sea. The diets of the gulls from the colonies furthest from each other differed, while the diets from the other colonies overlapped with each other. The spatial segregation and dietary similarities suggest that lesser black‐backed gulls foraged at different sites and utilized two main habitat types, although these were similar across foraging areas for all colonies except the single offshore island. The avoidance of intraspecific competition results in colony‐specific foraging patterns, potentially causing more intensive utilization of terrestrial foraging sites, which may offer more predictable and easily available foraging compared with the marine environment.     

An energetic correlate between colony size and foraging effort in seabirds, an example of the Adélie penguin Pygoscelis adeliae

Central-place foraging seabirds alter the availability of their prey around colonies, forming a "halo" of reduced prey access that ultimately constrains population size. This has been indicated indirectly by an inverse correlation between colony size and reproductive success, numbers of conspecifics at other colonies within foraging range, foraging effort (i.e. trip duration), diet quality and colony growth rate. Although ultimately mediated by density dependence relative to food through intraspecific exploitative or interference competition, the proximate mechanism involved has yet to be elucidated. Herein, we show that Adélie penguin Pygoscelis adeliae colony size positively correlates to foraging trip duration and metabolic rate, that the metabolic rate while foraging may be approaching an energetic ceiling for birds at the largest colonies, and that total energy expended increases with trip duration although uncompensated by increased mass gain. We propose that a competition-induced reduction in prey availability results in higher energy expenditure for birds foraging in the halo around large colonies, and that to escape the halo a bird must increase its foraging distance. Ultimately, the total energetic cost of a trip determines the maximum successful trip distance, as on longer trips food acquired is used more for self maintenance than for chick provisioning. When the net cost of foraging trips becomes too high, with chicks receiving insufficient food, chick survival suffers and subsequent colony growth is limited. Though the existence of energetic studies of the same species at multiple colonies is rare, because foraging metabolic rate increases with colony size in at least two other seabird species, we suggest that an energetic constraint to colony size may generally apply to other seabirds.     

Variations in the diet of introduced Norway rats (Rattus norvegicus) inferred using stable isotope analysis

The presence of introduced Norway rats Rattus norvegicus has raised concerns for the fate of the large least auklet Aethia pusilla colony situated at Sirius Point, Kiska Island, Alaska. Previous studies have documented extreme interannual variation in least auklet reproductive success and potential drastic population declines, both of which have been attributed to the varying abundance of, and predation by, Norway rats. A diet study would resolve the uncertainty that remains about the role of rats in the auklet's reproductive failure and the colony's decline. Our main objectives here were to quantify the variation in diet of introduced Norway rats and assess predation on least auklets. Using stable isotope analysis we document wide variability in rat diet dependent on location and provide direct evidence that Norway rats are preferentially preying on least auklets at Sirius Point. In conclusion, we hypothesize that the observed wide variability in rat diet will contribute to the persistence of rats on Kiska long after auklets have been extirpated. The persistence of rats enabled by their foraging plasticity will increase their effects by creating ecological traps within which prospecting individuals will fall and be depredated. This has large conservation consequences as it suggests that when seabirds are extirpated recolonization by prospecting birds is virtually impossible and island ecosystems will continue to be negatively affected and altered as long as introduced predators, such as rats, remain within them.     

Assessment of 17 new whiskered auklet (Aethia pygmaea) microsatellite loci in 42 seabirds identifies 5–15 polymorphic markers for each of nine Alcinae species

We isolated 17 microsatellite loci in the whiskered auklet (Aethia pygmaea) and tested them for amplification in 48 species from 13 seabird families (including 42 seabirds). Fifteen of these loci were also tested for polymorphism in 38 of the species, which included nine species of Alcinae (four auklets, Atlantic puffin, dovekie, razorbill and two murre species). On the average, nine loci were polymorphic per Alcinae species.     

Effects of air and water temperatures on resting metabolism of auklets and other diving birds

For small aquatic endotherms, heat loss while floating on water can be a dominant energy cost, and requires accurate estimation in energetics models for different species. We measured resting metabolic rate (RMR) in air and on water for a small diving bird, the Cassin's auklet (Ptychoramphus aleuticus), and compared these results to published data for other diving birds of diverse taxa and sizes. For 8 Cassin's auklets (~165 g), the lower critical temperature was higher on water (21 °C) than in air (16 °C). Lowest values of RMR (W kg?1) averaged 19% higher on water (12.14 ± 3.14 SD) than in air (10.22 ± 1.43). At lower temperatures, RMR averaged 25% higher on water than in air, increasing with similar slope. RMR was higher on water than in air for alcids, cormorants, and small penguins but not for diving ducks, which appear exceptionally resistant to heat loss in water. Changes in RMR (W) with body mass either in air or on water were mostly linear over the 5- to 20-fold body mass ranges of alcids, diving ducks, and penguins, while cormorants showed no relationship of RMR with mass. The often large energetic effects of time spent floating on water can differ substantially among major taxa of diving birds, so that relevant estimates are critical to understanding their patterns of daily energy use.     

Is the citrus-like plumage odorant of crested auklets (Aethia cristatella) a defense against lice?

Plumage odors may function as chemical defenses against ectoparasites in birds. We tested this hypothesis for crested auklets (Aethia cristatella), a species of colonial seabird that emits a very strong citrus-like odor from its plumage. This odorant contains known chemical repellents. We evaluated evidence for chemical defense in this species using two approaches. First, we exposed pigeon lice to the volatiles emitted by freshly plucked plumage and by whole specimens. Louse survivorship was compared between these treatments and two controls. Second, we compared louse abundance on crested auklets versus a closely related congener that nests in close association. Louse survivorship did not differ between crested auklet treatments and controls. Comparison of ectoparasite loads showed that crested auklets had significantly higher louse abundance than least auklets (A. pusilla), even after controlling for body size. Our results failed to support two expectations for chemical defense. Presence of the crested auklet plumage odorant in our experiments did not reduce louse life span. Presence of the aldehyde odorant in nature did not reduce louse abundance on crested auklets. Hence we conclude that the aldehyde odorant is not immediately lethal to lice at natural concentrations in plumage.     

Variation in breeding parameters of Eleonora’s falcon (Falco eleonorae) and factors affecting its reproductive performance

We gathered data for a four-year period (2004?C2007) on the nesting ecology and reproductive performance of Eleonora??s falcon in the Aegean Sea. We investigated in an indirect way the relation between clutch size and pre-laying food availability, the significance of site and pair quality on productivity, and the effects of habitat and intraspecific competition on breeding success. Overall, the species selected nest sites sheltered from sun exposure but not from the prevailing wind. Hatching, fledging, and breeding success rates were estimated at 64, 72%, and 60%, respectively. Fledglings per breeding pair ranged from 1.19 to 1.75, and fledglings per successful pair from 1.84 to 2.0, between the years. Productivity depended on parental care rather than nest-site quality. Breeding parameters varied significantly between the years, exhibiting a strong spatial yet local effect. Low success rates were recorded in specific colonies, which were attributed to adverse weather conditions and habitat degradation. Insect food availability prior to egg-laying, estimated via plant biomass, was positively correlated to clutch size. Meanwhile, low wind strengths in August, large distances from the mainland, and population size in the vicinity of the colonies had negative effects on breeding success. Considering data at the Mediterranean scale, a longitudinal trend is observed across the breeding range of this species, with breeding parameters slightly decreasing from west to east.     

Influence of density‐dependent competition on foraging and migratory behavior of a subtropical colonial seabird

Density‐dependent competition for food resources influences both foraging ecology and reproduction in a variety of animals. The relationship between colony size, local prey depletion, and reproductive output in colonial central‐place foragers has been extensively studied in seabirds; however, most studies have focused on effects of intraspecific competition during the breeding season, while little is known about whether density‐dependent resource depletion influences individual migratory behavior outside the breeding season. Using breeding colony size as a surrogate for intraspecific resource competition, we tested for effects of colony size on breeding home range, nestling health, and migratory patterns of a nearshore colonial seabird, the brown pelican (Pelecanus occidentalis), originating from seven breeding colonies of varying sizes in the subtropical northern Gulf of Mexico. We found evidence for density‐dependent effects on foraging behavior during the breeding season, as individual foraging areas increased linearly with the number of breeding pairs per colony. Contrary to our predictions, however, nestlings from more numerous colonies with larger foraging ranges did not experience either decreased condition or increased stress. During nonbreeding, individuals from larger colonies were more likely to migrate, and traveled longer distances, than individuals from smaller colonies, indicating that the influence of density‐dependent effects on distribution persists into the nonbreeding period. We also found significant effects of individual physical condition, particularly body size, on migratory behavior, which in combination with colony size suggesting that dominant individuals remain closer to breeding sites during winter. We conclude that density‐dependent competition may be an important driver of both the extent of foraging ranges and the degree of migration exhibited by brown pelicans. However, the effects of density‐dependent competition on breeding success and population regulation remain uncertain in this system.     

Using Seabird Habitat Modeling to Inform Marine Spatial Planning in Central California’s National Marine Sanctuaries

Understanding seabird habitat preferences is critical to future wildlife conservation and threat mitigation in California. The objective of this study was to investigate drivers of seabird habitat selection within the Gulf of the Farallones and Cordell Bank National Marine Sanctuaries to identify areas for targeted conservation planning. We used seabird abundance data collected by the Applied California Current Ecosystem Studies Program (ACCESS) from 2004–2011. We used zero-inflated negative binomial regression to model species abundance and distribution as a function of near surface ocean water properties, distances to geographic features and oceanographic climate indices to identify patterns in foraging habitat selection. We evaluated seasonal, inter-annual and species-specific variability of at-sea distributions for the five most abundant seabirds nesting on the Farallon Islands: western gull (Larus occidentalis), common murre (Uria aalge), Cassin’s auklet (Ptychorampus aleuticus), rhinoceros auklet (Cerorhinca monocerata) and Brandt’s cormorant (Phalacrocorax penicillatus). The waters in the vicinity of Cordell Bank and the continental shelf east of the Farallon Islands emerged as persistent and highly selected foraging areas across all species. Further, we conducted a spatial prioritization exercise to optimize seabird conservation areas with and without considering impacts of current human activities. We explored three conservation scenarios where 10, 30 and 50 percent of highly selected, species-specific foraging areas would be conserved. We compared and contrasted results in relation to existing marine protected areas (MPAs) and the future alternative energy footprint identified by the California Ocean Uses Atlas. Our results show that the majority of highly selected seabird habitat lies outside of state MPAs where threats from shipping, oil spills, and offshore energy development remain. This analysis accentuates the need for innovative marine spatial planning efforts and provides a foundation on which to build more comprehensive zoning and management in California’s National Marine Sanctuaries.     

Home ranges of introduced rats on Christmas Island: A pilot study

Understanding the spatial ecology of invasive rats (Rattus spp.) is necessary to inform management actions to reduce their impact on native flora and fauna. This study investigates home range sizes of exotic rats around seabird colonies and urban areas on Christmas Island, where rat predation is suspected to be adversely affecting fledgling success among local seabirds. It was hypothesised that rat home range sizes would be smaller in urban areas owing to more consistent food availability. Home ranges of male rats were significantly larger compared with their female counterparts, with male rats maintaining larger home ranges in urban areas compared with seabird colonies. Conversely, female rats had smaller home ranges in urban areas compared with seabird colonies. Our findings suggest a possible correlation between the spatial distribution of food resources and home range size. Additionally, the spatial distribution of breeding females across the landscape had a significant influence on the home ranges of male rats. These findings have important implications for proposed efforts to manage rat populations on Christmas Island, while also providing valuable information regarding the ecology of invasive rats on tropical islands.     

Size of ornament is negatively correlated with baseline corticosterone in males of a socially monogamous colonial seabird

The Goymann–Wingfield model predicts that glucocorticoid levels in social animals reflect the costs of acquiring and maintaining social status. The crested auklet is one of the few avian colonial species where a mutual ornament in males and females is used in both sexual and aggressive displays. Previous studies of the crested auklet support the notion that the crest ornament is a badge of status in this species. Here, we examined the relationship between the crest ornament size and the adrenocortical function in breeding crested auklets. Crest length was negatively correlated with corticosterone at baseline in males, but not in females. Baseline corticosterone in females (but not in males) was negatively correlated with body condition index. Although male and female crested auklets are monomorphic in their ornamental traits, our results suggest that the socially mediated physiological costs associated with status signaling may differ between the sexes.     

Habitat-related aggressive behaviour between neighbouring colonies of the polydomous wood ant Formica aquilonia

Ant species of the late succession stages usually have several queens and a polydomous colony structure, where several colonies occupy the same territory without competing. The wood ant Formica aquilonia is a good example of such a species in boreal forests. In this species, the lack of intraspecific competition may be caused by the stable environment and abundant food resources (e.g. excretions of tree-living aphids). We studied how habitat destruction, in the form of clear-cutting, affects aggressiveness between neighbouring colonies in F. aquilonia. Intercolonial relationships were more hostile in clear-cuts than in forest interiors. This aggression may be the result of increased intraspecific resource competition or alienation of neighbouring colonies caused by the loss of visual orientation cues. We suggest that the original polydomous relationships between colonies decline and colonies may start aggressive competition for the remaining resources.