Habitat constraints and spatial bias in seabird colony distributions

What governs the size and location of seabird colonies has long intrigued population ecologists. Previous analysis of the distribution of colonies of four European seabirds revealed a spatial bias - large colonies occurred farther apart than expected by chance alone - suggesting that intraspecific competition for food supplies during breeding may regulate colony size. Here we use computer intensive statistics to show that a similar spatial bias exists in three burrow-nesting seabirds - Cassin's auklet Ptychoramphus aleuticus , rhinoceros auklet Cerorhinca monoccrata , and ancient murrelet Synthliboramphus antiquns - nesting off the Pacific coast of Canada. Local habitat constraints explain much of the existing spatial bias: large colonies cannot fit on small islands, and large islands suitable for colonies tended to occur far apart. However, a residual spatial bias still remained for ancient murrelets and Cassin's auklets (but not rhinoceros auklets) after habitat constraints are built into the analysis, for which intraspecific food competition remains a plausible explanation.     

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.     

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.     

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.     

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.     

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.     

Polymorphic microsatellite DNA markers for the rhinoceros auklet (Cerorhinca monocerata)

Eight polymorphic microsatellite loci were isolated from the partial genomic library of the rhinoceros auklet Cerorhinca monocerata. The heterozygosities observed at the isolated loci using eight primer sets in 46 nestlings from one breeding colony in Japan ranged from 0.311 to 0.773, and all but one locus did not deviate from the Hardy–Weinberg expectations. Cross‐species amplification in the tufted puffin Fratercula cirrhata was successfully performed using six of the eight primer sets, indicating their applicability to this species.     

Metabolic and developmental responses of alcid chicks to experimental variation in food intake.

I tested whether the ability of chicks to suspend growth and developmental processes in response to food shortages is greater among alcids with food resources that fluctuate over short time periods than it is among close relatives with food that is continuously available. I examined changes in chick resting metabolic rate (RMR) in response to short-term food deprivation in horned and tufted puffins (intermittent food provisioning) and crested and parakeet auklets (continuous food provisioning). RMR was based on measurements of chick oxygen consumption rates (Vo2) under thermoneutral conditions. RMR of postabsorptive chicks scaled allometrically with body mass, and regression slopes were statistically indistinguishable among species. Mass-independent RMR of the same individuals decreased significantly after 48 h of food deprivation. The decrease in the mass-independent RMR was greater in puffins (46.8% in horned and 47.4% in tufted puffins) than in auklets (29.4% in crested and 23.7% in parakeet auklets). To test whether the observed decrease in RMR was due to less energy being allocated to growth, I examined developmental responses of horned and tufted puffins to experimental variation in rates of food intake. I found retarded growth rates in body mass, skeletal elements, and feathers in chicks experiencing low rates of food intake. The retardation of growth processes extended the developmental period. My findings suggest that developmental plasticity in juvenile alcids might be related to temporal variability of prey in oceanic environments.     

Molecular phylogenetics and ecological diversification of the transisthmian fish genus Centropomus (Perciformes: Centropomidae).

Phylogenetic relationships among the 12 recognized fish species in the New World genus Centropomus (Pisces, Centropomidae) were analyzed using allozyme electrophoresis and 618 bp of the mitochondrial DNA 16S ribosomal RNA (rRNA) gene. Molecular phylogenetic trees were generally consistent with previously published partial hypotheses based on morphological evidence. However, previously undefined sister group relationships between major species groups were resolved using molecular data, and phylogenetic hypotheses for Centropomus based on 16S rRNA sequences were better supported than were allozyme-based hypotheses. The high level of congruence among the trees inferred from the nuclear and mitochondrial characters provided a firm phylogenetic basis for analysis of ecological diversification and molecular evolution in the genus. Compared to basal Centropomus species, members of the most nested species group were significantly larger in body size and occupied a marine niche only peripherally utilized by their congeners. We also observed substitution rate heterogeneity among 16S rRNA lineages; in contrast to expectations based on "metabolic rate" and "generation interval" models, relative substitution rates were faster than expected for the group of large-bodied snooks. Using the Pliocene rise of the Central American isthmian marine barrier to calibrate rates of 16S ribosomal gene evolution in Centropomus, we found that the rates for the genus were similar to those reported for higher vertebrates. Analysis of the three sets of transisthmian geminate taxa in Centropomus indicated that two of the pairs were probably formed during the Pliocene rise of the isthmus while the third pair diverged several million years earlier.     

Phylogenetic relationships and divergence times of Charadriiformes genera: multigene evidence for the Cretaceous origin of at least 14 clades of shorebirds

Comparative study of character evolution in the shorebirds is presently limited because the phylogenetic placement of some enigmatic genera remains unclear. We therefore used Bayesian methods to obtain a well-supported phylogeny of 90 recognized genera using 5 kb of mitochondrial and nuclear sequences. The tree comprised three major clades: Lari (gulls, auks and allies plus buttonquails) as sister to Scolopaci (sandpipers, jacanas and allies), and in turn sister to Charadrii (plovers, oystercatchers and allies), as in previous molecular studies. Plovers and noddies were not recovered as monophyletic assemblages, and the Egyptian plover Pluvianus is apparently not a plover. Molecular dating using multiple fossil constraints suggests that the three suborders originated in the late Cretaceous between 79 and 102 Mya, and at least 14 lineages of modern shorebirds survived the mass extinction at the K/T boundary. Previous difficulties in determining the phylogenetic relationships of enigmatic taxa reflect the fact that they are well-differentiated relicts of old, genus-poor lineages. We refrain from suggesting systematic revisions for shorebirds at this time because gene trees may fail to recover the species tree when long branches are connected to deep, shorter branches, as is the case for some of the enigmatic taxa.     

Phylogeny and cryptic diversity in geckos (Phyllopezus; Phyllodactylidae; Gekkota) from South America's open biomes

The gecko genus Phyllopezus occurs across South America's open biomes: Cerrado, Seasonally Dry Tropical Forests (SDTF, including Caatinga), and Chaco. We generated a multi-gene dataset and estimated phylogenetic relationships among described Phyllopezus taxa and related species. We included exemplars from both described Phyllopezus pollicaris subspecies, P. p. pollicaris and P. p.przewalskii. Phylogenies from the concatenated data as well as species trees constructed from individual gene trees were largely congruent. All phylogeny reconstruction methods showed Bogertia lutzae as the sister species of Phyllopezus maranjonensis, rendering Phyllopezus paraphyletic. We synonymized the monotypic genus Bogertia with Phyllopezus to maintain a taxonomy that is isomorphic with phylogenetic history. We recovered multiple, deeply divergent, cryptic lineages within P. pollicaris. These cryptic lineages possessed mtDNA distances equivalent to distances among other gekkotan sister taxa. Described P. pollicaris subspecies are not reciprocally monophyletic and current subspecific taxonomy does not accurately reflect evolutionary relationships among cryptic lineages. We highlight the conservation significance of these results in light of the ongoing habitat loss in South America's open biomes.     

Distribution of Xantus' Murrelet Synthliboramphus hypoleucus at sea in the Southern California Bight, 1995–97

We radiomarked 153 Xantus' Murrelets Synthliboramphus hypoleucus captured at sea near Santa Barbara Island (SBI), the largest murrelet colony in the California Channel Islands, USA. We tracked these radiomarked murrelets in the Southern California Bight (SCB) off coastal southern California during the 1995-97 breeding seasons. In 1995 during mild El Niño conditions, the murrelets were distributed in non-upwelling areas. In 1996-97, they were distributed in dense patches, aggregating in cool upwelled waters near the northern Channel Islands or south of San Nicolas Island. Murrelets flew longer distances from SBI to foraging areas in 1997 (̄= 111 ± 44 km) than in 1996 (̄= 62 ± 25 km), but the distances they travelled did not differ between months (Apr and May) within years. Mean foraging distances from SBI were similar for 'incubating' murrelets (determined on the basis of repeated visits to SBI) and 'non-incubating' murrelets during the colony attendance period. We attributed the low return rate of radiomarked murrelets to SBI to the capture and marking of a large proportion of birds that were not actively incubating rather than to any adverse effects of radio attachment. We believe changes in murrelet foraging patterns between the 1970s and 1990s are associated with changes in prey resources in the SCB. Flexibility in the foraging strategies of these murrelets may be related to the highly variable marine environment at the southern end of the California Current Up welling System.     

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.     

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.     

Mitochondrial DNA phylogeny of North American field crickets: perspectives on the evolution of life cycles,songs, and habitat associations

North American field crickets (genus Gryllus) exhibit a diversity of life cycles, habitat associations, and calling songs. However, patterns of evolution for these ecological and behavioral traits remain uncertain in the absence of a robust phylogenetic framework. Analyses of morphological variation have provided few clues about species relationships in the genus Gryllus. Here we use comparisons of mitochondrial DNA restriction site maps for 29 individuals representing 11 species (including potential outgroups) to examine relationships among eastern North American field crickets. Initially chosen as likely outgroup taxa, the two European species of Gryllus do not obviously fall outside of an exclusively North American clade and (based on amount of sequence divergence) appear to have diverged from North American lineages at about the same time that major North American lineages diverged from each other. The egg-overwintering crickets comprise a strongly supported monophyletic group, but relationships among these three closely related species cannot be resolved. The mtDNA data are consistent with a single origin of egg diapause and do not support a model of recent life cycle divergence and allochronic speciation for Gryllus pennsylvanicus and G. veletis. The two crickets are not sister species, despite remarkable similarity in morphology, habitat, and calling song. This conclusion is consistent with published data on allozyme variation in North American field crickets. The habitat associations of eastern North American field crickets have been labile, but calling songs sometimes have remained virtually unchanged across multiple speciation events.     

Polemoniaceae phylogeny and classification: implications of sequence data from the chloroplast gene ndhF

The chloroplast gene ndhF was used to study phylogenetic relationships of the Polemoniaceae at two levels: among members of the Ericales and among genera of the family. Sequence data for interfamilial analyses consisted of 2266 bp for 14 members of the Ericales, including four species of the Polemoniaceae, plus three outgroup taxa. The Polemoniaceae were found to be related to Diospyros, Fouquieria, the Primulales, Rhododendron, and Impatiens, but relationships among taxa were generally not well supported. The precise position of the Polemoniaceae within the Ericales remains obscure. Data for intrafamilial analyses consisted of 1031 bp for 27 species of the Polemoniaceae, including at least one species from most genera of the family, plus five outgroup taxa. A single most parsimonious tree was identified. The analyses suggested that subfamily Cobaeoideae, excluding Loeselia, is monophyletic and that Huthia is sister to Cantua. Acanthogilia was sister to the remainder of subfamily Cobaeoideae. Subfamily Polemonioideae plus Loeselia formed four subclades that were strongly supported as monophyletic and represent the major lineages of the subfamily.     

Status,trends, and patterns of covariation of breeding seabirds at St Lazaria Island,Southeast Alaska, 1994–2006

Aim We examined patterns of covariation among piscivorous and planktivorous seabirds breeding at St Lazaria Island in order to evaluate their responses to interannual changes in sea surface temperature, a variable that affects marine food webs. In addition, we evaluated seabird population trends for responses to decadal‐scale changes in the marine ecosystem. Location St Lazaria Island, Sitka Sound, Alaska. Methods Established seabird monitoring protocols for the Alaska Maritime National Wildlife Refuge were followed in estimating population trends, the timing of nesting events and the reproductive success of eight species of seabirds between 1994 and 2006. Results  Population increases were noted for storm‐petrels (Oceanodroma furcata and O. leucorhoa), rhinoceros auklets (Cerorhinca monocerata) and glaucous‐winged gulls (Larus glaucescens). We found no population trend for pelagic cormorants (Phalacrocorax pelagicus), but it appeared that populations of common (Uria aalge) and thick‐billed (U. lomvia) murres and of tufted puffins (Fratercula cirrhata) declined. We detected no linear trends in either breeding chronology or reproductive success over the study period for any seabird. All species of piscivorous seabirds apparently responded similarly to environmental cues as there was a positive covariation among species in the timing of nesting. Piscivores tended to nest earlier, and most species had higher rates of reproductive success in years with relatively warm spring sea temperatures. In contrast, planktivorous Leach’s storm‐petrels (O. leucorhoa) tended to nest earlier when spring and summer sea temperatures were relatively cool. Clearly, seabirds at St Lazaria were responding to interannual changes in sea temperatures near the breeding colony, probably as a result of effects on the food webs. Main conclusions Every seabird species we monitored at St Lazaria exhibited significant population trends between 1994 and 2006. For most species there appeared to be a relationship between both the timing of nesting and reproductive rates and spring or summer sea surface temperatures. Responses at both decadal (populations) and interannual (timing and reproductive success) scales make seabirds useful candidates for helping to monitor change in the marine environment.     

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.     

Molting and Seasonal Bill-Plate Shedding in the Whiskered Auklet (Aethia pygmaea)

Feather molting and bill-late shedding were studied because of the unique features of the whiskered auklet biology; i.e., they continue to visit the colony after departure of their young. Like other auklets, the whiskered auklets begin to molt during breeding and do not lose their capacity for flight. The molt pattern of different wing feathers is adaptive and allows new feathers to be protected (when they are soft and could be easily injured) by old or full-grown new feathers during flight or feeding (diving) due to the different timing of the molt of primary feathers and their coverts. The possibility of combining breeding with molt appears to be related to the feeding features of the species. The species that feed on abundant and highly aggregated plankton are able to molt during breeding. The pattern of bill-plate shedding in the whiskered auklet is similar to that in the crested auklet.     

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.