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.     

The importance of Paracallisoma coecus and myctophid fishes to nesting fork-tailed and Leach's storm-petrels in the Queen Charlotte Islands, British Columbia

The diet of fork-tailed (Oceanodroma furcata) and Leach's storm-petrel(O.leucorhoa) was studied at Petrel Island, a small island northof Hippa Island in the Queen Charlotte Islands, British Columbiain 1983. The diet of the two storm-petrel species consistedmostly of the amphipod Paracallisoma coecus and fish. Most fisheswhich could be identified were myctophids, and may have beenacquired by storm-petrels at night. Fishes increased while P.coecusdeclined in importance as a food over the nestling season ofthe storm-petrels. The amount of oil also declined in the Leach'sstorm-petrel diet over the season. Since P.coecus contains muchoil, the seasonal decline of both P.coecus and oil was probablyassociated. Other less important foods were jellyfishes, copepods,isopods, euphausiids, shrimp, squid and octopus. Jellyfishes(Velella velella) only occurred in the diet of Leach's storm-petrels,which may relate to that storm-petrel feeding in open oceanwhere V. velella occurs most frequently. The most numerous copepodwas Neocalanus cristatus, which was only observed in the storm-petreldiet early on during the study period. An abyssal copepod, Bathycalanusbradyi, and the isopod Cirolana californiensis (tentativelyidentified, the latter only previously known from sea canyonsoff California) were surprising food items of storm-petrels.Most euphausiid prey consisted of Thysanoessa spinifera andEuphausia pacifica.     

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.     

Parental provisioning and predation risk in rhinoceros auklets (Cerorhinca monocerata): effects on nestling growth and fledging

We examined the effects of predation risk on the behavior ofrhinoceros auklets (Cerorhinca monocerata) breeding at PineIsland, British Columbia, in 1990. Provisioning parents in someareas of the colony risked predation by bald eagles (Haliacetusleucocephalus). Chicks in high and low predation risk areasof the colony hatched on approximately the same date, receivedsimilar amounts of food to 46 days of age, grew at the samerate, reached similar peak masses, and fledged at similar masses.However, chicks in high predation areas fledged at a youngerage than did chicks in low predation areas. These data are consistentwith the hypothesis that parents in high risk areas terminatedprovisioning several days before those in lower risk areas.Mass at fledging was inversely related to age at fledging inboth high and low risk areas. The regression line for the highrisk habitats lies below that from the low risk habitats, aspredicted by a model that examines optimal time of fledgingfrom the perspective of the parents. We conclude that risk ofpredation represents a significant cost of reproduction to somerhinoceros auklets and that individual auklets within the colonyvary their behavior according to predation risk.     

Spatial heterogeneity and genetic variation in the copepod Neocalanus cristatus along two transects in the North Pacific sampled by the Continuous Plankton Recorder

We present a macrogeographic study of spatial heterogeneityin an important subarctic Pacific copepod and describe the firstgenetic analysis of population structure using Continuous PlanktonRecorder (CPR) samples. Samples of Neocalanus cristatus werecollected at a constant depth of 7 m from two CPR tow-routes,(i) an east–west 6500-km transect from Vancouver Island,Canada to Hokkaido Island, Japan, and (ii) a north–southtransect of 2250 km from Anchorage, Alaska to Tacoma, Washington.Analysis of these samples revealed three features of the biologyof N. cristatus. First, N. cristatus undergoes small-scale dielvertical migration that is larger among stages CV–adult(3–6 times more abundant at 7 m at night), than stagesCI–CIV (only 2–4 times higher at night). Secondly,while there were no regions where N. cristatus did not appear,each transect sampled a few large-scale macrogeographic patches.Thirdly, an analysis of molecular variation, using a partialsequence of the N. cristatus cytochrome oxidase I gene, revealedthat 7.3% (P < 0.0001) of the total genetic variation amongN. cristatus sampled from macrogeographic patches by the CPRcould be explained by spatial heterogeneity. We suggest thatspatial heterogeneity at macrogeographic scales may be importantin plankton evolution.     

The impacts of introduced House Mice on the breeding success of nesting seabirds on Gough Island

Invasive species are the main threat to island biodiversity; seabirds are particularly vulnerable and are one of the most threatened groups of birds. Gough Island, a UNESCO World Heritage Site in the South Atlantic Ocean, is an Important Bird and Biodiversity Area, and one of the most important seabird colonies globally. Invasive House Mice Mus musculus depredate eggs and chicks of most seabird species on the island, but the extent of their impact has not been quantified. We used field data and bootstrapped normal distributions to estimate breeding success and the number of surviving chicks for 10 seabird species on Gough Island, and compared estimates with those of analogous species from predator‐free islands. We examined the effects of season and nest‐site location on the breeding success of populations on Gough Island, predicting that the breeding success of Gough birds would be lower than that of analogues, particularly among small burrow‐nesting species. We also predicted that winter‐breeding species would exhibit lower breeding success than summer‐breeding species, because mice have fewer alternative food sources in winter; and below‐ground nesters would have lower breeding success than surface nesters, as below‐ground species are smaller so their chicks are easier prey for mice. We did indeed find that seabirds on Gough Island had low breeding success compared with analogues, losing an estimated 1 739 000 (1 467 000–2 116 000) eggs/chicks annually. Seven of the 10 focal species on Gough Island had particularly high chick mortality and may have been subject to intense mouse predation. Below‐ground and winter breeders had lower breeding success than surface‐ and summer‐breeders. MacGillivray's Prion Pachyptila macgillivrayi, Atlantic Petrel Pterodroma incerta and Tristan Albatross Diomedea dabbenena are endemic or near‐endemic to Gough Island and are likely to be driven to extinction if invasive mice are not removed.     

Chick‐provisioning behaviour of two shearwaters breeding in south‐eastern Australia

The chick‐provisioning behaviour of the short‐tailed shearwater Puffinus tenuirostris and the wedge‐tailed shearwater Puffinus pacificus was investigated in a mixed colony on Montague Island, New South Wales, Australia, over two breeding seasons. This colony is located at opposite edges of the breeding distribution of the two species. Frequent weighing techniques were used to determine chick feeding frequency, feed timing, meal size, chick weight loss and indices of food conversion efficiency of the chicks. Short‐tailed shearwater parents fed their chicks larger more infrequent meals than wedge‐tailed shearwater parents. Short‐tailed shearwater chicks demonstrated higher food conversion efficiencies and lower weight loss than wedge‐tailed shearwater chicks, indicating either differences in diet or metabolic rates. The feeding frequency in wedge‐tailed shearwaters also fluctuated more widely than for short‐tailed shearwaters over the two breeding seasons. Despite the fact that the timing of the breeding cycle on Montague Island is almost identical for the two species, these differences in chick provisioning are probably a result of differences in prey type and location, so they may help explain variations in annual breeding success and limits to the distribution of the two species.     

Life history strategies of Calanus sinicus in the southern Yellow Sea in summer

Ecological and physiological features of the planktonic copepodCalanus sinicus in the southern Yellow Sea in summer were studiedto reveal its life history strategy. From the coastal shallowwaters to the central part of the southern Yellow Sea, a shiftof the stage composition occurs from being dominated by theegg-nauplius stage to being dominated by the fifth copepodite(CV) stage. Most CVs reside in the Yellow Sea Cold Water Mass(YSCWM), where both temperature and food abundance are low.CVs in the YSCWM have longer body lengths, heavier body weightsand higher carbon contents than those outside the YSCWM. Onboardincubations show that the development of CVs in the YSCWM issuspended. Energy conservation, development suspension and lackof diel vertical migration (DVM) behavior suggest a diapausestatus for the CVs in the YSCWM, although vertical distributionpatterns indicate the CV individuals are not fully synchronousin physiology and development. This adaptive oversummering strategywould help C. sinicus to live through the warm and food-limitedsummer in the central part of the southern Yellow Sea; bothlow temperature and low food supply are necessary for CV tomaintain the resting state in the YSCWM. Calanus sinicus exhibitsdifferent life history strategies in different regions of thesouthern Yellow Sea in summer.     

Distribution of Calanus species in Kongsfjorden, a glacial fjord in Svalbard

The distribution of Calanus species was investigated in Kongsfjordenin summer of 1996 and 1997. In both years Calanus finmarchicusand Calanus glacialis dominated, although the boreal C. finmarchicuswas more abundant than the Arctic C. glacialis in 1997. Thiscoincided with a 2°C higher water temperature at 50 m in1997, indicating stronger influence of Atlantic origin waterthat year. Advected Calanus finmarchicus occurred in deep andsubsurface layers of the outer fjord in 1996 (200 ind. m^-3,mainly CIII). A less abundant local population aggregated insurface layers of the inner fjord (100 ind. m^-3). Similarly,advected C. finmarchicus occurred in subsurface layers in 1997(446 ind. m^-3, mainly CIII and CIV) and a local population insurface layers (183 ind. m^-3, mainly CI). Calanus glacialisin 1996 aggregated as CII and CIII in the deep layers of theouter fjord (272 ind. m^-3), whereas CIII–CV were abundant(216 ind. m^-3) in cold surface waters of the inner fjord. In1997 C. glacialis (mostly CIII–CV) was more abundant inthe outer than in the inner part of the fjord (40 and 192 ind.m^-3, respectively). Within Kongsfjorden, Calanus finmarchicusneeds one year to complete its life cycle, whereas Calanus glacialisneeds two. Calanus hyperboreus seems to be an expatriate inthe fjord system.     

A Model for Evaluating Time Constraints on Short-term Reproductive Success in Altricial Birds

SYNOPSIS. We develop a simple single dimension model incorporatingthe time and energy commitments of breeding altricial birdsin an effort to understand the evolutionary constraints on parentalcare. We chose time as the dimension of preference, becauseit is a naturally bounded constraint, e.g., length of day orbreeding season. The utility of the model was evaluated by comparingsimulations of time allocation of various breeding scenariosfor the European starling (Sturnus vulgaris) to field data.The structure of the model may prove useful in determining theevolutionary constraints on parental care imposed by the developmentalpattern of chicks.     

Mass and date at departure affect the survival of Ancient Murrelet Synthliboramphus antiquus chicks after leaving the colony

I compared the timing of colony departure and body mass of 53 Ancient Murrelet Synthliboramphus antiquus chicks that were retrapped as adults in Haida Gwaii, British Columbia, with those of 3992 chicks not retrapped. If the probability of recapture is a measure of survival, survival was related to both mass and date. Chicks that left the colony at 26 g or less had a lower chance of survival than heavier chicks, and those that left after the median date of departure survived better in some years than those that left earlier. The effect of date was not related to a seasonal change in departure mass because chick mass declined with departure date. I suggest that, because of heavy adult mortality during breeding, the timing of breeding in the Ancient Murrelet is based on a compromise between the optimum dates for chick and adult survival. This is in contrast to evidence from other, nonprecocial, seabirds.     

Integrating conventional microscopy and molecular analysis to analyse the abundance and distribution of four Calanus congeners in the North Atlantic

Analysis of the demographic structure of Calanus species inthe North Atlantic presents particular difficulties due to theoverlapping spatial distributions of four main congeneric species(Calanus finmarchicus, Calanus helgolandicus, Calanus glacialisand Calanus hyperboreus). These species have similar morphologies,making microscopic discrimination only possible between someof the species at late copepodite or adult stages. However,molecular techniques now offer the possibility of screeningsignificant numbers of specimens and unambiguously identifyingthem to species, regardless of developmental stage. Unfortunately,the processing rate of specimens by molecular methods is stilltoo low to offer a realistic alternative to microscopy for analysisof samples from large field surveys. Here, we outline and testan approach involving the use of molecular methodology in conjunctionwith conventional microscopy to assess the species assignmentof developmental stage abundances of Calanus congeners. Ourstudy has highlighted many important methodological issues.First, it cannot be assumed that the species composition ishomogeneous across the development stages; applying proportionalspecies composition of adults to morphologically undistinguishableearlier development stages can result in error. The second importantconclusion is that prosome length may be a highly unreliablediscriminator of C. finmarchicus and C. glacialis.     

Molecular phylogeny of Neocalanus copepods in the subarctic Pacific Ocean, with notes on non-geographical genetic variations for Neocalanus cristatus

Monophylies of Neocalanus cristatus, Neocalanus plumchrus andNeocalanus flemingeri were revealed by nucleotide sequencesof mitochondrial 16S rRNA gene (410 bp) but not of nuclear 18SrDNA gene (1802 bp). Intraspecific variations in mitochondrial16S rRNA of N. cristatus collected from geographically distantregions were very low (<0.5%).     

The comparative breeding biology of Adelie and Chinstrap Penguins Pygoscelis adeliae and P. antarctica at Signy Island, South Orkney Islands

Adélie Penguins and Chinstrap Penguins breed in abundance at Signy Island, South Orkney Islands, which is near the northern breeding limit of the former but in the centre of the latter's breeding range. During the 1980–81 and 1981–82 breeding seasons daily checks were made to record laying date and interval between laying, egg weight and volume, incubation period and shifts by each parent, hatching date and interval between hatching, the date on which the young first creched and their age at the time, and fledging date and age. This was done for the eggs and chicks from 60 marked nests of each species. Adelies started breeding about one month before Chinstraps and showed other adaptations which may relate to early breeding and probably to the presence of ice around the breeding site. These adaptations included longer incubation shifts and a heavier and larger first egg which may promote more rapid brood reduction. Chinstraps had shorter incubation shifts and both eggs had similar weights and volumes. Although the later breeding of Chinstraps meant that chick rearing coincided with the availability of a better food source than that utilized by Adelies, Chinstraps were unable to cope with persistent heavy ice conditions in 1980–81 and were no more successful than Adelies in 1981–82. All previous breeding data for both species are reviewed and it is suggested that early breeding by Adelies may be a consequence of competition for food with Chinstraps but, by retaining adaptations to harsh environmental conditions, Adelies are well able to breed successfully. Conversely, Chinstrap Penguins appear mainly adapted to milder environmental conditions and it is unlikely that they could compete successfully with Adelie Penguins at breeding sites on the Antarctic Continent.     

Spatial and temporal variation in the diet of the Pacific sand lance Ammodytes hexapterus in waters off the coast of British Columbia,Canada

Variation in the diet of the Pacific sand lance Ammodytes hexapterus was examined in three years (2009–2011) at four sites in British Columbia, Canada. There were 12 major taxa of prey in diets, eight of which were Crustacea, with copepods being by far the dominant taxon in all 12 site‐years. Of the 22 copepod taxa recorded, only Calanus marshallae and Pseudocalanus spp. occurred in all collections, and these two calanoid species dominated diets in terms of frequency of occurrence and total numbers of prey (Pseudocalanus spp. in most collections), and total prey biomass (C. marshallae in all collections). Based on an index of relative importance, C. marshallae was the primary prey at the two southerly sampling sites (Pine and Triangle Islands) and Pseudocalanus spp. at the two northerly sites (Lucy Island and S'G ang Gwaay). Based on an index of dietary overlap, the species composition of the copepod component of A. hexapterus diets overlapped very strongly at the northerly and the southerly pairs of sites in both a cold‐water La Niña year (2009) and a warm‐water El Niño year (2010), but overall there was more homogeneity amongst all four sites in the La Niña year.     

Temperature, food and mate limitation of copepod reproductive rates: separating the effects of multiple hypotheses

Temperature, food and the availability of mates may all limitrates of egg production is freshwater diaptomids. We have usedthe reproductive phases of Diaptomus paliidus to develop anindex of food limitation (f) and an index of mate limitation(M), each of which responds independently of temperature. Theresponse of the food limitation index to high and low temperaturesand high and low food concentrations was examined in a two-wayfactorial experiment in the laboratory. The index was highlyresponsive to a change in food concentration, stable duringa change in temperature alone at the high food level and responsiveto the synergistic interaction of food and temperature effectswhen both factors were limiting. Laboratory data indicate thatthe f index should not be biased by mate limitation unless maledensities are 6.7 x 10^–l0 males 1^–1 or less. Themate limitation index was highly responsive to mate availability,and ranged from a low of 9–19 in the presence of abundantmales to 100 in tbe absence of males. Applications of the fand M indices to a natural population of D. paliidus over 6weeks indicated that both food and mate availability were limitingthe reproductive rates of the copepods during the sampling period. ¹Present address: Institute of Animal Resource Ecology, Universityof British Columbia, Vancouver, Canada V6T 1W5     

Consequences of foraging trip duration on provisioning behaviour and fledging condition of common murres Uria aalgae

We examine the provisioning constraints of a pursuit‐diving seabird in a cold ocean regime by comparing the behaviour of common murres Uria aalge rearing chicks at two colonies in the Northwest Atlantic during 1998‐2000. Funk Island is the largest (340,000–400,000 breeding pairs) and most offshore (60 km) colony of common murres in eastern Canada. Seventy‐five percent of the Northwest Atlantic population of common murres breeds on this island. Great Island is one island within the Witless Bay Ecological Reserve, which is the second largest breeding aggregation (100 000 breeding pairs) and is located near‐shore (2 km). The primary forage fish species in Newfoundland waters is capelin Mallotus villosus, which spawns on or near coastal beaches during summer. Therefore, the two study colonies differ in their distance to food resources and colony size. It is within this natural context that we compare: (1) prey types and frequency of delivery (amount of prey), (2) parental time budgets, and (3) the mass and condition (mass/wing length) of fledglings at both colonies. Similarly sized female capelin (100–150 mm) were delivered to chicks at both colonies. Foraging time per day per parent, a proxy of foraging effort, was similar at both colonies (Great Island: 5.1 h; Funk Island: 5.5 h), as was the percentage of time spent with mates (Great Island: 12.3%; Funk Island: 10.9%). Foraging trips, however, were longer at Funk Island (4.1 h) than at Great Island (2.9 h). This resulted in lower feeding rates of chicks (0.17 feeds per h) and poorer condition of fledglings (2.9 g/mm) at Funk Island compared to those at Great Island (0.22 feeds per h; 3.9 g/mm). We hypothesize that provisioning efforts are constrained at Funk Island by (1) distant food resources and increased competitor density, resulting in longer foraging trip durations and (2) the time spent paired with mates at the colony, which may reflect a minimum time required to maintain breeding sites due to higher breeding densities at Funk Island compared to Great Island. Demographic consequences of this poor fledgling condition at Funk Island are unknown, but fledglings may sufficiently accelerate growth at sea due to their closer proximity to an important nursery area. If fledgling survival is compromised, however, the lower potential for growth at Funk Island will impact the entire Northwest Atlantic population of common murres.     

Chick diet of dovekies Alle alle in Northwest Greenland

This paper reports the results of an investigation of dovekies (Alle alle) breeding at Hakluyt Island in the Thule District, Northwest Greenland. The paper presents a comprehensive examination of the chick diet of dovekies in the species' most important breeding area. Food delivered to nestlings consisted primarily of copepods Calanus hyperboreus and C. glacialis in the copepodite stages IV and V. Themisto libellula, other amphipods, decapods and fish larvae in small numbers were found in the chick diet. The zooplankton in the chick diet is presented both in terms of numbers and dry weight. The characteristic copepod of high-arctic waters, C. hyperboreus, was found to be the single most important species in the chick diet, contributing approximately 54% of the dry food weight. The food samples revealed low diversity, and the three species C. hyperboreus, C. glacialis and T. libellula made up 91.5% of the total zooplankton dry weight. The results from this paper are compared with previous dovekie-chick diet studies. Accepted: 14 June 2000     

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.     

COOPERATIVE BREEDING OF THE NORTHWESTERN CROW CORVUS CAURINUS IN BRITISH COLUMBIA

The behaviour of helpers at nests of Northwestern Crows was studied on Mandarte Island and Mitlenatch Island, British Columbia. Not all nests had a helper and there was only one helper per nest. Helpers participated in varying degrees in the defence of the territory and nest, feeding of the nestlings and fledglings and they cached food on the territory. Adult males fed helpers, and helpers obtained most of their food on the adults' territory. Adults with helpers laid larger clutches and produced more fledglings per nest than adults without helpers. It is suggested that cooperative breeding in the Northwestern Crow is of recent origin.