Haematology and metabolism of the blood of the little penguin, Eudyptula minor

1. We studied the haematology of the little penguin, Eudyptula minor, and measured red cell enzymes, and the rate of O2 and glucose consumption and CO2 and lactate production of whole penguin blood. 2. Little penguins have a very low RBC count (1.66 x 10(12) l-1) and very large red cells (MCV = 229 microns 3), resulting in a normal haemoglobin and haematocrit. 3. Despite the lower body temperature of penguins, the rate of O2 consumption by whole blood is close to that predicted from studies on other birds.     

Foraging behaviour of the little penguin, Eudyptula minor: initial results and assessment of instrument effect

We investigated the foraging behaviour of little penguins using a new archival electronic activity recorder which simultaneously measures speed and depth against time. We present the first integrated data of foraging behaviour of two little penguins, from which we were able to distinguish between several types of travelling and foraging behaviours. The little penguins foraged mainly within 15 m of the surface and travelled at speeds between 8 and 9 km h^-1. Using attachments ranging between 1.4 and 11.8% penguin cross-sectional area (0.1 and 6% penguin mass), and isotopic water, we also assessed the effects of carrying devices while foraging. Both water influx and metabolic rates were significantly lower in penguins carrying devices, compared to penguins foraging without devices attached. Even the relatively small attachments resulted in a decreased foraging efficiency and we suggest that there is no simple or fixed relationship between size of device and the effect on the bearer.     

Comparison of barometric and pneumotachographic measurements of resting ventilation in the little penguin (Eudyptula minor)

1. We compared the use of the barometric method and pneumotachography for measurement of ventilation in the little penguin Eudyptula minor. 2. Simultaneous use of both techniques showed the barometric method to give a reliable estimate of tidal volume. 3. Comparison of birds with and without masks for pneumotachography showed wearing masks to produce a significant increase in ventilation, principally through a raised respiratory frequency.     

Internal organ weights of the blue penguin Eudyptula minor

Abstract

This work presents a database of weights of normal internal organs for the blue penguin, Eudyptula minor. During a necropsy study of 213 adult and fledged immature blue penguins, external body measurements and internal organ weights were recorded. The relationship between organ weights and body weights shows that the liver and abdominal fat pad are most closely correlated with the body mass index, a quantitative indication of adequacy of condition.     

Isolation and characterization of polymorphic microsatellite loci for the study of paternity and population structure in the little penguin Eudyptula minor

We isolated and characterized eight novel microsatellite loci in the little penguin Eudyptula minor, using nonradioactive polymerase chain reaction‐based techniques to screen GA and GAAA repeats from enriched genomic DNA libraries. All eight loci were polymorphic and seven were variable in our main study population (mean H_E = 0.613, mean N_A = 7.14). Cross‐amplification using a microsatellite primer developed in Spheniscus demersus (African penguin) yielded one additional polymorphic locus. This locus combined with six of the little penguin loci is suitable for paternity assignment in little penguins (exclusion probability for seven unlinked loci = 0.993).     

Diving behaviour and energetics in breeding little penguins (Eudyptula minor)

We present data on the diving behaviour and the energetics of breeding little penguins in Tasmania, Australia. Using an 18 m long still water canal in conjunction with respirometry, we determined the energy requirements while diving. Using electronic devices measuring dive depth or swimming speed, we investigated the foraging behaviour at sea. Cost of Transport was calculated to be minimal at the speed the birds prefer at sea (1.8 m/s) and averaged 11.1 J/kg/m (power requirements at that speed: 20.0 W/kg). Metabolic rate of little penguins resting in water was found to be 8.5 W/kg. The externally-attached devices had no significant influence on the energy expenditure.
Foraging trips can be divided into four distinct phases with different diving behaviours. A mean of 500 dives was executed per foraging trip lasting about 18 hours with 60% of this time being spent swimming. The total distance travelled averaged 73 km per day, although foraging range was about 12km. Mean swimming speed of little penguins at sea was 1.8 m/s, maximum swimming speed was 3.3 m/s. More than 50% of all dives had maxima not exceeding 2 m. Maximum depth reached was 27 m. Mean dive duration was 21 s. There were inter-sex differences in diving behaviour as well as changes in foraging behaviour over the breeding period. Aerobic dive limits (ADL) in the wild were estimated between 42 and 50 s. From the swim canal experiments we derived an ADL of 44 s. Total oxygen stores were calculated to be 45 ml O₂/kg. Only 2% of all dives exceeded the ADL. FMRs at sea were calculated to be between 1280 and 1500 kJ/kg/d according to chick size. The yearly food requirements of a breeding little penguin amount to 114 kg.     

Diet segregation between two colonies of little penguins Eudyptula minor in southeast Australia

We studied foraging segregation between two different sized colonies of little penguins Eudyptula minor with overlapping foraging areas in pre‐laying and incubation. We used stomach contents and stable isotope measurements of nitrogen (δ¹⁵N) and carbon (δ¹³C) in blood to examine differences in trophic position, prey‐size and nutritional values between the two colonies. Diet of little penguins at St Kilda (small colony) relied heavily on anchovy while at Phillip Island (large colony), the diet was more diverse and anchovies were larger than those consumed by St Kilda penguins. Higher δ¹⁵N values at St Kilda, differences in δ¹³C values and the prey composition provided further evidence of diet segregation between colonies. Penguins from each colony took anchovies from different cohorts and probably different stocks, although these sites are only 70 km apart. Differences in diet were not reflected in protein levels in the blood of penguins, suggesting that variation in prey between colonies was not related to differences in nutritional value of the diet. Anchovy is currently the only available prey to penguins throughout the year and its absence could have a negative impact on penguin food supply, particularly at St Kilda where the diet is dominated by this species. While it is difficult to establish whether diet segregation is caused by inter‐ or intra‐colony competition or spatial differences in foraging areas, we have shown that colonies with broadly overlapping foraging ranges could have significant differences in trophic position, diet composition and prey size while maintaining a diet of similar nutritional value.     

Blue penguin (Eudyptula minor) nest distribution and breeding success on Otago Peninsula, 1992 to 1998

Abstract

Annual counts of nests with eggs or chicks (known nests) were made at blue penguin (Eudyptula minor) breeding sites on the Otago Peninsula in each November from 1994 to 1997. Although the population has doubled to an estimated 600 known nests over this period, the number of breeding sites on the Otago Peninsula has reduced since the 1970s. Breeding success at three areas at Taiaroa Head were monitored by regular nest checks in the breeding season from 1992 to 1998. At Taiaroa Head reproductive success ranged from 41 to 78% at the three sites during the seven‐year study and was generally higher for pairs nesting in nest boxes than for those in burrows. The percentage of breeding pairs that laid a second clutch after fledging at least one chick from their first clutch (double brooded) varied between seasons (0–48%) and was correlated with the date of the onset of breeding. Egg loss, possibly through predation by Norway rats (Rattus norvegicus), influenced the significantly lower reproductive success at one area (Area A) at Taiaroa Head during the 1996 season.     

Acoustic properties of male advertisement and their impact on female responsiveness in little penguins Eudyptula minor

We examined the relationship between acoustic properties of male advertising calls, male body size and female responses in little penguins Eudyptula minor . Larger males produced calls of lower frequency. Playback experiments indicated that females were more likely to respond to low or medium pitched calls, than to high ones (although only 28% of females responded vocally to playback). This may reflect the interest of unmated females in large to medium sized senders as potential mates, or indicate a stronger territorial response to large intruders.     

Stable isotope analysis as a tool to monitor dietary trends in little penguins Eudyptula minor

Long‐term dietary monitoring of seabirds can be used to relate population fluctuations to at‐sea events. Stomach flushing is a conventional dietary monitoring technique, but has a number of disadvantages. Stable isotope analysis (SIA) is a less invasive method that provides unbiased dietary information over a longer period. We evaluated stable isotope analysis as a potential tool for monitoring long‐term little penguin Eudyptula minor diet. We determined diet composition during the chick feeding stage using stomach flushing and SIA at three separate colonies, using spatial variation in diet as a surrogate for potential temporal variation. Bayesian isotopic mixing models were generated for blood and feathers to evaluate their ability to discriminate broad‐scale (fish, squid, crustaceans) and fine‐scale (individual prey species) diet composition. Differences in stable carbon and nitrogen isotope ratios were found between colonies: broad‐scale isotopic mixing models predicted different proportional contributions of broad taxa (fish, cephalopod, crustacean) to diet than was indicated by stomach samples, reflecting the bias incurred by one‐off stomach contents analysis. Fine‐scale isotopic mixing models predicted proportional contributions of prey items with less certainty. Blood isotopic mixing models had narrower confidence intervals than models for feathers, but trends in δ¹⁵N for feathers mirrored those for blood. Our results suggest that relying on stomach contents analysis to detect shifts in prey consumption in little penguins could be very misleading, resulting in a less‐than‐complete idea of total prey consumption. SIA of little penguin tissues could be used to monitor dietary shifts across dissimilar taxa that may affect population numbers, but would fail to detect shifts between fish species.     

Reproductive performance of little penguins Eudyptula minor in relation to year, age, pair-bond duration, breeding date and individual quality

We measured breeding performance of little penguins Eudyptula minor at Phillip Island, Victoria, Australia, during a 21-year period. All birds considered in this paper (n=307) were of known age (2–22 y) and sex, and most were of known, or closely-estimated, pair-bond status (1–8 mates per bird; pair-bond durations 1–13 y). Breeding dates and breeding performance varied markedly from year to year; measures of annual performance were not associated with early breeding. Measures of individual breeding performance (clutch-size, hatching success, chick masses and productivity) were related to early laying, parental age, and duration of pair-bond. Dependence of breeding performance on parental age was curvilinear, levelling off at about 8 y of age. Productivity declined significantly among birds older than 8 y; this decline was not due to events in the last year of breeding ("terminal illness"). Breeding performance increased with duration of pair-bond at least through y 5. Early breeding was significantly related to age and duration of pair-bond. Most of these relationships were stronger among males than among females, and many of them were not significant when females were considered alone. After controlling for other factors, breeding performance varied significantly among birds, but autocorrelations were low and limited to intervals of one year. Parental quality (defined for birds studied in six or more years as the individual bird term in a GLM for productivity controlling for other factors) was not correlated with lifespan or other demographic parameters, but high-quality birds were less prone to change mates and burrows than low-quality birds. We know of no previous study in which simultaneous effects of laying date, age and pair-bond duration on breeding performance were measured, while controlling for year, individual quality and terminal illness.     

Viral infections of little blue penguins (Eudyptula minor) along the southern coast of Australia

Serum antibodies to strains of avian paramyxovirus and flavivirus were detected in little blue penguins sampled at Port Campbell and Phillip Island, Victoria, Australia. No antibody to Newcastle disease virus (NDV) was detected in 267 sera collected, although one penguin captured for experimental studies had a hemagglutination-inhibition antibody titer of 2(4) to this virus. Experimental studies showed that the avian paramyxovirus designated APMV-IM and strain V4 of NDV were non-pathogenic for penguins, although the penguins could have been previously infected with these or similar virus strains. A flavivirus designated Saumarez reef virus, and an unnamed virus isolated from ticks on Macquarie Island, Southern Ocean were pathogenic causing disease and mortality in penguins inoculated with the viruses.     

Acoustic convergence between two nocturnal burrowing seabirds: experiments with a penguin Eudyptula minor and a shearwater Puffinus tenuirostris

The evolution of acoustic signals is influenced by environmental constraints. We studied two sympatric but unrelated seabirds: the Little Penguin Eudyptula minor and Short-tailed Shearwater Puffinus tenuirostris , to examine the degree to which similarities in their ecology had led to convergence in their calls. Both species nest in burrows in Southern Australia and, at night, are highly vocal and territorial. First, we analysed the physical characteristics of the territorial call. Secondly, we studied the transmission of calls through burrows and varying distances through vegetation. Thirdly, we used playback experiments of natural signals to demonstrate that the response disappears between 4–8 m, and of modified signals to understand the coding-decoding process linked to the territorial function of the call. The structure of the territorial calls of the two species clearly differs, but both species produce a succession of gaps in amplitude and frequency, and a high degree of redundancy. Our experiments show that, to decode the territorial message, birds pay attention only to parameters that are less degraded during propagation and ignore fine details of structure that are quickly degraded, even at relatively short distances (< 8 m). In both species, territorial information is mainly conveyed by the rhythmic succession of two sounds (syllables or subsyllables), birds paying attention to the FM structure of these successive sounds but not to the AM. This convergent coding is adaptive in that it reduces the possibility that the meaning may be distorted by interference from noise and acoustic screening.     

Increased survival and breeding performance of double breeders in little penguins Eudyptula minor,New Zealand: evidence for individual bird quality?

The little penguin Eudyptula minor is unique among penguin species in being able to fledge chicks from two clutches in one breeding season. Pairs laying two clutches in a given season make a higher reproductive investment, and may be rewarded by a higher reproductive success as they may raise twice as many chicks as pairs laying one clutch. The higher effort made by pairs laying two clutches could correlate negatively with survival, future reproductive performance or offspring survival, indicating a cost of reproduction. Conversely, a positive relationship between the number of clutches produced in a given breeding season and survival, future reproductive performance or offspring survival would indicate that birds laying two clutches belonged to a category of birds with higher fitness, compared to birds laying only one clutch in the season. In this study we used a long‐term data set taken from an increasing population of little penguins in Otago, SE New Zealand. We modelled the relationship between the number of clutches laid in a breeding season and survival probability, reproductive performance in the next breeding season and first year survival of offspring using capture‐recapture modelling.
Birds laying two clutches produced 1.7 times more fledglings during a breeding season than pairs laying one clutch. We found that birds laying two clutches had a higher probability of breeding in the following breeding season, a higher probability of laying two clutches in the following breeding season and a higher survival probability. There was no overall difference in post‐fledging survival between the young of birds producing one clutch and the young of birds producing two clutches. However, the survival of young of single clutch breeders declined with laying date, whereas the young of double clutch breeders had the same survival rate irrespective of laying date. For a subset of data with birds of known age, we found evidence that the probability of laying two clutches increased with age. However, there were also indications for differences among birds in the tendency to lay two clutches that could not be attributed to age. We tentatively interpret our results as evidence of quality difference among little penguin breeders.     

Few genetic differences between Victorian and Western Australian blue penguins,Eudyptula minor

Abstract

Blue penguins, Eudyptula minor, breeding on Penguin Island, Western Australia are considerably larger than other blue penguins in Australia. If genetic isolation is the cause, it may have implications for the conservation status of some blue penguin populations. We compared the sequences of two mitochondrial gene regions (cytochrome‐b and the control region) from Western Australian blue penguins with other populations of blue penguins from Australia and New Zealand. We found few differences between sequences from Western Australia, Phillip Island, Victoria and Otago, New Zealand, although all three differed considerably from other New Zealand blue penguins. Sequences for the control region from the Western Australian blue penguins and 30 more birds breeding at various Australasian sites provided further support for two major clades within Eudyptula; an Australian clade (including Otago) and a New Zealand clade.     

Burrow and mate fidelity in the Little Penguin Eudyptula minor at Lion Island, New South Wales, Australia

Although both mate and nest fidelity are thought to exist in Little Penguins Eudyptula minor , few studies have statistically assessed the relationship between both types of fidelity and breeding success. Observations of the breeding biology of the Little Penguin were carried out between 1990 and 1998 at Lion Island, New South Wales, Australia. The existence of fidelity and the effect of breeding success on subsequent fidelity were assessed for this population. Strong evidence for the existence of nest and mate fidelity was found with high rates of return to the same burrow and mate being recorded for both sexes. For all years of the study, the average number of female and male birds faithful to their nest was 76 and 79%, respectively, with only two noted cases of intraseasonal burrow swapping. The rate of divorce varied widely between years, with a range of 0–40%. A marginal association of a log linear model showed a highly significant relationship ( P  < 0.001) between success of a clutch in one year and fidelity to a mate in the following year. Consequently, parents successfully raising a fledgling in one year were more likely to reunite in the following year than were pairs that attempted unsuccessfully to breed. Pairs failing to fledge chicks were much more likely to separate owing to the absence of one parent in the following year. We were unable to detect any association between breeding success in one year and subsequent change of nest in the next.     

Influence of biological and ecological factors on hematological values in wild Little Penguins, Eudyptula minor

Clinical hematology and biochemistry are recognized as useful aids in health diagnosis in birds. The Little Penguin (Eudyptula minor) is endemic to Australia and numbers are declining due to a number of factors including declining health. This 2-year study on both Bowen and Lion Island populations aimed at assessing the potential causes of their declining health. Blood was collected from a total of 294 adult Little Penguins. R.B.C., Hb, P.C.V., M.C.V., M.C.H., M.C.H.C., W.B.C. and differential counts, thrombocytes and T.P.P. were measured. Multiple regression was used to identify relationships between hematological values and the following predictor variables: site, season, sex, time held prior to sampling and body condition. Values obtained showed some differences compared to the values for other penguin species reported in the literature. P.C.V., R.B.C., Hb, M.C.H.C. and T.P.P. showed major differences in comparison with a previous study on little penguins. Considering the influence of predictor variables, it appeared that: P.C.V., R.B.C., M.C.V., M.C.H., M.C.H.C. varied according to seasons; P.C.V., R.B.C., M.C.H., T.P.P. were correlated to body condition; P.C.V. was higher in birds on Bowen Island compared to birds on Lion Island; and T.P.P. was higher in females. These ecological and biological predictor variables have a strong influence upon hematological values. As a consequence, they must be taken into consideration when interpreting results of future studies.     

Survival analysis of Little Penguin Eudyptula minor chicks on Motuara Island, New Zealand

Chick survival of Little Penguins Eudyptula minor was studied on predator-free Motuara Island, Cook Strait, New Zealand (41̀05'S, 174̀15'E), in 1995 and 1996. We used the Kaplan-Meier estimator and robust Cox regression to estimate chick survival rate (pL se) at 0.325 pL 0.044, leading to an estimated survival from laying to fledging of 0.13 or a reproductive output of 0.26 chicks per pair and breeding attempt. Starvation posed the greatest mortality risk, followed by unknown factors and rain. Risk of death due to rain was restricted to the guard stage, whereas starvation occurred throughout the nesting period, though with a peak in the early guard stage. Significant seasonal differences in survival rate were detected in both years, but with reversed trends, survival decreasing with the season in 1995 and increasing in 1996. Failure of adults to relieve their partner on the nest after chicks hatched accounted for 16% mortality or 34% of all chick deaths. Differences in chick survival rate between nest types were significant in 1995, a year with high rainfall, but not in 1996. Nests in the base of hollow trees had the highest chick survival rate. Of chicks in open nests - a nest type that is unusual for this species - 5.4% fledged. Our results suggest that on Motuara Island good breeding sites are scarce and that the food supply has been poor during the years of this study.