Synchronized group association in little penguins, Eudyptula Minor.

Most seabirds form groups on land and at sea, but there is little information whether seabird groups are formed deliberately or randomly. We investigated whether little penguins formed groups composed of the same individuals when they crossed the beach each day over four breeding seasons (2001–2004) using an automated penguin monitoring system (APMS). We used an association matrix to determine the number of times any two birds crossed the APMS in the same group. The number of these group associations or ‘synchronized parade’ behaviour was determined for every possible pair of individuals, giving a total association value for each pair of birds during the postguard stage of the reproductive cycle. We concluded that a penguin group was composed of 5–10 individuals within 40-s intervals. Penguin groups were formed nonrandomly in years of high breeding success (2002 and 2003), but not in years of low breeding success (2001 and 2004). Age of birds was a significant factor in composition of groups. Little penguins with higher association values shared similar characteristics or ‘quality’, which in turn may increase the functional efficiency of their groups, especially if they are also foraging together. However, low association indices indicated that seeking the same associates was not a priority. It is costly for any animal to synchronize their attendance with the same individuals, so it could be beneficial to display synchronized parade behaviour in good breeding years but it could result in intraspecific competition for food during poor breeding years.

Time-budget and feeding behaviour of the squirrel glider (petaurus norfolcensis) in remnant linear habitat

The squirrel glider (Petaurus norfolcensis) occurs in forests and woodlands in eastern Australia. In Victoria it is now largely restricted to small, fragmented areas and is considered endangered. In this study, the time-budget, feeding behaviour and related habitat use of the squirrel glider were investigated in a linear remnant of roadside vegetation near Euroa, Victoria. Timed observations of three males and three females, fitted with radio-collars, were made in each of four seasons. Gliders were observed for a total of 53.2 h, during which they devoted 72% of time to foraging activities. Grooming accounted for 16% of observation time. Exudates associated with homopterous insects were the primary food items consumed throughout the year. Arthropods, nectar and pollen, and Acacia gum formed the remainder of the diet. The proportion of time devoted to harvesting these food items showed marked seasonal variation. The primary dependence on homopterous insect exudates in this study area contrasts with other investigations at sites of greater floristic diversity where nectar and pollen were the most important dietary resources. This highlights the need to obtain ecological information from the range of habitats occupied by a species. Large trees are a vital habitat component of remnant linear vegetation in this study area, providing gliders with critical foraging resources. Retention of such trees is essential for the longevity of glider populations.

Characterisation of proteins in the milk of fur seals

Milk protein composition was investigated throughout the lactation periods of the Australian fur seal (Arctocephalus pusillus doriferus) and Antarctic fur seal (Arctocephalus gazella). The mean protein content of the milk was found to be 10.9% and 10.6% respectively. The concentration of total protein did not change during lactation, although a decline in casein content of the milk in late lactation was apparent. Milk protein concentration during a foraging/suckling cycle of the Antarctic fur seal analysed at the time of arrival on shore, and 24 h and 72 h after arrival was 12.8%, 11.4% and 12.5% respectively. Re-feeding animals at 72 h resulted in a significant increase in milk protein content to 14.9%. Characterisation of milk protein by SDS-PAGE analysis revealed 5 casein and 10 major whey protein bands. Amino-terminal sequencing indicated that the majority of the whey fraction of the milk is β-lactoglobulin (β-LG). The limited amino acid sequence indicated 3 different β-LGs were secreted in the milk. Subsequently, RT-PCR was used to extend the sequence of one of the β-LGs and translation of the 464 bp fragment indicated that it shared 79% sequence identity with feline β-LG II.

The lactation cycle of the fur seal

The fur seal is a mammal with an unusual ability to turn its milk production on and off without significantly altering the gross morphology of the mammary gland. This atypical lactation cycle is due to the fact that maternal foraging and infant nursing are spatially and temporally separate (Bonner, 1984). Maternal care involves the suckling of offspring over a period of at least 4 months, but lactation can extend to more than 12 months. Following a perinatal fast of approximately 1 week, females depart the breeding colony to forage at sea and, for the remainder of lactation, alternate between short periods ashore suckling their young with longer periods of up to 4 weeks foraging at sea. Whilst foraging at sea, milk production in the fur seal mammary gland either ceases or is reduced (Arnould & Boyd, 1995b).

Fur seal adaptations to lactation: insights into mammary gland function

The fur seal (Arctocephalus spp. and Callorhinus spp., members of the pinniped family) is a mammal with the unusual capability to modulate its lactation cycle by turning milk production on and off without the typical mammalian regression and involution of the mammary gland. Lactation has evolved from constraints arising from the spatial and temporal separation of infant nursing and maternal foraging as the mother gives birth and feeds the pup on land while acquisition of nutrients for milk production occurs at sea. The lactation cycle begins with the female fur seal undergoing a perinatal fast of approximately 1 wk, after which time she departs the breeding colony to forage at sea. For the remainder of the long lactation period (116–540 days), the mother alternates between short periods ashore suckling the young with longer periods of up to 4 wk of foraging at sea. Milk production continues while foraging at sea, but at less than 20% the rate of production on land. Fur seals produce one of the richest milk reported, with a very high lipid content contributing up to 85% of total energy. This feature serves as an adaptation to the young's need to produce an insulating blubber layer against heat loss and to serve as an energy store when the mother is away foraging at sea. This atypical pattern of lactation means mothers have long periods with no suckling stimulus and can transfer high-energy milk rapidly while on land to minimize time away from foraging grounds. The absence of suckling stimulus and milk removal during foraging does not result in the onset of involution with associated apoptosis of mammary secretory cells and a subsequent progressive breakdown of the cellular structure of the mammary gland. The mechanisms controlling lactation in the fur seal mammary gland have been investigated using molecular and cellular techniques. These findings have shed light on the processes by which the unique features of lactation in the fur seal are regulated.

Individual variation in the pup attraction call produced by female Australian fur seals during early lactation

Otariid seals (fur seals and sea lions) are colonial breeders with large numbers of females giving birth on land during a synchronous breeding period. Once pups are born, females alternate between feeding their young ashore and foraging at sea. Upon return, both mother and pup must relocate each other and it is thought to be primarily facilitated by vocal recognition. Vocalizations of thirteen female Australian fur seals (Arctocephalus pusillus doriferus) were recorded during the breeding seasons of December 2000 and 2001, when pups are aged from newborns to one month. The pup attraction call was examined to determine whether females produce individually distinct calls which could be used by pups as a basis for vocal recognition. Potential for individual coding, discriminant function analysis (DFA), and classification and regression tree analysis were used to determine which call features were important in separating individuals. Using the results from all three analyses: F0, MIN F and DUR were considered important in separating individuals. In 76% of cases, the PAC was classified to the correct caller, using DFA, suggesting that there is sufficient stereotypy within individual calls, and sufficient variation between them, to enable vocal recognition by pups of this species.

Species-specific cell-matrix interactions are essential for differentiation of alveoli like structures and milk gene expression in primary mammary cells of the Cape fur seal (Arctocephalus pusillus pusillus)

Few models are in place for analysis of extreme lactation patterns such as that of the fur seals which are capable of extended down regulation of milk production in the absence of involution. During a 10–12 month lactation period, female fur seals suckle pups on shore for 2–3 days, and then undertake long foraging trips at sea for up to 28 days, resulting in the longest intersuckling bouts recorded. During this time the mammary gland down regulates milk production. We have induced Cape fur seal (Arctocephalus pusillus pusillus) mammary cells in vitro to form mammospheres up to 900 μm in diameter, larger than any of their mammalian counterparts. Mammosphere lumens were shown to form via apoptosis and cells comprising the cellular boundary stained vimentin positive. The Cape fur seal GAPDH gene was cloned and used in RT-PCR as a normalization tool to examine comparative expression of milk protein genes (αS2-casein, β-lactoglobulin and lysozyme C) which were prolactin responsive. Cape fur seal mammary cells were found to be unique; they did not require Matrigel for rapid mammosphere formation and instead deposited their own matrix within 2 days of culture. When grown on Matrigel, cells exhibited branching/stellate morphogenesis highlighting the species-specific nature of cell–matrix interactions during morphological differentiation. Matrix produced in vitro by cells did not support formation of human breast cancer cell line, PMC42 mammospheres. This novel model system will help define the molecular pathways controlling the regulation of milk protein expression and species specific requirements of the extracellular matrix in the cape fur seal.

Variation in maternal provisioning by lactating Antarctic fur seals: response to experimental manipulation in pup demand

An experiment involving the supplementary feeding of pups was conducted on Antarctic fur seals to investigate the factors influencing maternal foraging-attendance cycles and the differential use of nutritional resources for growth, maintenance and storage by pups. For 40% of the lactation period, male pups were given a supplement mimicking the chemical composition of Antarctic fur seal milk at a dose equivalent to 35% of the normal mass-specific milk energy intake for the species. Milk consumption, body composition and growth rates were monitored during and after the supplementary feeding period and maternal foraging-attendance cycles were monitored throughout lactation. During the supplementary feeding period, treatment pups (n=8) grew 32% faster and deposited greater adipose tissue stores than controls (n=8) but consumed the same amount of maternal-delivered milk. When supplementary feeding was stopped (timed to coincide with peak maternal milk yield in this species), treatment pups lost mass whereas control group pups continued to grow. Treatment pups weaned at a younger age (109 days) than control pups (116 days) but at the same mass (13 kg). Maternal attendance durations did not differ between the treatment and control groups throughout lactation. However, mothers of treatment pups had significantly shorter foraging trip durations (3.74 days) than mothers of control pups (4.74 days) during the period of supplementary feeding (there were no significant differences throughout the rest of lactation). These findings are in accordance with predictions of a marginal-value model of fur seal lactation behaviour.

Differential macrohabitat use by birds on an unregulated lowland floodplain of south-eastern Australia

In many lowland floodplains around the world, upriver interferences to flows (weirs, dams, off-takes) have led to much reduced frequency and duration of flooding. As a result, many floodplain wetlands are now inundated relatively rarely if at all. Given regulation of most lowland rivers in southeastern Australia, we assessed use of wetlands by birds in the essentially unregulated Ovens River in northeastern Victoria. Twelve sites (0.4-1.2 ha) were studied after flooding. Four sites were 'permanent billabongs', four were temporary wetlands and the other four were randomly selected woodland sites >60 m from the nearest water body (including the river) acting as 'control' or 'reference' sites. Aquatic birds were not recorded using woodland sites, but many species were differentially associated with either billabongs or temporary wetlands. A surprising number of non-aquatic birds either exclusively or differentially were associated with wetland sites compared with woodland sites. We concluded that heterogeneous macrohabitat will increase local avian biodiversity on lowland floodplains. Moreover, densities and diversity of non-aquatic, woodland species also increased with the presence of wetlands. Temporary wetlands were used differently from permanent billabongs by birds, especially in foraging methods. This suggests that the reinstatement of major flooding on heavily regulated floodplains would be ecologically advantageous for birds by providing foraging and breeding opportunities.

Body composition changes, metabolic fuel use and energy expenditure during extended fasting in subantarctic fur seal (Arctocephalus tropicalis) pups at Amsterdam Island

The fasting metabolism of 71- to 235-d-old subantarctic fur seal (Arctocephalus tropicalis) pups from Amsterdam Island, southern Indian Ocean, was investigated during the long foraging trips of their mothers. Body lipid reserves were proportionally greater in female than male pups and higher in postmoult (37%) than premoult (10%) animals. The mass-specific rate of mass loss did not differ between the sexes but was lower than observed in other species. Daily mass loss was estimated to 56% fat, 10% protein, and 34% water. The rate of protein catabolism (15 g d⁻¹) was negatively related to the size of initial lipid stores and accounted for 9% (±1%) of total energy expenditure. However, body composition changes during the fast were not equal between the sexes, with females relying more on protein catabolism than males (11% and 5% of total energy expenditure, respectively). Energy expenditure (270 kJ kg⁻¹ d⁻¹) and metabolic water production (11.5 mL kg⁻¹ d⁻¹) rates are the lowest reported for an otariid species. These results suggest that subantarctic fur seal pups greatly reduce activity levels to lower energy expenditure in addition to adopting protein-sparing metabolic pathways in order to survive the extreme fasts they must endure on Amsterdam Island.

Milk consumption, body composition and pre-weaning growth rates of Australian fur seal

Pre-weaning growth rates, body composition, milk consumption and mass gain efficiency were measured in Australian fur seal Arctocephalus pusillus doriferus pups born in two consecutive breeding periods. Australian fur seals have the highest birth mass of any fur seal species (male 8.3 kg; female 7.2 kg). While their absolute pre-weaning growth rate (male 62 g·day⁻¹; female 53 g·day⁻¹) is similar to that of other temperate latitude fur seals, they have the longest birth-mass doubling time of any otariid species (134–136 days). Daily milk consumption increased from 400 g·day−1 (5 MJ·day−1) after birth to 675 g·day⁻¹ (13.7 MJ·day⁻¹) at age 210 day. However, mean mass-specific milk consumption (41 g·kg⁻¹) is substantially lower than in other otariid species (58–70 g·kg⁻¹) and, combined with a low mass gain efficiency (0.12 g·g⁻¹), contributes to the low mass-specific growth rates observed. There were no significant differences in either absolute or mass-specific milk consumption between the sexes. Significant differences, however, were found between the sexes in the body composition of pups with females generally having larger body lipid stores than males for any given mass. Peak milk yield by Australian fur seal females is estimated at 0.60 MJkg−0.75, substantially less than in Antarctic fur seals. The low level of maternal energy transfer in Australian fur seals may reflect the relatively low marine productivity of their foraging areas.

Variation in plasma leptin levels in response to fasting in Antarctic fur seals (Arctocephalus gazella)

Plasma leptin levels were determined in 8 lactating female and 20 pup Antarctic fur seals (Arctocephalus gazella) during fasting periods of normal duration. Plasma leptin levels ranged from 1.35-3.19 ng·ml^-1 in lactating females and 1.79-4.80 ng·ml^-1 in pups and were not positively correlated with body mass or condition. A negative trend, however, was observed between plasma leptin levels and body condition in lactating females upon their arrival at the colony following a foraging trip (beginning of fast). In accordance with findings in other species, plasma leptin levels dropped significantly (P<0.02) in response to the 17-19% drop in body mass experienced by pups during fasting. In contrast, plasma leptin levels in lactating females increased during the first 24 h of fasting before decreasing throughout the remaining 48 h of the fast. This unexpected result could be due to the high level of energy expenditure by seals as they swim back to the colony (i.e. post-exercise response) or may be influenced by the intense suckling activity experienced by females during the onshore fasting periods. The results of this study support recent findings in other carnivore species which suggest the primary physiological role of leptin in these species may not necessarily be as a signal of the magnitude of body energy reserves.

Fasting metbabolism in Antarctic fur seal (Artocephalus gazella) pups

The metabolism of 52–73-day old Antarctic fur seal pups from Bird Island, South Georgia, was investigated during fasting periods of normal duration while their mothers were at sea foraging. Body mass decreased exponentially with pups losing 3.5–3.8% of body mass per day. Resting metabolic rate also decreased exponentially from 172–197 ml (O2)·min⁻¹ at the beginning of the fast and scaled to M_b^0.74 at 2.3 times the level predicted for adult terrestrial mammals of similar size. While there was no significant sex difference in RMR, female pups had significantly higher (F₁,₁₈=6.614, P<0.019) mass-specific RMR than male pups throughout the fasting period. Fasting FMR was also significantly (t₁₅=2.37, P<0.035) greater in females (823 kJ·kg⁻¹·d⁻¹) than males (686 kJ·kg⁻¹·d⁻¹). Average protein turnover during the study period was 19.3 g·d⁻¹ and contributed to 5.4% of total energy expenditure, indicating the adoption of a protein-sparing strategy with a reliance on primarily lipid catabolism for metabolic energy. This is supported by observed decreases in plasma BUN, U/C, glucose and triglyceride concentrations, and an increase in β-HBA concentration, indicating that Antarctic fur seals pups adopt this strategy within 2–3 days of fasting. Mean RQ also decreased from 0.77 to 0.72 within 3 days of fasting, further supporting a rapid commencement of protein-sparing. However, RQ gradually increased thereafter to 0.77, suggesting a resumption of protein catabolism which was not substantiated by changes in plasma metabolites. Female pups had higher TBL (%) than males for any given mass, which is consistent with previous findings in this and other fur seal species, and suggests sex differences in metabolic fuel use. The observed changes in plasma metabolites and protein turnover, however, do not support this.

Interactions between fur seals and a squid jig fishery in southern Australia

Interactions between Australian fur seals (Arctocephalus pusillus doriferus) and the Southern Squid Jig Fishery (SSJF) were investigated in April–May 2002. Details on the number of seals present, distance from the vessel, age and gender, and their behaviour were recorded using scan sampling techniques over 26 nights from eight vessels operating out of Portland and Port Fairy on the southwest coast of Victoria. Seals were observed on all nights but none were recorded in 30% of all (777) scan observations. Of the seals attending vessels at any one time (1.89 ± 0.24), 67% were involved in activities unrelated to jigging operations with the most common behaviour category being resting/grooming. Only 3.6% of observations involved seals targeting squid caught on jig lures whereas a further 29% were of foraging on squid within 40 m of the vessel. Damage to fishing gear attributable to seals was recorded on only three occasions. There was no evidence of negative impacts on seals from vessel operations. The majority of seals foraging on squid around vessels were adult females (71%) with the remainder being almost exclusively juvenile males. The current level of interactions between Australian fur seals and vessels in the SSJF appears minor.

Determinants of native avian richness in suburban remnant vegetation : implications for conservation planning

While urban areas are increasingly recognized as having potential value for biodiversity conservation, the relationship between biodiversity and the structure and configuration of the urban landscape is poorly understood. In this study we surveyed birds in 39 remnant patches of native vegetation of various sizes (range 1–107 ha) embedded in the suburban matrix in Melbourne, Australia. The total richness of species within remnants was strongly associated with the size of remnants. Remnant-reliant species displayed a much stronger response to remnant area than matrix-tolerant species indicating the importance of large remnants in maintaining representative bird assemblages. Large remnants are important for other ecological groups of species including migratory species, ground foraging birds and canopy foraging birds. Other landscape (e.g. amount of riparian vegetation) and structural components (e.g. shrub cover) of remnants have a lesser role in determining the richness of individual remnants. This research provides conservation managers and planners with a hierarchical process to reserve design and management in order to conserve the highest richness of native species within urban areas. First of all, conservation efforts should preferentially focus on the retention of larger remnants of native vegetation. Second, where possible, riparian vegetation should be included within reserves or, where it is already present, should be carefully managed to ensure its integrity. Third, efforts should be focused at maintaining appropriate habitat and vegetation structure and complexity.