California sea lions interfere with striped marlin hunting behaviour in multi-species predator aggregations.

The open ocean offers a suite of ecological conditions promoting the occurrence of multi-species predator aggregations. These mixed predator aggregations typically hunt large groups of relatively small and highly cohesive prey. However, the mechanisms and functions of these mixed predator aggregations are largely unknown. Even basic knowledge of whether the predator species' interactions are mutualistic, commensal or parasitic is typically missing. Moreover, recordings of attack and capture rates of marine multi-species predator aggregations, which are critical in understanding how and why these interactions have evolved, are almost completely non-existent owing to logistical challenges. Using underwater video, we quantified the attack and capture rates of two high-trophic level marine predators, California sea lions (Zalophus californianus) and striped marlin (Kajikia audax) attacking schools of fishes in the Southern California Current System, offshore the Baja California Peninsula. Recording over 5000 individual attacks across 13 fish schools, which varied in species, size and predator composition, we found that sea lions kleptoparasitized striped marlin hunts and reduced the frequency of marlin attacks and captures via interference competition. We discuss our results in the context of the phenotypic differences between the predator species and implications for a better understanding of multi-species predator aggregations. This article is part of the theme issue 'Mixed-species groups and aggregations: shaping ecological and behavioural patterns and processes'.

Within-litter differences in personality and physiology relate to size differences among siblings in cavies.

Many aspects of an animal's early life potentially contribute to long-term individual differences in physiology and behaviour. From several studies on birds and mammals it is known that the early family environment is one of the most prominent factors influencing early development. Most of these studies were conducted on highly altricial species. Here we asked whether in the highly precocial cavy (Cavia aperea) the size rank within a litter, i.e. whether an individual is born as the heaviest, the lightest or an intermediate sibling, affects personality traits directly after birth and after independence. Furthermore, we investigated whether individual states (early growth, baseline cortisol and resting metabolic rate) differ between siblings of different size ranks and assessed their relation to personality traits. Siblings of the same litter differed in personality traits as early as three days after birth. Pups born heaviest in the litter were more explorative and in general more risk-prone than their smaller siblings. Physiological state variables were tightly correlated with personality traits and also influenced by the size rank within litter, suggesting that the size relative to littermates constitutes an important factor in shaping an individual's developmental trajectory. Our data add valuable information on how personalities are shaped during early phases of life and indicate the stability of developmentally influenced behavioural and physiological traits.

Photoperiodic effects on reproductive development in male cavies (Cavia aperea).

Age at maturity is a particularly important life history parameter, as it predicts potential life time reproductive success in many small mammal species. Animals should therefore optimize the timing of maturation by reacting to environmental stimuli that predict future environmental conditions. Photoperiod often reliably predicts ecological conditions. Animals born into a photoperiod indicating favorable conditions (spring, summer) may mature earlier and at a lower weight than animals born into conditions indicating unfavorable conditions (autumn, winter). So far most work was done on small, altricial rodent species and we still lack knowledge about their precocial relatives. Precocial animals are born much further developed than altricials and might show less plasticity in their ontogenetic trajectory than the latter. We tested the influence of photoperiod simulated by increasing (spring) or decreasing (autumn) light in climate chambers on important life history parameters in a medium sized rodent, the highly precocial cavy (Cavia aperea). We wanted to determine whether photoperiod influences timing of maturation and early growth in male cavies and whether patterns of testosterone in blood are reflected by patterns of testosterone metabolites in feces.Males born into simulated spring grew faster and matured at an earlier age than males born into simulated autumn conditions. Patterns of testosterone in blood correlated with testosterone metabolites measured in feces. Male cavies strongly react to predictive photoperiod cues by adjusting growth and timing of maturation as we found previously for females, corroborating the importance of seasonal cues for adjustments of life history.

Mama's boy: sex differences in juvenile survival in a highly dimorphic large mammal, the Galapagos sea lion.

In many mammals, early survival differs between the sexes, with males proving the more fragile sex ["Fragile male (FM) hypothesis"], especially in sexually dimorphic species where males are the larger sex. Male-biased allocation (MBA) by females may offset this difference. Here, we evaluate support for the FM and MBA hypotheses using a dataset on Galapagos sea lions (Zalophus wollebaeki). We statistically model sex-specific survival as it depends on body mass and environmental conditions (sea surface temperature, SST, a correlate of marine productivity) at three developmental stages, the perinatal phase (1st month), the main lactation period (1st year), and the weaning period (2nd year). Supporting the FM hypothesis, we found that, early in life (1st month), at equal birth mass, males survived less well than females. During the remainder of the first year of life, male survival was actually less sensitive to harsh environmental conditions than that of females, contradicting the FM hypothesis and supporting the MBA hypothesis. During the second year of life, only male survival suffered with high SSTs as predicted by the FM hypothesis. At each developmental stage, observed survival rates were almost equal for both sexes, suggesting that mothers buffer against the inherent fragility of male offspring through increased allocation, thereby masking the differences in survival prospects between the sexes.

Field metabolic rate of lactating female Galápagos fur seals (Arctocephalus galapagoensis): the influence of offspring age and environment.

Galápagos fur seals, Arctocephalus galapagoensis, live in a seasonal environment which varies strongly in productivity from year to year. We measured how the field metabolic rates (FMR) of lactating females varied with season, pup age and year. Energy expenditure was measured using doubly labeled water (DLW) during the cold seasons of 1984 and 1985 in 9 mothers of 1-3-month-old pups and 5 mothers of yearlings, and during the 1986 warm season in 8 mothers of 6-month-old pups. Young pups gained 0.84% mass/day during the cold season, but larger pups during the warm season lost 1.25% mass/day. During the warm season, females had lower relative total body water than during the cold season suggesting higher fat content during the warm, less productive season, but the effect was even more marked when comparing different years of the study: fat content was high in 1984 and 1986 and low in 1985. The FMR of mothers varied from 134 to 167 W but did not show significant differences between any of the pup age-groups. Among the years of the study, FMR showed only a trend towards low energy expenditure of mothers of young pups in 1984. The mean FMR was lower than for other otariids. Mothers may limit energy expenditure independent of pup age and season to minimize their own risk of starvation in an environment of comparatively low productivity, varying unpredictably due to frequent El Niño events.

Lunar cycles in diel prey migrations exert a stronger effect on the diving of juveniles than adult Galápagos fur seals.

In our study of the development of diving in Galápagos fur seals, we analysed changes in diving activity and body mass trends over the lunar cycle. Based on previously observed lunar cycles in colony attendance patterns, we hypothesized a greater impact of prey migrations of deep scattering layer organisms on younger fur seals. Using electronic dive recorders, we determined that seals dived less and deeper on moonlit nights than at new moon, and incurred body mass losses. These changes in foraging over the lunar cycle correlate with the suppression of the vertical migration of prey by lunar light. All effects were more pronounced in juveniles than adult females, with greater relative mass loss during full moon, which must (i) negatively affect long-term juvenile growth rates, (ii) lengthen periods of maternal dependence, and (iii) contribute to the lowest reproductive rate reported for seals. This underlines the importance of studying ontogeny in order to understand life histories, and for determining the susceptibility of animal populations to fluctuations in food availability.

Are precocial young cheaper? Lactation energetics in the guinea pig.

Caviomorphs stand out from other small rodents by producing extremely precocial young. Pups begin to nibble solid food right after birth and can survive weaning when 5 d old. We tested whether these properties of young lead to a low cost of lactation in guinea pigs (Cavia porcellus). In contrast to altricial rodents, energy acquisition and expenditure of guinea pig mothers (litter size, three) was highest during the first half of the 23-d lactation period. Daily metabolizable energy intake at peak lactation was increased by 126% above nonreproducing level and was 65% of the predicted maximal intake. Peak daily energy expenditure was 1.9 times basal metabolic rate, and resting energy expenditure was elevated 26% above nonreproductive level. From the third day, young consumed solid food and intake increased linearly over the course of lactation. During lactation, total intake of litters through solid food consumption (assuming efficient digestion) amounted to 86% of the energy intake that mothers allocated to reproduction during that period. The pattern of energy expenditure during lactation in guinea pigs differs substantially from that of altricial rodents and is made possible because pups contribute through solid food intake. Overall energy demand on the mothers is not much lower than in altricial rodents, but lower peak energy demand during lactation in guinea pigs may reduce the coupling of reproduction to periods of high food abundance.

Ontogenetic changes in food intake and digestion rate of the herbivorous marine iguana (Amblyrhynchus cristatus, Bell).

Young reptiles have higher relative energy demands than adults, but the proposed ontogenetic changes in diet to fulfil these demands were not found in the algae-eating Galápagos marine iguanas on Santa Fé. Feeding and digestion rates were investigated to analyse how young achieve higher energy intake. Daily food intake of free ranging marine iguana hatchlings (6-11 months old) was about one third that of adults, but relative intake (g dry mass · g-1 wet mass · day-1) was four times higher in the hatchlings. During feeding experiments, relative daily food intake of hatchling marine iguanas was approximately three times higher than that of adults (0.042 vs 0.013 g dry mass · g-0.8 wet mass · day-1), and mean gut passage time was two times shorter (5 vs 10 days). The hatchlings also maintained high body temperatures (36.7° C) even under relatively cool day-time air temperatures of 32° C. Apparent digestibility of algal food measured both during feeding trials and by Mn2+ AAS (atomic absorption spectrometry) for free-ranging iguanas was 70%, independent of body size and temperature. The red algae prevailing in the diet were high in protein (30% dry mass) and energy (12.1 kJ/g dry mass). Diving iguanas had higher rates of energy intake than intertidal foragers, but daily intake was less. Maintenance of high body temperature enabled hatchlings to achieve high digestion rates and, combined with high relative intake, thus achieve sufficient energy intake for rapid growth despite higher mass specific metabolic rates. Estimates of biomass of marine iguanas and their algal food are given for a section of coastline on Santa Fé.

The complete primary structure of the marine Carnivora, galapagoes fur seal (Arctocephalus galapagoensis, Otariidae) hemoglobins.

The complete primary structure of the two hemoglobin components of the fur seal (Arctocephalus galapagoensis) is presented. The two components (HbI and HbII) occur in nearly equal amounts and have identical beta-chains; whereas the two alpha-chains (alpha I/alpha II) differ by six exchanges Ile/Val, Met/Thr, Ser/Ala, Pro/His, Lys/Gly, and Thr/Ala at positions 10, 34, 35, 50, 78, and 131, respectively. The components were isolated by DEAE-Sephacel chromatography and were separated into the globin chains by RP-HPLC on a column of Nucleocil-C4. The sequences have been determined by Edman degradation in liquid- and gas-phase sequencer, using the native chains and tryptic peptides. The sequences compared with those of other Carnivora species and an adult human globin chains. An identical beta-chain is found in fur seal and walrus, whereas larger differences were found between alpha I and alpha II compared to beta-chains.

Characterization of proteins and fatty acid composition in Galapagos fur seal milk. Occurrence of whey and casein protein polymorphisms.

1. Milk proteins of the Galapagos fur seal (Arctocephalus galapagoensis) were separated adequately into whey and casein fractions using bovine milk analysis methods. 2. In samples from days 5-30 of lactation 40% of the total proteins were whey and 60% caseins; in mid-lactation, day 150, 25% were whey and 75% casein proteins. 3. Electrophoretic and isoelectric focusing patterns of fur seal whey protein differed widely from bovine patterns, whereas those of caseins were similar. 4. Polymorphisms of fur seal whey and casein proteins were noted and did not seem related to different stages of lactation. 5. C-16 and C-18 fatty acids contributed about 70% of fatty acids; 63% of the total acids in milk fat were unsaturated.

Immobilisation of free-ranging Galapagos sea lions (Zalophus californianus wollebaeki).

Nine free-ranging Galápagos sea lions were immobilised for marking, using a combination of ketamine (3 to 5 mg per kg) and xylazine (0.5 to 1.0 mg per kg). The degree of effect depended largely on the animals' behavioural and physiological state before immobilisation (after parturition, high arousal, subject to aggression, etc.) All nine animals survived. Overheating in the immobilised state may quickly result in heart and circulatory failures.