Maternal investment in Antarctic fur seals: evidence for equality in the sexes?

Studies of the otariids (fur seals and sea lions), a highly sexually dimorphic group, have provided conflicting evidence of differential maternal expenditure in male and female offspring and, thus, suggestions that they conform to predictions of investment theory are equivocal. Since the mid-1970s, a diversity of research on Antarctic fur seals (Arctocephalus gazella) including studies of their reproductive ecology, lactation energetics, and foraging behaviour have been conducted at Bird Island, South Georgia that have resulted in one of the more complete and diverse data sets for any species of otariid. These long-term data were reviewed to determine whether there was any evidence to support that differential maternal expenditure occurred in Antarctic fur seals. Most of the data examined were collected during five consecutive austral summers from 1988 through 1992 and included years in which local food resources were abundant and scarce. We were unable to detect differences in the sex ratios of pups at birth or sex-biased differences in growth rates estimated from serial data, the number of foraging trips made, the duration of attendance ashore, diving behaviour, suckling behaviour, or milk consumption in any year and in the duration of foraging trips or age at weaning in 2 of 3 years. In addition, we found no evidence of greater reproductive costs between mothers with sons or daughters relative to their reproductive performance the following year. In contrast, sex-biased differences were only found in the duration of foraging trips in 1990, the age at weaning in 1988, and consistently in growth rates estimated from cross-sectional data. We suggest that differential maternal expenditure does not occur in Antarctic fur seals because male pups probably do not gain greater benefit from additional maternal expenditure than female pups. After weaning, males experience a period of rapid juvenile growth over 3–4 years during which time body mass nearly trebles. This growth will almost certainly be dependent upon available food resources then rather than on any maternal expenditure received over the first 4 months of life and, thus, the assumptions of the Trivers and Willard hypothesis are probably invalid for Antarctic fur seals. Received: 10 July 1996 / Accepted after revision: 3 March 1997     

Parent–offspring and sibling conflict in Galápagos fur seals and sea lions

Parent–offspring conflict theory is well supported by theoretical arguments. However, empirical observations are often difficult to interpret and have contradicted one of its most appealing predictions that parent and offspring should disagree over killing of nest or littermates. We present the first examples of deadly conflict between siblings of different cohorts. In Galápagos fur seals (Arctocephalus galapagoensis) and sea lions (Zalophus wollebaeki), mothers often wean their single offspring at 2 years. This leads to a situation where up to 23% of all pups are born while the older sibling is still being nursed. Younger siblings are disadvantaged by being born lighter than neonates without older still dependent siblings. Pups born while an older sib is still dependent grow less in early life (fur seal) and suffer increased early mortality (both species) through direct aggression or scramble competition with the older sibling. This effect is much stronger in years of high sea surface temperature (El Niño) indicating low marine productivity and if the older offspring is a male. In both species, mothers interfere aggressively in this conflict by defending the younger offspring. In years of El Niño, intense resistance to maternal aggression by the older offspring happens frequently in the fur seal. Such resistance against weaning can induce maternal neglect of the newborn. Given substantial year to year variation in offspring growth, maternal aggression forces weaning in the older sibling only if it has reached sufficient size to support itself by foraging. In Galápagos fur seals, pups with older siblings can either represent insurance against loss of older offspring or extra reproductive value.     

Sibling competition in guinea pigs (<Emphasis Type="Italic">Cavia aperea</Emphasis> f. <Emphasis Type="Italic">porcellus</Emphasis>): scrambling for mother’s teats is stressful

Lactation is the most energy-intense period in the life of a female mammal. This can cause severe conflict between mother and offspring over the duration of lactation but also between siblings over the amount of milk each pup gets from its mother. Thus, competitive interactions between siblings are expected, and competition is likely to increase with litter size, particularly in species where the number of offspring exceeds the number of teats. We studied sibling competition in the domestic guinea pig (Cavia aperea f. porcellus), which has two teats, but frequently bears litters of up to five pups. By cross-fostering we created non-competition (control) litters with two pups and competition litters with four pups and observed nursing behaviour on days 5, 10, 15 and 20 postpartum. Pups of larger litters had lower growth rates, indicating increased competition among siblings in these litters. Pups of larger litters had to wait longer for access to a teat and spent less time suckling than pups of smaller litters but ate more solid food instead. Additionally, we manipulated the individual short-term need of pups by separating half of the pups of each litter for 2 h from their mothers before observation. Within a litter, hungry pups achieved access to milk faster and spent more time suckling than non-hungry pups. Pups competed mostly by scramble competition. Aggressive interactions occurred only in large litters. Pups of large litters had higher cortisol levels than pups in small litters. These effects decreased with age as pups became increasingly independent of maternal milk. Pup behaviour appears to fit better with models of scramble competition than with those of honest signalling. This contribution is part of the special issue “Sibling competition and cooperation in mammals” (guest editors: Robyn Hudson and Fritz Trillmich).     

Effects of El Niño on distribution and reproductive performance of Black Brant

Climate in low-latitude wintering areas may influence temperate and high-latitude breeding populations of birds, but demonstrations of such relationships have been rare because of difficulties in linking wintering with breeding populations. We used long-term aerial surveys in Mexican wintering areas and breeding areas in Alaska, USA, to assess numbers of Black Brant (Branta bernicla nigricans; hereafter brant) on their principal wintering and breeding area in El Ni?o and non-El Ni?o years. We used Pollock's robust design to directly estimate probability of breeding and apparent annual survival of individually marked brant at the Tutakoke River (TR) colony, Alaska, in each year between 1988 and 2001. Fewer brant wintered in Mexico during every El Ni?o event since 1965. Fewer brant were observed on the principal breeding area following each El Ni?o since surveys began in 1985. Probability of breeding was negatively related to January sea surface temperature along the subtropical coast of North America during the preceding winter. Between 23% (five-year-olds or older) and 30% (three-year-olds) fewer brant nested in 1998 following the strong El Ni?o event in the winter of 1997-1998 than in non-El Ni?o years. This finding is consistent with life history theory, which predicts that longer-lived species preserve adult survival at the expense of reproduction. Oceanographic conditions off Baja California, apparently by their effect on Zostera marina (eelgrass), strongly influence winter distribution of brant geese and their reproduction (but not survival), which in turn affects ecosystem dynamics in Alaska.     

Transmission of food preferences in spiny mice (Acomys cahirinus) via nose-mouth interaction between mothers and weanlings

Summary Two laboratory experiments assessed the role of nose-to-head and nose-mouth investigation in the transmission of food preferences from Acomys cahirinus mothers to their offspring. Pups spent more time nosing and licking the nose-mouth area of mothers who ate a novel food (fresh potato or carrot) than of mothers who ate no novel food. A greater amount of the less-preferred novel food (potato) was subsequently eaten by pups whose mothers had eaten potato than pups whose mothers had eaten either carrot or familiar laboratory chow only. Mothers maintained a stereotypic posture during nose-mouth investigation, holding still with mouth slightly open, providing both a source of specific food cues and a context in which pups could become familiar with them. These data suggest an active role by Acomys mothers in the development of offspring food preferences.     

Costs and benefits of communal rearing to female prairie voles (<Emphasis Type="Italic">Microtus ochrogaster</Emphasis>)

Communal rearing of offspring may help mothers maximize their investment in offspring at a reduced cost to their own bodily condition, thus maximizing their potential for reproductive success. The objective of this study was to quantify the costs and benefits of communal rearing to prairie vole (Microtus ochrogaster) pups and mothers. Mothers were assigned to one of three social units: solitary mothers, singularly breeding groups (i.e. one mother and one non-reproductive sister) and plurally breeding groups (i.e. two lactating sisters). For each type of social unit, some replicates were provided with food ad libitum, while others were provided with limited food. The body mass of focal mothers (i.e. the first mother to produce a litter) was a significant predictor of pup growth. Regardless of food availability, litters of focal mothers in plurally breeding groups gained more weight than litters reared by solitary mothers. Pups reared in singularly breeding groups were intermediate in weight gain, but did not gain significantly more weight than solitary offspring. There was no difference in the body mass of focal mothers from each type of social unit, regardless of food availability. Within plurally breeding groups, the weight gain of the two litters and body mass of focal and second mothers did not differ. However, focal mothers from plurally breeding groups nursed fewer pups than solitary mothers and also fewer pups than their nestmates when food was limited. Our results suggest that plural breeding results in greater fitness to mothers than solitary and singular breeding.Communicated by E. Korpimäki     

Head start: Australian sea lion pups gain experience of adult foraging grounds before weaning

The extended lactation period of the Australian sea lion Neophoca cinerea is suggested to provide additional time for offspring to achieve nutritional independence. We examine the dive and movement development of pups at two age classes (6?C10?months) from two colonies (Lilliput and Olive Islands) in South Australia using archival GPS and time-depth recorders to investigate the degree of overlap with maternal home range. Older pups from Lilliput travelled significantly further and spent more time at sea than younger pups. At Olive Island, there was a similar increase in distance travelled and time spent at sea with age. Pups at both colonies started occupying adult female home range at 6?months. Australian sea lion pups can explore adult foraging habitat at least 8?months prior to weaning, allowing them to learn the location of suitable habitat and the skills required to hunt successfully.     

Parental investment and the secondary sex ratio in northern elephant seals

Summary Data on northern elephant seals, Mirounga angustirostris, bearing on sex ratio theory were collected at Año Nuevo, California, and other Californian and Mexican Islands, during the period 1967 to 1988. The mass of males exceeded that of females by 7–8% at birth and at weaning. The sex ratio was biased to males at birth (51.2%) and was near unity at weaning (49.6% males). The sex ratio did not vary as a function of maternal age or maternal mass except in 6-year-old females, who produced significantly more males. Although sons cost more to rear in energetic terms than daughters, and mothers were more successful weaning the latter, the sex of the pup reared exerted no significant effect on the mother's reproductive performance the following year or on her subsequent survival. These data suggest that parents invest equally in sons and daughters when investment is measured in terms of future reproduction (Fisher 1930) and provide no support for the theory of adaptive shifts in sex ratio (Trivers and Willard 1973). The small sex difference in mass due to maternal effort reflects the fact that females fast during lactation and all energy transferred is from limited body stores. Because of these circumstances, selection for superior condition at the end of the period of parental investment may act more strongly on pups, who have the opportunity to steal milk, than on their mothers.     

Lactation performance of guinea pigs (Cavia porcellus) does not respond to experimental manipulation of pup demands

To ensure proper development of young, mothers should react to offspring signals of need. Studies of such parent-offspring interaction often manipulated litter size to measure effects of changed offspring food demand. We used the extreme precociality of guinea pig (Cavia porcellus) pups to increase offspring demand without changing any other litter characteristic. As pups contribute to their own energy demands from birth by independent feeding, their food demand can be increased by withholding solid food early in lactation. We studied whether mothers reacted to increased food demand of offspring by enhanced parental care, especially by changes in the pattern of milk production and nursing. Pups deprived of solid food early in lactation grew more slowly and were in poorer body condition than pups in control litters, even after the former had access to solid food in late lactation. Mothers of deprived young reacted to offspring long-term need by maintaining nursing behaviour for longer than control mothers. However, this change in behaviour did not occur early in lactation when pup short-term need was greatest nor did it result in increased milk transfer at any time. Energy allocation of mothers measured by changes in their food intake, maternal body reserves, and milk production stayed the same whether offspring had early access to solid food or not. Thus mothers did not increase energy allocation to pups even though they apparently had information about the pups' poor state.     

The effect of pup vocalisations on food allocation in a cooperative mammal, the meerkat (Suricata suricatta)

In the meerkat (Suricata suricatta), a cooperative mongoose, pups follow potential feeders while the group is foraging and emit incessant calls when soliciting food from them. In contrast to a ’stationary’ brood of chicks, in which nestlings are fed at a fixed location, meerkat pups are ’mobile’ and become spread out. The question arises whether meerkat pups that experience different constraints to those facing chicks have evolved similar begging strategies. This paper describes the vocalisations that meerkat pups emit in the context of begging and investigates the influence of these calls on food allocation by older group members and on the behaviour of littermates. Meerkat pups use two types of calls when soliciting food from a potential feeder. The most common is a ’repeat’ call, which pups emit continuously when following an older forager over several hours a day. In addition, when a potential feeder finds a prey item, the pups next to it emit a bout of calls with increased calling rate, amplitude and fundamental frequency, termed ’high-pitched’ calls. Observations, together with playback experiments, showed that more prey was allocated to pups that called longer and more intensely. The pup closest to a feeder was almost always fed. The probability of emitting high-pitched calls did not depend on the time since a pup had received food, and the change from repeat to high-pitched calls occurred suddenly. The main function of the high-pitched call, therefore, does not appear to be to signal a pup’s hunger state. More likely, the two calls, in the context of begging, may be an adaptation to energetic constraints in a mobile feeding system. Pups, which are dispersed during foraging, may emit repeat calls over long periods to prevent potential feeders from eating all the prey themselves. At the moment a potential feeder finds prey, pups may give the more intense high-pitched calls to direct feeders to bring the food item to them and not to a littermate. Therefore, unlike the stationary feeding system where chicks emit one type of begging call when the feeder approaches the nest, meerkats, with a mobile feeding system, have evolved two discrete types of vocalisations in the context of begging. Received: 22 November 1999 / Revised: 1 July 2000 / Accepted: 17 July 2000     

Sexual dimorphism in sea lion pups: differential maternal investment,or sex-specific differences in energy allocation?

Proximal mechanisms underlying a faster growth rate in male compared to female California sea lion pups were investigated. Males are significantly larger at birth than females. Specifically, we asked if differential maternal investment contributed to enhanced male growth via: (1) larger mothers having disproportionately more male pups, (2) more time and energy put into foraging by mothers of male pups, and (3) greater milk production in mothers of male pups. We also considered four aspects of differential energy utilization and acquisition by male and female pups: (1) male pups attempting to save energy for growth by changes in behavior, (2) longer suckling bouts with mother and more sneak suckling of non-filial females by male pups, (3) lower maintenance costs in males via a lowered resting metabolic rate, and (4) increased assimilation efficiency in males. Our study showed that there are no differences in the size of females or length of foraging trips for mothers of male and female pups. Male pups received more milk from their mothers, but the difference was no longer significant when the larger body size of males was considered. There were no differences in either the activity budgets or suckling behavior of male and female pups. Male pups, however, did have lower resting metabolic rates than females. We conclude that enhanced male perinatal growth is a consequence of a larger size at birth, proportionally more milk from mothers to support the greater demands of larger body size, and lower maintenance costs due to a lower resting metabolic rate. Received: 28 April 1995/Accepted after revision: 25 July 1995     

Community change in the variable resource habitat of the abyssal northeast Pacific

Research capable of differentiating resource-related community-level change from random ecological drift in natural systems has been limited. Evidence for nonrandom, resource-driven change is presented here for an epibenthic megafauna community in the abyssal northeast Pacific Ocean from 1989 to 2004. The sinking particulate organic carbon food supply is linked not only to species-specific abundances, but also to species composition and equitability. Shifts in rank abundance distributions (RADs) and evenness, from more to less equitable, correlated to increased food supply during La Ni?a phases of the El Ni?o Southern Oscillation. The results suggest that each taxon exhibited a differential response to a sufficiently low dimension resource, which led to changes in community composition and equitability. Thus the shifts were not likely due to random ecological drift. Although the community can undergo population-level variations of one or more orders of magnitude, and the shape of the RADs was variable, the organization retained a significant consistency, providing evidence of limits for such changes. The growing evidence for limited resource-driven changes in RADs and evenness further emphasizes the potential importance of temporally variable disequilibria in understanding why communities have certain basic attributes.     

Weaning in the guinea pig (<Emphasis Type="Italic">Cavia aperea</Emphasis> f. <Emphasis Type="Italic">porcellus</Emphasis>): Who decides and by what measure?

Offspring should be selected to influence maternal effort to maximize their own fitness, whereas mothers are selected to limit investment in present progeny. In mammals, this leads to a conflict over the amount of milk provided and the timing of weaning. The intensity and time course of such conflict has so far mostly been investigated experimentally in altricial rodents. However, it is expected that offspring options for conflict will depend on developmental state. We therefore investigated in the highly precocial domestic guinea pig (Cavia aperea f. porcellus) who decides over nursing performance and weaning and how pup state influences these decisions. Specifically, we tested whether a threshold mass of pups predicts weaning time. By exchanging older litters against neonates and vice versa, we produced a situation in which females differed in lactational stage from the cross-fostered pups. Our results indicate that females decide about the timing of weaning, as cross-fostered younger pups were weaned at a much younger age than controls and older pups benefited from continuing lactation of foster mothers. Growth rates did not differ in the treatment groups, and different weaning ages resulted in differing weaning mass refuting the hypothesis that weaning is based on a threshold mass of offspring. This constitutes clear evidence that in a precocial rodent, the guinea pig, decisions about maternal care are primarily determined by maternal state and little influenced by pup state despite the extreme precociality of offspring. We suggest that precocial pups show little resistance to early weaning when food is abundant, as they reach sufficient nutritional independence by the middle of lactation to enable independent survival.     

Sex differences in mass loss rate and growth efficiency in Antarctic fur seal (Arctocephalus gazella) pups at Macquarie Island

We investigated the relationship between the mass gained by a pup during a period of maternal attendance (as an index of milk intake) and the duration of the preceding foraging trip in relation to the mass-specific rate of mass loss during fasting periods and the growth rate of Antarctic fur seal (Arctocephalus gazella) pups at Macquarie Island. We found that (1) serially weighed male pups grew significantly faster than females pups and that (2) fasting female pups lost mass at a significantly higher rate (2.55% day⁻¹) than male pups (2.12% day⁻¹) of the same mass; (3) during periods of maternal attendance, there were no intersexual differences in the amount of mass gained by pups of the same size, hence (4) female pups required a higher daily mass gain to grow at the same rate as male pups. Our results show that intersexual differences in growth rate may be accounted for by intersexual differences in mass-specific rate of mass loss, because females lost 0.42% more of their total mass per day (i.e. 4.2 g kg⁻¹ day⁻¹) compared with male pups of the same body mass. Despite intersexual differences in growth rates, our results indicate equality of maternal expenditure between the sexes. Intersexual differences in the rate of mass loss may be due to differences in the metabolic rate, activity level and/or body composition of male and female pups. Received: 19 August 1998 / Received in revised form: 19 April 1999 / Accepted: 19 April 1999     

Satellite tracking of Humboldt penguins (Spheniscus humboldti) in northern Chile

During the El Niño of 1982/1983, the Humboldt penguin population diminished dramatically in the whole distributional area of the species. Recovery of the population was slow since 1983 and it has been suggested that large numbers of Humboldt penguins die at sea, entangled in nets, or starve to death, even during non-“El Niño” years. We were able to determine for the first time, how Humboldt penguins on Pan de Azúcar Island (26°S; 72°W) utilize their marine habitat and where their feeding areas lie. For this purpose we employed two streamlined Argos satellite transmitters during the 1994/1995 and 1995/1996 breeding seasons, respectively. Mean travelling speed of Humboldt penguins during foraging trips was 0.94?m s^?1 and 50% of bird positions were located within 5?km of the island (90% within 35?km). Total area covered by Humboldt penguins foraging from Pan de Azúcar Island was 12?255?km². Satellite transmitters also recorded dive duration; penguins spent on average 7.8 to 9?h diving per foraging day but showed no preferences for particular feeding areas. Mean daily dive durations (4-d mean) recorded during the 1994/1995 breeding season were positively correlated between birds. Significant correlation between dive duration and sea surface temperature anomalies and negative correlation between dive duration and fishery landings at nearby Caldera harbour indicate that the 1994/1995 increase in foraging effort was a response to deteriorating prey availability. Sea surface temperatures during the 1995/1996 breeding season were colder than average, and we observed no trends in bird diving activities.     

Evidence of equal maternal investment in the sexes in the polygynous and sexually dimorphic grey seal (Halichoerus grypus)

In polygynous and sexually dimorphic mammals, parents may be expected to bias their investment towards sons because variation in reproductive success is usually higher among males than among females. Moreover, male reproductive success often depends on adult body size, which, in turn, may depend on the level of parental investment. We therefore predicted that in the grey seal (Halichoerus grypus), a polygynous and sexually dimorphic phocid seal, females should invest more in individual sons than in individual daughters. We found that male pups were born heavier than female pups, but that the growth rates and suckling behaviour were similar for the two sexes. The growth rates and the birth weights were not correlated for the pups of either sex. Mothers did not behave differentially towards offspring of the two sexes, except that mothers of male pups spent more time in visual contact with their pups. Male and female pups had similar activity levels and begged at similar rates. We argue that reports of equal expenditure on the two sexes can be accepted as evidence of equal investment, provided that three assumptions are fulfilled. First, parental care must be costly to the parent. Second, energy expenditure must be the most important component of parental investment. Third, there must be no negative correlation between maternal body condition and the ratio of sons to daughters produced. We argue that these assumptions are met in our study, and that our results provide evidence of equal maternal investment in the sexes in grey seals.     

Current indirect fitness benefits associated with philopatry in juvenile prairie voles

Summary Prairie vole (Microtus ochrogaster) family groups were examined to determine possible indirect fitness benefits from the presence of juvenile helpers at the natal nest. The reproductive performance of family groups retaining two juveniles was compared with that of families in which all juveniles were removed when the subsequent litter was born. Litter sizes at birth of the second litter and pup survival rates were the same in the two treatments. Offspring quality was affected by the presence of juveniles however. Pups reared with juveniles weighed 13% more at weaning than pups reared without juveniles. Pups also opened their eyes sooner when juveniles were present. Differences in growth and development may have been affected by the amount of time pups were alone in the nest since pups in families with juveniles were left alone less frequently than were pups without juveniles. Maternal behavior patterns were not affected by the presence of juveniles. In contrast, fathers in families with juveniles spent more time in non-parental behaviors such as feeding, drinking, and foraging. In families with large litters, mothers delivered a subsequent litter sooner if juveniles were present. Subsequent litter sizes were the same in both treatments. Overall, both infants and parents benefited from the presence of juveniles, suggesting that helping may enhance the helper's indirect fitness in multiple ways.     

Stable isotopes document winter trophic ecology and maternal investment of adult female southern elephant seals (<Emphasis Type="Italic">Mirounga leonina</Emphasis>) breeding at the Kerguelen Islands

Individual specialisation is widespread and can affect a population’s ecological and evolutionary dynamics. Whether intra-specific niche differences can influence reproductive investment was examined in a marine mammal, the southern elephant seal (Mirounga leonina), whose females were known to forage in two different areas during the austral winter. The study was conducted at Kerguelen Islands (49°21′S, 70°18′E), southern Indian Ocean, in late winter–early spring 2006. Pups were used as proxies of their mothers’ biology and combined information on their weaning mass (a proxy of females’ foraging success and short-term fitness) together with their blood δ¹³C value (a proxy of female foraging zone). First, the use of isotopic signature of pups was validated to study the female foraging ecology during their pre-breeding trip by demonstrating that δ¹³C and δ¹⁵N values of pups and their mothers were positively and linearly correlated. Then, blood samples were taken from a large number of newly-weaned pups, which were also weighed, to provide information at the population level. Estimated δ¹³C values of female seals encompassed a large range of values (from −23.7 to −19.1‰) with an unimodal frequency distribution, suggesting no contrasted foraging areas within the population. No significant relationship was found between pup weaning mass and their carbon signature, indicating no link between female foraging areas and maternal foraging success and investment. Finally, blood δ¹³C and δ¹⁵N values gave new insights into southern elephant seal ecology, suggesting that females mainly foraged north of the Polar Front where they preyed upon myctophid fish in late winter.     

Foraging behaviour of sympatric Antarctic and subantarctic fur seals: does their contrasting duration of lactation make a difference?

The duration of periods spent ashore versus foraging at sea, diving behaviour, and diet of lactating female Antarctic (Arctocephalus gazella, AFS) and subantarctic (A. tropicalis, SFS) fur seals were compared at Iles Crozet, where both species coexist. The large disparity in lactation duration (SFS: 10 months, AFS: 4 months), even under local sympatry, has led to the expectation that AFS should exhibit higher foraging effort or efficiency per unit time than SFS to allow them to wean their pups in a shorter period of time. Previous evidence, however, has not supported these expectations. In this study, the distribution of foraging trip durations revealed two types of trips: overnight (OFT, <1 day) and long (LFT, >1 day), in common with other results from Macquarie Island. However, diving behaviour differed significantly between foraging trip types, with greater diving effort in OFTs than in LFTs, and diving behaviour differed between fur seal species. OFTs were more frequent in SFS (48%) than in AFS (28%). SFS performed longer LFTs and maternal attendances than AFS, but spent a smaller proportion of their foraging cycle at sea (66.2 vs. 77.5%, respectively). SFS dove deeper and for longer periods than AFS, in both OFTs and LFTs, although indices of diving effort were similar between species. Diel variation in diving behaviour was lower among SFS, which foraged at greater depths during most of the night time available than AFS. The diving behaviour of AFS suggests they followed the nychthemeral migration of their prey more closely. Concomitant with the differences in diving behaviour, AFS and SFS fed on the same prey species, but in different proportions of three myctophid fish (Gymnoscopelus fraseri, G. piabilis, and G. nicholsi) that represented most of their diet. The estimated size of the most important fish consumed did not vary significantly between fur seal species, suggesting that the difference in dive depth was mostly a result of changes in the relative abundance of these myctophids. The energy content of these fish at Iles Crozet may thus influence the amount and quality of milk delivered to pups of each fur seal species. These results contrast with those found at other sites where both species coexist, and revealed a scale of variation in foraging behaviour which did not affect their effort while at sea, but that may be a major determinant of foraging efficiency and, consequently, maternal investment.     

Multi-season climate synchronized historical fires in dry forests (1650-1900), northern Rockies, U.S.A

Our objective was to infer the climate drivers of regionally synchronous fire years in dry forests of the U.S. northern Rockies in Idaho and western Montana. During our analysis period (1650-1900), we reconstructed fires from 9245 fire scars on 576 trees (mostly ponderosa pine, Pinus ponderosa P. & C. Lawson) at 21 sites and compared them to existing tree-ring reconstructions of climate (temperature and the Palmer Drought Severity Index [PDSI]) and large-scale climate patterns that affect modern spring climate in this region (El Ni?o Southern Oscillation [ENSO] and the Pacific Decadal Oscillation [PDO]). We identified 32 regional-fire years as those with five or more sites with fire. Fires were remarkably widespread during such years, including one year (1748) in which fires were recorded at 10 sites across what are today seven national forests plus one site on state land. During regional-fire years, spring-summers were significantly warm and summers were significantly warm-dry whereas the opposite conditions prevailed during the 99 years when no fires were recorded at any of our sites (no-fire years). Climate in prior years was not significantly associated with regional- or no-fire years. Years when fire was recorded at only a few of our sites occurred under a broad range of climate conditions, highlighting the fact that the regional climate drivers of fire are most evident when fires are synchronized across a large area. No-fire years tended to occur during La Ni?a years, which tend to have anomalously deep snowpacks in this region. However, ENSO was not a significant driver of regional-fire years, consistent with the greater influence of La Ni?a than El Ni?o conditions on the spring climate of this region. PDO was not a significant driver of past fire, despite being a strong driver of modern spring climate and modern regional-fire years in the northern Rockies.