Mother-pup separation and adoption in northern elephant seals

Summary The evolution of fostering behavior, parental care directed toward another's young, has been the focus of much recent interest. During a five-year study of northern elephant seals (Mirounga angustirostris) at Año Nuevo, California, we recorded the frequency of mother-pup separation, reunion, and adoption of orphaned pups in crowded and low-density breeding areas. While most females nursed their own pup exclusively until it was weaned, many females, especially young mothers (age 3–5 years), were unable to raise a pup successfully. In the crowded main breeding harem on Año Nuevo Island, 24 to 47 percent of the pups born each year were separated from their mothers from 1977 to 1980. Mother-pup separation and pup mortality were associated with the following inter-related factors: 1) female density; 2) weather and tidal conditions; 3) topographical features of the breeding areas (i.e., degree of exposure to high tides and surf); and 4) the proportion of young, maternally inexperienced females pupping in a particular area. Most motherpup separations were caused directly by 1) adult males moving through the harem; 2) pups wandering from their mothers; 3) female aggression; and 4) inclement weather. Most of the separations, as well as adoptions, occurred when pups were quite young. Mother-pup recognition appeared to be based on a combination of acoustic, visual, and olfactory cues, and most mother-pup reunions were effected by the female rather than her pup. On the main island breeding area, 572 orphans were marked. Of these, five percent relocated their mother, 27 percent were adopted or frequently cared for by foster mothers, and 68 percent were not adopted, or rarely fostered. The survival of an orphan was clearly contingent on the amount of care it received; most orphans which were not nursed or protected by females died before reaching 6 weeks of age. Frequently, an adopted orphan's foster mother was in the stage of lactation which corresponded closely to that of its own mother. The most common fostering event involved females that had lost their own pup and adopted a single orphan. Other pupless females attempted to steal a pup, cared for a pup while it was still with its mother, adopted a weaned pup, adopted two pups, or indiscriminately nursed any orphaned pup that approached. Some females kept their own pup in addition to fostering in alien pup. Most foster mothers were young and had little or no previous maternal experience. The formation of large, high-density breeding rookeries, due to a scarcity of suitable breeding sites, results in frequent mother-pup separations, especially during inclement weather and tidal conditions. Many opportunities for adoptive behavior are therefore presented, because of the great number of orphans and pupless females. Increased maternal experience appears to be a benefit associated with adoption. Some instances of fostering behavior may also be based on reproductive errors on the part of the foster mother.     

Harem choice and breeding experience of female southern elephant seals influence offspring survival

Female mammals can increase their lifetime fitness through modification of investment potential and by providing better rearing environments with improved breeding experience. We examined the relationships between reproductive fitness and the behavioural decisions that female southern elephant seals (Mirounga leonina) made during the breeding season. We examined whether mother age and breeding experience influenced reproductive success (measured as 1st-year survival probability), and whether there was a change in the choice of harem size with increasing age. Pups produced by young mothers had lower 1st-year survival probability than pups produced by older mothers. A significant increase in mean female mass with age required an analysis of both these effects on offspring survival. There was a significant positive effect of both female age and mass, and the interaction between the two, on 1st-year pup survival. The proportion of young mothers (<5 years old) decreased and the proportion of older mothers (>6 years old) increased with increasing harem size (harems surveyed from 1997 to 2001). Females chose larger harems in which to breed as they aged. Females demonstrated fidelity to breeding areas among successive breeding seasons, with older females displaying greater breeding-site fidelity than younger females. The mean number of previous breeding attempts per female within a harem (breeding experience) increased significantly with increasing harem size. Breeding females returned to breed later in the breeding season as they aged—we hypothesize that young, subordinate females gain a priority advantage by returning earlier. These results lend support to the hypothesis that there are fitness advantages, in terms of offspring survival, that are conferred to females that breed in successively larger harems with age. Potential mechanisms that select for females to improve their breeding conditions include improved mate selection and the avoidance of conspecific harassment in harems.Communicated by F. Trillmich     

Site fidelity of male Galápagos sea lions: a lifetime perspective

Knowledge about the distribution of resources can lead to the development of spatial preferences and long-term site fidelity. Individuals are expected to choose sites that best suit their needs. However, dominant individuals restrict movements of less competitive ones. Accordingly, one may expect spatial preferences to differ with regard to individual characteristics and to change over time. We investigated lifetime changes of site fidelity patterns with regard to reproductive success in male Galápagos sea lions (Zalophus wollebaeki). Showing a high degree of natal philopatry in the first 2 years of life, non-territorial males pass through a stage of fidelity to their natal colony where they develop preferences for areas outside prime breeding areas. Variation in the degree of spatial preferences was not associated with age or size, characteristics linked to an individual’s dominance status. For non-territorial males, roaming proved to be an adequate strategy to gain reproductive success. Only the most competitive males established territories in areas preferentially visited by females. They had a high probability to return to breeding areas where they successfully reproduced in previous seasons. Overall, the results reveal lifetime changes in site fidelity with regard to male status. The degree of site fidelity observed within the colony suggests familiarity and thus a high degree of tolerance among individuals using the same areas. This seems to facilitate attendance in the colony and thus the possibility to prospect for oestrus females.     

Size and seasonal influences on the foraging range of female grey seals in the northeast Atlantic

Evidence of segregation in foraging habitat has been demonstrated in some top marine predators, including cetaceans, pinnipeds and seabirds. However, most data are not adequate to assess differences relating to body size or seasonal influences. This has implications for quantitative modelling of population-level predator–prey interactions and ecosystem structure. We examined potential influence of body size and ‘fatness’ on the foraging trip characteristics of a top marine predator, the Atlantic grey seal (Halichoerus grypus), in southwest Ireland within the framework of optimal foraging theory to examine how female grey seals foraging behaviour varied with size, and across the period between moult and breeding. Larger seals undertook trips of greater duration and travelled further from haul-out sites than smaller seals. However, body fat was negatively associated with trip duration and extent. Seals spent more time at sea during the summer, but trips were shorter in extent, suggesting more localized foraging during this season.     

Offspring sex ratio in relation to female size in southern elephant seals,Mirounga leonina

Southern elephant seals Mirounga leonina display extreme sexual dimorphism. In addition females show great variation in size and stored resources at parturition. Therefore they present an excellent opportunity for examination of responses of sex ratio to resource availability. We studied the relationships between the size of southern elephant seal females at parturition and the size and sex of their pups at South Georgia over four breeding seasons. We found a large individual variation in maternal post-partum mass (range 296–977 kg, n=151). Larger mothers gave birth to larger pups, irrespective of the sex of their pup. Male pups were on average 14% larger than females at birth and consequently more costly to bring to parturition. Our results suggest that female southern elephant seals must weigh more than 300 kg if they are to breed at all, and more than 380 kg if they are to give birth to a male pup. Above this threshold the proportion of males among offspring rapidly increases with maternal mass, and stabilizes at a level not significantly different from parity. These results show that smaller females of southern elephant seals vary offspring sex ratio in a way that is consistent with theories on adaptive offspring sex ratio. A smaller mother with a male foetus may benefit from terminating her pregnancy and allocating the resources she saves to her own growth. She could then give birth to and raise a larger pup in the subsequent season.     

Life history consequences of variation in age at primiparity in northern elephant seals

Summary The age when female northern elephant seals, Mirounga angustirostris, bear their first young varies from 2 to 6 years. At Año Nuevo, California, a group of 77 females, primiparous at age 3, had a lower survivorship rate to each successive year up to age 8 than a group of 98 females that deferred initial pupping until age 4. The difference in survivorship appears to be due to the greater relative energetic costs of gestation and lactation incurred by the earlier breeding females during a period in their development when growth is rapid. An alternate hypothesis for the difference in survivorship — that young primiparous females are in poor condition from birth-is untenable; females that pupped early in life were larger at weaning age (a correlate of condition) than females that were primiparous 1 year later.Models based on the data show that differential survival of seals that vary in age at primiparity has important consequences for population growth and life history strategies. The effect of age at primiparity on the rate of increase of populations varies with colony density and juvenile survivorship. The optimal life history strategy for female elephant seals under most conditions existing today, including those at Ano Nuevo during the study period, is to bear the first offspring at age 4. Primiparity at age 3 is projected to be favored when harem density is very low and weaning success and juvenile survivorship are high; postponement of first breeding to age 5 is expected at high harem densities with intense competition for breeding space. Offprint requests to: B.J. Le Boeuf     

Movements and site fidelity of grass shrimp (<Emphasis Type="Italic">Palaemonetes pugio</Emphasis> and <Emphasis Type="Italic">P. vulgaris</Emphasis>) in salt marsh intertidal creeks

Grass shrimp are abundant residents of shallow-water habitats in US Atlantic and Gulf coast estuaries. Mark-recapture events determined the extent and direction of their movements in meso-tidal salt marsh creeks at North Inlet, South Carolina. Microwire-tagged Palaemonetes vulgaris demonstrated high fidelity for a dock in a subtidal channel. In the 48 days following release, 11 % of the 422 tagged shrimp were recaptured and 99 % of all recaptures were made there an average of 14 days post-release. Palaemonetes pugio, which favored the use of shallow intertidal areas, exhibited moderate fidelity for pools within creek beds with 53 % of the recaptured shrimp collected at the release site. Higher fidelity was observed for confluences of intertidal creeks and the subtidal channels. Of the 5565 shrimp released with coded microwire tags, 7.5 % were recaptured and 98 % of those recaptures were at release sites up to 53 days later. During another event, P. pugio released 235 m from their origin demonstrated relatively low fidelity for the new site and a lack of homing behavior. Of the 544 tagged shrimp, 3 % were recaptured; only 40 % of those recaptured were from the release site. Site fidelity is a mechanism by which previously discovered large and consistent differences in grass shrimp use of neighboring intertidal creek basins can be sustained over weeks, seasons, and years. Spatial variations and the tendency of these keystone organisms to resist relocation by tidal currents and occupy certain areas for extended periods should be considered in decisions about proposed alterations to creeks and shorelines.     

Sex ratio,monogamy and breeding success in the Midas cichlid (Cichlasoma citrinellum)

Summary The Midas cichlid (Cichlasoma citrinellum) is a large, aggressive monogamous fish from the Great Lakes of Nicaragua. It lives in an ecosystem where breeding sites are at a premium and where breeding success can be low. I tested the strength of the pair bond by manipulating the sex ratios in captive groups, using ratios of 1:1, 2:1, 4:1, 1:2 and 1:4 (:). No polygamy was seen at any of the ratios. Departure from a ratio of 1:1 decreased breeding success for those pairs that formed. Both sexes appeared to enforce the pair bond but did so in different ways: females attacked males and females equally but males attacked other males significantly more than they did females. These results, coupled with field observations, indicate the Midas cichlid is obligated to monogamy by the demands of competition for breeding sites, the need for two parents to protect the fry and by energetic constraints.     

Does male harassment of females contribute to reproductive synchrony in the grey seal by affecting maternal performance?

We investigated the possibility that male harassment of lactating females differed in relation to time of birth in the grey seal, Halichoerus grypus, on Sable Island, Nova Scotia. This was done by comparing the frequency of male disturbances, maternal performance and pup growth for females that either gave birth during the peak of the pupping season or after the peak. Of the females, 58% gave birth in a 7-day period near the beginning of the pupping period, when the operational sex ratio was 2–4 females per male. Late in the pupping period the operational sex ratio reversed to about 1 female for every 2 males. The relative frequency of disturbances by males was significantly greater for late-pupping mothers than for peak-pupping ones (1.9 vs. 1.4 encounters/h). Females that gave birth late also were disturbed by males 3 times more often than females that gave birth during the peak (3.4 vs. 1.1 % of observation time). Late-pupping mothers spent 22% less time suckling (4.0 vs. 5.1 % of observation time), had 30% slower growing pups (1.7 vs. 2.4 kg/d), and weaned pups that were 16% lighter (45.6 vs. 54.0 kg). The effect of birth time on pup mass gain and weaning mass was not attributable to factors such as maternal mass, pup birth mass or pup sex. We conclude that the reduced maternal performance is likely the result of the increased male harassment. As reduced weaning mass can lead to reduced juvenile survival, male harassment may have contributed to the enhanced reproductive synchrony in this species.     

Intra-colony relatedness and sociality in the allodapine bee Exoneura bicolor

Summary Overwintered and newly co-founded nests of Exoneura bicolor exhibit different grades of sociality. Intra-colony relatedness was estimated for adults and female brood in both nest types using allozyme data and a multiallelic estimator. The higher relatedness among female brood from overwintered nests compared to newly-founded nests is consistent with the difference in sociality between these colony types (semisocial versus quasisocial). However, intra-colony relatedness among adults is higher in cofounded nests than in overwintered nests, suggesting that differences in sociality are determined by benefit/cost ratios associated with non-reproductive strategies, rather than relatedness between interactants. Rapid egg-production in newly founded nests allows eggs to be stockpiled. This reduceds the tasks available to non-reproductives in early phases of colony development and limits the payoffs available for reproductive altruism. It is suggested that the social flexibility characteristic of allodapines is a consequence of communal progressive rearing which allows many benefits from cooperative nesting to be gained without worker sterility.     

Philopatry and mother-daughter associations in bushy-tailed woodrats: space use and reproductive success

Social groups in mammals are often based on overlapping generations of philopatric females, yet few researchers have examined fitness consequences of associations between females and their daughters. We examined survival and reproductive consequences of sharing a natal site with one's mother or daughter in wild bushy-tailed woodrats (Neotoma cinerea). Bushy-tailed woodrats reside on discrete rocky outcrops, and females tend to spend their entire reproductive life on their natal outcrop. Mothers and daughters that co-existed on the natal outcrop were closely associated in space. Juvenile females were more likely to survive their first winter to become breeding yearlings at their natal site if their mother was also present at the natal site during the winter. Juveniles survived equally well with or without their mother where densities of adult females were experimentally reduced, suggesting that females enhance their daughter's survival by facilitating their access to limited resources. Yearling females exhibited no reproductive costs from association with their mothers at the natal site during the breeding season; in fact, yearlings that bred at the natal site simultaneously with their adult mothers tended (P = 0.07) to experience fewer reproductive failures than did those that bred in the absence of their mothers. Mothers did not experience any detectable costs when sharing a natal site with a daughter. We conclude that bushy-tailed woodrats exhibit stable, fitness-enhancing associations among mothers and their philopatric daughters.     

Parental investment and pair-bond behavior among canvasback ducks (Aythya valisineria,Anatidae)

Summary Hypothetical patterns of male and female cumulative reproductive effort in canvasback ducks were used to predict the efforts of males and females in pair-bond reinforcing behavior through a breeding sequence. The predictions were that, at any point in time in a monogamous breeding system, given certain assumptions, (a) the sex investing less should be more active in reinforcing the pair bond, and (b) this relationship should shift within and between breeding attempts with the pattern of cumulative reproductive effort per brood by the male and female. Data on the behavior of newly-formed pairs were obtained by observing apparently paired but unmarked canvasbacks during spring migration. Later, 11 individuallymarked pairs were followed through breeding sequences, including repeated nesting attempts. Observed affiliative behavior fit the predictions reasonably well. That sex whose cumulative reproductive effort was less tended to be more active in maintaining proximity to its mate, initiating bouts of mutual display, and coordinating pair activities. Predictions were met in 14 of 24 cases, results were contrary in 3 cases, and in 7 cases trends were not statistically different (P>0.05). Shifts in relative male and female reinforcement efforts that coincided with major changes in investment were apparent. These results support the proposition that factors other than simple parental investment-related desertion costs might be important to an individual's behavioral decisions regarding pairbond reinforcement and ultimately mate abandonment. Discrepancies in a few predicted outcomes further suggest that males and females may use and balance various pair-reinforcement tactics differently through a breeding season.     

Impact of salt-marsh management on fish nursery function in the bay of Aiguillon (French Atlantic coast), with a focus on European sea bass diet

The Bay of Aiguillon is a national French Nature Reserve of great importance for birds. Recently, the managers of the Reserve (ONCFS-LPO) paid attention to the influence of saltmarshes management on the nursery function for fish feeding in creeks at high tide. A study carried out from March to July 2012 aimed to evaluate the use of saltmarshes by fish juvenile fraction according to the mowing intensity in salt marshes surrounding creeks: ceased, irregular or annual mowing. This community approach was completed by a focus on the European sea bass Dicentrarchus labrax individual diet, vacuity index and growth, and the biomass of a main potential prey (the amphipod Orchestia gammarella). Whatever the mowing intensity, the juvenile fraction was very high for the main species, which were the grey mullet Liza ramada, the European sea bass, undetermined clupeid and the European flounder Platichthys flesus. Adult and subadult for these species were anecdotic or totally absent. Despite very different biomasses of amphipods between mown and natural sites, vacuity index, prey composition and their relative abundance in the diet of European sea bass juveniles were little different, contrary to their hypothetical growth (i.e. when assuming site fidelity), which appeared higher in non-mown site. The low distance between sampling sites could allow fish exchange over time between optimal and suboptimal creeks to feed on, as a hypothesis to explain such results. Because mowing was subsidized by European Union (EU) to favour open habitats for geese and maintain an economic activity, indirect impacts on nursery for fish called into question the appropriateness of such agro-environmental measure on natural habitats, and related fish nursery function.     

Seals at sea: modelling seal distribution in the German bight based on aerial survey data

The Wadden Sea is an important habitat for harbour seals and grey seals. They regularly haul-out on sandbanks and islands along the coast. Comparably little is known about the time seals spend at sea and how they use the remainder of the North Sea. Yet, human activity in offshore waters is increasing and information on seal distribution in the North Sea is crucial for conservation and management. Aerial line transect surveys were conducted in the German bight from 2002 to 2007 to investigate the distribution and abundance of marine mammals. Distance sampling methodology was combined with density surface modelling for a spatially explicit analysis of seal distribution in the German North Sea. Depth and distance to coast were found to be relevant predictor variables for seal density. Density surface modelling allowed for a depiction of seal distribution in the study area as well as an abundance estimate. This is the first study to use aerial survey data to develop a density surface model (DSM) for a spatially explicit distribution estimate of seals at sea.

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Paternity analysis shows experience,not age,enhances mating success in an aquatically mating pinniped,the Weddell seal (<Emphasis Type="Italic">Leptonychotes weddellii</Emphasis>)

For polygynous mammals with no paternal care, the number of offspring sired is often the sole measure of male reproductive success. The potential for polygyny is highest when resources or other environmental factors such as restricted breeding sites force females to aggregate. In these circumstances, males compete intensely for females and mating success may vary greatly among males, further intensifying selection for those traits that confer an advantage in reproduction. Hence, determinants of male success in competition for females are likely to be under strong sexual selection. Paternity analysis was used in conjunction with measures of age, site fidelity, and behavior during the breeding season to assess variance in male breeding success in Weddell seals (Leptonychotes weddellii) breeding at Turtle Rock, McMurdo Sound (77.727S, 166.85E) between 1997 and 2000. Paternity could be assigned to 177 pups at relaxed or 80% confidence level or 111 pups at strict or 95% confidence levels. Weddell seals at Turtle Rock show a modest degree of polygyny with the greatest number of pups sired by any individual male in a single season equalling 5 or ∼10% of the pups born. Over four consecutive years, most (89.2%) males sired at least one pup. In a generalized linear model (GLM), age and the age first seen at the study site as an adult were unrelated to mating success, but adult experience, either site-specific or elsewhere in McMurdo Sound, over the reproductive life span of males explained nearly 40% of variance in total mating success with 80% confidence and 24% of variance at 95% confidence. While learning where females are likely to be may enhance male reproductive success, aquatic mating reduces the ability of males to monopolize females, and thereby increases equity in mating success.     

Feeding rates and diel vertical migration of copepods near South Georgia: comparison of shelf and oceanic sites

Seventeen Longhurst Hardy Plankton Recorder profiles were taken over a diel cycle in January 1990 to study the feeding of four major copepods over the South Georgia shelf. Ontogenetic changes in vertical migration were followed and feeding cycles determined by gut fluorometry for Calanoides acutus Stage CV, Calanus sinillimus CV and CVI, C. propinquus CV and Rhincalanus gigas CV and CVI. In common with a neighbouring oceanic site visited two weeks later and reported elsewhere, all four species had a diel cycle of feeding and migration. The vertical distributions of C. simillimus (all stages), R. gigas (nauplii) and Euphausia frigida (postlarvae) were similar at both sites, the night being spent within the chlorophyll maximum at 15 to 30 m. However, the biomass dominants, C. acutus and R. gigas, dwelt below the chlorophyll maximum, about 30 m deeper than their oceanic counterparts. Unlike the oceanic site, feeding at the shelf site was not restricted to darkness, but increased 6 to 10 h before nightfall and finished at dawn; the intervening period coincided with sinking and digestion. Daylight feeding may have been induced by the shorter night, lower light levels or greater food requirements at the shelf site, despite planktonic predators being over three times more abundant. Daily ration estimates for R. gigas at both sites were only 2% body carbon per day. These low values contrast with its smaller competirors, whose rations were in the range 5.6 to 27%.     

Ecological constraints on the behaviour of mother long-tailed macaques (Macaca fascicularis)

Summary The ecological consequence of the effects of different social conditions on the behaviour of females with infants were examined in two different-sized groups of long-tailed macaques (Macaca fascicularis) that inhabited adjacent home ranges in Gunung Leuser National Park in Indonesia. This study aims to test predictions of the within-group competition/predation avoidance hypothesis of social organization. Females, particularly those carrying an infant, are considered to experience the most direct effects of environmental constraints on fitness prospects. Data on maintenance behaviour, height in canopy and spatial position were collected for five mothers in a small group and eight mothers in a large group. Comparisons were made on the basis of group size and dominance rank. Mothers from the large group spent less time feeding on clumped fruits and more time foraging on dispersed food items. Lower-ranking mothers, undergoing the highest rate of within-group competition, foraged most. For macaques in the study area, predation risk is considered to decrease with height in the canopy. Mothers in the small group, assumed to be more susceptible to predation, remained substantially higher in the canopy than mothers in the large group. In both groups, mothers adjusted their height to their spatial position in the group. Lower-ranking mothers were more often found without neighbours, presumably to avoid feeding competition. As a result they stay higher in the canopy than higher-ranking mothers. The results clearly demonstrate that individual females in the same reproductive state and living in the same area, but in different social environments, adapted their behaviour according to the hypothesis. Correspondence to: D.R. Vos     

Assessing hypotheses about nesting site occupancy dynamics

Hypotheses about habitat selection developed in the evolutionary ecology framework assume that individuals, under some conditions, select breeding habitat based on expected fitness in different habitat. The relationship between habitat quality and fitness may be reflected by breeding success of individuals, which may in turn be used to assess habitat quality. Habitat quality may also be assessed via local density: if high-quality sites are preferentially used, high density may reflect high-quality habitat. Here we assessed whether site occupancy dynamics vary with site surrogates for habitat quality. We modeled nest site use probability in a seabird subcolony (the Black-legged Kittiwake, Rissa tridactyla) over a 20-year period. We estimated site persistence (an occupied site remains occupied from time t to t+1) and colonization through two subprocesses: first colonization (site creation at the timescale of the study) and recolonization (a site is colonized again after being deserted). Our model explicitly incorporated site-specific and neighboring breeding success and conspecific density in the neighborhood. Our results provided evidence that reproductively "successful" sites have a higher persistence probability than "unsuccessful" ones. Analyses of site fidelity in marked birds and of survival probability showed that high site persistence predominantly reflects site fidelity, not immediate colonization by new owners after emigration or death of previous owners. There is a negative quadratic relationship between local density and persistence probability. First colonization probability decreases with density, whereas recolonization probability is constant. This highlights the importance of distinguishing initial colonization and recolonization to understand site occupancy. All dynamics varied positively with neighboring breeding success. We found evidence of a positive interaction between site-specific and neighboring breeding success. We addressed local population dynamics using a site occupancy approach integrating hypotheses developed in behavioral ecology to account for individual decisions. This allows development of models of population and metapopulation dynamics that explicitly incorporate ecological and evolutionary processes.     

“Philandering” — a mixed mating strategy in the splendid fairy-wren Malurus splendens

Summary In a population of the splendid fairy-wren Malurus splendens, we describe 202 intrusions by solitary adult males in full breeding plumage into the territories of conspecifics. Intrusions were not secretive and although silent they involved conspicuous flight above the vegetation and several characteristic display elements, including petal-carrying. Fifteen intrusions were observed in the 2 months before eggs were laid, but the rest occurred during the period of nest-building and egg-laying when the breeding female was presumed to be fertile, and were by males from adjacent territories. A high level of extra-pair fertilization has previously been determined in this species and we interpret many of these intrusions as attempts by males to obtain extra-pair copulations (EPC); we call this activity philandering. The costs and benefits of EPC for males and females are discussed and the promiscuous mating strategy of M. splendens is contrasted with the persistent social pairing in stable groups of this long-lived cooperatively breeding species.Offprint requests to: I. Rowley     

Does temporal and spatial segregation explain the complex population structure of humpback whales on the coast of West Africa?

Humpback whales (Megaptera novaeangliae) in the Southeastern Atlantic Ocean (International Whaling Commission ‘Breeding Stock B’—BSB) are distributed from the Gulf of Guinea to Western South Africa. Genetic data suggest that this stock may be sub-structured, but it remains unknown if this is due to reproductive segregation. This paper evaluates the spatial and temporal population structure of BSB humpback whales using a combination of maternally and bi-parentally inherited markers. The genetic differentiation that we identify in this study could be due to a combination of (1) spatial and/or temporal segregation on breeding grounds in the greater Gulf of Guinea, (2) the possibility of maternally inherited site fidelity to specific feeding grounds and (3) the use of two generalized but exclusive migratory routes (coastal and offshore) between feeding and breeding areas. Further, photo-identification and genetic sampling efforts in other areas of the Sub-Saharan Western Africa winter range and targeted deployment of satellite tags would help to clarify some of the apparent complexity in the population structure of animals biopsied in this region.