Mother–offspring distances reflect sex differences in fine‐scale genetic structure of eastern grey kangaroos

Natal dispersal affects life history and population biology and causes gene flow. In mammals, dispersal is usually male‐biased so that females tend to be philopatric and surrounded by matrilineal kin, which may lead to preferential associations among female kin. Here we combine genetic analyses and behavioral observations to investigate spatial genetic structure and sex‐biased dispersal patterns in a high‐density population of mammals showing fission–fusion group dynamics. We studied eastern grey kangaroos (Macropus giganteus) over 2 years at Wilsons Promontory National Park, Australia, and found weak fine‐scale genetic structure among adult females in both years but no structure among adult males. Immature male kangaroos moved away from their mothers at 18–25 months of age, while immature females remained near their mothers until older. A higher proportion of male (34%) than female (6%) subadults and young adults were observed to disperse, although median distances of detected dispersals were similar for both sexes. Adult females had overlapping ranges that were far wider than the maximum extent of spatial genetic structure found. Female kangaroos, although weakly philopatric, mostly encounter nonrelatives in fission–fusion groups at high density, and therefore kinship is unlikely to strongly affect sociality.     

Synergistic effects of seasonal rainfall, parasites and demography on fluctuations in springbok body condition

1. Seasonality of rainfall can exert a strong influence on animal condition and on host-parasite interactions. The body condition of ruminants fluctuates seasonally in response to changes in energy requirements, foraging patterns and resource availability, and seasonal variation in parasite infections may further alter ruminant body condition. 2. This study disentangles the effects of rainfall and gastrointestinal parasite infections on springbok (Antidorcas marsupialis) body condition and determines how these factors vary among demographic groups. 3. Using data from four years and three study areas, we investigated (i) the influence of rainfall variation, demographic factors and parasite interactions on parasite prevalence or infection intensity, (ii) whether parasitism or rainfall is a more important predictor of springbok body condition and (iii) how parasitism and condition vary among study areas along a rainfall gradient. 4. We found that increased parasite intensity is associated with reduced body condition only for adult females. For all other demographic groups, body condition was significantly related to prior rainfall and not to parasitism. Rainfall lagged by two months had a positive effect on body condition. 5. Adult females showed evidence of a 'periparturient rise' in parasite intensity and had higher parasite intensity and lower body condition than adult males after parturition and during early lactation. After juveniles were weaned, adult females had lower parasite intensity than adult males. Sex differences in parasitism and condition may be due to differences between adult females and males in the seasonal timing of reproductive effort and its effects on host immunity, as well as documented sex differences in vulnerability to predation. 6. Our results highlight that parasites and the environment can synergistically affect host populations, but that these interactions might be masked by their interwoven relationships, their differential impacts on demographic groups, and the different time-scales at which they operate.     

Short‐term changes in body condition in relation to habitat and rainfall abundance in American redstarts Setophaga ruticilla during the non‐breeding season

In migratory birds, environmental conditions during the stationary period of the non‐breeding season are crucial to consider because they ultimately affect the fitness of individuals by influencing their subsequent migration, breeding success and survival. Although a few studies have investigated the influence of non‐breeding habitat on the capacity of individuals to cope with long‐term seasonal rainfall fluctuations, it remains unknown how habitat quality and variations in rainfall abundance–at a monthly scale–interact to affect non‐breeding condition of migrating birds. In this study, we examined the influence of monthly changes in rainfall abundance on body condition of non‐breeding female redstarts Setophaga ruticilla living either in a high quality habitat (mangrove) or in a low quality habitat (scrub). Body condition of both mangrove and scrub redstarts showed important variations over the study period, demonstrating for the first time that body condition of non‐breeding female redstarts can change rapidly in response to short‐term fluctuations in rainfall. Importantly, we found that female redstarts living in mangrove were usually in better condition during periods of low rainfall compared to females living in scrub. However, body condition did not differ between mangrove females and scrub females during an episode of frequent, heavy precipitation. Importantly, our study also demonstrated that the duration of a perturbation is an important determinant of body condition in redstarts since a prolonged drought resulted in similar low body condition for birds from both habitats. Age was not correlated with body condition whatever the habitat and the rainfall conditions. Our results demonstrate that high quality habitat can temporarily reduce the deleterious effect of a short‐term drought on body condition, but also, that a habitat of low quality does not constrain individuals when climatic conditions are optimal.     

Sexual segregation in western grey kangaroos: testing alternative evolutionary hypotheses

In sexually dimorphic ungulates, sexual segregation is hypothesized to have evolved because of sex-specific differences in body size and/or reproductive strategies. We tested these alternative hypotheses in kangaroos, which are ecological analogues of ungulates. Kangaroos exhibit a wide range of body sizes, particularly among mature males, and so the effects of body size and sex can be distinguished. We tested predictions derived from these hypotheses by comparing the distribution of three sex–sex size classes of western grey kangaroos Macropus fuliginosus , in different habitats, and the composition of groups of kangaroos, across seasons. In accordance with the predation risk-reproductive strategy hypothesis, during the non-breeding season, females, which were more susceptible to predation than larger males, and were accompanied by vulnerable young-at-foot, were over-represented in secure habitats. Large males, which were essentially immune to predation, occurred more often than expected in nutrient-rich habitat, and small males, which faced competing demands of predator avoidance and feeding, were intermediate between females and large males in their distribution across habitats. During the breeding season, females continued to be over-represented in secure habitats when their newly emerged pouch young were most vulnerable to predation. All males occupied these same habitats to maximize their chances of securing mates. Consistent with the social hypotheses, groups composed of individuals of the same sex, irrespective of body size, were over-represented in the population during the non-breeding season, while during the breeding season all males sought females so that mixed-sex groups predominated. These results indicate that body size and reproductive strategies are both important, yet independent, factors influencing segregation in western grey kangaroos.     

Demography of lions in relation to prey and habitat in the Maasai Mara National Reserve, Kenya

We studied lion demography in the Maasai Mara National Reserve between September 1990 and April 1992, with a special emphasis on the spatial and seasonal variation in demographic characteristics. Lion density (0.2–0.4 lions km^?2) and pride size (range 8–48) were high because of a high resident prey biomass (10 335 kg km^?2) augmented by migrant prey to 26 092 kg km^?2 in the dry season. Overall, their sex ratio was almost at parity and varied neither spatially nor seasonally. Sex ratio was even among subadults but skewed toward males and females among cubs and adults, respectively. This implies an increasing differential mortality of males with age through subadulthood. The age ratio varied seasonally because of a birth peak in March–June and an influx of subadults into the reserve during July–August, coincident with increases in migrant prey. The birth peak was apparently preceded by another peak in mating activity falling between November and May. Further research should investigate the precise causes of the biased cub sex ratio, low lion density in the Mara Triangle and the higher ratio of subadults in Musiara than in the Mara Triangle or Sekenani.     

Effect of rainfall on biomass,growth and dieback of pastures in an arid grazing system

The influence of rainfall and initial pasture biomass on growth and dieback of pastures in temporary enclosures over quarterly periods was examined from 1980 to 1985 at Menindee in western New South Wales. Rainfall during the previous 12 months had the most influential effect on biomass of grazed pasture, and accounted for 69% of the variance. During quarterly periods, when the biomass of pasture protected from grazing increased, rainfall accounted for 93% of the variance. For all quarterly periods (whether pasture biomass increased or decreased) current rainfall accounted for 72% of the variance in biomass change (growth minus senescence) and initial biomass accounted for a further 14%. The growth response to rainfall was similar in pastures grazed by a combination of sheep and kangaroos and in those grazed by kangaroos alone.     

Lifetime individual plasticity in body condition of a migratory bird

Body condition is an important trait that reflects the capacity of individuals to acquire food or resist disease, eventually allowing successful reproduction and survival. We first quantified the effects of condition on life history and a secondary sexual trait using long‐term lifetime data on condition in a migratory bird species, the barn swallow, Hirundo rustica. Second, we quantified sources of individual variation in condition by separating the within‐individual from the between‐individual components of variation of environmental condition and age on body condition. Therefore, we were able to partition variation as a result of selection and phenotypic plasticity. Within‐individual variation in body condition increased in early life until middle age (i.e. 3–4 years of age) in the two sexes followed by only a slight decrease in body condition during senescence in males but not in females. After accounting for age‐dependent variation, condition could be partitioned into a within‐individual plastic response to environmental conditions during migration and a nonplastic response (i.e. a between‐individual difference) to environmental conditions experienced in the African winter quarters. Specifically, there was a within‐individual increase in body condition as environmental conditions during migration improved in both males and females, independent of age. There was a between‐individual effect of condition found in the winter quarter in body condition of females, but not in males, which was attributed to the disappearance of females in poor body condition from the study population as a result of the higher natal dispersal of low‐quality females compared to high‐quality ones during years with favourable environmental conditions in the African winter quarters. Males and females also tended to be in better body condition during the warmer springs upon arrival at the breeding grounds. There was a temporal decline in female body condition during 1991–2007, whereas no significant trend was detected in males. Therefore, both intrinsic (e.g. age and sex) and extrinsic factors (e.g. climate) affected body condition. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 420–434.     

Female mate choice is not affected by mate condition in a fish with male care

Several studies have shown that mate choice based on condition leads to higher reproductive success of the choosing individual. Yet, a growing body of literature has failed to find support for mate choice based on mate condition, even when the choosing individual would clearly benefit from such a choice. This indicates that animals’ mate choice is often more complex than currently appreciated and that even well-founded expected preferences cannot be taken for granted. Using the broad-nosed pipefish, Syngnathus typhle, we manipulated male condition experimentally to explore whether it affects female mate choice. In this sex-role-reversed species, males care for the offspring in a specialised brood pouch. Males are the choosier sex, but given the opportunity, females are selective as well. During brooding, males can both provide embryos with nutrients and take up nutrients that originate from eggs deposited in the pouch, and embryo survival correlates positively with male condition. Together, this suggests that it would be beneficial for females to mate with males in high condition. However, we found no female preference for males in better condition. Thus, this study adds to the literature of mate choice that is unaffected by mate condition. Possible reasons for our result are discussed.     

The effects of a severe drought on the morphology of the adrenal glands and other indices of health in the red kangaroo Megaleia rufa (Desmarest) in north-western New South Wales*

Drought adversely affects health and reproduction in red kangaroos (Megaleia rufa) (Desmarest). There is a significant drop in body weight in adult kangaroos, and growth rates of pouch young are depressed. In adult males both lobes of the pituitary gland decrease in size. All zones in the adrenal glands become smaller and lipid stores increase. In adult females the posterior lobe of the pituitary increases in size and, although all zones in the adrenal glands shrink the evidence suggests continued stimulation of the Zonae fasciculata and glomerulosa. The decrease in size of the adrenal glands is accompanied by cortical folding and nodulation and other structural changes. Growth curves of pouch young are shown to fit the Johnson-Schumacher curve Y = A * exp [-b/(x + k)]. The data substantiate the findings of earlier workers but suggest that anoestrous female red kangaroos suckling small young may be better adapted to withstand drought than males.     

Endogenous nitrogen excretion by red kangaroos (Macropus rufus): effects of animal age and forage quality

Red kangaroos (Macropus rufus) are large (>20 kg) herbivorous marsupials common to arid and semiarid Australia. The population dynamics of red kangaroos are linked with environmental factors, operating largely through juvenile survival. A crucial period is the young-at-foot (YAF) stage, when juveniles have permanently left the mother's pouch but still take milk from a teat in the pouch. Forage quantity and quality have been implicated in drought-related mortalities of juvenile kangaroos. Here we compared how forage quality affected nitrogen (N) intake and excretion by YAF, weaned, and mature, nonlactating female red kangaroos. On high-quality forage (chopped lucerne hay, Medicago sativa) low in neutral-detergent fiber (43%+/-1%) and high in N (2.9%+/-0.1%), YAF and weaned kangaroos had ideal growth rates and retained 460-570 mg dietary N kg(-0.75) d(-1). But on poor-quality forage (chopped oaten hay, Avena sativa) high in neutral-detergent fiber (64%+/-1%) and low in N (0.9%+/-0.1%), YAF and weaned kangaroos could not sustain growth and were in negative N balance at -103+/-26 mg and -57+/-31 mg N kg(-0.75) d(-1), respectively. Notably, the YAF kangaroos excreted 64% of their truly digestible N intake from forage as nondietary fecal N (NDFN). By weaning age, the situation had improved, but the juveniles still lost 40% of their truly digestible N intake as NDFN compared with only 30% by the mature females. Our findings support field observations that forage quality, and not just quantity, is a major factor affecting the mortality of juvenile red kangaroos during drought.     

Sex-specific differences in body condition indices and seasonal mass loss in Tufted Puffins

ABSTRACT.   Reduced prey availability can affect the growth and survival of nestling seabirds. However, few studies have demonstrated similar effects on indices of adult body condition. We examined body condition and seasonal mass loss of breeding adult male and female Tufted Puffins ( Fratercula cirrhata ) at Chiniak Bay, Kodiak Island, Alaska, in 2004–2005. We determined sex using genetic analysis, developed a discriminant function to determine sex using morphometric measurements, and examined the body condition of adult males and females relative to the growth rates of their offspring. We found that morphological measurements were only moderately useful for sexing Tufted Puffins, with 74% of adults ( N = 176) correctly classified. We also found that the relationship between adult body mass and size differed between sexes and conclude that body condition indices must be calculated separately for each sex to avoid inter- and intrasexual bias. Body condition of male and female Tufted Puffins declined during the chick-rearing period. However, body condition of females did not differ between years, whereas male condition declined to a greater degree during 2004 when the mass of young at fledging was significantly lower. Although these results suggest that adult male Tufted Puffins sacrifice their own body condition in years of diminished nestling growth and females do not, reasons for this apparent intersexual difference in reproductive strategies remain unclear.     

The role of forage availability on diet choice and body condition in American beavers (Castor canadensis)

Forage availability can affect body condition and reproduction in wildlife. We used terrestrial and aquatic vegetation sampling, stable isotope analysis, and livetrapping to investigate the influence of estimated forage biomass on diet, body condition, and reproduction in American beavers (Castor canadensis) in the Namakan Reservoir, Voyageurs National Park, Minnesota, USA, May 2008–September 2009. Available terrestrial and emergent aquatic forage varied greatly among territories, but floating leaf aquatic forage was low in abundance in all territories. Variation in estimated biomass of available emergent and terrestrial vegetation did not explain variation in respective assimilated diets, but variation in floating leaf vegetation explained 31% of variation in assimilated floating leaf diets. No models using available vegetation explained variation in body condition. Body condition of individual females in spring did not affect kit catch per unit effort, and overwinter body condition of subadults and adults was similar between territories with and without kits. We found no evidence that available aquatic vegetation affected beaver body condition or fitness. Available forage may be above minimum thresholds to detect differences in diet choice or body condition. Other factors such as water level fluctuations or climatic variables may also explain variation in beaver body condition.     

Changes in body mass and feeding behaviour in male harbour seals, Phoca vitulina, in relation to female reproductive status

Loss of body mass has been used as an index of the cost of reproductive effort in mammals. We studied changes in body mass of male harbour seals, Phoca vitulina , during the breeding season on Sable Island, Nova Scotia, Canada. Individually marked subadult ( n = 21) and adult ( n = 22) males were captured at approximately six-day intervals throughout the breeding season. Adult males weighed an average of 108 kg ± 5·6 (S.E.) at initial capture, whereas subadults weighed an average of 76 ± 6·6 kg. The rate of mass loss by adult males did not differ from zero during the pre-mating period (i.e. the period without receptive females), but differed significantly from zero (-0·91 ± 0·007 kg/day) during the mating period (i.e. the period with receptive females). By contrast, the rate of mass change of subadults did not differ from zero in either period, indicating that food availability was unlikely to be responsible for the observed changes in adults. Adult males lost up to 24% of body mass during the breeding season. Examination of sera for the presence of chylomicrons (i.e. evidence of recent feeding) also suggested that adults stopped feeding during the mating period, while subadults did not. These results suggest that reproduction represents a significant energetic cost to adult male harbour seals.     

Using time-dependent models to investigate body condition and growth rate of the giant gartersnake

Identifying links between phenotypic attributes and fitness is a primary goal of reproductive ecology. Differences in within-year patterns of body condition between sexes of gartersnakes in relation to reproduction and growth are not fully understood. We conducted an 11-year field study of body condition and growth rate of the giant gartersnake Thamnophis gigas across 13 study areas in the Central Valley of California, USA. We developed a priori mixed effects models of body condition index (BCI), which included covariates of time, sex and snout–vent length and reported the best-approximating models using an information theoretic approach. Also, we developed models of growth rate index (GRI) using covariates of sex and periods based on reproductive behavior. The largest difference in BCI between sexes, as predicted by a non-linear (cubic) time model, occurred during the mating period when female body condition (0.014±0.001  se ) was substantially greater than males (−0.027±0.002  se ). Males likely allocated energy to search for mates, while females likely stored energy for embryonic development. We also provided evidence that males use more body energy reserves than females during hibernation, perhaps because of different body temperatures between sexes. We found GRI of male snakes was substantially lower during the mating period than during a non-mating period, which indicated that a trade-off existed between searching for mates and growth. These findings contribute to our understanding of snake ecology in a Mediterranean climate.     

川西平原大足鼠的种群生态学Ⅰ.种群动态和个体大小

我们利用标志重捕、夹捕解剖和笼养等三种方法于１９８９年１月至１９９５年６月对川西平原的大足鼠的种群密度、个体大小、生长和生物量进行了研究。种群密度最高１１．３只／ｈａ,最低０．７只／ｈａ,平均３．６只／ｈａ,并显示出与农作物相同的季节变动特征。雌雄性比由幼年组和亚成体组约１：１到成体组１：０．７８到老年组１：０．５３。雌雄个体的平均质量分别为１３６ｇ和１１４ｇ,两性的个体质量和后足长有明显差异,体长和耳长差异不明显。饲养的幼体及亚成体个体质量和尾长均持续增长,并可分别用Ｌｏｇｉｓｔｉｃ和对数方程较好地拟合。生物量则显示出比种群密度和个体质量更稳定的特征。     

Body mass, age and sexual maturity in short-beaked echidnas, Tachyglossus aculeatus

During the course of this 12 year field study body masses of 11 hatchling echidnas (Tachyglossus aculeatus multiaculeatus) and 25 pouch young between the ages of 5 and 60 days were recorded. Body mass increased from 0.3 to approximately 50 g in the first half of pouch life. It then quadrupled before young were placed in a burrow at 45 to 55 days of age. There was a positive correlation between the body mass of the female and that of her young at weaning. From 33 subadult echidnas located, tagged and radio tracked during this study, body masses of 10 were monitored to sexual maturity, i.e. when first encountered in a courtship train. Minimum age of sexual maturity ranged between 5 and 12 years. As subadults, there was no difference between mean body masses of males and females. At sexual maturity, mean body mass of females was significantly higher. No correlation was found between age at sexual maturity and body mass nor was there a significant difference in age of males and females at sexual maturity.     

Mate Switching in a Non-monogamous Species? The Case of the Common Quail (Coturnix coturnix)

We investigated the occutrence of mate switching in the common quail, a non-monogamous species with a temporal pair bond but without either male parental care or territoriality. The study was carried out throughout the breeding seasons of 1993–95 in Mas Esplugues (Catalonia, Spain) by monitoring 28 radio-tagged couples and 17 radio-tagged unpaired males. Agonistic interactions between males inside funnel traps containing a female were also recorded. Mate switching within the laying period occurred in 72% of the females, and new partners always had a higher body condition index than old partners. Agonistic interactions inside the funnel traps also showed that successful males had a better body condition index than losers. These results, along with the observed mate-guarding behaviour of females by their partners throughout the laying period, a highly male-biased sex ratio (five males per female), the lack of territoriality of males and the expected difficulties which unmated males experience in finding pairs, suggest that mate switching is not induced by paired males. Moreover, the constant inflow of new males observed throughout the fertile period of the female and the low costs stemming from mate change strongly support the idea that it is paired females who induce mate switching, in order to improve their fitness by mating with the best quality male available at every moment of their fertile period.     

Sexual selection on forelimb muscles of western grey kangaroos (Skippy was clearly a female)

Studies of sexual selection have tended to concentrate on obvious morphological dimorphisms such as crests, horns, antlers, and other physical displays or weapons; however, traits that show no obvious sexual dimorphism may nevertheless still be under sexual selection. Sexual selection theory generally predicts positive allometry for sexually selected traits. When fighting, male kangaroos use their forelimbs to clasp and hold their opponent and, standing on their tail, bring up their hind legs to kick their opponent. This action requires substantial strength and balance. We examined allometry of forelimb musculature in male and female western grey kangaroos (Macropus fuliginosus) to determine whether selection through male–male competition is associated with sex differences in muscle development. Forelimbs of males are more exaggerated than in females, with relatively greater muscle mass in males than the equivalent muscles in females. Furthermore, while muscles generally showed isometric growth in female forelimbs, every muscle demonstrated positive allometry in males. The significant positive allometry in male forelimb musculature, particularly those muscles most likely involved in male–male combat (a group of muscles involved in grasping: shoulder adduction, elbow flexion; and pulling: arm retraction, elbow flexion), clearly suggests that this musculature is subject to sexual selection. In addition to contributing to locomotion, the forelimbs of male kangaroos can also act as a signal, a weapon, and help in clasping, features that would contribute towards their importance as a sexually selected trait. Males would therefore benefit from well‐developed musculature of the arms and upper body during competition for mates. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 923–931.     

Site fidelity of Svalbard polar bears revealed by mark-recapture positions

Quantifying the degree of site fidelity in polar bears’ (Ursus maritimus) use of denning and mating areas in spring is of considerable interest for both basic and applied purposes. We analyzed 276 spring-to-spring movements (displacements) of 178 polar bears obtained from mark-recapture in the period 1987–2010 in Svalbard, Norway. Male and female subadults and adults showed site fidelity (only subadult females non-significantly) when their movements were compared to a scenario of random movements between all capture locations. The median observed displacement was 47.6 km (bootstrapped 95 % CI: 38.6–57.8 km), significantly smaller than the median potential displacement for random movements of 206.3 km (bootstrapped 95 % CI: 187.3–219.6 km). Subadult females tended to have the longest displacements, followed by adult males and adult females. However, large individual variation both in displacement distances and recapture frequency tended to blur age and sex differences. Analysis restricted to one movement for each bear showed that the difference between adult males and adult females was small and non-significant. This indicates that findings based on telemetry, which is almost always restricted to females, may be relatively representative of the whole adult Barents Sea population in the spring season.     

Female body condition and brood sex ratio in Yellow-legged Gulls Larus cachinnans

In the Yellow-legged Gull Larus cachinnans , males are the larger sex, and show more reproductive variance than females. We predicted that the proportion of male chicks in a brood should increase with female body condition. We investigated brood sex ratio by using DNA markers taken from samples of hatchlings or dead embryos, and female body condition using plasma cholesterol concentration as a reliable indicator. The brood sex ratio of females in good condition was male biased and the sex ratio of females in poor condition was female biased. This relationship was also significant in those nests where all the eggs laid were sexed. Thus, manipulation of embryo mortality cannot explain the biases reported in this study, suggesting that the sex ratio of the eggs was biased prior to laying. These results confirm that sex-ratio manipulation in gulls operates under natural conditions, and supports earlier experimental findings.