Genetic and ecological dynamics of species replacement in an arid-land river system

Museum records indicate that Hybognathus placitus was introduced into the Pecos River, New Mexico during the early 1960s. Approximately 10 years later, a congener, Hybognathus amarus, was extirpated from the system. We used microsatellite and mtDNA data, ecological data and modelling, and a computer simulation approach to reconstruct the history of invasion and species replacement. To identify the potential role of hybridization and introgression, we genetically screened H. amarus (n = 389) from the Rio Grande, New Mexico, and H. placitus (n = 424) from the Pecos River, New Mexico using four nuclear microsatellites and a partial fragment of the mtDNA ND4 gene. Assignment tests excluded hybridization as a primary factor in species replacement and suggested a role for interspecific competition. Genetic analyses showed that H. placitus were introduced into the Pecos River from at least two genetically distinct source populations in the Canadian and Red rivers, Oklahoma. Lotka-Volterra models of interspecific competition indicated that the number of founding individuals could have been as few as 20 for H. placitus to have competitively displaced H. amarus in the Pecos River in 10 to 15 generations. Observed differences of allele frequencies between source and founder populations indicated that between 32 and 115 H. placitus individuals founded the Pecos River. Genetic and ecological data suggest that interspecific competition could have led to species replacement in this arid-land river system.     

Introduced populations as genetic reservoirs for imperiled species: a case study of the Arkansas River Shiner (Notropis girardi)

The Arkansas River Shiner is a threatened species that has been extirpated throughout much of its native range (Arkansas River drainage) and remaining populations are imperiled. Prior to 1978, this species was accidently introduced to the Pecos River (Rio Grande drainage) via bait bucket, and has since persisted for over 30 years. Genetic data show that the Pecos River population maintains comparable levels of diversity at mitochondrial DNA and microsatellite loci relative to native range populations. Hence, we examined several factors that could be responsible for high introduced genetic diversity including (a) multiple introductions from genetically distinct sources (b) introduction of individuals from a genetically diverse source followed by rapid population expansion, (c) presence of life-history traits that foster propagule diversity and wide spatio-temporal demographic and genetic mixing; and (d) introduction to a suitable habitat in the non-native range. Our findings indicate Arkansas River Shiner was likely introduced from the Canadian River and subsequently experienced rapid population expansion that mitigated loss of diversity during the founding event. Threats to native Arkansas River Shiner have increased due to ongoing drought and water resource development, thus a finding of high diversity in the Pecos River suggests conservation significance of this non-native population. Further, it identifies the Pecos River as both a refuge for native endemic fishes and of genetic diversity of introduced, yet threatened, species.     

Do females adjust the sex of their offspring in relation to the breeding sex ratio?

When the adult sex ratio differs between years in local populations, but still is predictable between adjacent years, it has been proposed that the best strategy would be to bias the offspring sex ratio in favour of the rare sex. We tested this hypothesis using a data set of great reed warbler offspring, sexed by molecular techniques, that were collected over 11 breeding seasons at two adjacent reed marshes. Three important assumptions for this hypothesis are fulfilled in the studied great reed warbler population. First, a substantial proportion of great reed warblers are living in small local populations where sex ratio distortions would be sufficiently large and common. Second, breeding adults and their offspring return to breed in the local population to a high degree. Third, females have a possibility to assess the breeding sex ratio before laying their eggs. At our study site, the breeding sex ratio was positively correlated between successive years. However, contrary to our prediction, female great reed warblers seemed not to adjust their offspring sex ratio in relation to the local breeding sex ratio.     

Genetic effective size,Ne, tracks density in a small freshwater cyprinid,Pecos bluntnose shiner (Notropis simus pecosensis)

Genetic monitoring tracks changes in measures of diversity including allelic richness, heterozygosity and genetic effective size over time, and has emerged as an important tool for understanding evolutionary consequences of population management. One proposed application of genetic monitoring has been to estimate abundance and its trajectory through time. Here, genetic monitoring was conducted across five consecutive year for the Pecos bluntnose shiner, a federally threatened minnow. Temporal changes in allele frequencies at seven microsatellite DNA loci were used to estimate variance effective size (N_eV) across adjacent years in the time series. Likewise, effective size was computed using the linkage disequilibrium method (N_eD) for each sample. Estimates of N_e were then compared to estimates of adult fish density obtained from traditional demographic monitoring. For Pecos bluntnose shiner, density (catch‐per‐unit‐effort), N_eV and N_eD were positively associated across this time series. Results for Pecos bluntnose shiner were compared to a related and ecologically similar species, the Rio Grande silvery minnow. In this species, density and N_eV were negatively associated, which suggested decoupling of abundance and effective size trajectories. Conversely, density and N_eD were positively associated. For Rio Grande silvery minnow, discrepancies among estimates of N_e and their relationships with adult fish density could be related to effects of high variance in reproductive success in the wild and/or effects of supplementation of the wild population with captive‐bred and reared fish. The efficacy of N_e as a predictor of density and abundance may depend on intrinsic population dynamics of the species and how these dynamics are influenced by the landscape features, management protocols and other factors.     

Sex allocation and sex‐specific parental investment in an endangered seabird

Biased offspring sex ratio is relatively rare in birds and sex allocation can vary with environmental conditions, with the larger and more costly sex, which can be either the male or female depending on species, favoured during high food availability. Sex‐specific parental investment may lead to biased mortality and, coupled with unequal production of one sex, may result in biased adult sex ratio, with potential grave consequences on population stability. The African Penguin Spheniscus demersus, endemic to southern Africa, is an endangered monogamous seabird with bi‐parental care. Female adult African Penguins are smaller, have a higher foraging effort when breeding and higher mortality compared with adult males. In 2015, a year in which environmental conditions were favourable for breeding, African Penguin chick production on Bird Island, Algoa Bay, South Africa, was skewed towards males (1.5 males to 1 female). Males also had higher growth rates and fledging mass than females, with potentially higher post‐fledging survival. Female, but not male, parents had higher foraging effort and lower body condition with increasing number of male chicks in their brood, thereby revealing flexibility in their parental strategy, but also the costs of their investment in their current brood. The combination of male‐biased chick production and higher female mortality, possibly at the juvenile stage as a result of lower parental investment in female chicks, and/or at the adult stage as a result of higher parental investment, may contribute to a biased adult sex ratio (ASR) in this species. While further research during years of contrasting food availability is needed to confirm this trend, populations with male‐skewed ASRs have higher extinction risks and conservation strategies aiming to benefit female African Penguin might need to be developed.     

Effects of population structure and density on calf sex ratio in red deer (<Emphasis Type="Italic">Cervus elaphus</Emphasis>)—implications for management

There is ongoing interest to assess what factors affect offspring sex ratio, especially in ungulates. Wildlife managers might be interested in influencing this sex ratio for two reasons: either in order to limit population growth more effectively by reducing the proportion of females born or to increase revenues by a higher proportion of trophy bearing males in the population. While previous studies mostly focused on how maternal traits affect offspring sex ratio, we included here also male traits in our analysis. We achieved this by investigating data from 30 areas covering entire Lower Austria, collected over the past 12 years from both hunted red deer and those killed in road accidents. We focused our analyses on parameters that can be easily assessed by managers on the population and individual level, i.e. the numbers of animals culled in different age/sex classes and their body mass. We found that the proportion of females among calves increased with population density. Furthermore, we found that calf sex ratio (i.e. the proportion males among calves aged between 2 and 7 months) increased with increasing proportions of adult females and males older than 10 years, independent of the density effect. We conclude that wildlife managers interested in the effective reduction of red deer abundance and/or increasing the proportion of males among offspring should select a culling regime leading to a low population density dominated by adult, prime-aged females and males. This can be achieved by over-proportional removal of young females and warranting that a high number of strong males reach an age of at least 10 years.     

Sex Differences in Parietal Lobe Structure and Development

Structural magnetic resonance imaging studies provide evidence for sex differences in the human brain. Differences in surface area and the proportion of gray to white matter volume are observed, in particular in the parietal lobe. To our knowledge, no studies have examined sex differences in parietal lobe structure in younger populations or in the context of development. The present study evaluated sex differences in the structure of the parietal lobe in children aged 7 to 17 years. In addition, by adding a cohort of previously studied adults aged 18 to 50 years, sex differences in parietal lobe structure were examined across the age span of 7 to 50 years. Compared with the adult sample, the younger sample showed that the ratio of parietal lobe cortex to white matter was greater in female brains, but no sex differences in surface area. When examining the effects of age, surface area exhibited a significant sex-age interaction. In male brains, there was essentially no decrease in surfaces area over time, whereas in female brains, there was a significant decrease in surface area over time. These findings support the notion of structural sex differences in the parietal lobe, not only in the context of cross-sectional assessment but also in terms of differences in developmental trajectories.     

Estimating preharvest density,adult sex ratio,and fecundity of white‐tailed deer using noninvasive sampling techniques

Adult sex ratio and fecundity (juveniles per female) are key population parameters in sustainable wildlife management, but inferring these requires abundance estimates of at least three age/sex classes of the population (male and female adults and juveniles). Prior to harvest, we used an array of 36 wildlife camera traps during 2 and 3 weeks in the early autumn of 2016 and 2017, respectively. We recorded white‐tailed deer adult males, adult females, and fawns from the pictures. Simultaneously, we collected fecal DNA (fDNA) from 92 20 m × 20 m plots placed in 23 clusters of four plots between the camera traps. We identified individuals from fDNA samples with microsatellite markers and estimated the total sex ratio and population density using spatial capture–recapture (SCR). The fDNA‐SCR analysis concluded equal sex ratio in the first year and female bias in the second year, and no difference in space use between sexes (fawns and adults combined). Camera information was analyzed in a spatial capture (SC) framework assuming an informative prior for animals’ space use, either (a) as estimated by fDNA‐SCR (same for all age/sex classes), (b) as assumed from the literature (space use of adult males larger than adult females and fawns), or (c) by inferring adult male space use from individually identified males from the camera pictures. These various SC approaches produced plausible inferences on fecundity, but also inferred total density to be lower than the estimate provided by fDNA‐SCR in one of the study years. SC approaches where adult male and female were allowed to differ in their space use suggested the population had a female‐biased adult sex ratio. In conclusion, SC approaches allowed estimating the preharvest population parameters of interest and provided conservative density estimates.     

Superparasitism and sex ratio adjustment in a wasp parasitoid: results at variance with Local Mate Competition?

Anaphes nitens is a solitary parasitoid of the egg capsules of the Eucalyptus snout beetle, Gonipterus scutellatus. Some traits of its natural history suggest that Local Mate Competition (LMC) could account for sex ratio adjustment in this species. We tested whether males emerged early, a prerequisite for fully local mating, and investigated the occurrence and effect of superparasitism on adult size and pre-emergence mortality, factors that might influence sex ratio adjustment. We found in field-collected egg capsules that males emerged first. To investigate the effects of superparasitism on adult size, we compared the sizes of parasitoids that emerged early and late from egg capsules collected in the field, and from egg capsules parasitized and superparasitized in the laboratory. Superparasitism reduced parasitoid size, affecting females more strongly than males, and increased pre-emergence mortality. We estimated A. nitens sex ratio and parasitism rate in the field during 2 years in five localities and during 4 years in a sixth. Following LMC we expected an increase in sex ratio (proportion of males) with increasing parasitism rate (assumed to reflect parasitoid density). We found that sex ratio decreased from 0.38 when the parasitism rate was low (0-20%) to 0.21 when parasitism was high (80-100%). In contrast with field results, a laboratory experiment showed that: (1) at a low parasitism level sex ratio was clearly female biased (0.28+/-0.04), (2) at a high parasitism level sex ratio increased (0.40+/-0.07), (3) male larval survivorship was not lower than female survivorship, and (4) low-quality hosts (i.e. superparasitized) were allocated more males. We conclude that LMC cannot explain the sex ratio adjustment observed in the field, even at low parasitism rates, and alternative explications for highly female-biased sex ratios must be found. One such alternative is female-biased dispersal.     

Sex ratio at reproductive age: changes over the last century in the Italian population

The radical improvement in living conditions experienced in Italy during the last century caused a reduction in male extra-mortality during the prereproductive years. As a consequence, a progressive increase in the sex ratio at the beginning of the reproductive age (15-19 years) occurred, so that in recent times the sex ratio in the young adult population has approached the almost constant value of 1.06 observed at birth. We calculated that the sex composition would be the same in newborns and in young adults in about one generation: obviously, we have to assume that the sex differentials in mortality and migration are constant over time. The 1:1 equilibrium between sexes, which maximizes reproductive success, occurred in the 15-19 age group at the beginning of the century and shifted to the 30-35 age group in the 1990s. We compared the 1993-1995 sex ratios in different age groups in European Union countries and observed that in Italy as well as in other Mediterranean countries the numerical equality between sexes is reached at 30-35 years of age, while in north-central Europe it is reached later, approximately at the end of reproductive life.     

Sex-biased marine survival and growth in a population of coho salmon

Examination of historical records for coho salmon in Big Beef Creek, in western Washington, U.S.A., indicated that more adult males than females returned to spawn, and that the mean length of the females exceeded that of males. Sex-biased survival and faster growth among females are unusual among salmonids but precedented in some other coho salmon populations. To help determine the stage of life at which sex-biased mortality might occur, the sex-linked GH-ψH pseudogene was used to determine the sex of smolts emigrating from Big Beef Creek in 1995–1997. In each of the 3 years the sex ratio was indistinguishable from 50:50, indicating similar survival rates in fresh water, and implying that the male and female coho salmon follow different foraging strategies when they are at sea. The female strategy apparently results in greater mortality, but benefits survivors with greater size. The male strategy appears to allow greater survival at the cost of reduced size.     

Density-related changes in sexual selection in red deer.

In sexually dimorphic mammals, high population density is commonly associated with increased mortality of males relative to females and with female-biased adult sex ratios. This paper investigates the consequences of these changes on the distribution of male breeding success, the intensity of competition for females and the opportunity for sexual selection. After the red deer (Cervus elaphus L.) population of the North Block of Rum (Inner Hebrides) was released from culling, female numbers rose and male numbers declined, leading to an adult sex ratio of around one male to two females. This change was the result of increased mortality of males relative to females during the first two years of life; of increased emigration rates by young males; and of reduced immigration by males from outside the study area. The increasing bias in the adult sex ratio affected the timing of breeding as well as the distribution of mating success in males. As the adult sex ratio became increasingly biased towards females, the degree of skew in mating success (calculated across all harem-holders) increased, but mature males defended harems for shorter periods and a higher proportion of males held harems. In addition, a higher proportion of calves were fathered by immigrant males and the proportion fathered by males born in the study area declined. These results support the contention that, where high population density is associated with a female-biased adult sex ratio, competition for mates is likely to decline.     

Successful adult willow grouse are exposed to increased harvest risk

Age and sex ratios in bag records are frequently used as indices of population composition for harvested populations. However, vulnerability to harvest may differ by age and sex thereby producing bias in population estimates. We assessed whether age and sex affected vulnerability to harvest for willow grouse (Lagopus lagopus) where adult density and brood size were known in the harvested populations. We collected bag records during 2 days of controlled hunting in 4 areas in 2 years (2007 and 2008) in Jämtland county, Sweden. We found that vulnerability to harvest was different for chicks and adults, but not between male and female adults. Hunters encountered broods at a higher rate than single birds compared to personnel conducting pre-harvest counts along line transects. Furthermore, the probability of shooting a grouse was higher in encounters of broods than individual grouse. Proportionally, we calculated about a 50% probability of a hunter shooting either a chick or an adult independent of encountering a single bird or broods of 2–10 grouse. Increasing adult density also increased the vulnerability to harvest for adults relative to chicks, independent of the chick to adult ratio in the pre-harvest population. The different vulnerability of adults and chicks to harvest observed in this study will dampen variation in age classes in bag records compared to the population, and we caution against extrapolation of age ratios in bag records to harvested populations. © 2012 The Wildlife Society.     

Lion (Panthera leo) demographics in the south-western Kgalagadi Transfrontier Park

Changes in demographic features of the lion population in the south-western Kgalagadi Transfrontier Park cause conservation concern. The ratio between adult males and females changed from 1♂:2♀ to 1♂:1♀ over a 32-year period (1977–2010). This is atypical for undisturbed lion populations. We evaluated mechanisms that on their own or together could explain the trend by using both a discrete- and continuous-time capture–recapture model to analyse 2 years of individual-based observation data (May 2013–June 2015). Although most vital rates were within expected parameters, shifts in the sex structure remained similar to that reported for sub-adults previously. The population comprised of 26.9% cubs and juveniles (<2 years), 14.2% sub-adults (2–4 years) and 59% adults (>4 years). Litters had equal sex ratios at first detection. Birth rates were lower (0.44 cubs per female per year) than between 1998 and 2001 (0.69 cubs per female per year). The continuous model indicated a higher survival effect for female cubs, while both models showed higher survival rates/effects for sub-adult and adult females relative to males. However, the sex ratio for sub-adults was male-biased. The shift in demographic signals may associate with changes in age-related immigration rates of sub-adult males.     

Climate effects on offspring sex ratio in a viviparous lizard

1. Understanding individual and population responses to climate change is emerging as an important challenge. Because many phenotypic traits are sensitive to environmental conditions, directional climate change could significantly alter trait distribution within populations and may generate an evolutionary response. 2. In species with environment-dependent sex determination, climate change may lead to skewed sex ratios at hatching or birth. However, there are virtually no empirical data on the putative link between climatic parameters and sex ratios from natural populations. 3. We monitored a natural population of viviparous lizards with temperature-dependent sex determination (Niveoscincus ocellatus) over seven field seasons. Sex ratios at birth fluctuated significantly among years and closely tracked thermal conditions in the field, with the proportion of male offspring increasing in colder years. 4. This is the first study to demonstrate the effect of local climatic conditions (e.g. temperature) on offspring sex ratio fluctuations in a free-living population of a viviparous ectotherm. A succession of warmer-than-usual years (as predicted under many climate-change scenarios) likely would generate female-biased sex ratios at birth, while an increase in interannual variation (as also predicted under climate change scenarios) could lead to significant fluctuations in cohort sex ratios. If cohort sex ratio bias at birth leads to adult sex ratio bias, long-term directional changes in thermal conditions may have important effects on population dynamics in this species.     

Haematological profile in healthy urban population (8 to 70 years of age)

Haematological profile for 17 constituents of blood were determined in 998 healthy school children (8-19 years old) and 2246 healthy adult persons (20-70 years old) residing permanently or at least 5 years in a defined geographic region of Zagreb, Croatia. Physiological variations corresponding to age and sex were studied as the most important factors affecting biological variation in haematological constituents of blood. In our study erythrocytes, haemoglobin and haematocrit values were not sex dependent until the age of 13 after which the values were higher in men that in women. Sedimentation rate showed sex and age related differences in the adult age with higher values in women especially after 50 years. Total leukocyte count declined with age and in adults the values were slightly lower in women. Segmented neutrophil granulocytes showed the upward trend with age whereas the lymphocyte and monocyte counts declined. Women showed slightly higher platelet count in the adult age. Based on biological variation, we have estimated the reference intervals for 17 haematological constituents of blood in order to provide medically reliable evaluation of haematological laboratory results.     

Male life history and the unusual adult sex ratios of redfronted lemur, Eulemur fulvus rufus, groups

In group-living species, theoretical considerations indicate the existence of a fundamental conflict of interest between the sexes over the adult sex ratio within groups. Females may derive certain benefits from living with many males. Males, in contrast, should generally try to monopolize access to a group of females. Which sex ultimately controls adult group sex ratio is poorly known. We examined this conflict between the sexes in redfronted lemurs, Malagasy primates characterized by an unusual lack of female-biased adult sex ratios. Using various demographic and behavioural data from several groups collected over 6 years, we examined (1) the proximate determinants of this unusual sex ratio, (2) the temporal distribution of female fertile phases within groups as a determinant of male monopolization potential, (3) sources of between-group variation in the adult sex ratio, and (4) possible social benefits of the relatively high number of males for both sexes. Birth and mortality rates were not sex biased and males migrated considerably more frequently than females, providing no proximate explanation for the unusual sex ratio. However, certain life history traits (fast maturation, short interbirth intervals) may ultimately play a role because they act to facilitate joint group transfers of male coalitions. Despite a relatively small female group size and an associated high monopolization potential, female oestrous synchrony may prevent the formation of single-male groups. Reduced male group size seems to be the main predictor of take-over rate, and, thus, infanticide risk, suggesting that both sexes may benefit from the high number of coresident males, thereby considerably defusing the conflict of interest between the sexes.     

Male‐skewed adult sex ratio,survival, mating opportunity and annual productivity in the Snowy Plover Charadrius alexandrinus

Sex differences in adult mortality may be responsible for male‐skewed adult sex ratios and male‐skewed parental care in some birds. Because a surplus of breeding males has been reported in serially polyandrous populations of Snowy Plover Charadrius alexandrinus, we examined sex ratio, early‐season nesting opportunities, adult survival and annual reproductive success of a Snowy Plover population at Monterey Bay, California. We tested the hypotheses that male adult survival was greater than female survival and that a sex difference in adult survival led to a skewed adult sex ratio, different mating opportunities and different annual productivity between the sexes. Virtually all females left chicks from their first broods to the care of the male and re‐nested with a new mate. As a result, females had time to parent three successful nesting attempts during the lengthy breeding season, whereas males had time for only two successful attempts. Among years, the median population of nesting Plovers was 96 males and 84 females (median difference = 9), resulting in one extra male per eight pairs. The number of potential breeders without mates during the early nesting period each year was higher in males than in females. Adult male survival (0.734 ± 0.028 se) was higher than female survival (0.693 ± 0.030 se) in top‐ranked models. Annually, females parented more successful clutches and fledged more chicks than their first mates of the season. Our results suggest that in C. alexandrinus a sex difference in adult survival results in a male‐skewed sex ratio, which creates more nesting opportunities and greater annual productivity for females than for males.     

Food restricting young hamsters (Mesocricetus auratus) affects sex ratio and growth of subsequent offspring

Trivers and Willard (1973) predicted that stressed adult female mammals may enhance their fitness by skewing offspring sex ratios and maternal investment to favor daughters. The present study investigated whether stressing young hamsters might also influence sex ratio and growth of subsequent offspring. Control females received food ad libitum (A) on Days 1-50 postpartum (AA). Experimental females were food-restricted (R) either on Days 1-25 (RA), Days 26-50 (AR), or Days 1-50 (RR) postpartum. Subjects were mated when 91-95 days old. Litter sizes and survivorship (= % litters within a treatment that contained at least one pup), sex ratio (= % males), and pup weights in the next generation were recorded every fifth day from parturition until Day 25 postpartum. Control litters contained significantly more offspring at birth than did RR litters. Sex ratio was significantly higher at birth for AA litters than for the other treatments. Postpartum sex ratio within each group remained similar to that recorded at birth. RR litters contained significantly fewer pups compared to the other three treatments from Days 5-25. RR female pups weighed significantly more at birth than their counterparts in the other treatments. Weights of males at birth were similar in all treatments. By Day 25, both male and female RR pups weighed significantly less than control, AR, and RA pups. Food restriction early in life may have long-term consequences on sex ratio and pup growth in golden hamsters.     

Sexual dimorphism and intraspecific aggression,and their relationship to sex ratios in Caprella gorgonia Laubitz & Lewbel (Crustacea: Amphipoda: Caprellidae)

The amphipod Caprella gorgonia Laubitz & Lewbel is an obligate commensal on gorgonian octocorals. Its primary host is Lophogorgia chilensis (Verrill), found below 20 m.C. gorgonia breeds throughout the year, with wide fluctuations in abundance. Mating and oviposition follow molting. Sex reversal does not occur; two distinct sexes are present from the first instar after emergence from the brood pouch.Young males and females grow at approximately the same rate, but males are larger by a relatively constant increment. Males continue to grow at their original rate to a maximum size (about twice that of females). The growth rate of females is not limited by the onset of reproduction and brooding, but rather by an approach to maximum size when the rate is greatly reduced. Fecundity of females is not affected by size.The population sex ratio is about 1:3 (males:females), and about 1:4 among adults. The secondary sex ratio is 1:1. The post-emergence sex ratio bias is a result of heavier mortality among males. Sex ratios drop from 50% at emergence to 25% as females approach maximum size, then rise to 100% in larger size classes.Differential predation on males did not appear to be a source of any sex ratio bias. Adult males possess a “poison spine”, a puncturing weapon on the large second gnathopod, which functions in mating-related intraspecific combat with other males. Intraspecific male aggression during mating is a major cause of sex ratio bias. In the laboratory, increased density in breeding groups may affect mortality due to male aggression. In nature, adult sex ratios are negatively correlated with population density. The reproductive capacity of the population is not limited by a shortage of adult males, despite the low adult sex ratio.