Isolation and characterization of microsatellite loci in the great bustard,Otis tarda

With the aim to study population genetics of the endangered great bustard, Otis tarda, dinucleotide microsatellite loci were isolated using an adapted hybrid‐capture enrichment protocol. This work reports the characterization of a set of six polymorphic microsatellite markers within the great bustard (n = 52). Results from cross‐species amplifications in several other members of the family Otididae demonstrate that five primer pairs also successfully amplified homologous loci outside the species Otis tarda.     

Cultural transmission and flexibility of partial migration patterns in a long‐lived bird,the great bustard Otis tarda

Factors responsible for individual variation in partial migration patterns are poorly known, and identifying possible causes of these changes is essential for understanding the flexibility in migratory behavior. Analyzing 190 life histories of great bustards Otis tarda radio‐tagged in central Spain, we investigated the changes in migratory tendency across lifetime in this long‐lived bird, and how migratory flexibility is related to individual condition. In females migratory behavior was not fixed individually. For every age class there was a fraction of ca 15–30% of females that changed their migratory pattern between consecutive years. Migrant females tended to remain sedentary in years when they had dependent young to attend. These findings show that the female migratory tendency is a behaviorally flexible, condition‐dependent trait. Immature females usually acquired their migratory behavior by learning from the mother in their first winter or by social transmission from other migratory females in their second winter. As for immature males, their summer migratory behavior was not related to mother–offspring transmission, but learned from adult males. We found that their age‐related increase in migratory tendency was associated to a greater integration in flocks of migrant adult males. These results show that within the partial migration system, cultural transmission mechanisms, either mediated by kin or not, and individual condition, may contribute to shape the migratory tendency. Our study reinforces the view that the migratory behavior is an evolutionary complex trait conditioned by the interaction of individual, social and environmental factors. Particularly in long‐lived species with extended parental care, the inherited migration program may be shaped by mother–offspring and social transmission of migratory patterns.     

大鸨东方亚种遗传多样性的微卫星分析

为了探索东方亚种数量正在减少的原因和制定科学、有效的保护措施,利用微卫星DNA对大鸨东方亚种(Otistardadybowskii)的遗传多样性进行了研究。应用3个大鸨指名亚种的微卫星位点和13个波斑鸨的微卫星位点扩增了47个个体的基因组DNA,筛选出8个具有多态性的微卫星。其中有3个微卫星的多态性较低,其余5个微卫星的多态性较高。各位点的观察杂合度为0.0435-1.0000,平均杂合度(h)为0.6595;各位点多态信息含量(PIC)为0.0416-0.8520,平均为0.5497;有效等位基因数(E)为1.04-7.46,平均为3.61。4个位点符合HardyWeinberg平衡,4个位点偏离了HardyWeinberg平衡。多方面比较发现,大鸨东方亚种遗传多样性很低,且低于指名亚种,这可能由于其种群较小、历史遗传瓶颈作用、生境破碎化、分布地域紧缩等原因造成的。     

Proximate and ultimate causes of natal dispersal in the great bustard Otis tarda

Between 1991 and 1997 we studied the offspring independenceand juvenile dispersal in a wild population of great bustards(Otis tarda). Young males were independent and began their juveniledispersal at an earlier age (6–11 months) than young females(8–15 months). The juvenile dispersal period was longerand the distances reached farther in males than in females.Natal dispersal distances were also longer in males, all ofwhich dispersed from their natal areas and established as adultsat 5–65 km from their natal nests. In contrast, most femaleswere strongly philopatric, settling at 0.5–5 km from theirnatal nests. These marked sex differences in offspring independenceand dispersal may have evolved originally to maintain geneticdiversity and are probably reinforced through male competitionfor mates. Young males that had fed at higher rates and receivedmore feedings from their mothers during the early maternal dependenceperiod became indepthdent and tended to disperse earlier. Theyalso integrated earlier into adult male flocks and settled earlierat their definitive leks, which were closer to their natal sites,in areas of higher adult male density. None of these correlationswas found among young females. These results suggest that enhancedfood intake and maternal care of male offspring are vitallyimportant in increasing their competitive ability during theimmature period and probably also in their fitness as breedingadults. These results are in accordance with the selective valueof large size in males and suggest how this species might havereached such a marked sexual dimorphism in size.     

Satellite telemetry reveals long‐distance migration in the Asian great bustard Otis tarda dybowskii

The range of the great bustard stretches 10 000 km across Eurasia, one of the largest ranges of any threatened species. While movement patterns of the western subspecies of great bustard are relatively well‐understood, this is the first research to monitor the movements of the more endangered Asian subspecies of great bustard through telemetry and to link a breeding population of Asian great bustards to their wintering grounds. Using Argos/GPS platform transmitter terminals, we identified the annual movement patterns of three female great bustards captured at their breeding sites in northern Mongolia. The 4000 km round‐trip migration we have recorded terminated at wintering grounds in Shaanxi, China. This route is twice as long as has previously been reported for great bustards, which are among the heaviest flying birds. The journey was accomplished in approximately two months each way, at ground velocities of 48–98 km h^?1, and incorporated multiple and variable stopover sites. On their wintering grounds these birds moved itinerantly across relatively large home ranges. Our findings confirm that migratory behavior in this species varies longitudinally. This variation may be attributable to longitudinal gradients in seasonality and severity of winter across Eurasia. The distance and duration of the migratory route taken by great bustards breeding in Mongolia, the crossing of an international border, the incorporation of many stopovers, and the use of a large wintering territory present challenges to the conservation of the Asian subspecies of great bustard in this rapidly changing part of the world.     

Male sexual display and attractiveness in the great bustard Otis tarda: the role of body condition

Individual display rates and attractiveness of 17 radio-tagged, free-living great bustard males in north-western Spain were studied in relation to the degree of development of secondary sex traits, body condition, and age. Variance in display rate was significantly explained by body condition and length of moustache feathers, and neck development and display rate significantly accounted for the variance in male attractiveness. Age did not present any clear association with display rate or attractiveness, and it was significantly correlated only with neck development. These results suggest a total viability scenario of female mate choice in which the advertisement of global fitness is based on condition-dependent phenotypic and behavioural traits. Electronic Publication     

白洋淀流域大鸨越冬栖息地的适宜性评价

基于大鸨越冬期生境选择的相关研究成果,结合专家意见和近年来白洋淀流域大鸨的分布点记录,选择3项一级指标和13项二级指标,用以表征影响大鸨越冬生境选择的关键因子,并通过构建适宜性评价模型,对白洋淀流域大鸨越冬生境的质量进行了评价.结果表明:2005年,白洋淀流域内大鸨越冬适宜栖息地面积11907.25 km2,占流域总面积的34.1%;其中,最适宜生境面积4596.25 km2,仅占流域总面积的13.2%.研究区最适宜生境的空间分布相对集中,主要位于流域东部的白洋淀自然保护区及其周边(I区)和流域西南部的行唐、曲阳2县(II区).I区和II区中最适宜生境面积之和达2803.55 km2,占流域内最适宜生境面积的61.0%.为保护流域内大鸨的越冬生境,须重点针对上述2个区域的特点,分别采取适当措施加以保护.     

Genetic structure in Iberian and Moroccan populations of the globally threatened great bustard Otis tarda: a microsatellite perspective

Patterns of genetic structure and gene flow among populations help us understand population dynamics and properly manage species of concern. Matrilineal mtDNA sequence data have been instrumental in revealing genetic structure at the intraspecific level, but bi‐parentally inherited markers are needed to confirm patterns at the genome level and to assess the potential role of sex‐biased dispersal on gene flow, particularly in species where males are known to be the main dispersing sex. Here we use microsatellite loci to examine patterns of genetic structure across the range of the great bustard in Iberia and Morocco, an area representing 70% of the world population of this globally threatened species. We used population differentiation statistics and Bayesian analysis of population structure to analyse data from 14 microsatellite loci. These data provide greater resolution than mtDNA sequences, and results reveal the existence of three main genetic units corresponding to Morocco, the northeastern part of Spain, and the rest of the Iberian Peninsula. Our results, together with previous mtDNA data, confirm the genetic differentiation of the northern Africa population and the importance of the Strait of Gibraltar as a barrier to gene flow for both males and females, rendering the Moroccan population a separate management unit of high conservation concern.     

Display activity and seasonality of faecal sexual steroids in male great bustard (Otis tarda L.)

The non-invasive faecal sampling and RIA was used to measure faecal equivalents of testosterone (T), dehydroepiandrosterone (DHEA), oestradiol-17beta (E2) and progesterone (P4) in juvenile and adult great bustard males. Possible connections of diurnal and seasonal changes of sexual steroid levels and display activity were studied. Correlations were found between sexual steroid equivalent levels of faeces and display activity and agonistic behaviour in the different phases of annual cycle of adult males. In early display period increasing levels of androgens were measured, during main display period very high androgen dominance was observable against E2 and P4. During postnuptial moult strong T decrease and DHEA and P4 increase were detected. Elevation of E2 was measured during wintering. In juveniles level of DHEA was higher than level of T suggesting its importance in immature males. Decrease of T was detected between reproductive period and postnuptial moult and DHEA between reproduction and wintering, accompanying with E2 elevation. The inhibiting effect of inclement weather on gonad functions also was detected in our study. We suppose that the unexpected cold weather with strong wind depressed the levels of androgens both in juveniles and adults and the increase of faecal E2 was also detected.     

Sex-biased parasitism,seasonality and sexual size dimorphism in desert rodents

We investigated seasonality of gender differences in the patterns of flea infestation in nine rodent species to test if sex-biased parasitism in terms of mean abundance, species richness, prevalence and the level of aggregation (a) varies among hosts and between seasons, and (b) is linked to sexual size dimorphism. Sexual size differences were significant in both summer and winter in Acomys cahirinus, Gerbillus pyramidum and Meriones crassus, and in winter only in Acomys russatus, Gerbillus dasyurus, Gerbillus nanus and Sekeetamys calurus. Sexual size dimorphism was male biased except for A. russatus in which it was female biased. Manifestation of sexual differences in flea infestation was different among hosts between seasons. A significant effect of sex on mean flea abundance was found in six hosts, on mean flea species richness in five hosts and on prevalence in two hosts. Male-biased parasitism was found in summer in one host only and in winter in five hosts. Female-biased parasitism occurred in winter in A. russatus. Gender differences in the slopes of the regressions of log-transformed variances against log-transformed mean abundances occurred in three hosts. No relationship was found between sexual size dimorphism and any parasitological parameter in any season using both conventional regressions and the method of independent contrasts. Our results suggest that sex-biased parasitism is a complicated phenomenon that involves several different mechanisms.     

Factors causing long-term dynamics in the abundance of the Trans-Volga Great bustard (Otis tarda L.) population

The results of our survey of bustards in the southern part of the Saratov Trans-Volga region over an area of 12000 sq. km in 1990–2010 are presented; analysis of the abundance dynamics is performed; and the factors causing it are revealed. Data on the distribution of bustards over their biotopes in the period of premigration roaming are studied, and the demographic and spatial structure of the population is analyzed.     

Field determination of age in male great bustards (Otis tarda) in spring

During a long-term study of individually marked, free-living male great bustards captured as chicks and radio-tracked through several years in Spain, we studied the development with age of two secondary sex traits, the moustachial feathers and the neck plumage pattern. Juvenile males acquired full adult plumage between their fourth and seventh years. The main changes occurred at the neck, coinciding with the onset of sexual maturity. The grey colour typical of immature males turned to ivory white around the fourth to fifth spring, and a gradual increase was appreciated in adults in the brightness of the white colour of the upper neck and in the contrast between this and a progressively more intense chestnut brown at the neck base. Based on these changes, we proposed four neck plumage patterns that can be used to differentiate male age classes during the mating period. The development of moustachial feathers showed more interindividual variability and was not as useful as the neck plumage to estimate male age.     

Genetic variation and bill size dimorphism in a passerine bird, the reed bunting Emberiza schoeniclus

In passerine birds morphological differentiation in bill size within species is not commonly observed. Bill size is usually associated with a trophic niche, and strong differences in it may reflect the process of genetic differentiation and, possibly, speciation. We used both mitochondrial DNA (mtDNA) and nuclear microsatellites to study genetic variation between two subspecies of reed bunting, Emberiza schoeniclus schoeniclus and E.s. intermedia , along their distributional boundary in western Europe. These two subspecies are characterized by a high dimorphism in bill size and, although breeding populations of the two subspecies are found very close to each other in northern Italy, apparently no interbreeding occurs. The observed morphological pattern between the two subspecies may be maintained by geographically varying selective forces or, alternatively, may be the result of a long geographical separation followed by a secondary contact. MtDNA sequences of cytochrome b and ND5 (515 bp) showed little variation and did not discriminate between the two subspecies, indicating a divergence time of less than 500 000 years. The analysis of four microsatellite loci suggested a clear, although weak, degree of genetic differentiation in the large- and small-billed populations, as indicated by F _ST and R _ST values and genetic distances. The correlation between bill size and genetic distance between populations remained significant after accounting for the geographical distances between sampling localities. Altogether, these results indicate a very recent genetic differentiation between the two bill morphs and suggest that a strong selection for large bills in the southern part of the breeding range is probably involved in maintaining the geographical differentiation of this species.     

Sexual dimorphism,survival, and parental investment in relation to offspring sex in a precocial bird

Differential growth rate between males and females, owing to a sexual size dimorphism, has been proposed as a mechanism driving sex‐biased survival. How parents respond to this selection pressure through sex ratio manipulation and sex‐biased parental investment can have a dramatic influence on fitness. We determined how differential growth rates during early life resulting from sexual size dimorphism affected survival of young and how parents may respond in a precocial bird, the black brant Branta bernicla nigricans. We hypothesized that more rapidly growing male goslings would suffer greater mortality than females during brood rearing and that parents would respond to this by manipulating their primary sex ratio and parental investment. Male brant goslings suffered a 19.5% reduction in survival relative to female goslings and, based on simulation, we determined that a female biased population sex ratio at fledging was never overcome even though previous work demonstrated a slight male‐biased post‐fledging survival rate. Contrary to the Fisherian sex ratio adjustment hypothesis we found that individual adult female brant did not manipulate their primary sex ratio (50.39% male, n = 645), in response to the sex‐biased population level sex ratio. However, female condition at the start of the parental care period was a good predictor of their primary sex ratio. Finally, we examined how females changed their behavior in response to primary sex ratio of their broods. We hypothesized that parents would take male biased broods to areas with increased growth rates. Parents with male biased primary sex ratios took broods to areas with higher growth rates. These factors together suggest that sex‐biased growth rates during early life can dramatically affect population dynamics through sex‐biased survival and recruitment which in turn affects decisions parents make about sex allocation and sex‐biased parental investment in offspring to maximize fitness.     

Introducing the refined gravity hypothesis of extreme sexual size dimorphism

Background  

Explanations for the evolution of female-biased, extreme Sexual Size Dimorphism (SSD), which has puzzled researchers since Darwin, are still controversial. Here we propose an extension of the Gravity Hypothesis (i.e., the GH, which postulates a climbing advantage for small males) that in conjunction with the fecundity hypothesis appears to have the most general power to explain the evolution of SSD in spiders so far. In this "Bridging GH" we propose that bridging locomotion (i.e., walking upside-down under own-made silk bridges) may be behind the evolution of extreme SSD. A biomechanical model shows that there is a physical constraint for large spiders to bridge. This should lead to a trade-off between other traits and dispersal in which bridging would favor smaller sizes and other selective forces (e.g. fecundity selection in females) would favor larger sizes. If bridging allows faster dispersal, small males would have a selective advantage by enjoying more mating opportunities. We predicted that both large males and females would show a lower propensity to bridge, and that SSD would be negatively correlated with sexual dimorphism in bridging propensity. To test these hypotheses we experimentally induced bridging in males and females of 13 species of spiders belonging to the two clades in which bridging locomotion has evolved independently and in which most of the cases of extreme SSD in spiders are found.     

Breeding success of the Great Bustard Otis tarda in Zamora Province, Spain, in 1984

Great Bustards are still fairly abundant (5-2 nests/1000 ha) in Zamora. Overall fecundity was high (92% of adult females attempt to nest; 2–47 eggs/clutch; 89% egg fertility), but pre- and post-hatching mortality were also high (50% of eggs laid; 57% of nestlings hatched), due mainly to agricultural machinery but also to natural predators. Calculated annual yields per 100 adult females were: 227 eggs, 102 hatchlings, 44 fledglings. Production of 44 fledglings/y could sustain a stable adult population (mortality approx. 8%/y) if mortality of immatures is not more than 18–22%/y.     

Behavioural mechanisms that undermine species envelope models: the causes of patchiness in the distribution of great bustards Otis tarda in Spain

Despite the general success of species envelope models, capturing the fine-scale detail of patchiness in the distributions of some species is problematic. For great bustards in Spain, apparently suitable habitat patches remain unoccupied and cannot be distinguished from occupied patches in current distribution models. We consider philopatry and conspecific attraction as main behavioural mechanisms which could account for this patchiness, and then look for evidence of their influence on the distribution of great bustards across the whole of Spain. We compared the characteristics of habitat patches classed as suitable by a distribution model according to whether they were actually used or not. Occupied patches were larger than unoccupied patches and over-used in proportion to their size, suggesting aggregation and a metapopulation structure. Arguing that conspecific attraction may serve to transfer information about site history and environmental predictability (at least over a short time period), we compared the coefficients of variation in time-series of vegetation and climatic factors at occupied and unoccupied sites. Great bustards chose sites which were more environmentally stable at critical periods in the breeding cycle, "public information" that can only be gained from others rather than through sampling. There is thus evidence that both metapopulation dynamics and conspecific attraction influence the large scale distribution of great bustards in Spain. We discuss how alternative predictor variables and multi-stage analyses may help us to incorporate behavioural mechanisms into distribution models, but acknowledge that there are limits to the value of species envelope models for animal species making decisions.     

Factors driving extreme sexual size dimorphism of a sit-and-wait predator under low density

Sexual size dimorphism is often a likely outcome of the interplay between natural selection and sexual selection, with female size dictated primarily by natural selection that maximizes fecundity and male size by sexual selection that maximizes reproductive opportunities. Attention to male fitness has focused heavily on direct male-male conflict selecting for superior male size and/or fighting ability, although male reproductive traits vary immensely among animals. An alternative, advanced by Michael Ghiselin, posits highly mobile dwarf males as a strategy for finding relatively immobile females in low-density populations. Adult male crab spiders Misumena vatia , sit-and-wait predators, are strikingly smaller, much more active, and relatively longer-legged than their females. This size difference results largely from males having two fewer instars than females, which simultaneously results in marked protandry. Populations of M. vatia often were small and of low density, with a female-biased sex ratio and an operational sex ratio that changed strikingly over the season. Sexual selection through scramble competition (locating the female first) should favour this suite of characters in males of low-density populations. Although direct male-male contests favoured large males, the low densities of adult males and the dispersed, relatively immobile females led to low levels of direct intrasexual contest. Females exaggerated the problem of males in finding them by providing few cues to their presence, a pattern consistent with indirect mate choice. In addition to favouring high mobility, scramble competition favoured males that selected flowers attracting many prey, the sites most often occupied by females.     

Optimal climbing speed explains the evolution of extreme sexual size dimorphism in spiders

Several hypotheses have been put forward to explain the evolution of extreme sexual size dimorphism (SSD). Among them, the gravity hypothesis (GH) explains that extreme SSD has evolved in spiders because smaller males have a mating or survival advantage by climbing faster. However, few studies have supported this hypothesis thus far. Using a wide span of spider body sizes, we show that there is an optimal body size (7.4 mm) for climbing and that extreme SSD evolves only in spiders that: (1) live in high‐habitat patches and (2) in which females are larger than the optimal size. We report that the evidence for the GH across studies depends on whether the body size of individuals expands beyond the optimal climbing size. We also present an ad hoc biomechanical model that shows how the higher stride frequency of small animals predicts an optimal body size for climbing.