Density dependence in a recovering osprey population: demographic and behavioural processes

1. Understanding how density-dependent and independent processes influence demographic parameters, and hence regulate population size, is fundamental within population ecology. We investigated density dependence in growth rate and fecundity in a recovering population of a semicolonial raptor, the osprey Pandion haliaetus [Linnaeus, 1758], using 31 years of count and demographic data in Corsica. 2. The study population increased from three pairs in 1974 to an average of 22 pairs in the late 1990s, with two distinct phases during the recovery (increase followed by stability) and contrasted trends in breeding parameters in each phase. 3. We show density dependence in population growth rate in the second phase, indicating that the stabilized population was regulated. We also show density dependence in productivity (fledging success between years and hatching success within years). 4. Using long-term data on behavioural interactions at nest sites, and on diet and fish provisioning rate, we evaluated two possible mechanisms of density dependence in productivity, food depletion and behavioural interference. 5. As density increased, both provisioning rate and the size of prey increased, contrary to predictions of a food-depletion mechanism. In the time series, a reduction in fledging success coincided with an increase in the number of non-breeders. Hatching success decreased with increasing local density and frequency of interactions with conspecifics, suggesting that behavioural interference was influencing hatching success. 6. Our study shows that, taking into account the role of non-breeders, in particular in species or populations where there are many floaters and where competition for nest sites is intense, can improve our understanding of density-dependent processes and help conservation actions.     

Genetic consequences of a severe population bottleneck in the Guadalupe fur seal (Arctocephalus townsendi)

Population bottlenecks may lead to diminished genetic variability and correlative effects on fitness. The Guadalupe fur seal was nearly exterminated by commercial sealers during the late 18th and early 19th centuries. To determine the genetic consequences of this population bottleneck, we compared the variation at a 181 bp section of the mitochondrial DNA (mtDNA) control region from the bones of 26 prebottleneck fur seals versus variation in the extant population. We found 25 different mtDNA genotypes in the prebottleneck fur seals and only 7 genotypes among 32 extant fur seals, including only one of the ancient genotypes. These data demonstrate a substantial loss of genetic variability correlating with the recent population bottleneck. We also found from several genetic measures that the prehistoric population of Guadalupe fur seals was robust and that it had been increasing at some time during the late prehistoric period. Continued recovery of this species may, however, owe more to more immediate nongenetic factors, such as poaching and local availability of food resources during the breeding season and consequent effects on pup survival, than on the reduced genetic variability.     

Gastric intramural hematoma and hemoperitoneum in a captive northern fur seal

A 16-yr-old adult male northern fur seal (Callorhinus ursinus) was found dead in its outdoor pool in November 1995. The animal was maintained at Mystic Aquarium (Mystic, Connecticut, USA) from March 1980 to November 1995. Gross necropsy findings included hemoperitoneum and locally extensive gastric intramural hemorrhage that involved the posterior fundic, antral, and pyloric regions and extended into the duodenum. The gastric mural thickening grossly resembled hemangioma, and the gastric serosa was ruptured at the site of maximal mural expansion. In histologic sections of the stomach, a cribiform network of fibrin, which encompassed numerous variably-sized aggregates of closely packed erythrocytes, markedly expanded the submucosa. No vascular endothelium was identified in serial histologic sections of the expanded gastric submucosa stained with hematoxylin and eosin or immunohistochemically with antibodies to vimentin and Factor VIII-related antigen, establishing an absence of hemangioma. Carstairs' and Weigert's histochemical stains confirmed that the framework expanding the submucosa was fibrin. Although the appearance of the gastric wall resembled hemangioma, a population of neoplastic endothelial cells was not identified within the submucosal expansion of hemorrhage and fibrin, and microscopic evidence was most consistent with the diagnosis of gastric intramural hematoma. This lesion is a rare pathologic event that has not been reported in marine mammals, but one that should be included in diagnostic considerations of hemoperitoneum and gastric mural expansion.     

Evidence of natural reciprocal hybridisation between Pinus uliginosa and P. sylvestris in the sympatric population of the species

Peat-bog pine Pinus uliginosa Neumann has become extinct or rare in many parts of Europe. It is supposed to hybridize with P. sylvestris but the influence of hybridization to genetic erosion of the P. uliginosa gene pool is unknown. In the presented study, the crossability between P. uliginosa and P. sylvestris was analyzed in a sympatric population at Węgliniec reserve in Poland. The aim of the study was to prove natural hybridisation and to estimate the influence of this phenomenon on possible natural gene pool erosion of peat-bog pine. A sequence polymorphism in the trnF–trnL cpDNA region of P. uliginosa and P. sylvestris was used to develop the species diagnostic PCR-RFLP marker. The marker of paternally transmitted cpDNA was applied in haplotype analyses of the progeny from open-pollinated P. uliginosa seeds (collected in 2000–2002) and from P. sylvestris ones (in 2002). An inconsistency in species diagnostic cpDNA haplotypes of seedlings and parental trees was observable both for P. uliginosa and P. sylvestris (about 1% and 2% of hybrids seeds, respectively). The results prove the occurrence of reciprocal hybridisation between the species. The influence of hybridisation on the natural gene pool protection and measures to reintroduce peat-bog pine is discussed.     

Floral traits and isolation of three sympatric Aquilegia species in the Qinling Mountains, China

Floral isolation has been considered to be an important reproductive mechanism governing the species diversification in many genera. In a classic example Aquilegia, sympatric species from North America with diverse floral traits are generally associated with specialized pollinators that prohibit interspecific hybridization. It remains unclear whether species diversification in the genus from Eurasia is also maintained by floral isolation. We investigated floral phenology, floral characteristics and pollinators in three sympatric Aquilegia species (A. ecalcarata, A. incurvata and A. yabeana) in the Qinling Mountains, Shanxi Province, China from 2001 to 2005. The spurless A. ecalcarata flowers earlier than the other two species with nectar spurs but their floral phenology overlaps. Major pollinators of A. ecalcarata are syrphid flies while bumblebees are major for A. incurvata and A. yabeana. Therefore our observations confirm that mechanical isolation through differential pollinators could contribute reproductive isolation between spurless and spurred species, as demonstrated by studies from North America. Whether floral isolation plays a major role in the reproductive isolation between two spurred species (A. incurvata and A. yabeana), however, remains to be seen. Further studies are required to quantify the potential role of geographical isolation because they occupy different habitats.     

Two sympatric Byrsonima species (Malpighiaceae) differ in phenological and reproductive patterns

Sympatric and taxonomically related species may present pre- and/or postzygotic mechanisms for reproductive isolation. We compared the phenology and reproductive biology of Byrsonima intermedia and B. pachyphylla, two sympatric Malpighiaceae species, growing in a remnant of Cerrado in Central Brazil. Both species are evergreens and do not have an overlapping flowering period. In both species, dispersal occurs during the rainy season with low and intermediate overlap in the production of immature and mature fruits, respectively. Both species exhibit self-incompatibility (SI), or, in the case of B. intermedia, facultative SI, and, hence, depend on pollinators to compensate for the lack of spontaneous self-pollination. The flowers of the Byrsonima species were visited by fifteen species of bees that collected pollen and/or oil. Based on their more appropriate intrafloral posture, Epicharis flava and Centris varia were the main pollinators of both species, presenting more frequent visits and/or exhibiting trapline foraging behavior, tending to promote outcrossing. Temporal variation in flowering period and total or partial self-incompatibility seem to function as prezygotic isolation mechanisms that form barriers to gene flow between the studied species and probably avoid competition among pollinators. Both species make oil and pollen available to flower visitors and pollinators almost the entire year, but since Byrsonima intermedia has a long flowering time and is very abundant in the study area, it seems to be more important in maintaining flower-visiting bees.     

Dynamic influence of maternal and pup traits on maternal care during lactation in an income breeder, the antarctic fur seal

Life-history theory predicts that selection will favor optimal levels of parental effort that balance benefits of current reproduction with costs to survival and future reproduction. The optimal level of effort depends on parental traits, offspring traits, and provisioning strategy. Additionally, how these factors influence effort may differ depending on the stage of reproduction. The relative importance of maternal and offspring traits on energy allocation to offspring was investigated in known-age Antarctic fur seals Arctocephalus gazella across four stages of reproduction, using birth mass and milk-consumption measurements. Maternal traits were important during three of the four stages investigated, with larger females giving birth to larger pups and investing more in pups during perinatal and molt stages. Pup mass influenced maternal effort during the premolt stage, and provisioning strategy influenced postnatal maternal effort at all stages. Energy provided to the offspring during an attendance visit was positively related to the duration of the foraging-trip/visit cycle; however, when investment was controlled for trip/visit cycle duration, the overall rate of energy transfer was similar across trip durations. In addition to strong effects of maternal mass, pup traits affected energy allocation, suggesting that pup demand is important in determining maternal care. These findings emphasize the importance of considering state variables in life-history studies and suggest that timing of measurements of effort in species with long provisioning periods may influence conclusions and our ability to make comparisons of reproductive effort among species.     

Causes and consequences of fine-scale breeding dispersal in a female-philopatric species

The potentially confounded effects of factors affecting breeding dispersal have rarely been simultaneously examined. The consequences of breeding dispersal are even less studied, presenting a paradox: breeding dispersal seldom seems to improve breeding success, despite its presumed adaptiveness. We studied the causes and consequences of breeding dispersal in female-philopatric eiders (Somateria mollissima) in relation to the spatiotemporal predictability of nest success. Previous nest fate, breeding experience, and breeding density simultaneously affected breeding dispersal. Dispersal distances were longer among inexperienced breeders and after failed breeding. Individual dispersal distances decreased with increasing nest-site-specific breeding density, whereas island-specific nesting success peaked at intermediate densities. The fate of neighbouring nests (‘public information’) did not influence dispersal. Breeding dispersal was unrelated to subsequent hatching success, controlling for individual quality (body condition, breeding experience, previous nest fate), while it delayed hatch date, which is likely to impair reproductive success. This delay may result from the loss of acquired information of local breeding conditions, prolonging nest prospecting and establishment, also helping explain why breeding dispersal did not increase at high breeding densities, despite a potential reduction in nesting success. In long-lived species, however, dispersal-induced reductions in reproductive output in one season could be offset by improved parental survival prospects. Careful nest prospecting may be profitable, because overall nest success had a strong island-specific component but showed weak temporal variation, and successive individual nest fates were predictable between years. Once a safe nest site is found, females may breed at the same place successfully for many years.     

Selection and segregation distortion in a sex-differentiated population

We extend the classical model for selection at an autosomal locus in a sex-differentiated population to include segregation distortion. The equations remain the same, but the fitness parameters are interpreted differently and refer to alleles instead of genotypes. We derive conditions for internal and external stability of the equilibria, i.e., stability with respect to perturbations of alleles that are already present at equilibrium and stability with respect to invasion attempts by newly arising alleles. We show that, in a sex-differentiated population, external stability of an equilibrium can be judged on the basis of Shaw--Mohler criteria. Throughout, we compare the situation in populations with and without sex differentiation. Interestingly, internal stability is more difficult to achieve in a population without sex differentiation than in a population in which selection and segregation distortion are restricted to one sex. In a companion paper we show how the general results of the present paper can lead to new insights into specific systems such as the t complex of the house mouse.     

Maternal-larval population genetic traits in Stylophora pistillata, a hermaphroditic brooding coral species

Aspects of maternal-planula larval genetics in the monoecious scleractinian coral Stylophora pistillata (Red Sea, Eilat) were studied by amplified fragment length polymorphism (AFLP) methodology in two successive reproductive seasons. In total, 293 planulae and 10 adult colonies were analyzed. In June 2006, 147 planulae were collected from 10 shallow water colonies. In March, April and June 2007, 146 additional planulae were sampled from five of the ten 2006 sampled colonies. All AFLP products showed unalike band profiles indicating a fully sexual production pattern. We used 181 and 210 putative AFLP loci, of which the overall level of polymorphism in 2006 was 92 and 99 % in 2007 (respectively). Differences were also observed between 2006 and 2007 reproductive seasons in terms of total average gene diversity (0.191 vs. 0.247, respectively), suggesting fast turnover of sperm donor genotypes. In addition, increased numbers of potential sperm donor colonies in the vicinity of gravid females showed no impact on genetic differentiation levels in released larvae. UPGMA tree revealed clustering of maternal genotypes and their offspring, suggesting, as expected, high relatedness between planulae and their mothers. In addition, the average heterozygosity of each group of siblings was persistently lower than heterozygosity calculated for the respective maternal colony, suggesting the possibility of partial inbreeding. This trend of reduced genetic heterogeneity in Stylophora pistillata is an alarming sign for populations residing in the northern Red Sea coral reefs.     

Genetic variation and fine-scale population structure in American pikas across a human-modified landscape

Natural resource extraction can represent a major human modification to the landscape. Habitat reclamation is becoming an increasingly important strategy for abating the loss of biodiversity associated with these developments; however, the demographic and genetic consequences of colonizing artificial habitat remain unknown in many species. Here, we investigated the genetic consequences of landscape modifications for the American pika (Ochotona princeps) relative to two major developments in British Columbia, Canada: a large open-pit copper mine (Highland Valley Copper) under partial reclamation and a bisecting major highway (97C). We assessed microsatellite genotypic data for 109 individuals across 15 sites located either within the mine on artificial habitat or on adjacent natural habitat both north and south of the highway. There were no significant differences in levels of heterozygosity, allelic richness or inbreeding between natural (n?=?7) and artificial sites (n?=?8). However, pikas residing on artificial habitat exhibited significantly higher relatedness estimates. Bayesian clustering analyses revealed two distinct genetic units corresponding to north and south of the highway, with further substructure detected in the south. Likewise, high genetic friction was detected in the central region of the area, largely corresponding to the highway and modified landscape associated with the mine. At a finer scale, pairwise estimates of differentiation and migration rates suggest little gene flow may be occurring among sites across the sampling area, with some evidence for directional migration from artificial to natural sites. Overall, artificial habitat has been successful in promoting occupancy for American pikas, however, barriers to gene flow likely associated with resource extraction and road construction limit connectivity across the landscape.     

Phenotypic similarity and differentiation among sympatric cryptic species in a freshwater amphipod species complex

1. Cryptic species diversity poses evolutionary questions about its origin and maintenance, and ecological questions about the coexistence of seemingly identical species. 2. We examine patterns of morphological and life history differentiation in three sympatric cryptic species of freshwater amphipods within the Hyalella azteca species complex. These species are separated by extensive molecular evolution, but appear similar in phenotype. Species were collected from the littoral zone of a small kettle lake in Michigan, U.S.A., and identified to species by molecular genetic methods. 3. Two of the species were similar, differing only in female size, whereas the third was larger in body size and had larger clutches of smaller eggs than the other two. There were differences between the species also in pleon spine length and antennal segment number. 4. An analysis of allozyme variation among the cryptic species in three lakes suggests that the species are reproductively isolated within lakes. 5. We suggest that phenotypic similarity of these species is maintained by size‐selective predation by fish. The small, but statistically significant, differences in body size may form the axis for a tradeoff between resource exploitation and predator avoidance, a condition that can foster coexistence of phenotypically similar species.     

Nuclear and chloroplast genetic structure indicate fine-scale spatial dynamics in a neotropical tree population

Dicorynia guianensis is a canopy tree, endemic to the tropical rain forest of French Guiana. We compared generational and spatial genetic structure for maternally and biparentally inherited markers in two cohorts (adult and seedling) in order to infer processes shaping the distribution of genetic diversity. The study was conducted on a 40 ha study plot located at Paracou near Kourou, where 172 adults trees and 375 saplings were sampled. Aggregation of trees was therefore suggested at different distances, ranging from 100 to 400 m. There was a strong link between demographic and genetic spatial structures at small distances (less than 100 m) that is likely to be the consequence of restricted seed dispersal. Genetic differentiation was more pronounced between spatial aggregates than between cohorts. Despite the spatial differentiation, the species was able to maintain high levels of diversity for maternal genomes, suggesting rapid turnover of aggregates. Spatial autocorrelation was larger for chloroplast than nuclear markers indicating a strong asymmetry between pollen and seed flow. Fixation indices indicated a lower heterozygote deficiency for the adults, maybe because of gradual elimination of selfed trees. Genetic relatedness at lower distances was higher in adult trees than in saplings, as a result of generation overlapping in the adult cohort. Overall, our results confirm earlier biological knowledge about the dispersion mechanisms of the species, and lead to an enhanced role of spatial processes in the dynamics of genetic diversity of D. guianensis.     

Biological and environmental drivers of energy allocation in a dependent mammal, the Antarctic fur seal pup

Little is known about how variation in the pattern and magnitude of parental effort influences allocation decisions in offspring. We determined the energy budget of Antarctic fur seal pups and examined the relative importance of timing of provisioning, pup traits (mass, condition, sex), and weather (wind chill and solar radiation) on allocation of energy obtained in milk by measuring milk energy intake, field metabolic rate (FMR), and growth rate in 48 Antarctic fur seal pups over three developmental stages (perinatal, premolt, and molt). The relative amount of milk energy used for growth was 59.1% ± 8.1% during the perinatal period but decreased to 23.4% ± 15.5% and 26.0% ± 13.9% during the premolt and molt. This decrease was associated with a greater amount of time spent fasting, along with an increase in pup activity while the mother was at sea foraging. Average daily milk intake, pup mass, and condition were all important in determining how much energy was available for growth, but the amount of energy obtained as milk was the single most important factor determining pup growth. While mean mass-specific FMR did not change with developmental stage (range = 1.74-1.77 mL O(2)/g/h), the factors that accounted for variation in FMR did. Weather (wind chill and solar radiation) and pup traits (mass and condition) influenced mass-specific FMR, but these impacts varied across development. This study provides information about the factors influencing how offspring allocate energy toward growth and maintenance and improves our predictions about how a changing environment may affect energy allocation in pups.     

Within and between population variation in plant traits predicts ecosystem functions associated with a dominant plant species

Linking intraspecific variation in plant traits to ecosystem carbon uptake may allow us to better predict how shift in populations shape ecosystem function. We investigated whether plant populations of a dominant old-field plant species (Solidago altissima) differed in carbon dynamics and if variation in plant traits among genotypes and between populations predicted carbon dynamics. We established a common garden experiment with 35 genotypes from three populations of S. altissima from either Tennessee (southern populations) or Connecticut (northern populations) to ask whether: (1) southern and northern Solidago populations will differ in aboveground productivity, leaf area, flowering time and duration, and whole ecosystem carbon uptake, (2) intraspecific trait variation (growth and reproduction) will be related to intraspecific variation in gross ecosystem CO(2) exchange (GEE) and net ecosystem CO(2) exchange (NEE) within and between northern and southern populations. GEE and NEE were 4.8× and 2× greater in southern relative to northern populations. Moreover, southern populations produced 13× more aboveground biomass and 1.4× more inflorescence mass than did northern populations. Flowering dynamics (first- and last-day flowering and flowering duration) varied significantly among genotypes in both the southern and northern populations, but plant performance and ecosystem function did not. Both productivity and inflorescence mass predicted NEE and GEE between S. altissima southern and northern populations. Taken together, our data demonstrate that variation between S. altissima populations in performance and flowering traits are strong predictors of ecosystem function in a dominant old-field species and suggest that populations of the same species might differ substantially in their response to environmental perturbations.     

Parental genetic characteristics and hatching success in a recovering population of Lesser Kestrels

Decreased hatchability is a common consequence of inbreeding in oviparous organisms and it has been generally considered a useful measure of the effects of reduced genetic diversity on embryological development. Here, we examined the pattern of hatching failure in a wild population of the endangered Lesser Kestrel (Falco naumanni). Particularly, we first analyzed long-term changes of hatching failure over a 16-year study period (1991–2006), in which the study population experienced a concurrent demographic and genetic recovery, and then we determined the consequences of parental genetic characteristics on hatching success. Long-term data analyses revealed a significant decline of hatching failure over time, with annual average hatching failure decreasing from rates characterizing species that have passed a severe population bottleneck to levels generally reported for outbred bird populations. Partial purging of deleterious recessive alleles after the species population decline, the increase of heterozygosity over time reported in a previous study and/or the selection for efficient mechanisms of inbreeding avoidance could be responsible of the observed temporal pattern. In contrast to previous studies, we found no effect of parental genetic characteristics on hatching success. Even though we analyzed an extensive dataset, the 11 neutral markers typed may have had low power to detect such an association. Further, this analysis was limited to the last 5 years (2002–2006) of the whole study period, when DNA samples for genetic analyses were available. During these years, hatching rates were like those typically reported for non-inbred populations, suggesting that the absence of association could be explained by a reduction of the genetic load or consequence of the genetic recover reported in the study population in recent years.     

Fluctuating asymmetry in leaves and flowers of sympatric species in a tropical montane environment

Fluctuating asymmetry (FA) represents small, random variations in traits, presumably with bilateral symmetry, and is widely used as a tool to measure developmental instability in plants and animals. Because FA is a quick, simple and reliable measure, it has been frequently used for monitoring levels of environmental stress. This study investigated whether FA can be used as a predictor of individual developmental instability for four sympatric Melastomataceae species. To achieve that aim, 20 individuals of Trembleya laniflora, T. parviflora, Lavoisiera campos‐portoana and Tibouchina heteromalla were marked in southeastern Brazil and monitored before and during the flowering season. The FA index was calculated as the mean of the difference in the width or length between the left and right sides measured for each leaf or flower. All studied species exhibited asymmetry in the leaves and flowers, but the leaves of L. campos‐portoana and the petal width of T. heteromalla exhibited directional asymmetry, also an indicator of developmental instability. The highest level of leaf asymmetry was found in T. heteromalla and on flowers of L. campos‐portoana. None of the studied species exhibited a significant relationship between the FA level of the leaves and flowers on an individual basis, indicating that environmental and/or genetic sources of stress might act differently on different plant traits. For the studied species, measurements of FA can be suggested as useful tools to biomonitor levels of stress experienced by both leaves and flowers within the Melastomataceae family.     

Comparing competitive ability and associated metabolic traits between a resident and migratory population of bull trout against a non-native species

Juvenile bull trout Salvelinus confluentus from two geographically and ecologically distinct populations were compared with regard to their ability to compete with non-native brook trout Salvelinus fontinalis in an artificial stream, and with respect to their rates of oxygen consumption. Bull trout collected from a migratory population foraged more successfully against brook trout competitors than those from a resident population, capturing more of a limited amount of food items presented. The migratory population was also more aggressive (measured by the number of nips, chases and lateral threat displays) against brook trout competitors than the resident population. Bull trout from the migratory population had a higher oxygen consumption rate (203 mg O₂ kg · hr^-1) in the field than similar sized fish from the resident population (183 mg O₂ kg · hr^-1). These results suggest native bull trout have population-level variation in competitive ability against a non-native species and such competitive ability is positively associated with metabolism and migratory life history.