Spatial Association in a Highly Inbred Ungulate Population: Evidence of Fine-Scale Kin Recognition

We present the first evidence of fine-scale kin recognition, based on a continuous measure of relatedness, in ungulates. The spatial association between herdmates of a captive population of aoudad (Ammotragus lervia), where all the individuals are related, is analysed during resting time. Our goal was to estimate which factors influence individuals’ associations. The study population is highly inbred, although it does not show serious deleterious effects caused by consanguinity. It comprises a single captive herd, reproducing freely and in good conditions for more than 10 yr. It emerges that kin, measured as the coefficient of relationship between two given herdmates, is the main factor determining the spatial association (e.g. average distance) of male–male and female–female dyads, as more-related individuals tend to rest closer to each other than less-related ones. As for male–female dyads, individuals of a similar age tend to stay closer. To rule out any familiarity confounding effects, individuals’ cohabitation time in the herd was added as a random factor in the analyses. Concerning the type of dyad, mother–calf dyads are characterized by higher proximity than others, particularly during the suckling period, whereas males tend to stay closer to each other than females or male–female dyads, being also more kin-related. Female social rank does not influence spatial association between herdmates. These results are related to group composition of the species in the wild, which are characterized by intense mother–calf bonds and all-male groups that are probably kin-related. It is seen that adult male–female associations are not related to kinship, but to age similarity, which is in accord with the assumption that main family groups in the wild are formed by matrilineal lines, whereas males are the dispersing sex.supported by the project PBI-05-010 granted by Junta de Comunidades de Castilla-La Mancha.Peer reviewe

Female lesser kestrel (falco naumanni) also accepts extra-pair copulation

The lesser kestrel (Falco naumanni) is considered a socially monogamous species in which extra-pair copulation occurs at a low rate. A wide study of extra-pair mating behaviour carried out on this species found that mated females do not solicit extra-pair copulation and reject all copulation attempts elicited by extra-pair males. In fact, the infrequent instances of extra-pair fertilization found were attributed either to re-mating after divorce, or polygyny. Here I describe an observation indicating that females also promote and accept promiscuous mating, attaining and consummating extra-pair copulation. I suggest the possibility that promiscuous behaviour could vary among different populations in this species, as it has not yet been reported, in spite of the extensive work performed in this respect.The former Spanish Ministerio de Ciencia y Tecnología provided finance by means of the Research National Plan Project (REN 2002-03295). During this work the author has the benefit of a postgraduate fellowship of the Consejo Superior de Investigaciones Científicas and is currently supported by a pre-doctoral fellowship from the Junta de Comunidades de Castilla-La Mancha.Peer reviewe

Range expansion of an exotic ungulate (Ammotragus lervia) in southern Spain: ecological and conservation concerns

Evidence of aoudad Ammotragus lervia expansion in the southeastern quarter of the Iberian Peninsula is provided based on recent field surveys. Aoudad has become common in a limited region of the southeast of Spain since its introduction as a game species in Sierra Espuña Natural Park in 1970. Its adaptability enabled it to colonise nearby areas in a short period. Apart from this source of expansion, the increasing number of aoudads in Spanish private game reserves provided other centres of dispersion. In addition, aoudads were introduced on La Palma Island (Canary Islands), becoming a serious threat to endemic flora. Of great conservation concern is the species' potential as a competitor against native ungulates inhabiting the peninsula. Surveys conducted in southern Spain documented rapid colonization of new areas and established viable populations, consisting of adult males and females and the unequivocal presence of nursery groups, in the provinces of Alicante, Almería, Granada and Murcia. Also, aoudads have spread throughout the north and centre of La Palma. There are two main conservational concerns: the necessity of conducting detailed and reliable surveys in all potential regions where the species might expand, and the urgent need of changing current game policies in order to establish reliable controls on big game reserves to prevent animals from escaping.Peer reviewe

Mechanisms of colony selection by first-year Lesser Kestrels Falco naumanni

In colonial species, first-time breeders may use the number of settled conspecifics in colony selection, but such a relationship is confused by the correlation between colony size and nest-site availability. To distinguish conspecific attraction from neutral colony selection, we experimentally increased nest availability for first-year Lesser Kestrels Falco naumanni, allowing us to dissociate the number of vacant nest-sites from colony size at the arrival time of first-year birds. Under natural conditions, the number of first-year birds settling was positively correlated with both the number of philopatric and the total number of breeding pairs (colony size) already settled. However, the probability of occupation of experimentally manipulated nests by first-year birds was independent of colony size. In experimental colonies, the number of first-year birds settling was positively correlated with the number of manipulated nest-sites but not with the number of conspecifics. Overall, these results support a neutral colony selection by first-year Lesser Kestrels based on nest-site availability.This work received financial support from the projects PCI08-0130 of the Junta de Comunidades de Castilla-La Mancha(JCCM), CGL2005-05611-C02-02 and CGL2008-00095BOS of the Ministerio de Ciencia e Innovación. During this work, G.C. and J.O. were supported by predoctoral fellowships fromthe JCCM and the European Social Fund.Peer Reviewe

Increase of heterozygosity in a growing population of lesser kestrels

The lesser kestrel (Falco naumanni) suffered a sharp population decline over much of its European distribution range in the middle of the twentieth century. Still declining in some areas, the species has recently experienced a notable population recovery in certain regions. We examined the genetic diversity variation in a growing population of lesser kestrels from Central Spain over a 6-year period (2000–2005). The population studied showed a rapid demographic expansion, increasing in the number of both breeding pairs and colonies. Annual average heterozygosity and allelic diversity increased and genetic similarity between potential mates decreased over the study period. Several immigrants regularly arrived in the study area and introduced new alleles into the local population, pointing to immigration as the main cause contributing to the observed genetic recovery

Genetic consequences of natal dispersal in the colonial lesser kestrel

Dispersal is a life-history trait that plays a fundamental role in population dynamics, influencing evolution, species distribution, and the genetics and structure of populations. In spite of the fact that dispersal has been hypothesized to be an efficient behavioural mechanism to avoid inbreeding, the expected relationship between dispersal and mate relatedness still remains controversial. Here, we examine the genetic consequences of natal dispersal, namely the higher chance of obtaining genetically less similar mates as a result of moving from natal to breeding sites, in a lesser kestrel (Falco naumanni) population. Relatedness between individuals tended to decrease with distance between their breeding colonies, indicating that the study population follows an ‘isolation-by-distance’ pattern of spatial genetic structure. Such a fine-scale genetic structure generates a scenario in which individuals can potentially increase the chance of obtaining genetically less similar mates by dispersing over larger distances from their natal colony. Using dispersal information and genotypic data, we showed that mate relatedness decreased with natal dispersal distance, an effect that remained significant both while including and excluding philopatric individuals from the data set. These results, together with the well known detrimental consequences of reduced genetic diversity in the study population, suggest that dispersal may have evolved, at least in part, to avoid the negative fitness consequences of mating with genetically similar individuals.This work received financial support from the projects: CGL2005-05611-C02-02/BOS (Ministerio de Educación Ciencia) and PAI05-053 (Junta de Comunidades de Castilla-La Mancha). During this work, J.O. and G.C. were supported by predoctoral fellowships from the Junta de Comunidades de Castilla-La Mancha and the European Social Fund.Peer reviewe

The potential use of identification of skeletal remains for the early detection of Ammotragus: An exotic ungulate species in Southern Spain

The main aim of our work was to evaluate osteometry as a complementary tool for the early detection of Ammotragus lervia, an exotic ungulate, which currently shows an expanding trend in southeastern Spain. For this purpose, 142 metacarpi and 123 metatarsi from seven Iberian ungulate species were determined by means of a classification function. In a general way, this function works, but regarding related species (those very similar from a morphometrical view), correct determination was reached in only 44.4–90% of the cases. However, in these cases, we can use auxiliary criteria like sexually dimorphic traits, and reach 100% correct identification of bones.This work has been supported by an agreement between the Consejería de Medio Ambiente (Junta de Andalucía) and the University of Jaén.Peer reviewe

Evidence of subtle departures from Mendelian segregation in a wild lesser kestrel (Falco naumanni) population

Some alleles are inherited more frequently than expected from Mendel's rule. This phenomenon, known as transmission ratio distortion (TRD), is found in a broad variety of taxa, but it is thought to be unusual and occurs at a low frequency in any particular population. Here, we used seven microsatellite markers to search for possible TRD in a wild lesser kestrel (Falco naumanni) population. Among the nine alleles analysed with at least 200 known meioses for each sex, we found that two of them (156-AG5 in males and 362-FN1.11 in females) presented subtle (k=0.6) but significant departures from Mendelian segregation. Moreover, in a sample of 53 alleles with at least 15 known meioses, we found a positive correlation between their transmission rates and their frequencies in the population. To estimate the transmission scores for the loci and individuals, we developed a method that allowed us to discover that another locus, FP-46, showed significant TRD, despite the lack of a significant deviation from parity for the alleles considered individually. Finally, we found a consistent transmission bias both within loci and within individuals across loci. Inter-individual differences in TRD support the idea that distorters act over several loci that are evenly distributed across the whole genome, particularly in individuals bearing the distorter alleles. Overall, these findings suggest that TRD might be a more widespread phenomenon than previously revealed by analyses at the allele level.This work has been supported by el Ministerio de Ciencia e Innovación (projects: CGL2005-05611-C02-02/BOS and CGL2008-00095/BOS) and Junta de Comunidades de Castilla-La Mancha (project: PCI08-0130–3954). JO is supported by a postdoctoral JAE-Doc (CSIC) contract and GC by a pre-doctoral fellowship from the Junta de Comunidades de Castilla-La Mancha/European Social Fund (to GC).Peer Reviewe

Genetic characterization of avian malaria (Protozoa) in the endangered lesser kestrel, Falco naumanni

We genetically analyzed avian malaria (Protozoa) isolated from lesser kestrels (Falco naumanni) breeding in La Mancha, Central Spain. A total of 586 adult individuals were screened for blood parasites using a very efficient polymerase chain reaction approach that amplifies a partial segment (498 bp) of the cytochrome b gene of avian malaria of the genera Haemoproteus and Plasmodium. The prevalence of Plasmodium was 8.2%, and the prevalence of Haemoproteus was 4.1%. Sequence analyses revealed six unique lineages of avian malaria, three Plasmodium (LK5, LK6, RTSR1) and three Haemoproteus (LK2, LK3, LK4). According to sequence divergence, these lineages seem to correspond to at least three different species, although all recovered lineages could be independent evolutionary units. The third most common lineage (RTSR1) has been previously retrieved from two other avian host species, including a resident African bird species and a trans-Saharan migrant passerine, suggesting that lesser kestrels could acquire this Plasmodium lineage at their winter quarters in Africa.This work received financial support from the projects: CGL2005-05611-C02-02/BOS (MEC) and PAI05-053 (JCCM). During this work, J.O and G.C. were supported by predoctoral fellowships from the Junta de Comunidades de Castilla—La Mancha and the European Social Fund.Peer reviewe

No relationship between individual genetic diversity and prevalence of avian malaria in a migratory kestrel

Insight into the genetic basis of malaria resistance is crucial for understanding the consequences of this parasite group on animal populations. Here, we analyse the relationship between genotypic variation at 11 highly variable microsatellite loci and prevalence of three different lineages of avian malaria, two Plasmodium (RTSR1, LK6) and one Haemoproteus (LK2), in a wild population of the endangered lesser kestrel (Falco naumanni). Although we used a large sample size (584 typed individuals), we did not find any significant association between the prevalence of the studied parasite lineages and individual genetic diversity. Although our data set is large, the 11 neutral markers typed may have had low power to detect such association, in part because of the low parasite prevalence observed (less than 5% of infected birds). However, the fact that we have detected previous correlations between genetic diversity and other traits (ectoparasitism risk, fecundity) in the study population using the same panel of neutral markers and lower sample sizes suggests that other factors could underlie the absence of such a similar correlation with avian malaria. Differences in the genetics of the studied traits and in their particular basis of inbreeding depression (dominance vs. overdominance) may have led to malaria prevalence, but not other traits, being uncoupled with individual genetic diversity. Also, we cannot discard the possibility that the absence of association was a consequence of a low pathogenic effect of these particular malaria lineages on our lesser kestrel population, and thus we should not expect the evolution of genetic resistance against these parasites.This work received financial support from the projects: CGL2005-05611-C02-02/BOS (Ministerio de Educación Ciencia) and PAI05-053 (Junta de Comunidades de Castilla-La Mancha). During this work J.O. and G.C. were supported by predoctoral fellowships from the Junta de Comunidades de Castilla-La Mancha and the European Social Fund.Peer reviewe