Divergence, convergence, and the ancestry of feral populations in the domestic rock pigeon

Domestic pigeons are spectacularly diverse and exhibit variation in more traits than any other bird species [1]. In The Origin of Species, Charles Darwin repeatedly calls attention to the striking variation among domestic pigeon breeds-generated by thousands of years of artificial selection on a single species by human breeders-as a model for the process of natural divergence among wild populations and species [2]. Darwin proposed a morphology-based classification of domestic pigeon breeds [3], but the relationships among major groups of breeds and their geographic origins remain poorly understood [4, 5]. We used a large, geographically diverse sample of 361 individuals from 70 domestic pigeon breeds and two free-living populations to determine genetic relationships within this species. We found unexpected relationships among phenotypically divergent breeds as well as convergent evolution of derived traits among several breed groups. Our findings also illuminate the geographic origins of breed groups in India and the Middle East and suggest that racing breeds have made substantial contributions to feral pigeon populations.     

Population genomic,olfactory, dietary,and gut microbiota analyses demonstrate the unique evolutionary trajectory of feral pigs

Domestication is an intriguing evolutionary process. Many domestic populations are subjected to strong human-mediated selection, and when some individuals return to the wild, they are again subjected to selective forces associated with new environments. Generally, these feral populations evolve into something different from their wild predecessors and their members typically possess a combination of both wild and human selected traits. Feralisation can manifest in different forms on a spectrum from a wild to a domestic phenotype. This depends on how the rewilded domesticated populations can readapt to natural environments based on how much potential and flexibility the ancestral genome retains after its domestication signature. Whether feralisation leads to the evolution of new traits that do not exist in the wild or to convergence with wild forms, however, remains unclear. To address this question, we performed population genomic, olfactory, dietary, and gut microbiota analyses on different populations of Sus scrofa (wild boar, hybrid, feral and several domestic pig breeds). Porcine single nucleotide polymorphisms (SNPs) analysis shows that the feral population represents a cluster distinctly separate from all others. Its members display signatures of past artificial selection, as demonstrated by values of F_ST in specific regions of the genome and bottleneck signature, such as the number and length of runs of homozygosity. Generalised F_ST values, reacquired olfactory abilities, diet, and gut microbiota variation show current responses to natural selection. Our results suggest that feral pigs are an independent evolutionary unit which can persist so long as levels of human intervention remain unchanged.     

Disentangling composite colour patterns in a poison frog species

A phylogenetic approach was performed to infer whether variation in conspicuous colour-patterns of a poison frog (Dendrobatidae: Dendrobates tinctorius ) has evolved neutrally or under selection. Colour and pattern were split into components that were separately analysed and subsequently re-grouped via principal component analysis. This revealed four different 'displayed' factors on the dorsal and lateral views versus one 'concealed' factor on the ventral view. Based on the assumption that current patterns of trait variation contain information about the evolutionary history of the phenotype, we correlated these trait components to a neutrally evolving gene fragment (cytochrome b ). The concealed factor was significantly correlated with the marker fragment, which identified it as having evolved under genetic drift. Noncorrelation of all displayed factors with the marker may indicate the evolution of colour patterns on dorsum and flanks under selection. In our example, colour pattern should therefore be regarded as a multicomponent signal system.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 433–444.     

Die Taube – eine Erfolgsgeschichte. Vom Liebling der Götter zur Eroberung der Städte

The pigeon – a success story The pigeon is one of the first animals that man domesticated and it holds a special place in man's affections. The pigeon is far more than an ordinary bird since it is a symbol of different gods and human virtues. In the beginning, the pigeon used to be an ordinary biological creature until it found it's way into man's spiritual world. These associations contribute to man's love and admiration of pigeons. The feral pigeon is a wild descendant of dovecote pigeons and domestic breeds. All experiences up to now lead to the conclusion that reduction of pigeon populations can only be achieved by reducing their food supply. Because of the enormous reproduction rate of the pigeons, killing attempts without any simultaneous food reduction do not have any influence on the number of birds in the population.     

Parasites from feral pigeons as a health hazard for humans

Feral pigeons live in almost every large city in the world. The feeding of pigeons by humans, deliberately or accidentally, allows the build‐up of large populations that can cause a variety of problems, primarily the fouling of buildings and monuments by their droppings. Excreta of wild birds are a well‐known source of pathogenic microorganisms that can cause infection in man. The most important ectoparasites of feral pigeons are the red blood mite, Dermanyssus gallinae, and the pigeon tick, Argas reflexus, both of which can migrate into human living space when they lose their natural hosts. The bites of the red blood mite are irritating but harmless, whereas the pigeon tick can cause severe health problems to predisposed persons. After repeated bites from A. reflexus, people can develop IgE‐mediated (type I) allergy, which in extreme situations may lead to life‐threatening symptoms of anaphylactic shock. So far, one fatality due to A. reflexus deriving from feral pigeons has been reported. In the case of parasitic infestations of humans, the source must be removed by excluding pigeons from further breeding and by disinfection of the environment of pigeon‐breeding sites.     

Phylogeography of the last surviving populations of Rhodian and Anatolian fallow deer (Dama dama dama L., 1758)

The European fallow deer ( Dama dama dama ) is one of the most widespread cervids, and its distribution has been heavily affected by man. At present, only one wild autochthonous population is reputed to survive in Anatolia, but its census size is dramatically decreasing. This means that a significant portion of the ancestral genetic diversity of this taxon is seriously threatened. In the present study, a portion of the mitochondrial DNA (D-loop) in 37 D. d. dama specimens from three Mediterranean sites, Turkey, the island of Rhodes (Greece), and the Italian Peninsula, and seven individuals of the Persian fallow deer, Dama dama mesopotamica , was sequenced, and the results from the data analysis are interpreted in light of current archaeozoological and biogeographical knowledge. We conclude that: (1) D. d. mesopotamica is strongly differentiated from D. d. dama , confirming the results of previous genetic studies and (2) the Rhodian populations of D. d. dama , founded by humans in Neolithic times, possess a set of mitochondrial lineages, found in no other study populations. The persistence of these haplotypes is particularly significant because human-mediated processes (e.g. domestication) usually result in genetic depletion and erosion of an ancestral genetic pool. In the case of the Rhodes' population of fallow deer, we hypothesize that, during the foundation of this population, humans unknowingly preserved a remarkable portion of the original genetic diversity of the source population(s).  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 835–844.     

Incipient speciation driven by phenotypic plasticity? Evidence from sympatric populations of Arctic charr

Recent models suggest that the existence of environmentally induced polymorphisms within a single population (especially those related to foraging) facilitates the process of evolutionary divergence within a single gene pool by generating distinct phenotypic modes that are exposed to differential selection. In order to test a prediction of the phenotypic plasticity model of divergence, we used a well-documented polymorphism to disentangle the relative effects of morph and rearing environment in generating phenotypic variance. We reared first-generation offspring of two sympatric morphs of Arctic charr Salvelinus alpinus in the laboratory and compared their head morphology with that of their wild parents. Morphological characters with a known functional role in foraging were highly plastic. Rearing environment accounted for the largest component of the variation in expressed phenotype, but this environmental effect overlaid a clear (but small) genetic effect. We conclude that phenotypic plasticity has played a significant role in the evolution of this trophic polymorphism, but that the evolutionary process has progressed to the point that the gene pool is now segregated.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 611–618.     

A new diploid species of Hieracium sect. Pannosa (Asteraceae) from Bulgaria

Hieracium petrovae Vladimirov & Szeląg sp. nov. , a new diploid (2 n  = 18) species in H. sect. Pannosa Zahn, is described and illustrated from the Rhodope Mountains, South Bulgaria, and compared with related taxa. It grows in relict habitats in crevices of limestone rock together with many Balkan endemics. H. petrovae is morphologically similar to taxa from the H. pannosum , H. pilosissimum and H. heldreichii groups (collective species sensu Zahn) to some of which it is a presumed ancestral species.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 150 , 261–265.     

Evidence for asymmetric migration load in a pair of ecologically divergent stickleback populations

Gene flow between ecologically divergent populations can prevent local adaptation, resulting in lower mean fitness and directional selection within a population. Such maladaptation should tend to be stronger in populations receiving a relatively larger fraction of immigrants. We test this expectation by comparing the strength of selection in a pair of three-spine stickleback populations in adjoining but unequal-sized lake basins in British Columbia. A larger deeper basin is connected to a smaller shallower basin by a short channel that allows extensive migration between populations. The two basins contain distinct habitats and prey communities, and stickleback stomach contents and isotope ratios differ accordingly. Trophic morphology is correlated with diet, so we would expect these ecological differences to be accompanied by morphological divergence. However, high gene flow appears to constrain adaptive divergence: microsatellites indicate that the two basins represent a single panmictic gene pool, and phenotypic divergence is very subtle. As a result, fish in the smaller lake basin are subject to persistent directional selection towards a more benthic phenotype, whereas the larger population exhibits no significant selection. The results illustrate the potentially asymmetrical effect of migration-selection balance, and its effect on fitness within populations.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 273–287.     

Size-dependent mating strategies and the risk of cannibalism

The role of sexual selection in determining the nature and direction of sexual size dimorphism may depend upon the timing of sexual selection, and this may also influence the variation in male size. For example, selection through sperm competition favours smaller males in the highly sexually size dimorphic orb-weaving spider Nephila edulis , whereas larger males are better able to exclude their smaller rivals from the central hub of the web where mating takes place. We investigate experimentally the role of body size and hub tenure in determining male fertilization success when males of different sizes compete for a single female over a 24-h period that includes a period of darkness. Our results confirm that small and large males obtain similar paternity share but that, in contrast with previous studies, hub tenure does not translate into greater paternity share. Unexpectedly, smaller males are at greater risk of postmating sexual cannibalism than larger males, suggesting that natural selection through sexual cannibalism may place a lower limit on male size.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 355–363.     

Introducing cultivated trees into the wild: Wood pigeons as dispersers of domestic olive seeds

Animals may disperse cultivated trees outside the agricultural land, favoring the naturalization or, even, the invasiveness of domestic plants. However, the ecological and conservation implications of new or unexplored mutualisms between cultivated trees and wild animals are still far from clear. Here, we examine the possible role of an expanding and, locally, overabundant pigeon species (Columba palumbus) as an effective disperser of domestic olive trees (Olea europaea), a widespread cultivated tree, considered a naturalized and invasive species in many areas of the world. By analyzing crop and gizzard content we found that olive fruits were an important food item for pigeons in late winter and spring. A proportion of 40.3% pigeons consumed olive seeds, with an average consumption of 7.8 seeds per pigeon and day. Additionally, most seed sizes (up to 0.7 g) passed undamaged through the gut and were dispersed from cultivated olive orchards to areas covered by protected Mediterranean vegetation, recording minimal dispersal distances of 1.8–7.4 km. Greenhouse experiments showed that seeds dispersed by pigeons significantly favored the germination and establishment in comparison to non-ingested seeds. The ability of pigeons to effectively disperse domestic olive seeds may facilitate the introduction of cultivated olive trees into natural systems, including highly-protected wild olive woodlands. We recommend harvesting ornamental olive trees to reduce both pigeon overpopulation and the spread of artificially selected trees into the natural environment.     

Protecting buildings against feral pigeons

Feral pigeons (Columba livia, Gmelin 1789) cause different problems for building owners when using structures for daytime perching, sleeping, and breeding. Problems include fouling of building facades and pavements, transmission of allergens and pathogenic microorganisms, and infestations with ectoparasites emanating from breeding sites. Owners are primarily interested in keeping away unwanted pigeons from their property. Pest control companies offer different deterrent systems, of widely varying efficacy, for proofing buildings against feral pigeons. A better solution is avoiding attractive structures during building design or subsequent alterations of existing structures used by feral pigeons. With our study, we elaborate the relevant structural data to help to maintain a building free of pigeons. We performed experiments with free ranging feral pigeons in a feral pigeon loft in the City of Basel, Switzerland. The maximum outlet width a pigeon is not able to pass through is 4 cm; the respective outlet height is 5 cm and a pigeon-safe square opening is not larger than 6 × 6 cm. The maximum ledge width a pigeon is not able to sit on is 4 cm. The pigeon-safe angle of inclination for smooth construction materials (tinplate, glass, plastics) is 25°, for medium rough materials (wood, plane concrete) 35°, and for rough materials (sandstone, rough concrete) at least 50°. Additionally, we studied the behavioral strategies used by feral pigeons to surmount our experimental constructional restrictions, ledge width, and ledge inclinations. Our data provide the essential data to prevent feral pigeons from using building structures.     

Convergent body flattening in a clade of tropical rock-using lizards (Scincidae: Lygosominae)

The repeated occurrence of similar morphologies in organisms from similar habitats provides good evidence of convergent selection, and convergent patterns of evolutionary change. In lizards, a flattened morphology has often been noted; however, whether this trait is convergent in specific habitats has never been tested using phylogenetic methods. The present study examined patterns of morphological convergence in 18 species of tropical Lygosomine skinks from three broad habitat categories (generalist, leaf litter-dwelling, and rock-using species). In general, although there where relatively few morphological differences of species from different habitats, phylogenetic analyses revealed that rock-using species have consistently and repeatedly evolved a dorsoventrally flattened head and body. The adaptive basis of this flattened morphology is consistent with both biomechanical predictions of performance (e.g. climbing locomotion) and ecology (e.g. use of rock crevices, camouflage) of species that occupy rocky habitats.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 399–411.     

A simple method for measuring colour in wild animals: validation and use on chest patch colour in geladas (Theropithecus gelada)

Adaptive hypotheses about colour variation are widespread in behavioural ecology, and several methods of objective colour assessment have been proposed and validated for use in a wide variety of taxa. However, to date, the most objective and reliable methods of assessing colour are not readily applied to wild animals. In the present study, we present a simple method for assessing colour in unrestrained, wild subjects using digital photography. The method we describe uses a digital camera, a colour standard, and colour analysis software, and can be used to measure any part of the visible colour spectrum. We demonstrate that the method: (1) is accurate and precise across different light conditions; (2) satisfies previous criteria regarding linearity and red, green, and blue equality; and (3) can be independently validated visually. In contrast with previous digital methods, this method can be used under natural light conditions and can be readily applied to subjects in their natural habitat. To illustrate this, we use the method to measure chest colour in wild geladas ( Theropithecus gelada ). Unique among primates, geladas have a red patch of skin on their chest and neck, which, for males, is thought to be a sexually selected signal. Offering some support to this hypothesis, we found differences in chest 'redness' for males across different age groups, with males in their reproductive prime exhibiting the reddest chests.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 231–240.     

A new species of Cryptolepis (Apocynaceae: Periplocoideae) from Kenya and Tanzania, Africa

A new species of Cryptolepis is described from Kenya and Tanzania. It is a tropical forest climber and its unique second corona whorl of deeply cleft lobes distinguishes it from all other Cryptolepis species.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 343–346.     

Palaeoclimatic variation, adaptation and biogeography of inversion polymorphisms in natural populations of Drosophila robusta

Studies of natural and sexual selection in wild populations of Drosophila have historically provided strong inference for the maintenance of inversion polymorphism. Analysis of geographical variation in the Drosophila robusta chromosomal data collected over more than 50 years from 133 natural populations across eastern North America has confirmed several north–south and east–west clines in the frequencies of some gene arrangements and linked arrangement combinations. Patterns of geographical variation, including several north–south clines, revealed by regression and spatial autocorrelation analyses are concordant with palaeoclimatic shifts, Pleistocene glaciations and historical changes in the composition of North American forest communities. Because D. robusta is a sap-breeder, using the microbe-infested sap exudates of a number of deciduous tree species in which they carry out their life cycle, shifts in climate and palaeovegetation types since the formation of the eastern deciduous forests in the Miocene are hypothesized to be major factors influencing patterns of inversion polymorphisms across the range of this drosophilid species. In areas where sharp deviations in frequencies have been observed, particularly in the mid-western and western portions of the range, these divisions parallel historical geographical disjunctions in the species range that have yet to promote divergence and species formation despite the long history of D. robusta in North America.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 395–411.     

Eumelanin‐based colouration reflects local survival of juvenile feral pigeons in an urban pigeon house

Urbanisation introduces deep changes in habitats, eventually creating new urban ecosystems where ecological functions are driven by human activities. The higher frequency of some phenotypes in urban vs rural/wild areas has led to the assumption that directional selection in urban habitats occurs, which may thereby favour some behavioural and physiological traits in urban animal populations compared to rural ones. However, empirical evidence of directional selection on phenotypic traits in urban areas remains scarce. In this study we tested whether eumelanin‐based colouration could be linked to survival in two urban populations of the feral pigeon Columba livia. A number of studies in different cities pointed out a higher frequency of darker individuals in more urbanised areas compared to rural ones. To investigate whether directional selection through survival on this highly heritable trait could explain such patterns, we conducted mark–recapture studies on two populations of feral pigeons in highly urbanized areas. We predicted that darker coloured individuals would exhibit higher survival and/or philopatry (integrated into ‘local survival’) than paler coloured ones. No difference in local survival was found between adults of different colouration intensities. However, on one site, we found that darker juveniles had a higher local survival probability than light ones. Juvenile local survival on that site was also negatively correlated with the number of chicks born. This suggests the existence of colour‐ and/or density‐dependent selection processes acting on juvenile feral pigeons in urban environments, acting through differential mortality and/or dispersal.     

Two approaches for the selection of Relevant Genetic Units for Conservation in the narrow European endemic steppe plant Boleum asperum (Brassicaceae)

Several methods based on population biology, biogeography, ecology, and genetics have been traditionally used for the identification of units for conservation below the species level. We use a combination of two methods based on population genetic structure estimators and on probabilities of loss of rare alleles to identify the Relevant Genetic Units for Conservation (RGUCs). The aims were to assess the genetic diversity and population structure of the endemic steppe plant Boleum asperum (Brassicaceae), and to determine how many and which populations significantly represent the total genetic diversity and the rarest allelic variation. Despite the high amplified fragment length polymorphism genetic diversity values detected in B. asperum ( h _T = 0.744), caused probably by its hexaploidy and allogamy, moderate spatial genetic differentiation was detected among populations (< 20%) and geographical ranges (> 13%), suggesting the existence of an ancestral continuous distribution range that was fragmented into separate 'islands' in more recent historical times. Five RGUCs, accounting for the 85.10% of the total genetic variation and representative of the entire geographical distribution of the species, were selected for in situ conservation. Ex situ conservation is proposed to complement the preservation of B. asperum . This method of objective selection of populations may be applied to other candidate taxa for conservation with prior adjustment of the threshold values of diversity required for effective protection of each particular species.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 341–354.     

Tempo and constraint of adaptive evolution in Escherichia coli (Enterobacteriaceae, Enterobacteriales)

The utility of maintaining distinct macroevolutionary and microevolutionary theory has long been debated. Although population and quantitative genetics provide an extensive list of microevolutionary forces that might explain macroevolutionary trends, studies of these processes are temporally limited and may fail to fully explain macroevolutionary patterns. To understand the relationship between the macroevolutionary pattern and microevolutionary forces, we must first understand how different populations respond to a given novel environment over hundreds or even thousands of generations. This study details the tempo of fitness gain over 2000 generations in four replicate lineages from each of five different ancestral Escherichia coli clones. Adaptive tempo was measured in the evolved lineages and ancestry was a significant source of variation in that tempo. Microevolutionary theory suggests that adaptive tempo should be proportional to the distance from an optimum phenotype. Demographic fitness measures allowed estimation of the ancestral distance from an optimum in the present study. Ancestral distance from an optimum was significantly related to adaptive tempo but it did not account for all of the observed variation. This suggests the existence of both ancestor and clade specific constraints. Understanding the role of such constraints is critical to both microevolutionary and macroevolutionary theory.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 403–411.     

Habitat-specific differences in thermal plasticity in natural populations of a soil arthropod

When populations experience substantial variation in environmental conditions, they may evolve phenotypic plasticity in response to these varying selection pressures. Evolutionary theory predicts differentiation in the level of phenotypic plasticity among different habitats. We evaluated temperature-induced phenotypic responses in juvenile growth rate in natural populations of the springtail Orchesella cincta , inhabiting forest and heathland. These habitats typically co-occur but differ strongly with respect to, for example, thermal regime, relative humidity, and structure. Offspring of females from the two habitats were reared at different temperatures in climate rooms and the temperature response of juvenile growth rate and egg size was measured. We found a habitat-specific difference in plasticity of juvenile growth rate. The reaction norms of the forest populations were steeper than the reaction norms for heath populations at two replicated sampling sites. Egg weight itself was demonstrated to be a plastic trait with a higher egg weight at low temperatures, but the thermal response did not differ between habitats. We conclude that these populations have diverged due to strong local natural selection. Our results support the argument that the level of phenotypic plasticity itself can be under selection and that differentiation in reaction norms can occur even in neighbouring habitats with no barrier to gene flow.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 265–271.