Interlocality variation in speed of moult in the Citril Finch Serinus citrinella

The speed of avian moult has been related to body condition, and this to habitat quality. These facts suggest a relationship between habitat quality and moult speed. We looked for evidence of such a relationship in the Citril Finch Serinus citrinella in the Pyrenees, where, because of a Mediterranean influence, there is an important bioclimatic contrast between north- and south-facing slopes of the same mountain. This contrast strongly affects the general body condition of birds there (higher body mass, fat score and survival on north-facing slopes). Citril Finches in the higher quality area moulted more rapidly (measured as residual wing raggedness) than those in the lower quality area, the two sampling localities being only 5 km apart. As the birds initiated moult at the same time in the two locations, this difference in speed cannot have resulted from a differential effect of photoperiod. This stresses the importance of habitat selection, even at very small scales, for the completion of moult.     

Inter-locality variation in reproductive success of the citril finch Serinus citrinella

Habitat quality is generally thought to affect breeding success. We tested this effect comparing differences in clutch size and reproductive success between citril finch Serinus citrinella sub-populations closely located (<5 km) but differing in habitat quality, within the Port del Comte mountain, in the Catalonian Pre-Pyrenees. We found that birds in the low quality area (Bofia) showed significantly lower hatching, breeding and nesting success than finches in the high quality area (Vansa). These differences in reproductive success fit well with recently found differences in citril finch body mass, fat score, diet, survival rate and speed of moult between these two localities.     

Macrogeographic variations in food choice of mainland citril finches Carduelis [citrinella] citrinella versus insular Corsican (citril) finches Carduelis [citrinella] corsicanus

We studied variations in diet composition in sub-populations of mainland citril finches Carduelis [citrinella] citrinella in European mountain chains, compared to closely related insular Corsican (citril) finches Carduelis [citrinella] corsicanus on several Mediterranean Islands with a special emphasis on key food plants used during the breeding period. We found that citril finch sub-populations of the Pre-Pyrenees and the Black Forest rely mostly on mountain pine and dandelion seeds as main food resources during the breeding season. In contrast, insular Corsican finches on the Mediterranean islands of Capraia and Corsica fed mainly on various herb species. Finches in Corsica fed predominantly on shepherd’s purse, while birds from Capraia were choosing rosemary as their main food plant during the breeding period. These observations confirmed the previously expected difference in food choice between the two forms due to the observed increased niche breadth on Corsica. However, Corsican finches on Sardinia showed a food choice more similar to that of the citril finch populations, in that they fed predominantly on black pine seeds during breeding season. Thus we were not able to find clear behavioural differences in diet composition between all studied sub-populations of citril and Corsican finches. To conclude, food choice of both forms is highly adaptive and, in the different breeding areas, distinct key plants may be of special importance during the breeding season. The preference for these plants is likely to be linked to their abundance and their energetic and nutritional content.     

Geographical differentiation, acoustic adaptation and species boundaries in mainland citril finches and insular Corsican finches, superspecies Carduelis [citrinella]

Aim In birds, differentiation of acoustic characters is an important mechanism of reproductive isolation that may lead to an ethological–acoustic barrier, resulting in the formation of new species. We examined acoustic variation in mainland citril and insular Corsican finch populations, with the aim of assessing the degree of acoustic differentiation between both members of the superspecies Carduelis [citrinella] and documenting possible variation between local subpopulations that are geographically isolated. Location We chose study sites throughout the geographical ranges of citril and Corsican finches. For the citril finch, we obtained samples from the Black Forest (Germany), the Cevennes (France) and the Pyrenees (Spain); for the Corsican finch, we obtained samples from the islands Capraia and Sardinia (Italy) and Corsica (France). Methods We analysed frequent contact calls and elements of the perch song. Vocalization patterns of the study populations were compared by means of discriminant and hierarchical cluster analyses. Results There were significant differences in vocalization characteristics of perch songs and contact calls, which permitted unambiguous discrimination of citril and Corsican finch populations. However, we also detected significant differences in contact calls between mainland citril finch subpopulations. There was a pattern of clinal variation in vocalization: short, steeply modulated signals in the northern part of the geographical range (Black Forest) and long, shallowly modulated signals in the southern part (Pyrenees). Main conclusions Acoustically, mainland citril and insular Corsican finches separate well in their contact calls and perch songs. However, variation in the two vocalization patterns between subpopulations of mainland citril finches indicates that acoustic characteristics can evolve very quickly, not only on islands but also on the mainland. Local habitat differences may play a crucial role in the rapid evolution of these signals under full or partial isolation of small subpopulations. To judge the importance of signal variation as a pre‐mating isolating barrier, future studies will have to determine whether members of the distinct subpopulations are able to match their signals to each other if they re‐meet, and whether intraspecific species recognition is still possible.     

Genetic differences among mainland and insular forms of the Citril Finch Serinus citrinella

The Citril Finch Serinus citrinella breeds in several mountains of central and southern Europe and on Corsica, Sardinia and Tuscany islands. Several studies have emphasized differences in phenotype and vocalization, and some authors consider mainland and insular forms to be two distinct species. A genetic approach based on the sequences of a mitochondrial DNA fragment was used to estimate genetic distance separating several populations. Individuals from the Alps and the Pyrenees belong to one group and the individuals from Corsica to the second. These results, discussed together with data on ecology, biometrics and vocalization, suggest keeping only one polytypic species which includes two distinct forms.     

Local selection modifies phenotypic divergence among Rana temporaria populations in the presence of gene flow

In ectotherms, variation in life history traits among populations is common and suggests local adaptation. However, geographic variation itself is not a proof for local adaptation, as genetic drift and gene flow may also shape patterns of quantitative variation. We studied local and regional variation in means and phenotypic plasticity of larval life history traits in the common frog Rana temporaria using six populations from central Sweden, breeding in either open‐canopy or partially closed‐canopy ponds. To separate local adaptation from genetic drift, we compared differentiation in quantitative genetic traits (Q_ST) obtained from a common garden experiment with differentiation in presumably neutral microsatellite markers (F_ST). We found that R. temporaria populations differ in means and plasticities of life history traits in different temperatures at local, and in F_ST at regional scale. Comparisons of differentiation in quantitative traits and in molecular markers suggested that natural selection was responsible for the divergence in growth and development rates as well as in temperature‐induced plasticity, indicating local adaptation. However, at low temperature, the role of genetic drift could not be separated from selection. Phenotypes were correlated with forest canopy closure, but not with geographical or genetic distance. These results indicate that local adaptation can evolve in the presence of ongoing gene flow among the populations, and that natural selection is strong in this system.     

Adaptive differentiation in floral traits in the presence of high gene flow in scarlet gilia (Ipomopsis aggregata)

Plant–pollinator interactions are thought to be major drivers of floral trait diversity. However, the relative importance of divergent pollinator‐mediated selection vs. neutral processes in floral character evolution has rarely been explored. We tested for adaptive floral trait evolution by comparing differentiation at neutral genetic loci to differentiation at quantitative floral traits in a putative Ipomopsis aggregata hybrid zone. Typical I. aggregata subsp. candida displays slender white tubular flowers that are typical of flowers pollinated by hawkmoths, and subsp. collina displays robust red tubular flowers typical of flowers pollinated by hummingbirds; yet, hybrid flower morphs are abundant across the East Slope of the Colorado Rockies. We estimated genetic differentiation (F_ST) for nuclear and chloroplast microsatellite loci and used a half‐sib design to calculate quantitative trait divergence (Q_ST) from collection sites across the morphological hybrid zone. We found little evidence for population structure and estimated mean F_ST to be 0.032. Q_ST values for several floral traits including corolla tube length and width, colour, and nectar volume were large and significantly greater than mean F_ST. We performed multivariate comparisons of neutral loci to genetic correlations within and between populations and found a strong signal for divergent selection, suggesting that specific combinations of floral display and reward traits may be the targets of selection. Our results show little support for historical subspecies categories, yet floral traits are more diverged than expected due to drift alone. Non‐neutral divergence for multivariate quantitative traits suggests that selection by pollinators is maintaining a correlation between display and reward traits.     

Extranuclear differentiation and gene flow in the finite island model

Use of sequence information from extranuclear genomes to examine deme structure in natural populations has been hampered by lack of clear linkage between sequence relatedness and rates of mutation and migration among demes. Here, we approach this problem in two complementary ways. First, we develop a model of extranuclear genomes in a population divided into a finite number of demes. Sex-dependent migration, neutral mutation, unequal genetic contribution of separate sexes and random genetic drift in each deme are incorporated for generality. From this model, we derive the relationship between gene identity probabilities (between and within demes) and migration rate, mutation rate and effective deme size. Second, we show how within- and between-deme identity probabilities may be calculated from restriction maps of mitochondrial (mt) DNA. These results, when coupled with our results on gene flow and genetic differentiation, allow estimation of relative interdeme gene flow when deme sizes are constant and genetic variants are selectively neutral. We illustrate use of our results by reanalyzing published data on mtDNA in mouse populations from around the world and show that their geographic differentiation is consistent with an island model of deme structure.     

Historical demographic dynamics underlying local adaptation in the presence of gene flow

The range of a species is the result of the relative contribution of spatial tracking of environmental requirements and adaptation to ecological conditions outside the ancestral niche. The appearance of novel habitats caused by climatic oscillation can promote range expansion and accompanying demographic growth. The demographic dynamics of populations leave a signal in \ patterns. We modeled three competing scenarios pertaining to the circumstance of a range expansion by the Karoo Scrub‐Robin into newly available habitat resulting from the increasing aridification of southern Africa. Genetic variation was contrasted with the theoretical expectations of a spatial range expansion, and compared with data of a putative adaptive trait. We infer that this bird likely colonized the arid zone, as a consequence of adaptive evolution in a small peripheral population, followed by an expansion with recurrent exchange of migrants with the ancestral populations.     

Neutral gene flow in the presence of a selected gene with random or assortative mating

Neutral gene flow in an island model in the presence of a gene subject to selection has been analysed. The selection regime on the gene is such that the maintenance of a stable interpopulation differentiation at this locus is allowed for very low values of the migration rate. Mating at this locus may be random or assortative. Many models of zygotic or prezygotic isolating mechanisms with unifactorial inheritance are represented by this genetic model. Two general solutions for migration rates tending toward zero have been obtained for the case of migration preceding selection and vice versa. It is found that the zygotic isolating mechanisms and one type of prezygotic isolating mechanisms have the same qualitative and quantitative effects for the same values of selection coefficients and of recombination frequencies between the selected gene and the neutral gene. A second type of prezygotic mechanism behaves in a different way: the reduction of gene flow caused by such mechanisms is also a function of the “mating coefficients” of the various genotypes.     

A framework for comparing processes of speciation in the presence of gene flow

How common is speciation‐with‐gene‐flow? How much does gene flow impact on speciation? To answer questions like these requires understanding of the common obstacles to evolving reproductive isolation in the face of gene flow and the factors that favour this crucial step. We provide a common framework for the ways in which gene flow opposes speciation and the potential conditions that may ease divergence. This framework is centred on the challenge shared by most scenarios of speciation‐with‐gene‐flow, i.e. the need for coupling among different components of reproductive isolation. Using this structure, we review and compare the factors favouring speciation with the intention of providing a more integrated picture of speciation‐with‐gene‐flow.     

Local differentiation and gene dispersal distances in a Danish population of Armaria maritima

In a continuously distributed population of the self-incompatible species Armeria maritima , nine artificially delimited subpopulations were sampled in a transect with a length of 1300 m. Genetic variation was quantified with enzyme electrophoresis, which revealed five polymorphic loci. The genotypic distribution within subpopulations accorded to Hardy-Weinberg proportions, but revealed a Wahlund effect at one of the loci when the total population was considered. The Wahlund effect was supported by significant differences in allele frequencies among the nine subpopulations. The differentiation was low with a F_ST value of 0.028, which is much smaller than the previously reported value of 0.18 for 17 geographically separated Danish populations of this species. The low differentiation of the subpopulations was also revealed by an assignment test. In this test, the multilocus genotype of an individual is used to assign it to the subpopulation it has the highest likelihood to appear in. With this test, a lower percentage of individuals were assigned to the subpopulation of origin (24) than was seen with assignment of individuals to populations in Denmark (34). No abrupt changes in allele frequencies were observed. A spatial autocorrelation analysis suggested a breeding patch size of 500–700 m in diameter along the transect.     

IAA-induced xylem differentiation in the presence of colchicine

In internodal stem explants of Coleus blumei (BENTH.) colchicine(0.35 %) blocked IAA-induced wound vessel member formation butpermitted abnormal xylogenesis associated with the vascularbundles. Xylem elements differentiated in the presence of colchicinepossessed abnormal asymmetrical secondary wall sculpturing.Colchicine (0.01 %) permitted wound vessel ember formation withthe same type of abnormal wall sculpturing. These results suggestthe partial inactivation or reorientation of the microtubulesassociated with microfibril deposition in prexylem cells bycolchicine. ¹ Fulbright Professor of Botany, 1967–68. Permanent address:Department of Biological Sciences, University of Idaho, Moscow,Idaho, 83843, U. S. A.     

Absence of Mate Guarding in the Yellowhammer (Emberiza citrinella)?

The mate guarding behaviour of male yellowhammer (Emberiza citrinella) was studied with special reference to the effects of age, body size (tarsus length) and coloration of males. Measurements of intra-pair distance do at the most provide evidence for relatively lax mate guarding. On the other hand, patterns of male song activity and inter-male aggression were more in agreement with the predictions of the mate guarding hypothesis. The reasons for the comparatively low mate guarding intensity in the yellowhammer may be that males do not need to guard their mates intensely. Age differences were found in song and aggressive activity, older males singing and fighting the most. Size had no effect on guarding behaviour. Coloration was correlated with inter-male aggressiveness and conflict initiation propensity. Less colourful males fought the most in the pre-fertile period of their mates, whereas colourful and old males fought the most during the fertile period. This suggests that coloration may be an indicator of individual fighting and guarding ability.     

Local intracerebral implants of estrogen masculinize some aspects of the zebra finch song system

The song system of zebra finches is sexually dimorphic: the volumes of the song control nuclei and the neurons within these nuclei are larger in males. The song system of hatching female zebra finches is masculinized by systemic treatment with estrogen. We investigated the locus of this estrogen action by using microimplants of estradiol benzoate (EB). We implanted female zebra finch nestlings 10–13 days old with Silastic pellets containing approximately 2 μg EB at one of several sites: near the higher vocal center (HVC), in the brain distant from HVC, or in the periphery either under the skin of the breast or in the peritoneal cavity. Controls were either unimplanted or implanted near HVC with Silastic pellets without hormone. The brains were fixed by perfusion at 60 days, and the volumes of the song control regions as well as the sizes of individual neurons were measured. Neurons in HVC were lerger (more masculine) in the HVC-implanted group than in other groups, which did not differ among themselves. The size of neurons in the robust nucleus of the archistriatum (RA) and the lateral magnocellular nucleus ofthe neostriatum (lMAN) were inversely correlated with the distance of the EB pellet to HVC; neurons in RA and lMAN were larger when the EB pellets were closer to HVC. This result suggests that implants near HVC were at or near a site of estrogen action. To our knowledge, this is the first demonstration that localized brain implants of estrogen cause morphological masculinization in any species. 1994 John Wiley & Sons, Inc.     

Population differentiation and nuclear gene flow in the Dominican anole (Anolis oculatus)

Allele frequency data from nuclear microsatellite loci were used to investigate patterns of nuclear gene flow and population structure in the morphologically variable Dominican anole (Anolis oculatus). All six loci used proved to be highly polymorphic, with an average of 18.8 alleles per locus. Test for Hardy-Weinberg equilibrium revealed small numbers of heterozygote deficiencies at single loci in single populations and consistent patterns of increasingly significant heterozygote deficiency in global tests across populations and loci. No significant relationship between FST and patristic distances estimated from mitochondrial DNA sequences was detected and estimates of FIS were significantly higher in females than in males, indicating that gene flow may be sex-biased and mediated mainly by male migration. A highly significant correlation between linearized FST and loge (geographical distance) indicates that geographical proximity is a significant factor in the genetic structure of A. oculatus populations. However, levels of gene flow between morphologically differentiated parapatric populations are frequently seen to be relatively high. This supports the hypothesis of natural selection being the driving force behind the development and maintenance of morphological variation and shows that adaptive differentiation may be maintained despite the homogenizing influence of gene flow. Generally, the morphologically variable populations of A. oculatus are seen to be poor candidates for in situ speciation, but an exceptional case on the west coast of Dominica indicates that isolation resulting from vicariant events may lead to rapid differentiation at both mitochondrial and nuclear loci. This provides a possible mechanism for anole speciation on other Caribbean islands.     

Sexual differentiation of behavior in the zebra finch: effect of early gonadectomy or androgen treatment

Treatment of nestling zebra finches with estradiol benzoate (EB) has been shown to masculinize singing in females and demasculinize copulatory behavior in males, suggesting that sexual differentiation of these behaviors is under hormonal control such that testicular hormones induce the capacity for song and ovarian hormones suppress the capacity for mounting. Two experiments were carried out to obtain a more complete picture of sexual differentiation in this species. In Experiment 1, nestlings were injected daily for the first 2 weeks after hatching with testosterone propionate (TP), dihydrotestosterone propionate (DHTP), or a combination of DHTP and EB. As adults, birds were gonadectomized and implanted with TP prior to testing, then tested again after implantation with EB. Singing was not increased in females by any of the treatments. The only effect of either TP or DHTP given alone was defeminization of female proceptive behavior by DHTP. Thus androgens appear to have less influence than estrogens on sexual differentiation of behavior in this species. The combination of DHTP and EB demasculinized mounting in males. In Experiment 2, nestlings were gonadectomized at 7-9 days of age and implanted with TP prior to testing in adulthood. Early gonadectomy had little effect on later behavior; early castrated males sang, danced, and copulated normally and early ovariectomized females neither sang nor mounted.     

Local human pressures influence gene flow in a hybridizing Daphnia species complex

Anthropogenic environmental changes are considered critical drivers of the genetic structure of populations and communities through, for example, the facilitation of introgressive hybridization between syntopic species. However, the mechanisms by which environmental perturbations trigger changes in the genetic structure of populations and communities, such as the processes that determine the directionality of hybridization and patterns of mitochondrial introgression over many generations, remain largely unexplored. In this study, the changes in genetic structure of hybridizing members of the Daphnia longispina species complex were reconstructed over the last 100 years for three large temperate lakes under strong anthropogenic pressures via palaeogenetic analyses of resting egg banks. Drastic changes in the genetic structure of the Daphnia community, associated with hybridization events between D. longispina and D. galeata and subsequent introgression, were detected in Lakes Geneva and Bourget. In Lake Bourget, these changes were induced by the successful establishment of D. galeata with rising phosphorus levels and reinforced by the sensitivity of D. longispina to fish predation pressure. In Lake Geneva, the pattern of hybridization during eutrophication is more likely a function of the original taxonomic composition of the species complex in this lake. Lakes seem to require at least a meso‐oligotrophic status to allow D. galeata populations to establish and accordingly no D. galeata genotypes were found in the egg bank of oligotrophic Lake Annecy. In contrast to the generally assumed pattern of unidirectional hybridization in this species complex, bidirectional hybridization was recorded in Lakes Geneva and Bourget. Our results also demonstrate complex genetic trajectories within this species complex and highlight the irreversibility of changes in the genotypic architecture of populations driven by local human pressures. Finally, we show that extensive hybridization and introgression do not necessarily result in a large and homogenous hybrid swarm.