The geography of mammalian speciation: mixed signals from phylogenies and range maps

The importance of geographic isolation in speciation has been debated since the 19th century. Since the beginning of the 20th century, the consensus has been that most speciation involves divergence in allopatry. This consensus was based largely on decades of observations by naturalists and verbal arguments against speciation without isolation. Recent attempts to quantify the importance of allopatric versus sympatric speciation using comparative methods called "age-range correlation" (ARC) suggest that allopatric speciation is more common than sympatric speciation. However, very few taxa have been studied and there are concerns about the adequacy of the methods. We propose methodological improvements including changes in the way overlap between clades is quantified and Monte Carlo methods to test the null hypothesis of no relationship between phylogenetic relatedness and geographic range overlap. We analyze 14 clades of mammals, chosen because of the availability of data and the consensus among mammalogists that speciation is routinely allopatric. Although data from a few clades clearly indicate allopatric speciation, divergence with gene flow is plausible in others and many results are inconclusive. The relative rarity of significant correlations between phylogenetic distance and range overlap may have three distinct causes: (1) post-speciation range changes, (2) relative rarity of range overlap, and (3) a mixture of geographic modes of speciation. Our results support skepticism about ARC's power for inferring the biogeography of speciation. Yet, even if few clades provide clear signals, meta-analytic approaches such as ARC may set bounds on the prevalence of alternative modes of speciation.     

Dynamics of the genetic demographic structure of the Kursk province population: migration processes

Marriage records from parish books of the second half of the 19th century and marriage records of 1967-1970 and 1993-1995 obtained from registry offices were used to analyze the dynamics of genetically significant parameters of migration and marriage structure with respect to spouses' birthplaces in populations of different hierarchical levels in the Kursk oblast. It was found that, among the persons contracting marriage (both males and females), the proportions of those who were born in the same population and those who were born in any population of the Kursk oblast decreased by about one third and one fifth, respectively, for the 130-year period. In rural and small urban populations, the coefficients of marriage migration in the 19th century were an order of magnitude lower than in the 20th century. The immigration to urban populations was maximum in the late 1960s (m = 0.745 in small towns and m = 0.680 in Kursk), and that to rural populations, in the 1990s (m = 0.344). In both urban and rural populations, the mean distance between the spouses' birthplaces has increased by several times for the period studied. The endogamy level has decreased approximately twofold: from 0.797 to 0.380 in Kursk, from 0.897 to 0.419 in small towns, and from 0.958 to 0.440 in rural districts. The marriage assortativeness with respect to birthplace was maximum in the late 19th century (K = 0.393-0.491) and minimum in the 1960s (K = 0.155-0.246). The increase in genetic diversity of the urban population of the Kursk oblast due to migration has been slowing down since the late 20th century, whereas the outbreeding level is still increasing in rural populations.     

An attempt at the historical reconstruction of the genetic demographic structure of the Moscow population at the turn of the 20th century]

The genetic demographic structure of the Moscow population at the turn of the 20th century was studied based on the data from parish books and census records. The main sources of population gene pool replenishment were analyzed, and migration coefficients and the main parameters of the model of isolation by distance were estimated. Data on so-called quasigenetic markers (ethnicity and birthplace) were used to reveal intrapopulation heterogeneity, which facilitates the adaptation of migrants to a new ethnic and cultural environment. The spatial subdivision was analyzed with the use of GST statistics. Muscovites exhibited a considerable positive assortative mating with respect to birthplace. The results of this study provide the necessary historical perspective for predicting the current genetic demographic trends in the Moscow population. It was shown that the co-efficients of marriage migration were almost the same (0.7 < m < 0.8) in the late 19th and mid-20th centuries; however, these values were two times greater than in the late 20th century. This decrease in marriage migration was accompanied by a threefold increase in the radius of centripetal migration and a threefold decrease in the level of isolation by distance. It was determined that the increase in the ethnic and genetic diversity of the Moscow population in the 20th century had started in the 1860s.     

Dynamics of the Genetic Demographic Structure of the Kursk Oblast Population: Migration Processes

Marriage records from parish books of the second half of the 19th century and marriage records of 1967–1970 and 1993–1995 obtained from registry offices were used to analyze the dynamics of genetically significant parameters of migration and marriage structure with respect to spouses' birthplaces in populations of different hierarchical levels in the Kursk oblast. It was found that, among the persons contracting marriage (both males and females), the proportions of those who were born in the same population and those who were born in any population of the Kursk oblast decreased by about one third and one fifth, respectively, for the 130-year period. In rural and small urban populations, the coefficients of marriage migration in the 19th century were an order of magnitude lower than in the 20th century. The immigration to urban populations was maximum in the late 1960s (m = 0.745 in small towns and m = 0.680 in Kursk), and that to rural populations, in the 1990s (m = 0.344). In both urban and rural populations, the mean distance between the spouses" birthplaces has increased by several times for the period studied. The endogamy level has decreased approximately twofold: from 0.797 to 0.380 in Kursk, from 0.897 to 0.419 in small towns, and from 0.958 to 0.440 in rural districts. The marriage assortativeness with respect to birthplace was maximum in the late 19th century (K = 0.393–0.491) and minimum in the 1960s (K= 0.155–0.246). The increase in genetic diversity of the urban population of the Kursk oblast due to migration has been slowing down since the late 20th century, whereas the outbreeding level is still increasing in rural populations.     

Multiple forms of selection shape reproductive isolation in a primate hybrid zone

Speciation occurs when populations diverge and become reproductively isolated from each other. Natural selection is commonly accepted to play a large role in this process, and it has been widely assumed that reproductive isolation often results as a by‐product of divergence driven by adaptation in allopatry. When such populations come into secondary contact, reinforcement can act to strengthen reproductive isolation, but the frequency and importance of this process are still unknown. Here, we explored genomic signatures of selection in allopatry and sympatry for loci associated with reproductive isolation using a natural primate hybrid zone. By analysing reduced‐representation sequencing data, we quantified admixture and population structure across a howler monkey hybrid zone and examined the relationship between locus‐specific differentiation and introgression. We detected extensive admixture that was mostly limited to the narrow contact zone. Loci with reduced introgression into the heterospecific genomic background (the pattern expected for loci associated with reproductive isolation due to selection against hybrids) were significantly more differentiated between allopatric parental populations than loci with neutral and increased introgression, supporting the hypothesis that reproductive isolation is a by‐product of divergence in allopatry. Further, loci with reduced introgression showed greater differentiation in sympatry than in allopatry, suggesting a role for reinforcement. Thus, our results reflect multiple forms of selection that have shaped reproductive isolation in this system. We conclude that reproductive isolation may have initially been driven by divergence in allopatry, but later reinforced by divergent selection in sympatry.     

Effects of species invasion on population dynamics,vital rates and life histories of the native species

Invasions occurring in natural environments provide the opportunity to study how vital rates change and life histories evolve in the presence of a competing species. In this work, we estimate differences in reproductive traits, individual growth trajectories, survival, life histories and population dynamics between a native species living in allopatry and in sympatry with an invasive species of the same taxonomic Family. We used as a model system marble trout Salmo marmoratus (native species) and rainbow trout Oncorhynchus mykiss (non-native) living in the Idrijca River (Slovenia). An impassable waterfall separates the stream into two sectors only a few 100 meters apart: a downstream sector in which marble trout live in sympatry with rainbow trout and an upstream sector in which marble trout live in allopatry. We used an overarching modelling approach that uses tag-recapture and genetic data (>2,500 unique marble and rainbow trout were sampled and genotyped) to reconstruct pedigrees, test for synchrony of population dynamics and model survival and growth, while accounting for individual heterogeneity. The population dynamics of the two marble trout populations and of rainbow trout were synchronous. We found higher prevalence of younger parents, higher mortality and lower population density in marble trout living in sympatry with rainbow trout than in marble trout living in allopatry. There were no differences in the average individual growth trajectories between the two marble trout populations. Faster life histories of marble trout living in sympatry with rainbow trout are consistent with predictions of life history theory.     

Tree-ring growth shows that the significant population decline in Norway began decades before the Black Death

The Black Death (1349–1350 in Norway) is often cited as the cause of a severe population decline and building hiatus in the middle of the 14th century. This paper analyses this hypothesis by matching the Black Death with human and environmental impacts on tree-ring growth. The number of buildings dated by dendrochronology in Norway shows a dramatic decline several decades before the plague. In Norway, the building hiatus, which has parallels in several other places in Europe, dates from the late-13th century almost to the 16th century. The first dated houses built after the plague date from the 15th century and many of the logs have exceptionally wide tree rings compared to timber from other periods. Assuming the rapid growth was because of an open landscape, the trees are likely to have grown on infields of farms abandoned due to the 14th century population decline. Since many of these fast-growing trees germinated in the early-14th century and the number of dated buildings drops dramatically several decades before the plague, the Black Death can hardly be the only reason for the population decline in Norway and one plausible explanation is that some environmental impact occurred decades earlier. The dendroclimatological evidence of cold and wet summers in the years before the plague is suggestive, but historical sources also pinpoint famine due to crop failure. They also tell of farms being abandoned several decades before the plague and mention periods of heavy rainfall and famine in the early-14th century.     

Two centuries of the Scandinavian wolf population: patterns of genetic variability and migration during an era of dramatic decline

The grey wolf (Canis lupus) was numerous on the Scandinavian peninsula in the early 19th century. However, as a result of intense persecution, the population declined dramatically and was virtually extinct from the peninsula by the 1960s. We examined historical patterns of genetic variability throughout the period of decline, from 1829 to 1979. Contemporary Finnish wolves, considered to be representative of a large eastern wolf population, were used for comparison. Mitochondrial DNA (mtDNA) variability among historical Scandinavian wolves was significantly lower than in Finland while Y chromosome variability was comparable between the two populations. This may suggest that long-distance migration from the east has been male-biased. Importantly though, as the historical population was significantly differentiated from contemporary Finnish wolves, the overall immigration rate to the Scandinavian peninsula appears to have been low. Levels of variability at autosomal microsatellite loci were high by the early 1800s but declined considerably towards the mid-20th century. At this time, approximately 40% of the allelic diversity and 30% of the heterozygosity had been lost. After 1940, however, there is evidence of several immigration events, coinciding with episodes of marked population increase in Russian Karelia and subsequent westwards migration.     

Latest developments of dendro-archaeology in the French Alps: Assessing changes in mountain forests exploitation over the last millennium

Dendrochronological studies are carried out on rural buildings in the French Alps for twenty years. In this contribution, we examine an extensive dataset of larch timbers (n = 1294) that were dendrochronologicaly dated at 139 buildings. This material yielded felling dates spanning the 11th century to the 20th century and contributed to improve our knowledge of human occupation and exploitation of mountain forests during the second millennium CE. From the 11th to the 14th century, rural populations increasingly exploited older forests. The crises of the 14th century led to a significant decrease in construction and by extension in tree cutting. The beginning of the 15th century marks a resumption of construction and felling which is continuous until the beginning of the 20th century. There is a relationship between the age of harvested trees, the number of dates and population fluctuations, which corroborates findings at the European scale. Regional trade probably influenced the stem diameters used in constructions in the immediate vicinity of the forests. Logging regulations can be perceived through variations in the age of harvested trees. The 14th century gap in felling dates would also mark a transition period in the exploitation of mountain forests and by extension a transformation of the economy.     

Palatine torus in the Greenlandic Norse.

Physical anthropologists have long been intrigued by the distinctive oral tori expressed by the medieval Norse populations of Iceland and Greenland. To assess the temporal and spatial variation of one form of oral tori, palatine torus, observations were made on all available Greenlandic Norse skeletons, as well as on samples of medieval Icelanders and Norwegians. In terms of temporal variation, 12th to 14th century (medieval) Greenlanders from the Eastern and Western settlements exhibited higher frequencies and more pronounced expressions of palatine torus compared with early 11th century Greenlanders. The early Greenlandic sample closely approximated the medieval Icelandic and Norwegian samples for total torus frequency, although the Norwegians exhibited the trait to a less pronounced degree. As degree of expression is the most distinctive aspect of torus variation among the Norse, some combination of environmental factors, including increased masticatory stress and chronic undernutrition, probably accounts for most of the difference between settlement period and medieval Greenlanders. Although palatine torus may be hereditary in part, environmental factors play a significant role in the expression of this trait.     

The Duration of Exercise Tolerance in Terms of Circulatory Indices in the Inhabitants of the Sakha Republic (Yakutia) within a Ten-Year Period

The duration of exercise tolerance of the circulatory system in different ethnic populations of Yakutia early in and at the end of the last decade of the 20th century (1988–1989 and 1998–1999) was studied. Compared to the native population (Yakuts), those born in Yakutia (Europoids) demonstrated a higher tolerance at both stages of the study. It was found that the work capacity and exercise tolerance significantly decreased in both ethnic groups during the last decade of the 20th century.     

Observations on the history of Dutch physical stature from the late-Middle Ages to the present

In the late-Middle Ages and at the onset of the early modern period, the Dutch population was taller than in the first half of the 19th century. This inference is partially based on skeletal evidence, mainly collected by the Dutch physical anthropologist George Maat and his co-workers. A spectacular increase in Dutch heights began in the second half of the 19th century and accelerated in the second half of the 20th century. At the end of the 20th century, the Dutch became tallest in the world.     

Evaluating the effects of historic bottleneck events: an assessment of microsatellite variability in the endangered,North Atlantic right whale

Commercial exploitation reduced the North Atlantic right whale (Eubalaena glacialis) population from c. 12,000 in the 11th century to around 300 by the 21st century. We examine the effect of this population decline on levels of genetic variation at 16 microsatellite loci and contrast levels of variability to that in a closely related species (E. australis). Of the 13 loci developed from the E. glacialis genome, 100% were polymorphic in E. australis. In contrast, nine loci were polymorphic in E. glacialis and four were fixed. Both allelic diversity (A) and heterozygosity (H) were significantly lower in E. glacialis than E. australis (A = 3.2 +/- 2.6 vs. A= 6.9 +/- 3.3, P < 0.001; H= 0.31 +/- 0.25 vs. H= 0.72 +/- 0.23, P < 0.001, respectively). Bottleneck anlayses indicate that the population is in mutation-drift equilibrium and that a genetic bottleneck did not occur during the most recent decline (18th-20th centuries). Nevertheless, low frequency alleles are relatively uncommon in E. glacialis, suggesting that genetic variability has been reduced in this population. Possible origins of low genetic variability are discussed, including the slow but continual erosion of alleles during the 800-year period of decline.     

Parallel evolution and ecological selection: replicated character displacement in spadefoot toads

Ecological character displacement—trait evolution stemming from selection to lessen resource competition between species—is most often inferred from a pattern in which species differ in resource-use traits in sympatry but not in allopatry, and in which sympatric populations within each species differ from conspecific allopatric populations. Yet, without information on population history, the presence of a divergent phenotype in multiple sympatric populations does not necessarily imply that there has been repeated evolution of character displacement. Instead, such a pattern may arise if there has been character displacement in a single ancestral population, followed by gene flow carrying the divergent phenotype into multiple, derived, sympatric populations. Here, we evaluate the likelihood of such historical events versus ongoing ecological selection in generating divergence in trophic morphology between multiple populations of spadefoot toad (Spea multiplicata) tadpoles that are in sympatry with a heterospecific and those that are in allopatry. We present both phylogenetic and population genetic evidence indicating that the same divergent trait, which minimizes resource competition with the heterospecific, has arisen independently in multiple sympatric populations. These data, therefore, provide strong indirect support for competition''s role in divergent trait evolution.     

Allopatry increases the balance of phylogenetic trees during radiation under neutral speciation

The shape of a phylogenetic tree is defined by the sequence of speciation events, represented by its branching points, and extinctions, represented by branch interruptions. In a neutral scenario of parapatry and isolation by distance, species tend to branch off the original population one after the other, leading to highly unbalanced trees. In this case the degree of imbalance, measured by the normalized Sackin index, grows linearly with species richness. Here we claim that moderate values of imbalance for trees with large number of species can occur if the geographic distribution involves more than one deme (allopatry) and speciation is parapatric within demes. The combined values of balance (normalized Sackin index) and species richness provide an estimate of how many demes were involved in the process if it happened in such neutral scenario. We also show that the spatial division in demes moderately slows down the diversification process, portraying a neutral mechanism for structuring the branch length distribution of phylogenetic trees.     

Density compensation in allopatric populations of the pygmy shrewSorex minutus on Gotland and the outer hebrides: evidence for the effect of interspecific competition

Pygmy shrewsSorex minutus occur allopatrically in Ireland, the Outer Hebrides and Gotland in Sweden, and sympatrically with common shrewsS. minutus on the European mainland and in England. Competition theory redicts higher population density in allopatric populations as a consequence of relaxed competition. Here this prediction is tested by comparing allopatric populations of pygmy shrews in the Outer Hebrides and in Gotland with sympatric populations on the Swedish mainland. Population densities were higher in allopatry than in sympatry. Lower summed densities in allopatry, which is predicted by niche theory, was found on the Outer Hebrides, but not on Gotland. Size distributions of carabids, which are important food items, could not explain the differences in population densities. Since both avian predators and snakes are present in all areas, population density of pygmy shrews during autumn in concluded to be regulated primarly by competition with common shrews in areas of sympatry.     

Molecular genetic examination of the polymorphic Arctic charr Salvelinus alpinus of Thingvallavatn, Iceland

Thingvallavatn, Iceland contains two sympatric morphotypes (benthic and limnetic) of Arctic charr Salvelinus alpinus. Each morphotype is composed of two morphs and these differ markedly in ecology, behaviour and life history. We used molecular genetic approaches to test whether (i) genetic heterogeneity exists among morphs and (ii) if morphs arose in allopatry and came into secondary contact or arose sympatrically within the lake through genetic segregation and/or phenotypic plasticity. Direct sequencing of 275 bp of the mitochondrial DNA (mtDNA) control region, mtDNA restriction fragment length polymorphisms and single locus minisatellite analyses detected insufficient variation to test our hypotheses. Analysis of multilocus minisatellite band sharing detected no significant differences between morphs within the same morphotype. However, significant differences among morphs belonging to different morphotypes suggest some genetic heterogeneity in Thingvallavatn charr. Limnetic charr from Thingvallavatn were more similar to sympatric benthic charr than to allopatric limnetics from two other Icelandic lakes. This suggests that the Thingvallavatn morphs arose sympatrically within the lake rather than in allopatry followed by secondary contact.     

Biotic disturbance in expanding subarctic forests along the eastern coast of Hudson Bay

* The past and present occurrence of insect disturbance on white spruce (Picea glauca) trees was evaluated at their northern range limit on the eastern coast of Hudson Bay, and its effects on tree growth and population dynamics studied. * Three sites were sampled along an altitudinal gradient. Ring-width chronologies and stem analysis were used to evaluate tree growth. The occurrence of holes in the bark, of resin pockets and blue-stain fungi, and ring-width evidence for growth releases were used to assess the impact of bark beetle. * The white spruce population was established at these sites in the 17th century. Since their establishment, the spruce trees have developed a tree growth form, except at the uppermost site, where severe growth suppression occurred in the 19th century. Bark beetle and blue-stain fungi occurred with different timing and intensity. Their highest occurrence, associated with high mortality rates, was at the lowest site in the late 20th century. In the uppermost sites, biotic disturbance has occurred since the 18th century, associated with evidence for mechanical disturbance. * The simultaneous arrival of white spruce in the area resulted in a synchronous onset of spruce beetle activity driven by tree ageing. Unfavourable climatic conditions affected tree growth severely in the most exposed sites.     

The biological standard of living in 19th century Mexico and in the American West

During the mid-19th century, the United States acquired Texas and large parts of Mexican territory with the vast Mexican-born population. This paper considers the biological standard of living of the part of this population that was incarcerated in American prisons. We use their physical stature as a proxy for their biological welfare. These data confirm earlier results which showed that adult heights tended to stagnate in Mexico during the late-19th century despite considerable social and political turmoil. While there is some evidence of a decline in height among youth, the decline is slight (<1 cm). As in other 19th century samples, farmers were the tallest. Americans were taller than Mexican prisoners by about 2 cm.     

Beta-thalassemia genes in French-Canadians: haplotype and mutation analysis of Portneuf chromosomes.

beta-Thalassemia minor occurs at approximately 1% frequency in French-Canadians--in families residing in Portneuf County (population approximately 40,000) of Quebec province. We found eight different RFLP haplotypes at the beta-globin gene cluster in 37 normal persons and in 12 beta-thalassemia heterozygotes from six families. beta-Thalassemia genes in these families associated with two haplotypes only: Mediterranean I and Mediterranean II. There were two different beta-thalassemia mutations segregating in the Portneuf population: an RNA processing mutation (beta(+)IVS-1,nt110) on haplotype I (five families) and a point mutation leading to chain termination (beta(0) nonsense codon 39) on haplotype II (one family). The distribution of 5' haplotypes on normal beta A Portneuf chromosomes compared with other European populations was most similar to that in British subjects (data for French subjects have not yet been reported). Genealogical reconstructions traced the ancestry of carrier couples to settlers emigrating from several different regions of France to New France in the 17th century. These findings indicate genetic diversity of a greater degree among French-Canadians than recognized heretofore.