Three sympatric forms of Arctic charr Salvelinus alpinus complex (Salmoniformes,Salmonidae) from Lake Kamkanda,Northern Transbaikalia

We studied morphology, size and age structure, growth, feeding, and variation at microsatellite loci of three forms of Arctic charr Salvelinus alpinus complex (dwarf, small, large) from mountain Lake Kamkanda in River Olekma basin, northern Transbaikalia. The forms differ in meristic and morphometric characters, external appearance and size. The small form distinctly differs from the dwarf and large forms in higher number and length of gill rakers. The forms differ in growth rate; however, differences in growth between the dwarf and the small forms are not as large as between sympatric dwarf and small charr from other Transbaikalian lakes. The large form is heterogeneous in growth rate. The small form matures one year earlier than the dwarf form and has a shorter life span. The dwarf form is a benthophage, the small form is a planktophage, and the large form is a predator. The dwarf form spawns in September, while the small form spawns in November-December, and there is no overlap in their spawning time. The three forms have clear genetic differences, which support their reproductive isolation. It is assumed that the three forms of Arctic charr originated within Lake Kamkanda on the basis of trophic polymorphism and spawning time displacement and attained a high degree of morphological and genetic divergence.     

Two Reproductive Life History Types of Kokanee, Onchorynchus nerka, Exhibit Multivariate Morphometric and Protein Genetic Differentiation

Two reproductive types of kokanee are found in Okanagan Lake, British Columbia: one form that spawns in streams, and another that spawns approximately 2–4 weeks later along beaches of the lake. We examined the levels and patterns of genetic and morphometric variation among three populations (1 beach and 2 stream populations) to better understand life history differentiation. We assayed allozyme variation at 74 loci and identified 15 P_0.095 loci. Average F_ST was 0.041 among the three samples; the two stream-spawning populations grouped together in genetic distance analyses. We examined multivariate morphometric variation and fin size/shape variation using 35 truss characters. Populations did not sort morphologically by spawning type in principal component and relative warp analyses. Instead both analyses indicated that one stream-spawning population (Peachland Creek) was significantly more robust and had shallower caudal fins in comparison to the other samples. Second, clear multivariate shape differences between males and females were identified involving jaw size, mid-body dimensions, and caudal fin shape. A simple bivariate plot of tail `forkness' against fork length demonstrated that males had more forked tails and females had flattened tails. Level of differentiation of secondary sexual characteristics did not vary between the spawning types, although the beach-spawning population and one stream-spawning population did not show the strong, gender-discriminating variability in tail forkness. Although these two reproductive life history types of kokanee form discrete genetic groups, environmental differences are apparently insufficient to effect consistent differences in body shape, or fin size and shape between stream and beach-spawning morphs.     

Forest fragmentation and plant reproductive success: a case study in four perennial herbs

We assessed the effects of habitat fragmentation on reproductive success in natural populations of four forest herbs with differing life-history traits and whose distribution patterns appeared to be negatively affected by decreased habitat size and/or increased isolation: Carex sylvatica, Galium odoratum, Sanicula europaea and Veronica montana. Our aims were to test (1) whether habitat size and isolation are positively correlated with population size and isolation, respectively, (2) whether plant reproductive success, a major component of plant fitness, is reduced in small and/or isolated populations when also accounting for differences in habitat quality (edaphic conditions, light intensity) and the effects of plant size, and (3) whether species with different life histories are affected differently. There were significant positive relationships between habitat and population size and between habitat and population isolation in some, but not all of the species. We mostly found no negative effects of small population size or isolation on reproduction. However, reproductive success was reduced in small populations of Sanicula, and this effect was independent of differences in plant size and environmental conditions. The reduced fecundity in small populations may be a consequence of the Allee-effect, a possible mechanism being pollen limitation. Furthermore, the proportion of flowering ramets was reduced in small and isolated populations of Galium, which may have been caused by changes in population structure. Lastly, we found some evidence for largely outcrossing, non-clonal species to be more sensitive to reductions in population size, at least in terms of their reproductive success.     

Chromosomal variation segregates within incipient species and correlates with reproductive isolation

Reproductive isolation is a critical step in the process of speciation. Among the most important factors driving reproductive isolation are genetic incompatibilities. Whether these incompatibilities are already present before extrinsic factors prevent gene flow between incipient species remains largely unresolved in natural systems. This question is particularly challenging because it requires that we catch speciating populations in the act before they reach the full‐fledged species status. We measured the extent of intrinsic postzygotic isolation within and between phenotypically and genetically divergent lineages of the wild yeast Saccharomyces paradoxus that have partially overlapping geographical distributions. We find that hybrid viability between lineages progressively decreases with genetic divergence. A large proportion of postzygotic inviability within lineages is associated with chromosomal rearrangements, suggesting that chromosomal differences substantially contribute to the early steps of reproductive isolation within lineages before reaching fixation. Our observations show that polymorphic intrinsic factors may segregate within incipient species before they contribute to their full reproductive isolation and highlight the role of chromosomal rearrangements in speciation. We propose different hypotheses based on adaptation, biogeographical events and life history evolution that could explain these observations.     

Captive breeding of two insular populations of Pachyrhynchus sarcitis (Coleoptera: Curculionidae) from Lanyu and Babuyan Islands

Endemic species of oceanic islands are vulnerable due to their geographical isolation and small population sizes. For endangered island species, captive breeding program is an important strategy for conservation and sustainable management. Pachyrhynchus sarcitis is a flightless weevil decorated with colourful markings and distributed exclusively on several islets of Southeast Asia including Lanyu, Ludao, and Babuyan Islands. The life history of threatened Pachyrhynchus species of these islands was poorly known. This study reared P. sarcitis from Lanyu and Babuyan Islands for the first time in the laboratory using their host plant (Leea guineensis). The two island populations showed significant differences in instar numbers, in which the weevils from Lanyu Island had a higher instar number regardless the length of developmental duration. The adult body size of both sexes of the Babuyan population became smaller under laboratory condition; whereas Lanyu population in the laboratory did not show this trend. The differences in larval development might suggest local adaptation to the host plants or other life history associated characteristics which requires further research.     

Genetic,Ecological and Morphological Divergence between Populations of the Endangered Mexican Sheartail Hummingbird (Doricha eliza)

The Mexican Sheartail (Doricha eliza), an endangered hummingbird, is endemic to Mexico where two populations have a disjunct distribution. One population is distributed along the northern tip of the Yucatan Peninsula whereas the other is mostly restricted to central Veracruz. Despite their disjunct distribution, previous work has failed to detect morphological or behavioral differences between these populations. Here we use variation in morphology, mtDNA and nuDNA sequences to determine the degree of morphological and molecular divergence between populations, their divergence time, and historical demography. We use species distribution modeling and niche divergence tests to infer the relative roles of vicariance and dispersal in driving divergence in the genus. Our Bayesian and maximum likelihood phylogenetic analyses revealed that Doricha eliza populations form a monophyletic clade and support their sister relationship with D. enicura. We found marked genetic differentiation, with reciprocal monophyly of haplotypes and highly restricted gene flow, supporting a history of isolation over the last 120,000 years. Genetic divergence between populations is consistent with the lack of overlap in environmental space and slight morphological differences between males. Our findings indicate that the divergence of the Veracruz and Yucatan populations is best explained by a combination of a short period of isolation exacerbated by subsequent divergence in climate conditions, and that rather than vicariance, the two isolated ranges of D. eliza are the product of recent colonization and divergence in isolation.     

Comparison of phylogeographic structure and population history of two Phrynocephalus species in the Tarim Basin and adjacent areas

An aridification of the Tarim Basin and adjacent areas since middle Pleistocene has produced significant genetic structuring of the local fauna. We examined the phylogeographic patterns, population structure and history of Phrynocephalus axillaris and Phrynocephalus forsythii using a mitochondrial fragment ND4-tRNA(LEU). Phylogenetic hypotheses were constructed using maximum parsimony and Bayesian inference, and the divergence times of major lineages were estimated by BEAST. Population structure and history were inferred by nested clade analysis, neutrality tests, mismatch distribution, and isolation by distance analysis. The two species might have experienced different evolutionary history throughout their current distribution. For P. forsythii, a vicariant event, as a consequence of geological isolation and desert expansion, might have produced the significant divergence between the Tarim and the Yanqi populations. For P. axillaris, populations of the Yanqi, Turpan and Hami Basins might have been established through dispersal during demographic expansion. Climatic fluctuations caused alternate expansion and shrinkage of rivers and oases several times, which likely led to habitat fragmentation for both species. Interaction between vicariance, dispersal and habitat fragmentation produced the current distribution and genetic diversity. The observed difference between the two species may be due partially to their different reproductive modes (ovoviviparous vs. oviparous).     

Reproductive investment and somatic growth rates in longear sunfish

Synopsis Allocation of energy to current reproduction at the expense of other functions, such as growth, can limit future reproductive potential. This cost of reproduction is a central concept of life history theory but has been difficult to verify in comparative field studies. Three levels of comparison of growth rates and reproductive investments were evaluated within and among populations of longear sunfish,Lepomis megalotis. All three demonstrated high levels of reproductive investment associated with reduced somatic growth. Within populations of central longear sunfish there are precociously mature sneaker makes and later maturing parental makes; sneakers have greater gonadosomatic index (GSI) values and slower somatic growth rates than parental makes. Between subspecies of longear sunfish grown under common conditions, there are differences in age at maturity and in the level of physiological reproductive investment that are associated with distinct differences in growth rates. Between populations of central longear sunfish inhabiting different sites, there are differences in the level of reproductive investment that are also associated with differences in somatic growth. Each comparison produced evidence that trade-offs occur between these life history traits, supporting the hypothesis that there is a cost of reproduction among male sunfish and suggesting that differences in strategies of reproductive investment contribute to variation in somatic growth.     

Small edge populations at risk: genetic diversity of the green lizard (<Emphasis Type="Italic">Lacerta viridis viridis</Emphasis>) in Germany and implications for conservation management

Edge and central populations can show great differences regarding their genetic variation and thereby also in their probability of extinction. This fact might be of great importance for the conservation strategies of endangered species. In this study we examine the level of microsatellite variability within three threatened edge populations of the green lizard subspecies Lacerta viridis viridis (Laur.) in Brandenburg (Germany) and compare the observed variation to other edge and central populations within the northern species range. We demonstrate that the northernmost edge populations contain less genetic variation in comparison to the central population. However, there were no observable significant differences to the other edge population included in this study. Surprisingly, we observed a high genetic differentiation in a small geographical range between the three endangered populations in Brandenburg, which can be explained by processes like fragmentation, isolation, genetic drift and small individual numbers within these populations. We also detected unique genetic variants (alleles), which only occurred in these populations, despite a low overall genetic variation. This study demonstrates the potential of fast evolving markers assessing the genetic status of endangered populations with a high resolution. It also illustrates the need for a comparative analysis of different regions within the species range, achieving a more exact interpretation of the genetic variation in endangered populations. This will aid future management decisions in the conservation of genetic diversity in threatened species.     

Phenotypic integration of life history and morphology: an example from three-spined stickleback, Gasterosteus aculeatus L.

Although typically life history and morphology are studied separately, they may evolve most often in concert. Therefore, a full understanding of the evolution of fish life-history patterns may require exploration of the evolutionary interplay between components of life history and other aspects of phenotype. One of the most promising approaches to understanding phenotypic integration is population comparison. This approach is particularly effective when ancestralderived relationships are understood and when multiple populations can be inferred to have evolved derived character states independently. Here we provide an example of this approach using five allopatric populations of freshwater three-spined stickleback ( Gasterosteus aculcatus L.) that have diverged in response to differences in selection regimes among the lakes they inhabit. We demonstrate a relationship between one aspect of reproductive life history, clutch volume and relative body shape. The differences are consistent with those predicted on the basis of differences in trophic habit and overall body form. Finally, we discuss the value of particular groups of fish for use in comparative studies and explore the kinds of evolutionary issues that can be addressed through population comparison.     

Is isolation by adaptation driving genetic divergence among proximate Dolly Varden char populations?

Numerous studies of population genetics in salmonids and other anadromous fishes have revealed that population structure is generally organized into geographic hierarchies (isolation by distance), but significant structure can exist in proximate populations due to varying selective pressures (isolation by adaptation). In Chignik Lakes, Alaska, anadromous Dolly Varden char (Salvelinus malma) spawn in nearly all accessible streams throughout the watershed, including those draining directly to an estuary, Chignik Lagoon, into larger rivers, and into lakes. Collections of Dolly Varden fry from 13 streams throughout the system revealed low levels of population structure among streams emptying into freshwater. However, much stronger genetic differentiation was detected between streams emptying into freshwater and streams flowing directly into estuarine environments. This fine‐scale reproductive isolation without any physical barriers to migration is likely driven by differences in selection pressures across freshwater and estuarine environments. Estuary tributaries had fewer larger, older juveniles, suggesting an alternative life history of smolting and migration to the marine environment at a much smaller size than occurs in the other populations. Therefore, genetic data were consistent with a scenario where isolation by adaptation occurs between populations of Dolly Varden in the study system, and ecological data suggest that this isolation may partially be a result of a novel Dolly Varden life history of seawater tolerance at a smaller size than previously recognized.     

Divergence in juvenile growth and life history in two recently colonized and partially isolated chinook salmon populations

In 47 families of New Zealand chinook salmon, Oncorhynchus tshawytscha derived from two populations, differing in mean spawning date by 11–17 days and in juvenile life history, neither mean ovum weight nor mean time to hatch differed between the two populations, but substantial differences in mean family weight were apparent from 6 months after fertilization. Differences in growth rates from 12 to 24 months were relatively small, suggesting that most of the divergence in size and growth occurred during the first 6 months of life. There were differences in mean weight for families spawned 11–17 days apart, the approximate interval by which peak spawning date differs between the two populations, but these did not persist beyond 7 months. Differences between the two populations are consistent with their natural life history. The slower growing population experiences cooler temperatures during stream residence and is dominated by fish which spend a year in fresh water before seaward migration, whereas the faster growing population normally experiences warmer temperatures and migrates to sea in the first year of life. These results provide further evidence of life history evolution in partially isolated chinook salmon populations within 90 years ( c. 30 generations) of becoming established.     

Population differences in host use by a seed-beetle: local adaptation, phenotypic plasticity and maternal effects

For insects that develop inside discrete hosts, both host size and host quality constrain offspring growth, influencing the evolution of body size and life history traits. Using a two-generation common garden experiment, we quantified the contribution of maternal and rearing hosts to differences in growth and life history traits between populations of the seed-feeding beetle Stator limbatus that use a large-seeded host, Acacia greggii, and a small-seeded host, Pseudosamanea guachapele. Populations differed genetically for all traits when beetles were raised in a common garden. Contrary to expectations from the local adaptation hypothesis, beetles from all populations were larger, developed faster and had higher survivorship when reared on seeds of A. greggii (the larger host), irrespective of their native host. We observed two host plant-mediated maternal effects: offspring matured sooner, regardless of their rearing host, when their mothers were reared on P. guachapele (this was not caused by an effect of rearing host on egg size), and females laid larger eggs on P. guachapele. This is the first study to document plasticity by S. limbatus in response to P. guachapele, suggesting that plasticity is an ancestral trait in S. limbatus that likely plays an important role in diet expansion. Although differences between populations in growth and life history traits are likely adaptations to their host plants, host-associated maternal effects, partly mediated by maternal egg size plasticity, influence growth and life history traits and likely play an important role in the evolution of the breadth of S. limbatus’ diet. More generally, phenotypic plasticity mediates the fitness consequences of using novel hosts, likely facilitating colonization of new hosts, but also buffering herbivores from selection post-colonization. Plasticity in response to novel versus normal hosts varied among our study populations such that disentangling the historical role of plasticity in mediating diet evolution requires the consideration of evolutionary history.     

Life history of an invasive and unexploited population of Nile tilapia (Oreochromis niloticus) and geographical variation across its native and non-native ranges

Although the life history traits of Nile tilapia, Oreochromis niloticus have been studied since the early 20^th century, the potential range of life history parameters in unexploited populations and geographical variability in life history traits are still poorly understood. We explored life history traits (age composition, growth rate, mortality, size, and age at maturity) of an invasive and unexploited population in the Tabaru River, Yonaguni-jima Island, southwestern Japan, through comparisons with exploited populations across the species’ global distribution. Analysis of sectioned otoliths from 307 fish revealed that growth and maximum age were sexually dimorphic (females growing less but having greater longevity). Large-scale comparisons with exploited populations revealed that the unexploited Tabaru River population had a greater life span than exploited populations in other regions, but the growth rate was in the middle of the range of observed values. Although a high variation in life history parameters was observed among populations (L _∞, K, maximum age), we found no significant variation in life history traits by latitude or between African and non-African populations. Such a combination of long life span and high variability in life history traits in response to environmental and fishing pressures may aid the success of non-native Nile tilapia in various environments.     

Life histories of the powan,Coregonus lavaretus (L.) (Salmonidae,Coregoninae) of Loch Lomond and Loch Eck

Four aspects of the life histories of the two populations of powan Coregonus lavaretus (L.) in Scotland are described: growth (Eck powan are shorter and with greater year to year variance than Lomond); sexual maturation (Eck powan mature younger, but at similar weight to Lomond); spawning (timing in Eck varies, but is consistent in Lomond); and recruitment/mortality (fecundity, sex ratios, and mortality also vary in the short term). Short term differences between the physiological ecology of the populations can be ascribed to the size and topography of the lochs. Long term differences are more difficult to account for, and are more important in that they may signal changes in sustainability. Conservation of powan must be considered in terms of their synecological relationships, not in isolation.     

Remarkable life history polymorphism may be evolving under divergent selection in the silverleaf sunflower

Substantial intraspecific variation in life history is rare and potentially a signal of incipient ecological speciation, if variation is driven by geographically heterogenous natural selection. We present the first report of extensive life history polymorphism in Helianthus argophyllus, the silverleaf sunflower, and examine evidence for its evolution by divergent selection. In 18 populations sampled from across the species range and grown in a common garden, most quantitative traits covaried such that individuals could be assigned to two distinct life history syndromes: tall and late flowering with small initial flowerheads, or short and early flowering with larger initial flowerheads. Helianthus argophyllus exhibits regional genetic structure, but this population structure does not closely correspond with patterns of phenotypic variation. The early‐flowering syndrome is primarily observed in populations from coastal barrier islands, while populations from the nearby mainland coast, although geographically and genetically close, are primarily late flowering. Additionally, several traits are more differentiated among regions than expected based on neutral genetic divergence (Q_ST > F_ST), including the first principal component score corresponding with life history syndrome. This discordance between patterns of phenotypic and genetic variation suggests that divergent selection is driving genetic differences in life history across the species range. If so, the silverleaf sunflower may be in early stages of ecological speciation.     

Evidence for association of chromosomal form and development time from complex clines and geographic races in the grasshopper Caledia captiva (Orthoptera: Acridadae)

Evidence for an association between chromosomal form and development time in the grasshopper Caledia captiva (F.) was obtained through comparison of two geographic taxa and analysis of a complex latitudinal cline within one of the taxa. Northern populations of the Moreton taxon possess a metacentric genome and are slow-developing. In contrast, the Torresian taxon, distributed throughout northern, coastal Australia, a region of pronounced seasonality in rainfall, and southern populations of the Moreton taxon, which inhabit a region of pronounced seasonality in temperature, Soth have an acrocentric genome and are fast-developing. The convergence of chromosomal form and development time between Torresian and southern Moreton populations appears to be driven by convergence in life history. Seasonality limits grasshoppers to one generation per year and favours fast development. The transition between relatively acrocentric southern Moreton populations and relatively metacentric northern Moreton populations is gradual but not monotonic. Instead, a shift to a bivoltine life history in the middle of the transect occurs and is associated with shifts in both development time and chromosomal form. These results imply an adaptive role for chromosomal form, although the causative link between chromosomal variation and variation in development time remains to be established.     

Molecular genetic evidence for reproductive isolation between sympatric populations of smelt Osmerus in Lake Utopia, south-western New Brunswick, Canada

Two morphologically and ecologically distinct forms of smelt, Osmerus, reside sympatrically in Lake Utopia, south-western New Brunswick, Canada. The ‘normal-sized’ form matures at greater than 200 mm standard length, averages about 31–33 gill rakers, and spawns in lake outlets. By contrast, the ‘dwarf-sized’ form matures at less than 150 mm standard length, averages 34–36 gill rakers, and spawns in small streams 3–5 weeks later than the normal form. We tested whether these sympatric forms represented ecological polymorphism within a single population or two reproductively isolated demes by assaying variation within and between forms by mitochondrial DNA (mtDNA) restriction site and nuclear minisatellite DNA analyses. Analysis of smelt mtDNA with twelve restriction enzymes resolved ten composite genotypes (differing by an average 0.27% sequence divergence) which differed markedly in frequency between the forms. Net percentage sequence divergence between the forms was O.l6%. A Wagner parsimony/ bootstrapping analysis of the restriction site presence/absence matrix, however, suggested that there were no significant distinctions between dwarf and normal smelt based on the phylogeny of composite genotypes. Hybridization studies of genomic DNA digests with a minisatellite probe indicated both that nuclear restriction fragment differentiation and the frequency of specific fragments differed significantly between the forms. Significant genetic differentiation between the sympatric forms demonstrates that they are distinct gene pools and reproductively isolated. Our molecular evidence for reproductive isolation between dwarf and normal smelt in Lake Utopia, coupled with the persistent morphological and ecological differentiation between them, argues strongly that they are behaving as distinct species. The Lake Utopia Osmerus populations provide further illustration of the potential for rapid differentiation to the level of biological species in postglacial environments.     

Allozyme divergence and evolution in the genusLens

The genusLens includes 5 taxonomic species:L. culinaris is cultivated andL. orientalis, L. odemensis, L. ervoides, andL. nigricans are wild. All the species are annual and almost exlusively selfers. The wild lentils are distributed over a large geographical area and form small disjunct populations which are composed of a small number of plants. 67Lens populations were assayed electrophoretically for 9 enzyme systems; 15 enzymic genes with 37 alleles were identified. The genetic distances (D) measured between the pairs of populations indicated a significantly greater similarity between populations belonging to the same taxonomic species. Assuming the populations represent a random sample of the variability in each of the species the genetic distances (D) between the 5 taxa were calculated. The shortest genetic distance was found betweenL. orientalis andL. culinaris. Another significant feature of the data is the apparent isolation ofL. nigricans from the other 4 species. The genetic distances between theLens species are compared to the patterns of crossability barriers between them.     

Persistent postmating,prezygotic reproductive isolation between populations

Studying reproductive barriers between populations of the same species is critical to understand how speciation may proceed. Growing evidence suggests postmating, prezygotic (PMPZ) reproductive barriers play an important role in the evolution of early taxonomic divergence. However, the contribution of PMPZ isolation to speciation is typically studied between species in which barriers that maintain isolation may not be those that contributed to reduced gene flow between populations. Moreover, in internally fertilizing animals, PMPZ isolation is related to male ejaculate—female reproductive tract incompatibilities but few studies have examined how mating history of the sexes can affect the strength of PMPZ isolation and the extent to which PMPZ isolation is repeatable or restricted to particular interacting genotypes. We addressed these outstanding questions using multiple populations of Drosophila montana. We show a recurrent pattern of PMPZ isolation, with flies from one population exhibiting reproductive incompatibility in crosses with all three other populations, while those three populations were fully fertile with each other. Reproductive incompatibility is due to lack of fertilization and is asymmetrical, affecting female fitness more than males. There was no effect of male or female mating history on reproductive incompatibility, indicating that PMPZ isolation persists between populations. We found no evidence of variability in fertilization outcomes attributable to different female × male genotype interactions, and in combination with our other results, suggests that PMPZ isolation is not driven by idiosyncratic genotype × genotype interactions. Our results show PMPZ isolation as a strong, consistent barrier to gene flow early during speciation and suggest several targets of selection known to affect ejaculate‐female reproductive tract interactions within species that may cause this PMPZ isolation.