Evolutionary diversification of opercle shape in Cook Inlet threespine stickleback

We investigated the evolution of a large facial bone, the opercle (OP), in lake populations of the threespine stickleback that were founded by anadromous ancestors, in Cook Inlet, Alaska. Recent studies characterized OP variation among marine and lake populations and mapped a quantitative trait locus with a large influence on OP shape. Using populations from diverse environments and independent evolutionary histories, we examined divergence of OP shape from that of the anadromous ancestor. We report preliminary evidence for divergence between benthic and generalist lake ecotypes, necessitating further investigation. Furthermore, rapid divergence of OP shape has occurred in a lake population that was founded by anadromous stickleback in the 1980s, which is consistent with divergence of other phenotypic traits and with OP diversification in other lake populations. By contrast, there has been limited evolution of OP shape in a second lake population that may have experienced a genetic bottleneck early in its history and lacks genetic variation for OP divergence. Taken together, the results obtained from these two populations are consistent with studies of other stickleback phenotypic traits that implicate ancestral variation in postglacial adaptive radiation of threespine stickleback in fresh water.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 832–844.     

PARTIAL REPRODUCTIVE ISOLATION OF A RECENTLY DERIVED RESIDENT‐FRESHWATER POPULATION OF THREESPINE STICKLEBACK (GASTEROSTEUS ACULEATUS) FROM ITS PUTATIVE ANADROMOUS ANCESTOR

We used no‐choice mating trials to test for assortative mating between a newly derived resident‐freshwater population (8–22 generations since founding) of threespine stickleback (Gasterosteus aculeatus) in Loberg Lake, Alaska and its putative anadromous ancestor as well as a morphologically convergent but distantly related resident‐freshwater population. Partial reproductive isolation has evolved between the Loberg Lake population and its ancestor within a remarkably short time period. However, Loberg stickleback readily mate with morphologically similar, but distantly related resident‐freshwater stickleback. Partial premating isolation is asymmetrical; anadromous females and smaller resident‐freshwater males from Loberg Lake readily mate, but the anadromous males and smaller Loberg females do not. Our results indicate that premating isolation can begin to evolve in allopatry within a few generations after isolation as a correlated effect of evolution of reduced body size.     

Microgeographical diversification of threespine stickleback: body shape–habitat correlations in a small, ecologically diverse Alaskan drainage

Adaptive radiations are a major source of evolutionary diversity in nature, and understanding how they originate and how organisms diversify during the early stages of adaptive radiation is a major problem in evolutionary biology. The relationship between habitat type and body shape variation was investigated in a postglacial radiation of threespine stickleback in the upper Fish Creek drainage of Cook Inlet, Alaska. Although small, the upper Fish Creek drainage includes ecologically diverse lakes and streams in close proximity to one another that harbour abundant stickleback. Specimens from ancestral anadromous and derived resident freshwater populations differed substantially and could be distinguished by body shape alone, suggesting that the initial stages of adaptation contribute disproportionately to evolutionary divergence. Body shape divergence among resident freshwater populations was also considerable, and phenotypic distances among samples from freshwater populations were associated with habitat type but not geographical distance. As expected, stream stickleback from slow-moving, structurally complex environments tended to have the deepest bodies, stickleback from lakes with a mostly benthic habitat were similar but less extreme, and stickleback from lakes with a mostly limnetic habitat were the most shallow-bodied, elongate fish. Beyond adapting rapidly to conditions in freshwater environments, stickleback can diversify rapidly over small geographical scales in freshwater systems despite opportunities for gene flow. This study highlights the importance of ecological heterogeneity over small geographical scales for evolutionary diversification during the early stages of adaptive radiation, and lays the foundation for future research on this ecologically diverse, postglacial system.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 139–151.     

Phenotypic divergence among threespine stickleback that differ in nuptial coloration

By studying systems in their earliest stages of differentiation, we can learn about the evolutionary forces acting within and among populations and how those forces could contribute to reproductive isolation. Such an understanding would help us to better discern and predict how selection leads to the maintenance of multiple morphs within a species, rather than speciation. The postglacial adaptive radiation of the threespine stickleback (Gasterosteus aculeatus) is one of the best‐studied cases of evolutionary diversification and rapid, repeated speciation. Following deglaciation, marine stickleback have continually invaded freshwater habitats across the northern hemisphere and established resident populations that diverged innumerable times from their oceanic ancestors. Independent freshwater colonization events have yielded broadly parallel patterns of morphological differences in freshwater and marine stickleback. However, there is also much phenotypic diversity within and among freshwater populations. We studied a lesser‐known freshwater “species pair” found in southwest Washington, where male stickleback in numerous locations have lost the ancestral red sexual signal and instead develop black nuptial coloration. We measured phenotypic variation in a suite of traits across sites where red and black stickleback do not overlap in distribution and at one site where they historically co‐occurred. We found substantial phenotypic divergence between red and black morphs in noncolor traits including shape and lateral plating, and additionally find evidence that supports the hypothesis of sensory drive as the mechanism responsible for the evolutionary switch in color from red to black. A newly described third “mixed” morph in Connor Creek, Washington, differs in head shape and size from the red and black morphs, and we suggest that their characteristics are most consistent with hybridization between anadromous and freshwater stickleback. These results lay the foundation for future investigation of the underlying genetic basis of this phenotypic divergence as well as the evolutionary processes that may drive, maintain, or limit divergence among morphs.     

Sexual dimorphism of head morphology in three‐spined stickleback Gasterosteus aculeatus

This study examined sexual dimorphism of head morphology in the ecologically diverse three‐spined stickleback Gasterosteus aculeatus. Male G. aculeatus had longer heads than female G. aculeatus in all 10 anadromous, stream and lake populations examined, and head length growth rates were significantly higher in males in half of the populations sampled, indicating that differences in head size increased with body size in many populations. Despite consistently larger heads in males, there was significant variation in size‐adjusted head length among populations, suggesting that the relationship between head length and body length was flexible. Inter‐population differences in head length were correlated between sexes, thus population‐level factors influenced head length in both sexes despite the sexual dimorphism present. Head shape variation between lake and anadromous populations was greater than that between sexes. The common divergence in head shape between sexes across populations was about twice as important as the sexual dimorphism unique to each population. Finally, much of the sexual dimorphism in head length was due to divergence in the anterior region of the head, where the primary trophic structures were found. It is unclear whether the sexual dimorphism was due to natural selection for niche divergence between sexes or sexual selection. This study improves knowledge of the magnitude, growth rate divergence, inter‐population variation and location of sexual dimorphism in G. aculeatus head morphology.     

Geographical variations in freshwater populations of the three-spined stickleback,Gasterosteus aculeatus,in Japan

This paper describes an investigation of morphological variation among six freshwater populations (dominantly low-plated morph) of the three-spined stickleback,Gasterosteus aculeatus, in Japan. Such populations are mainly distributed in restricted water areas within a band from Ise to Wakasa Bays, the most constricted part of Honshu Island. According to some differences in morphological variation, i.e. plate morph, number of lateral plates, body size, body shape and body colour, they were classified into two types corresponding to occurrence in the Ibi River and Lake Biwa water systems. The former type was monomorphic dominated by lowplated morphs in spring-fed water, whereas the latter was a dimorphic population consisting of low- and partially-plated morphs. My results suggest that the plate morph type is not correlated with climate nor predation but is related to geological isolation patterns during the course of the landlocking process. Conversely, variations in meristic (number of dorsal and anal fin rays and gill rakers) and morphometric (body shape and body colour) characteristics may have been related to different environmental conditions. This study also provides supporting evidence that the freshwater three-spined stickleback is a distinct species from the anadromous stickleback,G. aculeatus.     

Changes in morphological characteristics of an introduced population of the threespine stickleback <Emphasis Type="Italic">Gasterosteus aculeatus</Emphasis> in Lake Towada,northern Japan

A freshwater population of the threespine stickleback Gasterosteus aculeatus occurs at an elevation of 400m in Lake Towada in the mountains of northern Japan. Morphological characteristics of each sex of this population were studied between 1985 and 1992. The 1985 fishes were larger in standard length than even anadromous populations, as well as all other freshwater populations, so the large body size of this population was unusual for the species. The population was also characterized by large ratios of head length, eye diameter, and gill raker length, and a higher ovary volume in 1992 than in 1985, but it was not readily characterized by meristic characters. The population may have been introduced within the last 20 years (probably in the early 1980s). The large body size of the population may not reflect a defense against predators because coexisting fishes were not important predators on threespine stickleback adults. It is supposed that the change of body size was induced by environmental changes in prey abundance in the lake and/or by the longevity of fishes.     

Isolation between sympatric anadromous and resident threespine stickleback species in Mud Lake,Alaska

A sympatric pair of anadromous and resident freshwater threespine stickleback species (Gasterosteus aculeatus species complex) occurs in Mud Lake in the Matanuska-Susitna Valley, Alaska. The two forms differ in an array of morphological traits, including traits associated with predator defense (e.g., spine lengths) and trophic ecology (e.g., number of gill rakers). Mud Lake is only the third lake reported to have anadromous stickleback (which have a complete row of lateral plates) coexisting with low-plated resident stickleback in the absence of intermediate partially plated fish. Microhabitat and seasonal isolation appear to contribute to reproductive isolation between the two forms.     

Frequency of Ectodysplasin alleles and limited introgression between sympatric threespine stickleback populations

The threespine stickleback (Gasterosteus aculeatus) is primitively an anadromous or resident marine species but has repeatedly colonized fresh water, where predictable phenotypic divergence usually occurs rapidly. A conspicuous element of this divergence is change of the number and position of lateral armor plates from about 33 that cover the entire flank (complete) to <10 anterior plates (low). This difference is caused primarily by variation at the Ectodysplasin (Eda) locus. The low Eda allele appears to be rarer in two geographically adjacent anadromous populations from Cook Inlet, Alaska than in most marine or anadromous populations reported from elsewhere, and there is no evidence of elevated gene flow for Eda between anadromous and resident lake threespine stickleback populations that breed in sympatry. However, the two anadromous populations are divergent for the frequencies of two complete Eda alleles. It is not clear how monomorphic low-plated freshwater populations in Cook Inlet have almost invariably acquired ancestral low Eda alleles from anadromous ancestors in which this allele appears to be extremely rare.     

Twenty years of body shape evolution in a threespine stickleback population adapting to a lake environment

Analysis of contemporary evolution can provide important insights into the pattern and rate of phenotypic evolution. The threespine stickleback population in Loberg Lake was exterminated in 1982, and a new population was founded between 1983 and 1989 by anadromous stickleback. The body shape of the Loberg Lake population resembled that of anadromous populations in 1990, although it had diverged markedly by 1992. Between 1992 and 2009, the population evolved more slowly to resemble typical lake populations in the region, diverging approximately 68% of the distance separating its putative ancestor and the original native population by 2009. Temporal evolution is the main source of variation, although spatial heterogeneity, armour phenotype, and allometry contribute significant variation. There was no significant effect of ancestral phenotypic shape covariance on the evolutionary trajectory of this population. Temporal variation in the Loberg Lake population provides a rare glimpse into the evolutionary response of a complex trait to natural selection after a major habitat shift. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 817–831.     

Variation in female life-history traits among Alaskan populations of the threespine stickleback, Gasterosteus aculeatus L. (Pisces: Gasterosteidae)

Life-history characteristics of female threespine stickleback (Gasterosteus aculeatus) were examined in 12 populations, 11 freshwater and one anadromous, within the Cook Inlet region of Alaska. Because this area has been deglaciated during the last 20 000 years, the freshwater populations are recently derived, probably independendy, from the local marine or anadromous stickleback. Freshwater threespine stickleback have undergone considerable morphological evolution within this region, apparently in response to environmental factors including predatory regimes and environmental productivity. Our freshwater study populations were selected to sample this range of morphological variation in order to determine whether life-history traits and morphologies have followed similar evolutionary trajectories. Freshwater populations could be categorized generally into one of three ecomorphotypes: those inhabiting relatively productive lakes having one or more piscivorous fishes present, and in which the stickleback exhibit a fully developed pelvic girdle; those inhabiting low-calcium lakes that lack piscivorous fishes, and in which the pelvic structures are incomplete; those living in streams with piscivorous fishes, in which the stickleback have fully developed pelvic girdles. The anadromous population constituted a fourth ecomorphotype that lives in marine waters, and is robusdy armored. The freshwater populations showed considerable variation in all life-history traits assessed, and this variation generally corresponded to our ecomorphological classifications. Nevertheless, within each ecomorphotype there was sufficient variation to suggest that morphological and life-history traits may not always respond in the same manner in response to the same selective regime.     

Migrate or stay: terrestrial primary productivity and climate drive anadromy in Arctic char

A shift in the magnitude and timing of animal migrations is one of the most documented ecological effects of climate change. Although migrations are largely driven by spatial variation in resource gradients, few studies connect expected changes in primary production with geographic patterns in migratory behavior. Here, we link lake primary production to the occurrence of sea migrations in the partially anadromous salmonid Arctic char (Salvelinus alpinus L.). We compiled presence/absence records of anadromous char populations spanning productivity and temperature gradients along the Norwegian coast. The probability of anadromy decreased with increasing migration distance, maximum slope of the migration route and lake productivity. There was a significant interaction between lake productivity and migration distance. The negative effect of longer migration distances was more severe in lakes with higher productivity, indicating reduced relative profitability of migration with increased feeding opportunities in freshwater. Lake productivity was mainly driven by terrestrial primary production in the catchment. We predicted future distributions of anadromous char given downscaled temperature and precipitation changes projected by two different emission scenarios and global climate models (GCMs). Projected increases in temperature and precipitation in 2071–2100 increased terrestrial primary production and, compared to the control scenario (1961–1990), decreased the range of anadromous populations. The prevalence of anadromy decreased by 53% in the HadAm3H GCM with the A2 emission scenario, 61% in HadAm3H with the B2 scenario and 22% in ECHAM4 with the B2 scenario. Cross‐ecosystem studies (e.g., terrestrial to freshwater) are critical for understanding ecological impacts of climate change. In this case, climate‐driven increases in terrestrial primary production are expected to increase primary production in lakes and ultimately reduce the prevalence of anadromy in Arctic char populations.     

Community structure affects trophic ontogeny in a predatory fish

While most studies have focused on the timing and nature of ontogenetic niche shifts, information is scarce about the effects of community structure on trophic ontogeny of top predators. We investigated how community structure affects ontogenetic niche shifts (i.e., relationships between body length, trophic position, and individual dietary specialization) of a predatory fish, brown trout (Salmo trutta). We used stable isotope and stomach content analyses to test how functional characteristics of lake fish community compositions (competition and prey availability) modulate niche shifts in terms of (i) piscivorous behavior, (ii) trophic position, and (iii) individual dietary specialization. Northern Scandinavian freshwater fish communities were used as a study system, including nine subarctic lakes with contrasting fish community configurations: (i) trout‐only systems, (ii) two‐species systems (brown trout and Arctic charr [Salvelinus alpinus] coexisting), and (iii) three‐species systems (brown trout, Arctic charr, and three‐spined sticklebacks [Gasterosteus aculeatus] coexisting). We expected that the presence of profitable small prey (stickleback) and mixed competitor–prey fish species (charr) supports early piscivory and high individual dietary specialization among trout in multispecies communities, whereas minor ontogenetic shifts were expected in trout‐only systems. From logistic regression models, the presence of a suitable prey fish species (stickleback) emerged as the principal variable determining the size at ontogenetic niche shifts. Generalized additive mixed models indicated that fish community structure shaped ontogenetic niche shifts in trout, with the strongest positive relationships between body length, trophic position, and individual dietary specialization being observed in three‐species communities. Our findings revealed that the presence of a small‐sized prey fish species (stickleback) rather than a mixed competitor–prey fish species (charr) was an important factor affecting the ontogenetic niche‐shift processes of trout. The study demonstrates that community structure may modulate the ontogenetic diet trajectories of and individual niche specialization within a top predator.     

Two morphological types in the reproductive stock of three-spined stickleback,Gasterosteus aculeatus,in Lake Harutori,Hokkaido Island

Synopsis Two morphological types of the trachurus form (completely plated morph) of three-spined stickleback, Gasterosteus aculeatus, are found in Lake Harutori, Kushiro, east district of Hokkaido Island. For convenience, the two forms are referred to as ‘large type’ and ‘small type’ on the basis of body length (discrimination at 70 mm in length). The two types were examined for morphometric, meristic and reproductive characteristics. They differ in body length, the number of lateral plates, means of the 2nd dorsal spine length/body length and of the pelvic spine length/body length, the relationship between body length and head length, between body length and gonad weight, between body weight and gonad weight, and between body length and ovarian egg number; significant differences were present for each sex. These two types were compared with the anadromous stickleback migrating into the freshwater area near Lake Harutori to breed. The anadromous fish was morphologically much more similar to the large type than the small type. It is suggested that the large type is also an anadromous form and the small type is the permanent freshwater resident.     

Effects of Partially Anadromous Arctic Charr (Salvelinus alpinus) Populations on Ecology of Coastal Arctic Lakes

Little research has been conducted on effects of iteroparous anadromous fishes on Arctic lakes. We investigated trophic ecology, fish growth, and food web structure in six lakes located in Nunavut, Canada; three lakes contained anadromous Arctic charr (Salvelinus alpinus) whereas three lakes did not contain Arctic charr. All lakes contained forage fishes and lake trout (Salvelinus namaycush; top predator). Isotope ratios (δ¹³C, δ¹⁵N) of fishes and invertebrates did not differ between lakes with and without anadromous Arctic charr; if anadromous Arctic charr deliver marine-derived nutrients and/or organic matter to freshwater lakes, these inputs could not be detected with δ¹³C and/or δ¹⁵N. Lake trout carbon (C):nitrogen (N) and condition were significantly higher in lakes with Arctic charr (C:N = 3.42, K = 1.1) than in lakes without Arctic charr (C:N = 3.17, K = 0.99), however, and ninespine stickleback (Pungitius pungitius) condition was significantly lower in lakes with Arctic charr (K = 0.58) than in lakes without Arctic charr (K = 0.64). Isotope data indicated that pre-smolt and resident Arctic charr may be prey for lake trout and compete with ninespine stickleback. Linear distance metrics applied to isotope data showed that food webs were more compact and isotopically redundant in lakes where Arctic charr were present. Despite this, lake trout populations in lakes with Arctic charr occupied a larger isotope space and showed greater inter-individual isotope differences. Anadromous Arctic charr appear to affect ecology and feeding of sympatric freshwater species, but effects are more subtle than those seen for semelparous anadromous species.     

Heterozygosity and phylogenetic relationship of Japanese threespine stickleback (<Emphasis Type="Italic">Gasterosteus aculeatus</Emphasis>) populations revealed by microsatellite analysis

Heterozygosity and phylogenetic relationships were analyzed using eight microsatellite loci for nine freshwater and one anadromous populations of threespine stickleback Gasterosteus aculeatus from northern Japan. Fis values and Hardy–Weinberg tests indicated that nine of the populations were in heterozygote deficit. Furthermore, on-going isolation of most of the populations by artificial destruction of population connection and habitat deterioration suggested that nearly all of the populations are likely to be in a threatened condition. Phylogenetic analysis revealed that there were two distinctive phylogenetic groups, each of which was composed of two and three freshwater populations. These populations included in the groups except ones recently established were located in inland waters and isolated from anadromous populations, so that they may have been able to keep unique genetic features. Therefore, it is necessary to protect as many populations as possible within the local population network including these inland populations. On the other hand, for freshwater populations located near the river mouth but recently isolated from anadromous populations by weir construction, it is important to remove or ameliorate the barrier and recover the gene flow. Microsatellite markers are important tools to monitor the population network and gene flow.     

Recent parallel divergence in body shape and diet source of alewife life history forms

Recent work suggests that juvenile alewives (Alosa pseudoharengus) share similar phenotypes among independently derived landlocked (freshwater resident) populations. Based on this observation, it is possible that the alewife life history forms represent a case of parallel adaptive divergence. To further evaluate this hypothesis, we describe patterns of body shape divergence between anadromous and landlocked alewife life history forms using geometric morphometrics. Our results suggest that body shape differs significantly between juveniles of the alewife forms: anadromous fish were more robust, with larger heads and deeper caudal peduncles, while landlocked fish from three independently isolated populations were more fusiform with thinner caudal peduncles and smaller heads. These differences matched population level dietary patterns, which suggest that anadromous fish consumed more littoral resources than landlocked fish. Finding consistent differences across populations of the same form supports the notion that landlocked alewives have diverged from their anadromous ancestors in a parallel manner, in response to pressures associated with being isolated in freshwater lakes. Comparing alewife phenotypes to expectations from the literature suggests that neither migration distance of the population, nor the relative availability of habitats in each lake, are likely drivers of the pattern we report. Instead, the pattern is consistent with the hypothesis that divergence between alewife forms results from the distinct effects of each form on its zooplankton prey.     

Evolutionary history of an adaptive radiation in species pairs of threespine sticklebacks (Gasterosteus): insights from mitochondrial DNA

Species pairs of threespine stickleback, Gasterosteus aculeatus, co-exist in several lakes in the Strait of Georgia, southwestern British Columbia. One species, ‘benthics’ is robust-bodied and is morphologically and behaviourally specialized for benthivory. The other species, ‘limnetics’ is specialized for planktivory in open-water habitats of the lakes. We examined mitochondrial DNA restriction site variation in benthic and limnetic sticklebacks as well as in solitary freshwater, anadromous (sea-run), and marine populations to test: (i) if benthic and limnetic pairs have evolved only once or multiple times (parallel evolution) and (ii) if the species have evolved sympatrically, or allopatrically from ‘double invasions’ of lakes by ancestral anadromous/marine sticklebacks. Stickleback mtDNA comprised a single clade with a low (mean = 0.40%) degree of sequence divergence among the 77 haplotypes resolved. Most nucleotide diversity (97%) was found within (rather than among) populations of anadromous/marine sticklebacks whereas most diversity (77%) was found among populations in freshwater sticklebacks. Significant differences in haplotype frequencies were found between benthics and limnetics in three of the four species pair lakes examined, but in all cases the pairs within lakes were characterized by unique assemblages of closely related haplotypes. Hierarchical clustering of divergence estimates suggested that comparable species from different lakes have originated independently in all lakes because in no case did comparable species from different lakes cluster together. Divergent species within lakes tended to be more closely related to one another than to species in other lakes and there were two cases were benthics and limnetics within a particular lake were monophyletic. In two of the four two-species lakes, limnetics were less divergent from putative ancestral anadromous/marine stickleback as predicted by the double invasion hypothesis, but in the two other lakes benthics were less divergent. Our data argue strongly that the species pairs have evolved independently in each lake were they now co-exist. Further, in two lakes our data are consistent with the species having evolved by sympatric divergence, but allopatric divergence followed by introgression of mtDNA that has obscured ancestral relationships cannot be discounted completely. Finally, despite remaining uncertainty about the geography of speciation, the species appear to have evolved in the face of gene flow arguing that natural selection acting on trophic ecology has been a major component of ecological speciation in sticklebacks.     

Phylogeography and adaptation genetics of stickleback from the Haida Gwaii archipelago revealed using genome‐wide single nucleotide polymorphism genotyping

Threespine stickleback populations are model systems for studying adaptive evolution and the underlying genetics. In lakes on the Haida Gwaii archipelago (off western Canada), stickleback have undergone a remarkable local radiation and show phenotypic diversity matching that seen throughout the species distribution. To provide a historical context for this radiation, we surveyed genetic variation at >1000 single nucleotide polymorphism (SNP) loci in stickleback from over 100 populations. SNPs included markers evenly distributed throughout genome and candidate SNPs tagging adaptive genomic regions. Based on evenly distributed SNPs, the phylogeographic pattern differs substantially from the disjunct pattern previously observed between two highly divergent mtDNA lineages. The SNP tree instead shows extensive within watershed population clustering and different watersheds separated by short branches deep in the tree. These data are consistent with separate colonizations of most watersheds, despite underlying genetic connections between some independent drainages. This supports previous suppositions that morphological diversity observed between watersheds has been shaped independently, with populations exhibiting complete loss of lateral plates and giant size each occurring in several distinct clades. Throughout the archipelago, we see repeated selection of SNPs tagging candidate freshwater adaptive variants at several genomic regions differentiated between marine–freshwater populations on a global scale (e.g. EDA, Na/K ATPase). In estuarine sites, both marine and freshwater allelic variants were commonly detected. We also found typically marine alleles present in a few freshwater lakes, especially those with completely plated morphology. These results provide a general model for postglacial colonization of freshwater habitat by sticklebacks and illustrate the tremendous potential of genome‐wide SNP data sets hold for resolving patterns and processes underlying recent adaptive divergences.     

Adaptation and constraint in a stickleback radiation

The evolution of threespine sticklebacks in freshwater lakes constitutes a well‐studied example of a phenotypic radiation that has produced numerous instances of parallel evolution, but the exact selective agents that drive these changes are not yet fully understood. We present a comparative study across 74 freshwater populations of threespine stickleback in Norway to test whether evolutionary changes in stickleback morphology are consistent with adaptations to physical parameters such as lake depth, lake area, lake perimeter and shoreline complexity, variables thought to reflect different habitats and feeding niches. Only weak indications of adaptation were found. Instead, populations seem to have diversified in phenotypic directions consistent with allometric scaling relationships. This indicates that evolutionary constraints may have played a role in structuring phenotypic variation across freshwater populations of stickleback. We also tested whether the number of lateral plates evolved in response to lake calcium levels, but found no evidence for this hypothesis.