Life-history variations in the fluvial sculpin, Cottus nozawae (Cottidae), along the course of a small mountain stream

Life-history variations in male and female fluvial sculpins, Cottus nozawae, were studied in a small mountain stream in Hokkaido, Japan, primarily by using capture-mark-recapture methods. At three study areas established along the stream course, the majority of marked sculpins were recaptured in their original location over one or more years, indicating their long-term occupation of each restricted habitat area. Sculpin densities increased toward the upstream habitats, whereas individual growth rates were more rapid downstream. In both sexes, sculpins distributed downstream matured at a larger body size and later in life than upstream sculpins, clearly demonstrating a clinal variation in these respects. A comparison of life-history variations in C. nozawae with those in amphidromous C. hangiongensis suggests that intrapopulational life-history variations in the former might be environmentally induced, and that one of the most important determinants for the variations in Cottus species might be population density.     

High levels of genetic variation in natural populations of marine lower invertebrates

The predictions of neutralist and selectionist hypotheses have been tested many times in the past, but mostly using data only from organisms such as vertebrates, with generally low to average heterozygosities. The more recent discovery of particularly high levels of genetic variation in marine sponges and coelenterates provides an opportunity to use data from such species to contribute further to the understanding of the determinants of heterozygosity in natural populations. Therefore, 23 species of sponges and coelenterates from temperate, tropical and boreal waters were analysed by gel electrophoresis for an average of 14.3 enzyme loci per species. Mean heterozygosity values for each species were unusually high, ranging between 0.106 and 0.401. The means and variances of the heterozygosity estimates showed reasonable correlation with neutralist predictions (with both the stepwise mutation and the infinite alleles models). Population sizes were generally difficult to estimate with any confidence, but, for one sponge species for which this was possible, levels of heterozygosity again were similar to neutralist predictions, although the same was not apparently true for three species of sea anemone. No differences were found between heterozygosity levels of tropical and temperate species of sponges and coelenterates, thus apparently contradicting the selectionist ‘trophic resource stability’ and ‘temporal environmental variation’ hypotheses. Conversely, however, the consistently high levels of genetic variation found in coelenterates and sponges may be argued to be related to common biological characteristics, such as sessile life, great evolutionary ‘age’, limited ability to disperse and probable low homoeostatic capability. Our results seem, overall, to agree well with neutralist expectations for species with large, stable population sizes. Also, the mean heterozygosities, their variances and the observed and expected proportions of polymorphic loci seem to fit well with predictions based on the neutralist hypothesis. However, the selectionist ‘environmental grain’ and the ‘shifting balance’ hypotheses fit the data equally well. As with much earlier work, the problems in distinguishing between the various predictions of selectionist or neutralist ideas make it both difficult and unwise to draw definite conclusions.     

Comparative ecology of prickly sculpin,Cottus asper,and coastrange sculpin,C. aleuticus,in the Eel River,California

Synopsis We documented species' distributions, size structure of populations, abundance in mainstem and tributary streams, habitat use, and diets of prickly sculpin, Cottus asper, and coastrange sculpin, C. aleuticus, in the Eel River drainage of California, to determine the processes allowing coexistence of these very similar fishes. We observed prickly sculpins at 43 sites and coastrange sculpins at 34. The species co-occurred at 26 sites. Young-of-year coastrange sculpins were only observed within 42 km of the ocean, but young-of-year prickly sculpins were present throughout the species range. Mean, maximum, and minimum lengths of coastrange sculpins were positively correlated with distance from the ocean but no significant relationships were found for prickly sculpins. Absolute abundance of both species was highest in mainstem habitat (prickly sculpins = 0.6 sculpins m^–2 and coastrange sculpins = 0.4 sculpins m^–2) . Tributary densities of both species tended to be less than 0.1 sculpins m^–2. The species inhabited very similar habitats and had very similar diets. Coastrange sculpin populations in upstream areas were maintained by immigration from downstream areas in contrast with prickly sculpin populations that produced young-of-year fish throughout their range. Densities were probably not high enough for interspecific interactions to be important. The factors limiting the upstream distribution of the species may include high water temperatures, stability of the stream bed, and behavior of the fish. In the past, the range of sculpins within the Eel River drainage probably fluctuated with changing physical conditions. Recent introductions of exotic species that compete with and prey upon sculpins, and ongoing human activities in the drainage could result in major reductions in the distribution and abundance of one or both species.     

PCR primers for polymorphic microsatellite loci in the plant‐ant Azteca ulei cordiae (Formicidae: Dolichoderinae)

Twelve microsatellite loci were isolated from the Peruvian tropical plant‐ant, Azteca ulei cordiae (Hymenoptera: Dolichoderinae). High levels of within‐population variation were observed at most loci, with number of alleles ranging from four to 18, and heterozygosity from 0.118 to 1 per population sample. Cross‐species amplification of these loci was also successfully tested in several other species of the same ant genus Azteca.     

Does habitat specialization shape the evolutionary potential of wild bird populations?

Because specialist species evolved in more temporally and spatially homogeneous environments than generalist species, they are supposed to experience less fluctuating selection. For this reason, we expect specialists to show lower overall genetic variation as compared to generalists. We also expect populations from specialist species to be smaller and more fragmented, with lower neutral genetic diversity. We tested these hypotheses by investigating patterns of genetic diversity along a habitat specialization gradient in wild birds, based on estimates of heritability, coefficients of variation of additive genetic variance, and heterozygosity available in the literature. We found no significant effect of habitat specialization on any of the quantitative genetic estimators but generalists had higher heterozygosity. This effect was mainly a consequence of the larger population size of generalists. Our results suggest that evolutionary potential does not differ at the population level between generalist and specialist species, but the trend observed in heterozygosity levels and population sizes may explain their difference in susceptibility to extinction.     

Mixed-species brooding between two Baikal sculpins: field evidence for intra- and interspecific competition for reproductive resources

In an examination of the distribution of nesting spaces of two Baikal sculpins, Paracottus kneri and Cottus kessleri , along two transects offshore at Bolshye Koty, a total of 43 P. kneri nests, 32 C. kessleri nests and one Batrachocottus baicalensis nest was found. Males of these species established breeding nests under overhanging stones in the same manner as most freshwater sculpins, but nests of each species were clearly partitioned in separate zones. Cottus kessleri nests were found where the lake floor was mostly covered with overhanging stones. Paracottus kneri nests, however, were found where the lake floor was moderately covered with overhanging stones. Two mixed-species brooding nests with a C. kessleri guarding male were found in an area bounded by the two zones. Each of these nests comprised one P. kneri egg mass and several C. kessleri egg masses, guarded by a C. kessleri male. The nest distribution of these fishes appears to reflect intra- and interspecific competition for overhanging stones as a reproductive resource rather than interspecific difference of nest selection.     

Population demography and heterozygosity–fitness correlations in natural guppy populations: An examination using sexually selected fitness traits

Heterozygosity–fitness correlations (HFCs) have been examined in a wide diversity of contexts, and the results are often used to infer the role of inbreeding in natural populations. Although population demography, reflected in population‐level genetic parameters such as allelic diversity or identity disequilibrium, is expected to play a role in the emergence and detectability of HFCs, direct comparisons of variation in HFCs across many populations of the same species, with different genetic histories, are rare. Here, we examined the relationship between individual microsatellite heterozygosity and a range of sexually selected traits in 660 male guppies from 22 natural populations in Trinidad. Similar to previous studies, observed HFCs were weak overall. However, variation in HFCs among populations was high for some traits (although these variances were not statistically different from zero). Population‐level genetic parameters, specifically genetic diversity levels (number of alleles, observed/expected heterozygosity) and measures of identity disequilibrium (g2 and heterozygosity–heterozygosity correlations), were not associated with variation in population‐level HFCs. This latter result indicates that these metrics do not necessarily provide a reliable predictor of HFC effect sizes across populations. Importantly, diversity and identity disequilibrium statistics were not correlated, providing empirical evidence that these metrics capture different essential characteristics of populations. A complex genetic architecture likely underpins multiple fitness traits, including those associated with male fitness, which may have reduced our ability to detect HFCs in guppy populations. Further advances in this field would benefit from additional research to determine the demographic contexts in which HFCs are most likely to occur.     

Population genetics of haplodiploid insects

Genic variation of seven species of Hymenoptera is described, using electrophoretic techniques. The heterozygosities range from 0.033 to 0.084. An average heterozygosity is calculated for 23 species of haplodiploid insects, and this value is significantly different from the same value for 18 Drosophila species or for 24 diploid insect species (including Drosophila). The niche width-genetic variation hypothesis is rejected as an explanation. A comparison of selection models and neutral models shows that both hypotheses are capable of explaining the data.     

DEVELOPMENTAL NOISE,PHENOTYPIC PLASTICITY,AND ALLOZYME HETEROZYGOSITY IN DAPHNIA

Previous theories and studies have postulated negative correlations between allozyme heterozygosity and developmental noise and between heterozygosity and phenotypic plasticity. We examined these relationships for morphological and life-history traits of Daphnia magna in four independent experiments using two different Moscow populations and one German population. Clones were raised under a range of food levels or individual densities. Heterozygosity was scored at five allozyme loci in two experiments and at three loci in two others. Relative differences in developmental noise among clones with different heterozygosity levels were estimated as the pooled residual variation from an analysis of variation that removed the effects of macroenvironment, clones, and their interaction. Plasticity was measured as the amount of macroenvironmental variation plus genotype-by-environment interaction variation. We found a positive correlation between developmental noise and heterozygosity, although this correlation varied among traits and experiments. This result contradicts most previous claims about these relationships. In contrast, we found that phenotypic plasticity and heterozygosity were negatively correlated for some traits. Developmental noise and phenotypic plasticity were correlated for only two traits in two different experiments. This trait-specific relationship is in concordance with previous studies. Our results could not be explained by effects of developmental time, a previously hypothesized mechanism. We propose several explanations for our results and the disparate results of others that do not require that heterozygosity be the actual cause of variation in developmental noise.     

Biochemical polymorphisms and phenetic relationships in rodents of the genusCtenomys from Southern Brazil

Protein variation at 20 loci was analyzed by starch gel electrophoresis in population samples of Ctenomys flamarioni, C. torquatus, C. sp., and C. minutus collected in 25 localities of Southern Brazil. Results show that these four species exhibit higher levels of genetic variability (He = 0.11-0.17) than those reported for most other fossorial rodents. Estimates of similarity coefficients indicate that C. minutus and C. sp. (S = 0.91) are the closest species, while C. flamarioni (S = 0.77) is the most distant from the others. The data presented here support the hypothesis of a relatively recent disjunction of C. minutus from C. sp. Within-species similarity indices are of the same order of magnitude as those found between species. The relatively high levels of heterozygosity observed are not in accordance with the view that subterranean taxa should be less variable than aboveground species due to the uniformity of their environment.     

Population genetic structure of two rare tree species (Colubrina oppositifolia and Alphitonia ponderosa,Rhamnaceae) from Hawaiian dry and mesic forests using random amplified polymorphic DNA markers

Hawaiian dry and mesic forests contain an increasingly rare assemblage of species due to habitat destruction, invasive alien weeds and exotic pests. Two rare Rhamnaceae species in these ecosystems, Colubrina oppositifolia and Alphitonia ponderosa, were examined using random amplified polymorphic DNA (RAPD) markers to determine the genetic structure of the populations and the amount of variation relative to other native Hawaiian species. Relative variation is lower than with other Hawaiian species, although this is probably not a consequence of genetic bottleneck. Larger populations of both species contain the highest levels of genetic diversity and smaller populations generally the least as determined by number of polymorphic loci, estimated heterozygosity, and Shannon's index of genetic diversity. Populations on separate islands were readily discernible for both species as were two populations of C. oppositifolia on Hawai'i island (North and South Kona populations). Substructure among Kaua'i subpopulations of A. ponderosa that were ecologically separated was also evident. Although population diversity is thought to have remained at predisturbance levels, population size continues to decline as recruitment is either absent or does not keep pace with senescence of mature plants. Recovery efforts must focus on control of alien species if these and other endemic dry and mesic forest species are to persist.     

Aspects of the ecology of the prickly sculpin,Cottus asper Richardson,a persistent native species in Clear Lake,Lake County,California

Synopsis The biology of the prickly sculpin was investigated in Clear Lake, Lake County, California in order to determine how it has persisted in the face of introductions of numerous exotic species when most other native species have declined in abundance or have become extinct. Sculpins over 15 mm SL inhabited all types of benthic habitats in the lake, while post larval sculpins were pelagic when the postlarvae of exotic species were absent. The feeding ecology of sculpins was distinct from the other fishes in the lake in that they fed largely on amphipods and chironomid midge larvae regardless of the time of year, time of day, or habitat. Sculpins were uncommon in the stomachs of piscivorous fishes, except juvenile largemouth bass (Micropterus salmoides). It is concluded that prickly sculpins have persisted in Clear Lake in part because they are ecologically distinct from the exotic species and are not preyed upon by them to any great extent, and in part because they have managed to survive other man-related perturbations of this ecosystem.     

ALLOZYME VARIATION IN OLD WORLD RACES OF SORGHUM BICOLOR (POACEAE)

A survey of allozyme variation in cultivated races of sorghum (Sorghum bicolor) was undertaken. Eight plants each of 83 accessions representing the five primary races and five of the intermediate races of sorghum were analyzed for 15 enzyme systems encoded by 27 loci. Low levels of variation were found within accessions, which is typical of self-pollinating species. Little variation was also found among accessions. Compared with other cereals, S. bicolor is depauperate in allozyme variation. We found an average of 1.81 alleles per locus with a mean expected heterozygosity of 0.008 for the accessions and total panmictic heterozygosity of 0.093. Only 9% of the variation present was found within accessions, while 91% was among accessions. Most of the variation present is attributable to differences in geographic origin of the accessions rather than racial differences. Western and eastern Africa have the highest levels of total heterozygosity (0.108 and 0.088, respectively), while southern Africa has the lowest (0.008). Principal component analysis revealed continuous variation among races and geographic regions with the accessions failing to segregate into discrete racial or geographic clusters.     

Effects of the natural tidal cycle and artificial temperature cycling on Hsp levels in the tidepool sculpin Oligocottus maculosus

The rocky intertidal zone is characterized by a predictable cycle of environmental change cued by the ebb and flow of the tides. Tidepools are thus an excellent environment in which to determine whether predictability of environmental change can entrain an endogenous rhythmicity in heat shock protein (Hsp) levels. In this study, we monitored changes in Hsp mRNA and protein levels that occurred over the tidal cycle in tidepool sculpins and investigated whether there was an endogenous tidal rhythm in Hsp expression that persisted once the sculpins were transferred to a stable environment. Fluctuations in the tidepool environment increased hsc70, hsp70, and hsp90 mRNA levels, which translated into increased Hsc/Hsp70 and Hsp90 protein levels; however, this was not due to an endogenous tidal rhythm in Hsp levels because sculpins held under constant conditions did not show any rhythmicity in the expression of these genes. By exposing sculpins to an artificial temperature cycling regime that mimicked the temperature changes of a mid-intertidal pool, we were able to account for the direct role of temperature in regulating Hsp expression. However, there are additional extrinsic factors that likely integrate with temperature and result in differences between the hsp induction profiles that were observed in sculpins inhabiting their natural environment and those in cycling conditions in the laboratory.     

Growth and age of staghorn sculpins of the genus <Emphasis Type="Italic">Gymnocanthus</Emphasis> (Cottidae) in Peter the Great Bay and adjacent waters of Primorye (Sea of Japan)

Age and growth have been studied in four species of staghorn sculpins of the genus Gymnocanthus inhabiting Primorye (the Russian sector of the Sea of Japan). Large black edged and purplegray sculpins, G. herzensteini and G. detrisus, inhabited deeper waters compared with threaded sculpin G. pistilliger and (especially) whip sculpin G. intermedius. The size differences between different sculpin species could already be revealed in the diameter of mature oocytes and the length of hatching larvae and fries passing to the benthic phase of development. The maximum size in these species correlated with their maximum age. G. herzensteini, G. detrisus, G. pistilliger, and G. intermedius could attain the age of no less than 16+, 15+, 9+, and 7+, respectively. The gains in fish of the same age were also greater in species attaining greater size values. In all the species, females lived longer than males and were larger than males of the same age. Throughout the life cycle, the growth rate gradually came down; the decrease was especially pronounced during the period of sexual maturation. In the southern part of the geographic range, the specimens of purplegray sculpin attained smaller size and showed lower growth rate compared with the fish from the northern part of the range. The threaded sculpin demonstrated the inverse relationships. In general, females were dominating in samples of staghorn sculpins; however in small and medium size groups, the proportion of males was greater, which was due to their smaller, compared with females, absolute gains. In elder groups, the proportion of males gradually went down, as they were dying earlier than females.     

Genetic variability and large scale differentiation in two species of littorinid gastropods with planktotrophic development, Littorina littorea (L.) and Melarhaphe (Littorina) neritoides (L.) (Prosobranchia: Littorinacea), with notes on a mass occurrence of M. neritoides in Sweden

A planktotrophic larval development suggests a relatively high gene flow over long distances. Both Littorina littorea and Melarhaphe neritoides have an egg capsule and planktotrophic larvae which taken together are pelagic over a period of 4–8 weeks. In the absence of differential selection one would expect low levels of genetic differentiation over large distances in these two species. In this study of allozyme variation, low levels of differentiation were found over thousands of kilometres (Norway to Spain for L. littorea and Sweden to Greece for M. neritoides). This supports the hypothesis of two species with a high dispersal potential. A second expectation from neutral theory is that effective population size is positively correlated with average levels of genetic variation within species. In light of the generally high densities of local populations of these species and the high interpopulation migration rate, both L. littorea and M. neritoides may be considered as having high N_e s. Contrary to neutral expectation, L. littorea revealed very low levels of heterozygosity over its whole European range (mean H_exp= 0.015, 15 loci), while average heterozygosity of M. neritoides (H_exp= 0.084, 11 loci) was no more than in other littorinid species. This paper also reports the occurrence of M. neritoides on the Swedish west coast in 1988, 89 and 90, and two factors which may have promoted this unusually large invasion are discussed.     

Comparative nuclear and mitochondrial genome diversity in humans and chimpanzees

Restriction mapping and sequencing have shown that humans have substantially lower levels of mitochondrial genome diversity (d) than chimpanzees. In contrast, humans have substantially higher levels of heterozygosity (H) at protein-coding loci, suggesting a higher level of diversity in the nuclear genome. To investigate the discrepancy further, we sequenced a segment of the mitochondrial genome control region (CR) from 49 chimpanzees. The majority of these were from the Pan troglodytes versus subspecies, which was underrepresented in previous studies. We also estimated the average heterozygosity at 60 short tandem repeat (STR) loci in both species. For a total sample of 115 chimpanzees, d = 0.075 +/0 0.037, compared to 0.020 +/- 0.011 for a sample of 1,554 humans. The heterozygosity of human STR loci is significantly higher than that of chimpanzees. Thus, the higher level of nuclear genome diversity relative to mitochondrial genome diversity in humans is not restricted to protein-coding loci. It seems that humans, not chimpanzees, have an unusual d/H ratio, since the ratio in chimpanzees is similar to that in other catarrhines. This discrepancy in the relative levels of nuclear and mitochondrial genome diversity in the two species cannot be explained by differences in mutation rate. However, it may result from a combination of factors such as a difference in the extent of sex ratio disparity, the greater effect of population subdivision on mitochondrial than on nuclear genome diversity, a difference in the relative levels of male and female migration among subpopulations, diversifying selection acting to increase variation in the nuclear genome, and/or directional selection acting to reduce variation in the mitochondrial genome.      

Analysis of the relationship between allozyme heterozygosity and fitness in the rare Gentiana pneumonanthe L.

Especially for rare species occurring in small populations, which are prone to loss of genetic variation and inbreeding, detailed knowledge of the relationship between heterozygosity and fitness is generally lacking. After reporting on allozyme variation and fitness in relation to population size in the rare plant Gentiana pneumonanthe, we present a more detailed analysis of the association between heterozygosity and individual fitness. The aim of this study was to test whether increased fitness of more heterozygous individuals is explained best by the ‘inbreeding’ hypothesis or by the ‘overdominance’ hypothesis. Individual fitness was measured during 8 months of growth in the greenhouse as the performance for six life-history parameters. PCA reduced these parameters to four main Fitness Components. Individual heterozygosity was scored for seven polymorphic allozyme loci. For some of these loci (e.g. Aat3, Pgm1 and 6Pgdh2) heterozygotes showed a significantly higher relative fitness than homozygotes. To test the inbreeding model, regression analyses were performed between each Fitness Component and the number of heterozygous loci per individual. Multiple regressions with the adaptive distance of five loci as independent variables were used to test the overdominance model. Only the inbreeding model was a statistically significant explanation for the relationship between heterozygosity and fitness in G. pneumonanthe. The number of heterozygous loci was significantly negatively correlated with the coefficients of variation of three of the six initially measured fitness parameters. This suggests a lower developmental stability among more homozygous plants and may explain the higher phenotypic variation in small populations of the species observed earlier. The importance of the results for conservation biology is discussed.     

The osmorespiratory compromise in sculpins: impaired gas exchange is associated with freshwater tolerance

We acclimated two species of sculpin, the freshwater prickly sculpin (Cottus asper) and the closely related marine Pacific staghorn sculpin (Leptocottus armatus) to freshwater ( approximately 0 g/L), brackish water (15 g/L), and seawater (30 g/L) for at least 4 wk and examined the relationships between respiration, ion regulation, gill morphology, and freshwater tolerance. The prickly sculpin successfully acclimated to all three salinities and did not experience appreciable changes in plasma osmolality, [Cl-], or mortality. Gill Na+/K+-ATPase activity was lowest in prickly sculpins acclimated to freshwater, their native salinity, and increased during acclimation to seawater. Furthermore, prickly sculpins acclimated to freshwater had a 30% higher P(crit) than fish acclimated to brackish water or seawater; P(crit) is the environmental P(O2) below which an animal can no longer maintain a routine (.-)M(O2), and an increase in P(crit) represents a compromise of respiratory gas exchange. The higher P(crit) observed in prickly sculpins acclimated to freshwater is likely a consequence of their having small, relatively thick gills that increase in thickness (by approximately 1 microm) during freshwater exposure. In contrast, the marine Pacific staghorn sculpin successfully acclimated to brackish water and seawater, but high mortality (25%) was observed after 3 wk of exposure to freshwater. Pacific staghorn sculpins exposed to freshwater suffered significant, 15%-20%, reductions in plasma osmolality and [Cl-], and these losses in plasma ions resulted in a 1.4-fold increase in gill Na+/K+-ATPase activity. Pacific staghorn sculpins have large, thin gills that are not modified in response to salinity acclimation, and as a result, these animals show no respiratory compromise during freshwater acclimation, as evidenced by the lack of change in P(crit), but show significant ion regulatory disturbance. Overall, this study suggests that gill thickening and the resulting respiratory compromise are necessary for freshwater tolerance in sculpins.     

A benthic predatory fish does not cause selection on armour traits in three‐spined stickleback Gasterosteus aculeatus (Gasterosteiformes: Gasterosteidae)

Predation can promote divergence between prey populations and contribute to ecological speciation. In theory, predators can also constrain prey population divergence. In coastal British Columbia, Canada, Gasterosteus aculeatus (three‐spined stickleback) species pairs only occur in lakes with a single species of predatory fish: Oncorhynchus clarkii (the cutthroat trout). Similar lakes containing additional predatory fish species (Cottus asper, prickly sculpins; Oncorhynchus mykiss, rainbow trout) contain only single species of morphologically intermediate stickleback, suggesting that these predators prevent the coexistence of stickleback species pairs. We conducted a mesocosm experiment to investigate how prickly sculpins might constrain divergence, by quantifying their impact on survival and natural selection on antipredator (armour) traits in F₂ stickleback from a cross between ecologically divergent populations. We tested three hypotheses: (1) sculpin predation on sticklebacks reduces survival in a way that could result in their exclusion from certain niches; (2) sculpins compete with stickleback; (3) sculpins respond to prey vulnerabilities in similar ways to cutthroat trout, tending to constrain rather than to enhance divergence. We found that sculpins significantly reduce stickleback survival, that their presence per se does not reduce growth in stickleback, and that predation did not result in selection on any of the armour traits measured, or on gill raker length, which is an important trophic trait. These results tend to refute hypotheses (2) and (3), while supporting hypothesis (1). © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 877–885.