Characterization of Charr Chromosomes Using Fluorescence in Situ Hybridization

The chromosomal locations of several families of tandem repetitive DNA sequences and the 5S rDNA were determined using fluorescence in situ hybridization (FISH) in the five North American charr species: Salvelinus namaycush, S. fontinalis, S. alpinus, S. malma, and S. confluentus. The pattern of hybridization of three centromeric repetitive sequences previously isolated from S. namaycush and S. alpinus was unique in each species. Dual-color FISH experiments showed that in several species many of the centromeres had the EcoRI-DraI family in addition to either the AluI-RsaI type A or type B families. The EcoRI-DraI family which was found only at the centromeres of acrocentric chromosomes in S. namaycush, S. fontinalis and S. malma was also found at centromeres of selected metacentrics in S. alpinus (one pair) and S. confluentus (four pairs) whose chromosomes have undergone additional centric fusions compared to the other species. The locations of 5S rDNA sequences were different in each species except for the two most closely related (S. alpinus and S. malma). Two whole-arm chromosome paint probes, one specific for the short and the other for the long arm of the lake charr sex chromosomes, hybridize to the same chromosome pair in all species. Results with other paint probes suggest that independent centric fusions have occurred in S. alpinus and S. confluentus which is consistent with the phylogenetic tree obtained previously for Salvelinus with cytogenetic and DNA data.     

The utilization of a Pacific salmon Oncorhynchus nerka subsidy by three populations of charr Salvelinus spp.

The L_F‐at‐age trajectories differentiated two populations of Dolly Varden charr Salvelinus malma and a population of Arctic charr Salvelinus alpinus from the eastern end of Iliamna Lake, Alaska. Salvelinus malma from the Pedro Bay ponds were the smallest for a given age, followed by Salvelinus alpinus from the lake, and S. malma from the Iliamna River were much larger. The utilization of a large sockeye salmon Oncorhynchus nerka subsidy by the three Salvelinus spp. populations was then investigated by comparing diet data and mixing model (MixSIR) outputs based on carbon and nitrogen stable isotopes. Stomach contents indicated that both S. malma populations fed on O. nerka products, especially eggs and larval Diptera that had scavenged O. nerka carcasses, whereas S. alpinus fed on a variety of prey items such as three‐spined sticklebacks Gasterosteus aculeatus and snails. Stable‐isotope analysis corroborated the diet data; the two S. malma populations incorporated more O. nerka‐derived nutrients into their tissues than did S. alpinus from the lake, although all populations showed substantial utilization of O. nerka‐derived resources. Salvelinus alpinus also seemed to be much more omnivorous, as shown by stable‐isotope mixing models, than the S. malma populations. The dramatic differences in growth rate between the two S. malma populations, despite similar trophic patterns, indicate that other important genetic or environmental factors affect their life history, including proximate temperature controls and ultimate predation pressures.     

Genetic Differentiation of Three Sympatric Charr Species from the Genus SalvelinusInferred from PCR–RFLP Analysis of Mitochondrial DNA

We examined differentiation of three sympatric charr species Dolly Varden charr Salvelinus malmaWalbaum, white-spotted charrS.leucomaenisPallas, and Levanidov charrS. levanidovi Chereshnev, Scopetz, Gudkov. Using restriction fragment length polymorphism analysis (RFLP), three mitochondrial DNA fragments (ND1/ND2, ND5/ND6, and Cytb/D-loop) amplified in PCR were compared. The divergence of mtDNA nucleotide sequences betweenS. levanidovi and S. leucomaenis was about 9.7%, between S. levanidovi and S. malma, 8.7%, and between S. malma andS. laecomaenis, 7.5%. These results indicate genetic isolation of the species examined and support the earlier suggestion on closeness of Levanidov charr to the common ancestor of the genus Salvelinus.     

Comparative catches and food habits of dolly varden and arctic charrs,Salvelinus malma and S. alpinus,at Karluk,Alaska, in 1939–1941

Synopsis For many years the seriousness of the predations of dolly varden (Salvelinus malma) upon the eggs, alevins and juveniles of Pacific salmons (Oncorhynchus) has been a controversial subject. Over $ 300,000 was spent in Western Alaska from 1920 to 1941 for bounties on dolly varden in the belief that they were serious salmon predators. The writter undertook this study because of an intense desire to evaluate the need and justification for such a bounty system. The incidental examination of stomach contents of charrs which were captured for tagging or for tag recovery purposes from Karluk Lake and Karluk River on Kodiak Island, Alaska, and from Shelikof Strait and Uyak Bay during the summer months of 1939, 1940 and 1941 presented an excellent opportunity to pursue such a project. Stomachs were collected and contents analyzed from 1,992 arctic charr (Salvelinus alpinus) and 2,565 dolly varden charr (Salvelinus malma), taken by beach seine, gill net, fyke net, hook and line and by weir trap from Karluk Lake; from 956 dolly varden taken by seine and weir trap from the Karluk River; and from 462 dolly varden taken by beach seine and commercial salmon traps in Uyak Bay. Among the more than 5,000 charr stomachs examined, only 42 were found to contain salmon smolts, parr or alevin. Examination of 500 stomachs taken from dolly varden charr captured in the lower Karluk River at the height of the red salmon (Oncorhynchus nerka) smolt migration in May of 1939 revealed very little evidence of predation on the smolts in fresh water. There was some evidence that this relationship may have changed after the migrating charr had become adjusted to salt water, although the examination of 460 stomachs of dolly varden captured in salt water revealed nothing particularly incriminating in its role as a salmon predator. The fact that the downstream migration of the dolly varden reached its peak before that of the red salmon smolts was considered significant. Examination of the 4,500 charr stomachs at Karluk Lake in the months from April through September led to the discovery that two distinct species of charrs were present, one, the dolly varden, primarily stream-inhabiting and anadromous, and the other, the arctic charr, primarily lake-inhabiting and nonanadromous. Although red salmon eggs constituted 32% of the total volume of food ingested by both species of charr at Karluk Lake, there was considerable evidence that they were consumed in salvage or scavenger feeding, and there was practically no evidence that they were consumed as a result of predatory feeding. Only five, or one-tenth of one percent, of the thousands of charr stomachs examined at Karluk Lake contained red salmon parr or alevin.A summary of the contrasts in the feeding and migratory habits and physical characteristics of Salvelinus malma and Salvelinus alpinus led to the conclusion that alpinus was a more serious potential salmon predator than malma at Karluk Lake. In view of the fact that alpinus was seldom reached by the predator-control program whereas malma was being systematically destroyed only because it was more easily captured due to its migratory tendencies, it was recommended that the predator control program be terminated in Karluk waters.Published post mortem in shortened and slightly edited version. Reprint requests to the Editor.     

Speciation and genetic divergence of three species of charr from ancient Lake El'gygytgyn (Chukotka) and their phylogenetic relationships with other representatives of the genus Salvelinus

The diversity of phenotypically different and often reproductively isolated lacustrine forms of charrs of the genus Salvelinus represents a substantial problem for taxonomists and evolutionary biologists. Based on the analysis of variability of ten microsatellite loci and two fragments of mitochondrial DNA (control region and cyt‐b gene), the evolutionary history of three charr species from Lake El'gygytgyn was reconstructed, and phylogenetic relationships between the main representatives of the genus were revealed. Three species from Lake El'gygytgyn were strongly reproductively isolated. Long‐finned charr described previously as Salvethymus svetovidovi, an ancient endemic form in the lake, originated 3.5 Mya (95% Bayesian credible intervals: 1.7, 6.1). Placement of this species in the phylogenetic tree of Salvelinus was not determined strictly, but it should be located in the basal part of the clade Salvelinus alpinus – S. malma species complex. The origin of small‐mouth charr S. elgyticus and Boganida charr S. boganidae in Lake El'gygytgyn was related to allopatric speciation. Their ancestors were represented by two glacial lineages of Taranets charr S. alpinus taranetzi from Asia. In Lake El'gygytgyn, these lineages entered into secondary contact postglacially. A revision of the main phylogenetic groups within the Salvelinus alpinus – S. malma complex is conducted. The Boganida charrs from Lakes El'gygytgyn and Lama (Taimyr) belong to different phylogenetic groups of Arctic charr and should not be regarded as a single species S. boganidae. Using the charrs from Lakes El'gygytgyn and Lama as a case study, we show that a model of sympatric speciation, which seemed more probable based on previous empirical evidence, was rejected by other data.     

The most variable vertebrate on Earth

In the search for the most variable non-human vertebrate on Earth, intraspecific variation of ten variable traits was compared among ten highly variable species. Mammals, birds and many reptiles, amphibians and fishes were excluded because most of the variation is among, and not within species. The focus was on northern fishes, where high intraspecific variation is well documented. The ten selected species were European whitefish Coregonus lavaretus, chinook salmon Oncorhyncus tshawytscha, sockeye salmon O. nerka, rainbow trout O. mykiss, atlantic salmon Salmo salar, brown trout S. trutta, arctic charr Salvelinus alpinus, brook charr S. fontinalis, dolly varden charr S. malma and threespine stickleback Gasterosteus aculeatus. Variation included not only size and phenotype, but also ecology, behaviour and life history. The traits were geographic range, migration, habitat, adult size, colour, body form, polymorphism, diet, reproduction and genetics. Arctic charr came on top in the final ranking, followed by dolly varden charr and rainbow trout. The two least variable were chinook salmon and threespine stickleback. It is proposed that arctic charr, which is also the northernmost fish on Earth, has evolved its unique variability in range, size, phenotype, ecology and life history by adapting to the extreme and highly unpredictable ecological conditions of arctic and other northern lakes for many glacial periods.     

Fish growth hormone genes: Divergence of intron sequence in charrs of Salvelinus genus

Sequences of the two large introns (C and D) from two paralogous growth hormone genes, GH1 and GH2, were compared in eight charr species of the Salvelinus genus (Osteichthes, Salmonidae). It was demonstrated that the rates of intron divergence in these two genes were remarkably different. Introns in the GH1 gene appeared to be more conservative, while the rate of intron variations was considerably higher in the GH2 gene. These data suggest that noncoding regions of nuclear genes under the selective pressure. The lower congruence of phylogenetic scheme constructed based on an analysis of the GH1 introns compared to that based on the GH2 data, as well as with the traditional views on the evolution of charr species, also favors the supposition on differences in the forces and directions of the selective pressure on the introns.     

Redd superimposition by introduced rainbow trout,oncorhynchus mykiss, on native charrs in a japanese stream

Spawning redd superimposition of introduced, spring-spawning rainbow trout,Oncorhynchus mykiss, on native, fall-spawning Dolly varden,Salvelinus malma, and white-spotted charr,S. leucomaenis, were examined in a small stream in Hokkaido, Japan. The stream reaches in which Dolly Varden and white-sported charr redds were observed in fall 1997 greatly overlapped with the reaches in which rainbow trout redds were recorded in spring 1998. Spawning microhabitats were also similar between trout and the two charr species. Thirteen and 3% of Dolly Varden and white-spotted charr redds, respectively, were superimposed by rainbow trout redds. The eggs or alevins in the disturbed charr redds were potentially damaged because charrs were not likely to have emerged from the redds before the superimposition occurred. In sufficiently great abundance, introduced rainbow trout may negatively impact native charr populations by dislodging the latter’s spawning redds.     

Life-history styles and somatic allocation in iteroporous arctic charr and semelparous pink salmon

Synopsis RNA/DNA ratios were used to estimate recent somatic allocation of spawning arctic charr,Salvelinus alpinus, and pink salmon,Oncorhynchus gorbuscha. Smaller arctic charr morphs had lower ratios than larger morphs. Larger male pink salmon had lower ratios than smaller males. Thus recent somatic allocation (anabolic metabolic activity and growth) is coupled with cumulative lifetime growth in iteroparous arctic charr, but is uncoupled in semelparous pink salmon. These data suggest that somatic investment in these species is being differentially effected by the energetic costs of reproduction.     

Distribution of two congeneric charrs in streams of Hokkaido Island,Japan: considering multiple factors across scales

Salvelinus leucomaenis (white-spotted charr) and S. malma (Dolly Varden) are distributed throughout Hokkaido Island, Japan, but sites where they occur in sympatry are rare. In general, S. malma inhabit upstream reaches and S. leucomaenis extend downstream to the ocean. Factors influencing their distribution were analyzed at four spatial scales ranging from the whole island to individual stream pools. At the island scale, S. leucomaenis were found in the warmer south-west region and at lower altitudes elsewhere, whereas S. malma were found in the colder north-east and at higher altitudes. At a regional scale, the downstream limit of S. malma and upstream limit of S. leucomaenis shifted to lower altitude from south-west to north-east across the island, coincident with the decrease in temperature. Further analysis showed that transition points from S. leucomaenis or sympatry to S. malma in individual watersheds were closely related to an index of cumulative mean monthly temperatures exceeding 5°C. However, at the scale of a single watershed, the transition occurred at different altitudes, gradients, and temperatures in two tributaries, apparently because stream discharge, habitat, and disturbances from floods interacted with these abiotic factors to limit distribution. The two charr species developed interspecific dominance hierarchies in individual pools, and there was strong complementary density compensation among stream pools that could be explained by interspecific competition but not by differences in habitat. However, patterns at watershed and regional scales suggested that interspecific competition interacts with temperature in complex ways. We conclude that the importance of various abiotic and biotic factors in shaping Hokkaido charr distributions depends on the scale at which they are viewed.     

Genetic divergence of sympatric charrs of the genus <Emphasis Type="Italic">Salvelinus</Emphasis> from Nachikinskoe Lake (Kamchatka)

We studied genetic differentiation of two charr species, Dolly Varden Salvelinus malma malma Walbaum and resident lacustrine charr Salvelinus sp., which sympatrically inhabit Nachikinskoe Lake (the Bol’shaya River basin) in southwestern Kamchatka Peninsula. Using restriction analysis (RFLP), three mitochondrial DNA fragments (ND1/ND2, ND5/ND6, and Cyt b/D-loop) amplified in polymerase chain reaction (PCR) were compared. The divergence of the mtDNA sequences between Salvelinus sp. and S. malma malma was 2.8%; Salvelinus sp. and S. taranetzi, 0.36%; Salvelinus sp. and S. krogiusae, 0.21%; Salvelinus sp. and S. alpinus, 3.0%. These results point to reproductive isolation of charrs in Nachikinskoe Lake and support the earlier suggestion on a close relationship between Salvelinus sp., S. taranetzi, and S. krogiusae.     

Spatial and temporal variation in invertebrate consumer diets in forested and herbaceous wetlands

Lake Kronotskoye (the Kronotsky Biosphere State Reserve, south-eastern Kamchatka Peninsula) contains three closely related resident morphotypes charrs, which are considered to be either independent species (white charr Salvelinus albus, longhead charr Salvelinus kronocius, Schmidt’s charr Salvelinus schmidti) or a united lacustrine-riverine charrs, represented by several phenotypes. Salvelinus malma malma is isolated from the lake charr populations by an upstream migration barrier in the Kronotskaya River, which flows out of the lake. We examined the level of genetic variability and divergence of mtDNA both between charrs from Kronotsky Lake and between resident lacustrine charrs and the hypothetical ancestor species S. m. malma. The RFLP-PCR analysis was used to examine six regions (ND1/ND2, ND3/ND4L/ND4, ND5/ND6, COI/COII/A8, A8/A6/COIII/ND3, and Cytb/D-loop), comprising ~80% of the mtDNA. Significantly different levels of diversity were found among the populations of lacustrine charrs. S. albus and S. schmidti had the highest indices of mtDNA diversity among the investigated populations from the different habitats. Heterogeneity tests revealed highly significant differentiation among lake populations and among riverine (Kronotskaya River) and lake (Lake Kronotskoye) populations of charrs, indicating their reproductive isolation. Hierarchical analysis of molecular variance revealed the following regularities of diversity distribution: the high proportion of interpopulation variation (93.25%) and low but statistically significant subdivision between charr populations (6.75%, P < 0.001). Results of the present study suggest that the populations of S. albus, S. kronocius, S. schmidti belong to the S. m. malma phylogenetic group. The divergence value of mtDNA of resident charrs (0.0357–0.0010%) does not exceed the intraspecific variability of S. m. malma. The analysis of the mtDNA haplotypes genealogy of charrs showed that the low values of nucleotide divergence reflect a short period from the beginning of divergence from the ancestral lineages and are due to ancestral polymorphism, as well as to haplotype exchange between the diverged phylogenetic groups as a result of introgressive hybridization.     

Isolation and cross‐familial amplification of 41 microsatellites for the brook charr (Salvelinus fontinalis)

The brook charr (Salvelinus fontinalis; Osteichthyes: Salmonidae) is a phenotypically diverse fish species inhabiting much of North America. But relatively few genetic diagnostic resources are available for this fish species. We isolated 41 microsatellites from S. fontinalis polymorphic in one or more species of salmonid fish. Thirty‐seven were polymorphic in brook charr, 15 in the congener Arctic charr (Salvelinus alpinus) and 14 in the lake charr (Salvelinus namaycush). Polymorphism was also relatively high in Oncorhynchus, where 21 loci were polymorphic in rainbow trout (Oncorhynchus mykiss) and 16 in cutthroat trout (Oncorhynchus clarkii) but only seven and four microsatellite loci were polymorphic in the more distantly related lake whitefish (Coregonus clupeaformis) and Atlantic salmon (Salmo salar), respectively. One duplicated locus (Sfo228Lav) was polymorphic at both duplicates in S. fontinalis.     

The sequence of the isoepoxydon dehydrogenase gene of the patulin biosynthetic pathway in <Emphasis Type="Italic">Penicillium</Emphasis> species

Interest in species of the genus Penicillium is related to their ability to produce the mycotoxin patulin and to cause spoilage of fruit products worldwide. The sequence of the isoepoxydon dehydrogenase (idh) gene, a gene in the patulin biosynthetic pathway, was determined for 28 strains representing 12 different Penicillium species known to produce the mycotoxin patulin. Isolates of Penicillium carneum, Penicillium clavigerum, Penicillium concentricum, Penicillium coprobium, Penicillium dipodomyicola, Penicillium expansum, Penicillium gladioli, Penicillium glandicola, Penicillium griseofulvum, Penicillium paneum, Penicillium sclerotigenum and Penicillium vulpinum were compared. Primer pairs for DNA amplification and sequencing were designed from the P. griseofulvum idh gene (GenBank AF006680). The two introns present were removed from the nucleotide sequences, which were translated to produce the IDH sequences of the 12 species for comparison. Phylogenetic relationships among the species were determined from rDNA (ITS1, 5.8 S, ITS2 and partial sequence of 28S rDNA) and from the idh nucleotide sequences minus the two introns. Maximum parsimony analysis showed trees based on rDNA and idh sequences to be congruent. It is anticipated that the genetic information obtained in the present study will aid in the design of probes, specific for patulin biosynthetic pathway genes, to identify the presence of these mycotoxigenic fungi. The U.S. Government's right to retain a non-exclusive, royalty-free license in and to any copyright is acknowledged.     

The extinction of Arctodiaptomus alpinus (Copepoda) following the introduction of charr into a small alpine lake Dvojno Jezero (NW Slovenia)

In 1991 about twenty-five age 2+ specimens of the Arctic charr (Salvelinus alpinus (L.) (length between 100 and 130 mm) were introduced into the fishless small mountain lake Dvojno Jezero (= Twin Lake) (area 0.5+0.4 ha; max. depth 8 m, altitude 1670 m) in NW Slovenia. The first spawning of the Arctic charr was observed in autumn 1994 and regularly each year thereafter. Fish of different sizes were observed from 1995 onward each year but a fish census was never carried out. Autumn zooplankton samples, collected before the introduction of Arctic charr, contained adults (including ovigerous females) of two copepod species: Cyclops abyssorum tatricus and Arctodiaptomus alpinus. No planktonic Cladocera were found. Seven years after the introduction of fish standing crop of zooplankton declined 100-fold. The zooplankton contained only a few copepodites (IV and V) of C. a. tatricus. Simultaneously, water transparency declined and the concentration of chlorophyll a increased and filamentous green algae and picoplankton became abundant in the littoral zone of both lakes.     

Diel Habitat Partitioning by Bull Charr and Cutthroat Trout During Fall and Winter in Rocky Mountain Streams

We used underwater observation to determine diel habitat partitioning between bull charr, Salvelinus confluentus, and cutthroat trout, Oncorhynchus clarki, during fall and winter (0.1–8.3°C) in two Rocky Mountain streams that differed in habitat availability. The majority (>70%) of both species emerged from concealment cover at night, though bull charr exhibited a greater tendency for nocturnal behavior than cutthroat trout. Differences in day and night counts were most pronounced at temperatures <3°C, when very few fish of either species were observed in the water column during the day, but both species were common at night. Both species used concealment cover of large woody debris and boulder substrate crevices in deep pools during the day. At night, fish emerged from cover and habitat use shifted to shallow water with low cover. Microhabitat partitioning among species and size classes occurred at night, cutthroat trout moving into shallower, faster water that was farther from cover compared to bull charr. Smaller fish of both species occupied focal positions in slower, shallower water closer to the substrate than larger fish. Large, mixed-species aggregations also were common in beaver ponds both day and night. High variation in diel and site-specific winter habitat use suggests the need for caution in developing habitat suitability criteria for salmonids based solely on daytime observations or on observations from a few sites. Our results support the need to incorporate nocturnal habitat use and partitioning in studies of salmonid ecology.     

Takvatn Through 20 Years: Long-term Effects of an Experimental Mass Removal of Arctic Charr, Salvelinus Alpinus, From a Subarctic Lake

Between 1984 and 1989, the experimental removal of 31 tons (666000 fish) of stunted Arctic charr, Salvelinus alpinus, from Takvatn in northern Norway, had strong effects on the populations of Arctic charr, brown trout, Salmo trutta, and three-spined sticklebacks, Gasterosteus aculeatus. The littoral catch per unit effort (CPUE) of charr had decreased by 90% in 1990 and then increased to about 50% of the initial level by 1994 while the pelagic CPUE had decreased to zero. Growth in both charr and trout greatly improved when the charr density had decreased, and large fish of both species appeared in the catches. These large fish became predators on small charr in the littoral zone. The incidence of trout increased from below 1% to 15% from 1988 to 1999 after a brief peak at 30% in 1992 and 1993. The charr population attained a bimodal size distribution and did not return to the stunted state during the 10 years following the intensive fishing period. The mass removal experiment showed that it is possible to change the structure of a charr population by intensive fishing. Predation on small charr from cannibals and large trout was probably essential for maintaining the new population structure. An increase in the growth of young charr from 1995 to 1997 was related to a high consumption of Daphnia and Eurycercus. Rapid changes in the growth of charr followed the density fluctuations in sticklebacks, which show large annual variations in this system; the rapid changes in charr growth were probably caused by variations in the competition intensity for cladoceran prey between young charr and sticklebacks. Twenty years of data has provided important information, but even more time is needed to follow the long-term trends in northern lakes such as Takvatn.     

Sorghum laxiflorum and S. macrospermum, the Australian native species most closely related to the cultivated S. bicolor based on ITS1 and ndhF sequence analysis of 25 Sorghum species

Australian species make up seventeen of the worlds twenty-five recognised species of Sorghum, with the genus separated into five sections: Eu-sorghum, Chaetosorghum, Heterosorghum, Para-sorghum and Stiposorghum. Whereas the genetic relationships within section Eu-sorghum are well known, little is known about the genetic relationships and crossabilities outside the primary genepool. We made a detailed investigation of phylogenetic relationships within Sorghum to identify wild species most closely related to cultivated sorghum (with outgroups Zea mays and Saccharum officinarum). The ribosomal ITS1 gene of ten species and the chloroplast ndhF gene from nineteen species were sequenced. Independent and combined analyses of the ITS1 and ndhF data sets were undertaken. The Eu-sorghum species were resolved into a strongly supported lineage by all three analyses, and included the Australian natives S. laxiflorum and S. macrospermum in the ITS1 and combined analyses. All remaining sorghum species were resolved into a second well-supported lineage in the combined analyses, although some internal relationships within this second lineage remain unresolved. We identified S. laxiflorum and S. macrospermum as the Australian species most closely related to cultivated sorghum. Our data support a reduction in the number of subgeneric sections from five to three: Eu-sorghum (unchanged); a combined Chaetosorghum/Heterosorghum to reflect the very close relationship between these two species; and a combined Para-sorghum/Stiposorghum section, thereby removing the unclear taxonomic and genetic boundaries between these species.     

A test of the hypothesis of pheromone attraction in salmonid migration

Synopsis We tested the hypothesis that anadromous salmonids are guided on their homeward migration by population-specific pheromones. Our findings do not support the hypothesis. Wild migrant Arctic charr,Salvelinus alpinus, from Ikarut River, Labrador were transferred and held in a tributary previously uninhabited by anadromous fish. None of the charr migrating up Ikarut River entered the tributary after fish were transferred. Similarly, migrant charr, which were caught in Ikarut River and released in the tributary below the captive fish, did not remain in the tributary. We re-evaluated the data which have been used to uphold the concept of pheromone attraction in salmonid migration and concluded that support for the hypothesis is unsubstantiated.     

Molecular Systematics and Evolution of the Growth Hormone Introns in the Salmoninae

DNA sequence data was collected for the C and D introns in the duplicate growth hormone loci (GH1 and GH2) from Brachmystax lenok, two subspecies of Hucho hucho, Hucho (Parahucho) perryi, Salmo salar, Salmo trutta, Acantholingua ohridana (Salmothymus), six species of Salvelinus, eight species of Oncorhynchus including O. masou, and three outgroups including Thymallus thymallus, Coregonus artedi, and Coregonus clupeaformis. Phylogenetic analyses were performed using maximum parsimony and maximum likelihood (PAUP, version 4.08beta) with gaps as missing data and as a fifth base. B. lenok was basal in all of the trees and all of the other genera were monophyletic with the exception that A. ohridana always placed within Salmo, and H. hucho sp. often placed with B. lenok. The GH1 introns supported a sister relationship between Oncorhynchus and Salvelinus, while the combined GH2 introns were ambiguous at this node. This result contrasts with trees based on morphology and the ribosomal ITS1 sequences that support a sister relationship between Salmo and Oncorhynchus. The only estrogen response element (ERE) in the gene is found in the C intron and has mutated in GH2 in all of the species except B. lenok. The ERE element in GH1 has undergone another mutation in all of the species except for B. lenok, and members of the two genera Salvelinus and Oncorhynchus. Thus these latter two genera are the only ones with a difference in expression of GH1 and GH2 in the presence of estrogen. Differences in selective pressure on the introns in the duplicate genes in different taxa could account for the conflicting results obtained in the phylogenetic analysis.