Temperature‐dependent competition between juvenile salmonids in small streams

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Karyological differences of the Northern Dolly Varden Salvelinus malma malma and the white char Salvelinus albus from the Kamchatka River basin

The karyotypes of northern Dolly Varden and white char, sympathrically inhabiting the Kamchatka River basin, were studied. The karyotype of Dolly Varden was stable: 2n = 78 and NF = 98 + 2, while in white char, polymorphism and mosaicism for the chromosome number were revealed: 2n = 76-79, NF = 98 + 2. Using a routine chromosome staining technique, the karyotype of white char (2n = 78) was shown to be identical to that of Dolly Varden. In both karyotypes, similar sets of marker chromosomes were present: two pairs of submetacentric (SM), one pair of submeta-subtelocentric (SM-ST), one pair of large acrocentric (A), and one pair of large sub-telocentric (ST) chromosomes. However, the karyotypes of Dolly Varden and white char differed in the number and location of nucleolus organizer regions (NORs). In Dolly Varden, single NORs located in the telomeric regions of the marker SM-ST chromosomes were observed. In white char, NORs were multiple and located both in the telomeric regions of the marker SM-ST chromosomes and on the short and long arms of large ST chromosomes. The identical marker chromosomes indicate considerable phylogenetic relatedness between Dolly Varden and white char from the Kamchatka River basin. Variation in NORs provides evidence for the reproductive isolation of these chars and their species status.     

Variability of microsatellite DNA and genetic differentiation of populations in the resident form of Dolly Varden Salvelinus malma krascheninnikovi from Sakhalin

Variability in 11 microsatellite loci is followed for nine populations of the resident southern form of Dolly Varden Salvelinus malma krascheninnikovi from Sakhalin. In all studied populations, the level of genetic differentiation is significant with global values of F _st and R _st 0.182 and 0.329, respectively. On the NJ tree, two clusters are separated significantly: the former is represented by the populations from southwestern Sakhalin (the Novoselka River basin), and the latter includes all populations from southern and eastern Sakhalin. Abrupt size variations (bottleneck events) are suggested for several studied populations, and 50% reductions of allelic richness and average heterozygosity are observed in two lacustrine populations of the Novoselka River basin in comparison to the riverine populations of this basin. The results suggest an occurrence of a comparatively recent gene flow between the populations of resident Dolly Varden from Sakhalin and absence of reproductive isolation between the anadromous and resident forms. The degradation of the majority of Dolly Varden populations can be expected in the near future, and conservation projects should be developed.     

Karyological Differences between Northern Dolly Varden Salvelinus malma malma and White Char Salvelinus albusfrom the Kamchatka River Basin

The karyotypes of northern Dolly Varden and white char, sympatrically inhabiting the Kamchatka River basin, were studied. The karyotype of Dolly Varden was stable: 2n= 78 andNF= 98 + 2, while in white char, polymorphism and mosaicism for the chromosome number were revealed: 2n= 76–79, NF= 98 + 2. Using a routine chromosome staining technique, the karyotype of white char (2n= 78) was shown to be identical to that of Dolly Varden. In both karyotypes, similar sets of marker chromosomes were present: two pairs of submetacentric (SM), one pair of submeta-subtelocentric (SM-ST), one pair of large acrocentric (A), and one pair of large subtelocentric (ST) chromosomes. However, the karyotypes of Dolly Varden and white char differed in the number and location of nucleolus organizer regions (NORs). In Dolly Varden, single NORs located in the telomeric regions of the marker SM-ST chromosomes were observed. In white char, NORs were multiple and located both in the telomeric regions of the marker SM-ST chromosomes and on the short and long arms of large ST chromosomes. The identical marker chromosomes indicate considerable phylogenetic relatedness between Dolly Varden and white char from the Kamchatka River basin. Variation in NORs provides evidence for the reproductive isolation of these chars and their species status.     

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

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

Female‐biased migration of stream‐dwelling Dolly Varden in the Shiisorapuchi River, Hokkaido, Japan

The first evidence for female‐biased migration in a partially migratory stream‐dwelling salmonid the Dolly Varden Salvelinus malma , a phenomenon well known in sea‐run and lake‐run populations, is presented. Dolly Varden in the Shiisorapuchi River in central Hokkaido, Japan, used both tributaries, of which there are many, and the main stem, but spawned only in tributaries. The size structures of Dolly Varden (≥age 1 + years) in tributaries were unimodal (<100 mm fork length, L _F) during non‐spawning seasons but changed to bimodal during spawning seasons (lower mode <110 mm, upper mode >120 mm L _F). Mature individuals were observed in both modal groups. From the trapping and census data, the small group appeared to be tributary resident and the large group main stem migrant. Males were common in both resident and migrant components. Most females, however, migrated to the main stem to mature, indicating female‐biased migration.     

Asymmetric competition drives lake use of coexisting salmonids

To what degree are population differences in resource use caused by competition and the occupation of adjacent positions along environmental gradients evidence of competition? Habitat use may be the result of a competitive lottery, or restricted by competition. We tested to what extent population differences in habitat use of two salmonids, cutthroat trout (Oncorhynchus clarki) and Dolly Varden charr (Salvelinus malma) were influenced by interspecific competition. We hypothesized that the depth distribution of Dolly Varden charr would be affected by competition from the more littoral and surface-oriented cutthroat trout, and that the depth distribution of cutthroat trout would be little affected by competition from Dolly Varden charr. Sympatric populations of cutthroat trout and Dolly Varden charr were created by reciprocal transfers of previously allopatric populations in two experimental lakes. We found evidence of asymmetric competition, as Dolly Varden charr were displaced from littoral habitats when sympatric with cutthroat trout, whereas cutthroat trout remained unaffected by the presence of Dolly Varden charr. Evolved differences between the species, and differences between experimental lakes, also contributed to population differences in habitat use, but asymmetric competition remained as the main driver of different depth distributions in sympatry.     

Karyotype and divergence of stream Dolly Varden from the Southern Sakhalin

The karyotype of stream Dolly Varden inhabiting a tributary of the Belaya River (the basin of Naiba River, southern Sakhalin) was determined (2n = 82 and NF = 98 + 2). According to the main characteristics (chromosome number and arm number, the presence of a pair of marker submeta-subtelocentric chromosomes with nucleolus organizer regions (NORs), one pair of large acrocentric chromosomes, and one pair of subtelocentric chromosomes), this karyotype is identical to the karyotype of anadromous southern Dolly Varden from Salvelinus malma krasheninnikovi of Primorye and Japan. However, in most stream Dolly Varden individuals, additional active nucleolus organizer regions (NORs) located in telomeric and paracentric regions of two to three pairs of acrocentric chromosomes were revealed. It is suggested that the stream and anadromous southern forms of Dolly Varden are evolutionarily related NORs that are silent in the anadromous souther form are active in the stream form. Possible causes of these differences in NOR activity are discussed.     

On the Origin of the Lacustrine Charr <Emphasis Type="Italic">Salvelinus alpinus</Emphasis> Complex from the Kolyma and Sea of Okhotsk Basins

Morphological and genetic characteristics are used for the analysis of the origin of charrs from the lakes of the Kolyma basin (Chuk, Gek, Gulyaevskoe-4, Gulyaevskoe-6, and Lenkovoe) and the Sea of Okhotsk basin (Lake Chistoe). In addition to these samples, the fragments of the control region of mitochondrial DNA (CR mtDNA, 550-bp) and exon 2 of recombination activating gene 1 (RAG1, 899-bp) are sequenced in the charrs from lakes Dzhul’etta and Cherechen’ (Kolyma basin) and lakes Elikchanskie, Bol’shoi Mak-Mak, Ueginskoe (Sea of Okhotsk basin), Ulakhan-Silyan-Kyuel’ (Yana River basin) as well as in northern Dolly Varden from the Kamchatka River. The charr from Lake Chistoe is represented by northern Dolly Varden. In the charrs from lakes Chuk, Gek, Gulyaevskoe-4, Gulyaevskoe-6, Lenkovoe, Cherechen’, Elikchanskie, Bol’shoi Mak-Mak, and Ueginskoe, the haplotypes of mtDNA of the Bering group (a haplogroup of northern Dolly Varden) are revealed. Based on the morphological and RAG1 data, populations of charrs from these lakes belong to the Salvelinus alpinus complex. A transfer of mtDNA of northern Dolly Varden to the charrs from the Kolyma and Sea of Okhotsk basins occurred during the last postglacial expansion and subsequent hybridization. Based on the results of morphological and genetic analysis, the charrs from the Kolyma and Sea of Okhotsk basins cannot be unambiguously referred to the phylogenetic groups of Eurasian Arctic charr or Taranets charr. The presence of mtDNA haplotypes of the Arctic group (a haplogroup of Taranets charr) in the populations of lakes Dzhul’etta, Maksi, and Ueginskoe shows a possibility of their belonging to the group of Taranets charr.     

Evidence for divergence between sympatric stone charr and Dolly Varden along unique environmental gradients in Kamchatka

Here, we describe the biological and ecological differences between partially anadromous Dolly Varden and riverine stone charr distributed in the Kamchatka River middle course. Endemic stone charr, being the ambush predator, is defined by the accelerated growth and prolonged lifespan, robust body, and large mouth, as well as specific marble coloration since an early age. The significant restriction in gene flow between the stone charr and benthos-eating Dolly Varden was supported by an allelic distribution of eight microsatellite loci. Herewith, the identity of mitochondrial DNA control region and intron sequences of growth hormone and metallothionein genes confirms a relatively recent diversification of the groups within the single basin. We suggest the key role of the specific environment for reproduction isolation of stone charr. It occupies coniferous taiga zone with dark-colored water, whereas Dolly Varden spawns uppercourse in the zone of stunted, windblown forest. Conifer litter decomposition products could drive natural selection of stone charr via metabolism acceleration and resistance to toxicants. The metabolic difference could promote piscivory and development of the specific morphological peculiarities of the stone charr. Therefore, we present the first evidence in favor of trophic-based sympatric diversification in a river basin at high latitudes.     

Metapopulation processes and persistence in remnant water vole populations

We examined the spatial distribution of water vole populations in four consecutive years and investigated whether the regional population processes of extinction, recolonisation and migration influence distribution and persistence. We examined how such regional processes are influenced by spatial variation in habitat quality. In addition, we assessed the relevance of metapopulation concepts for understanding the dynamics of species that deviate from classical metapopulation assumptions and developing conservation measures for them. Populations were patchy and discrete, and the patchy distribution was not static between years. Population turnover occurred even in the absence of predatory mink, which only influenced the network of populations at the end of the study. Most populations were clustered close together in the upper tributaries. Local population persistence was predominantly influenced by population size: large populations were more persistent. Recolonisation rates were influenced by isolation and habitat quality. The isolation estimates which best explained the distribution of water vole populations incorporated straight‐line distances, suggesting water voles disperse overland. The distribution of recolonised sites indicated that dispersing voles actively selected habitat on the basis of its quality. Water voles depart from some of the assumptions made by frequently used metapopulation models. In particular there is no clear binary distinction between suitable and non‐suitable habitat. Accounting for variation in habitat quality before investigating temporal changes in population distribution allowed us to demonstrate that the key metapopulation processes were important. The significance of regional population processes relative to local population processes may have increased in declining, fragmented populations compared to pristine regional populations. We hypothesise that although mink predation is likely to eventually cause regional extinction in many areas, metapopulation processes have delayed this decline. Consequently, conservation measures should take into account mink predation rates and regional population processes, before considering aspects of habitat quality.     

Potential fragmentation and loss of thermal habitats for charrs in the Japanese archipelago due to climatic warming

1. The upper thermal limits of the present distributions of two charr species, Dolly Varden, Salvelinus malma , and white-spotted charr, S. leucomaenis , in streams of the Japanese archipelago were examined using groundwater temperature as an index of thermal condition. The lower limits of the altitudinal distributions of Dolly Varden and white-spotted charr were delineated, respectively, by 8 and 16 °C groundwater isotherms.
2. The potential impact of future climatic warming on the geographical distribution, habitat extent and population fragmentation of each species was predicted at both the full archipelago and individual catchment levels.
3. For Dolly Varden, analysis at the full archipelago level indicated a loss of 27.6, 67.2, 79.6 and 89.6% of the current geographical range, respectively, for a 1, 2, 3 and 4 °C increase in mean annual air temperature. The present distribution area of white-spotted charr would likewise reduce by 4.1, 20.5, 33.8 and 45.6%, respectively.
4. Based on the analyses of three individual catchments, one for Dolly Varden and two for white-spotted charr, the lower habitat boundaries for the two charr species could be expected to rise increasingly to higher elevations in each catchment as warming proceeded. As a consequence, there would be large reductions in mean habitat area, with increasing habitat fragmentation followed by localized extinctions of the two species.     

Potential fragmentation and loss of thermal habitats for charrs in the Japanese archipelago due to climatic warming

1. The upper thermal limits of the present distributions of two charr species, Dolly Varden, Salvelinus malma , and white-spotted charr, S. leucomaenis , in streams of the Japanese archipelago were examined using groundwater temperature as an index of thermal condition. The lower limits of the altitudinal distributions of Dolly Varden and white-spotted charr were delineated, respectively, by 8 and 16 °C groundwater isotherms.
2. The potential impact of future climatic warming on the geographical distribution, habitat extent and population fragmentation of each species was predicted at both the full archipelago and individual catchment levels.
3. For Dolly Varden, analysis at the full archipelago level indicated a loss of 27.6, 67.2, 79.6 and 89.6% of the current geographical range, respectively, for a 1, 2, 3 and 4 °C increase in mean annual air temperature. The present distribution area of white-spotted charr would likewise reduce by 4.1, 20.5, 33.8 and 45.6%, respectively.
4. Based on the analyses of three individual catchments, one for Dolly Varden and two for white-spotted charr, the lower habitat boundaries for the two charr species could be expected to rise increasingly to higher elevations in each catchment as warming proceeded. As a consequence, there would be large reductions in mean habitat area, with increasing habitat fragmentation followed by localized extinctions of the two species.     

Spatial and temporal variation in patch occupancy and population density in a model system of an arctic Collembola species assemblage

We employed an experimental model system to investigate the mechanisms underlying patterns of patch occupancy and population density in a high arctic assemblage of Collembola species inhabiting a sedge tussock landscape on Svalbard. The replicate model systems consisted of 5 cores of the tussocks (habitat patches) imbedded in a barren matrix. Four of the patches were open so that animals could migrate between them, while there was one closed patch per system to test the effect of migration on extinction rate. There were model systems of two types: one with long and one with short inter‐patch distances to test the effect of patch isolation on colonisation and extinction rates. Each of the four most common collembolan species at the field site were introduced to two open patches per system (source patches), with the other two functioning as colonisation patches for the species. The experiment was run in an ecotrone over three identical, simulated arctic summers separated by winters of 3 weeks. Six replicates of systems with short and long inter‐patch distances were sampled at the end of each summer. The species varied markedly in their performance in both open arenas and closed patches, indicating differential responses to patch humidity, consistent with their differential distribution along the moisture gradient in the field site. The extinction – colonisation dynamics differed markedly between species as predicted from our field studies. This could partly be ascribed to differential dispersal and colonisation ability, but also to different tolerance to spatially variable patch quality and/or tendency for aggregative behaviour. Three of the species exhibited dynamics that superficially resemble what could be expected from classical metapopulation dynamics. However, there was a striking discrepancy between what would be expected from the effect of migration on the extinction rate of isolated patches (in particular closed patches) and the observed rates. Thus, metapopulation processes, such as stochastic colonisation and extinction events due to demographic stochasticity, were relatively unimportant compared to other sources of spatial variability among which subtle differences in patch quality are probably most important. We discuss the value of combining field studies with model system experiments, in particular when habitat quality cannot easily be measured in the field. However, our field and laboratory studies also emphasise the need for a thorough knowledge of species‐specific life history traits for making biologically sound interpretations based on both observational and experimental data.     

Life history traits and abundance can predict local colonisation and extinction rates of freshwater mussels

1. A critical need in conservation biology is to determine which species are most vulnerable to extinction. Freshwater mussels (Bivalvia: Unionacea) are one of the most imperilled faunal groups globally. Freshwater mussel larvae are ectoparasites on fish and depend on the movement of their hosts to maintain connectivity among local populations in a metapopulation. 2. I calculated local colonisation and extinction rates for 16 mussel species from 14 local populations in the Red River drainage of Oklahoma and Texas, U.S. I used general linear models and AIC comparisons to determine which mussel life history traits best predicted local colonisation and extinction rates. 3. Traits related to larval dispersal ability (host infection mode, whether a mussel species was a host generalist or specialist) were the best predictors of local colonisation. 4. Traits related to local population size (regional abundance, time spent brooding) were the best predictors of local extinction. The group of fish species used as hosts by mussels also predicted local extinction and was probably related to habitat fragmentation and host dispersal abilities. 5. Overall, local extinction rates exceeded local colonisation rates, indicating that local populations are becoming increasingly isolated and suffering an ‘extinction debt’. This study demonstrates that analysis of species traits can be used to predict local colonisation and extinction patterns and provide insight into the long‐term persistence of populations.     

Flexible seaward migration of Dolly Varden <Emphasis Type="Italic">Salvelinus malma</Emphasis> in the Shiretoko Peninsula,Hokkaido, Japan

The migratory history of Dolly Varden Salvelinus malma was investigated in 30 streams in the Shiretoko Peninsula, Hokkaido, Japan, which is in the southern part of their distribution. Captured Dolly Varden were classified into two categories by their body coloration: silvery fish were thought to be an anadromous form, and non-silvery fish a fluvial form that remains in freshwater throughout its life. Forty-seven of 1,739 Dolly Varden collected in the spring and ten of 2,126 Dolly Varden collected in the autumn were silvery fish. Twenty silvery fish and 120 non-silvery fish were examined by analysis of otolith Sr:Ca ratio. Three silvery fish from autumn and 15 non-silvery fish had migrated to salt water or brackish water. The other specimens of silvery fish were regarded as smolt or pre-smolt before seaward migration. The present study suggests that Dolly Varden in the Shiretoko Peninsula have a flexible migratory history, where some of the non-silvery fish utilize marine environments while some individuals follow an anadromous life history.     

Local and Regional Determinants of Colonisation-Extinction Dynamics of a Riparian Mainland-Island Root Vole Metapopulation

The role of local habitat geometry (habitat area and isolation) in predicting species distribution has become an increasingly more important issue, because habitat loss and fragmentation cause species range contraction and extinction. However, it has also become clear that other factors, in particular regional factors (environmental stochasticity and regional population dynamics), should be taken into account when predicting colonisation and extinction. In a live trapping study of a mainland-island metapopulation of the root vole (Microtus oeconomus) we found extensive occupancy dynamics across 15 riparian islands, but yet an overall balance between colonisation and extinction over 4 years. The 54 live trapping surveys conducted over 13 seasons revealed imperfect detection and proxies of population density had to be included in robust design, multi-season occupancy models to achieve unbiased rate estimates. Island colonisation probability was parsimoniously predicted by the multi-annual density fluctuations of the regional mainland population and local island habitat quality, while extinction probability was predicted by island population density and the level of the recent flooding events (the latter being the main regionalized disturbance regime in the study system). Island size and isolation had no additional predictive power and thus such local geometric habitat characteristics may be overrated as predictors of vole habitat occupancy relative to measures of local habitat quality. Our results suggest also that dynamic features of the larger region and/or the metapopulation as a whole, owing to spatially correlated environmental stochasticity and/or biotic interactions, may rule the colonisation – extinction dynamics of boreal vole metapopulations. Due to high capacities for dispersal and habitat tracking voles originating from large source populations can rapidly colonise remote and small high quality habitat patches and re-establish populations that have gone extinct due to demographic (small population size) and environmental stochasticity (e.g. extreme climate events).     

Effects of habitat size and isolation on species immigration–extinction dynamics and community nestedness in a desert river system

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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.     

Matrix Matters:?Differences of Grand Skink Metapopulation Parameters in Native Tussock Grasslands and Exotic Pasture Grasslands

Modelling metapopulation dynamics is a potentially very powerful tool for conservation biologists. In recent years, scientists have broadened the range of variables incorporated into metapopulation modelling from using almost exclusively habitat patch size and isolation, to the inclusion of attributes of the matrix and habitat patch quality. We investigated the influence of habitat patch and matrix characteristics on the metapopulation parameters of a highly endangered lizard species, the New Zealand endemic grand skink (Oligosoma grande) taking into account incomplete detectability. The predictive ability of the developed zxmetapopulation model was assessed through cross-validation of the data and with an independent data-set. Grand skinks occur on scattered rock-outcrops surrounded by indigenous tussock (bunch) and pasture grasslands therefore implying a metapopulation structure. We found that the type of matrix surrounding the habitat patch was equally as important as the size of habitat patch for estimating occupancy, colonisation and extinction probabilities. Additionally, the type of matrix was more important than the physical distance between habitat patches for colonisation probabilities. Detection probability differed between habitat patches in the two matrix types and between habitat patches with different attributes such as habitat patch composition and abundance of vegetation on the outcrop. The developed metapopulation models can now be used for management decisions on area protection, monitoring, and the selection of translocation sites for the grand skink. Our study showed that it is important to incorporate not only habitat patch size and distance between habitat patches, but also those matrix type and habitat patch attributes which are vital in the ecology of the target species.