Polymorphism in lake trout in Great Bear Lake: intra‐lake morphological diversification at two spatial scales

Great Bear Lake is the most northerly lake of its size and provides unique opportunities for intraspecific diversification. Despite increasing attention to intraspecific polymorphism, several knowledge gaps remain (e.g. determining the extent of intraspecific diversification in large relatively pristine lakes and at which spatial scale it can occur). We focused on geographical patterns of morphological differentiation within lake trout (Salvelinus namaycush) to describe two levels of intralake diversification in Great Bear Lake. We used a combination of geometric and traditional linear measurements to quantify differences in body shape, head shape, and fin and body lengths among 910 adult lake trout from the five distinct arms of Great Bear Lake. Although head and fin linear measurements discriminated the three common morphotypes at the whole‐lake level, inter‐arm variation in body shape was observed within each morphotype. A comparison of genetic and morphological distance matrices revealed the lack of an association between the two sets of data, although both comparisons revealed an association in the inter‐arm variation patterns among morphotypes, suggesting a phenotypically plastic response to distinct environments. The whole‐lake and inter‐arm morphological variation observed within lake trout demonstrates the importance of considering scale, especially across large lakes that exhibit marked complexity and a variety of freshwater habitats. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 114 , 109–125.     

Temporal constraints on the potential role of fry odors as cues of past reproductive success for spawning lake trout

Deciding where to reproduce is a major challenge for most animals. Many select habitats based upon cues of successful reproduction by conspecifics, such as the presence of offspring from past reproductive events. For example, some fishes select spawning habitat following odors released by juveniles whose rearing habitat overlaps with spawning habitat. However, juveniles may emigrate before adults begin to search for spawning habitat; hence, the efficacy of juvenile cues could be constrained by degradation or dissipation rates. In lake trout (Salvelinus namaycush), odors deposited by the previous year's offspring have been hypothesized to guide adults to spawning reefs. However, in most extant populations, lake trout fry emigrate from spawning reefs during the spring and adults spawn during the fall. Therefore, we postulated that the role of fry odors in guiding habitat selection might be constrained by the time between fry emigration and adult spawning. Time course chemical, physiological, and behavioral assays indicated that the odors deposited by fry likely degrade or dissipate before adults select spawning habitats. Furthermore, fry feces did not attract wild lake trout to constructed spawning reefs in Lake Huron. Taken together, our results indicate fry odors are unlikely to act as cues for lake trout searching for spawning reefs in populations whose juveniles emigrate before the spawning season, and underscore the importance of environmental constraints on social cues.     

Behavioural responses of female lake trout Salvelinus namaycush to male chemical stimuli and prostaglandin F2α

Male olfactory cues may guide aggregation on spawning reefs, mate evaluation and synchronized gamete release in lake trout Salvelinus namaycush, but a lack of information on the source and identity of the cues precludes direct tests of their function. Using a two-channel flume, we found ovulated female lake trout increased time spent in the channel treated with spermiated male-conditioned water, urine and bile but not synthesized prostaglandin F_2α. We suggest future efforts to characterize male olfactory cues focus on urine and postulate that bile acids contribute to its behavioural activity.     

Application of otolith analyses to investigate broad size distributions of young yellow perch in temperate lakes

Otolith analyses were used to back-calculate young-of-the-year (YOY) yellow perch Perca flavescens hatch-date estimates to interpret broad length distributions observed within a small Great Lake, Lake St. Clair, and a small inland lake, Lake Opinicon, during 1998. For the Great Lake, the earliest observed hatch date occurred 2 weeks after suitable water temperatures and latest hatch dates occurred the same week temperatures were considered too warm for spawning. For the inland lake, the earliest hatch date occurred 4 weeks after suitable water temperatures and the latest hatch dates occurred 2 weeks after the water temperatures were considered too warm for spawning. It is inferred that spawning in each lake had a duration of >9 weeks. This suggests that natural perch populations can protract their spawning season opportunistically under the appropriate environmental cues. During 1998, these cues involved a shortened winter, earlier spring, and slow warming to typical summer temperatures, caused by the El Niño-Southern Oscillation. Time of YOY hatch determined the absolute opportunity for growth and resulted in a match or mismatch with optimal foraging conditions and contributed to the development of the observed YOY length distributions.     

Lake trout (<Emphasis Type="Italic">Salvelinus namaycush</Emphasis>) spawning behaviour: the evolution of a new female strategy

A video camera inside an underwater housing was used to record the spawning activities of lake trout, Salvelinus namaycush, in Kushog Lake, Ontario (Canada). Contrary to the commonly accepted belief describing lake trout as the only salmonine to spawn exclusively at night, the recordings were taken during the day. Lake trout spawning behaviour is described in detail and compared with other salmonine species. The loss of female “nest-building” and assumption of a novel itinerant strategy during which females travel across the breeding grounds accompanied by a group of males, is considered to be an important change in the evolution of the lake trout’s unique mode of spawning.     

Suitability of habitat for spawning lake trout

We provide further insight into the reproductive ecology and spawning requirements of lake trout. New comparative information about substrate characteristics, sediment transport, quality of interstitial water at spawning substrates, and the role of temperature in site selection and time of spawning is given for lakes Simcoe and Manitou (Ontario) and Seneca Lake (New York). Spawning lake trout commonly use stable lag deposits derived from glacial sediments, or relict features such as fans, bars or submerged talus slopes. Artificial breakwaters of broken material may also provide suitable substrates. Optimal particle sizes range from 4 to 10 cm diameter but larger materials to 30 cm are also successfully utilized for spawning. The transport of finer particulates by wind generated water movements may limit the suitability of some substrates and successful spawning sites are usually remote from depositional effects. Successful embryo development is associated with low nutrient conditions, with high dissolved oxygen (>7 mg L^-1) and with low un-ionized ammonia (<12.5 g L^-1) in the interstitial water of spawning substrates. Shallow-water spawning appears to be the common strategy of colonizing lake trout. Some deepwater spawning in the Great Lakes may reflect initial colonization in shallow-water and adaptation to later increases in water level, but some may also reflect unique behavioural and physiological adaptations. Temperature is an important cue, and many wild and hatchery stocks spawn at 8 to 13 °C with latitudinal shifts in the actual time of spawning. These requirements are summarized as a dichotomous key for evaluation of approaches to restoration of lost or damaged lake trout stocks.Presented at the Conference on Rehabilitation of Lake Trout in the Great Lakes: A Critical Assessment (sponsored by the Great Lakes Fishery Commission, Ann Arbor, Michigan, January 10–14, 1994).     

Among‐individual diet variation within a lake trout ecotype: Lack of stability of niche use

In a polyphenic species, differences in resource use are expected among ecotypes, and homogeneity in resource use is expected within an ecotype. Yet, using a broad resource spectrum has been identified as a strategy for fishes living in unproductive northern environments, where food is patchily distributed and ephemeral. We investigated whether specialization of trophic resources by individuals occurred within the generalist piscivore ecotype of lake trout from Great Bear Lake, Canada, reflective of a form of diversity. Four distinct dietary patterns of resource use within this lake trout ecotype were detected from fatty acid composition, with some variation linked to spatial patterns within Great Bear Lake. Feeding habits of different groups within the ecotype were not associated with detectable morphological or genetic differentiation, suggesting that behavioral plasticity caused the trophic differences. A low level of genetic differentiation was detected between exceptionally large‐sized individuals and other piscivore individuals. We demonstrated that individual trophic specialization can occur within an ecotype inhabiting a geologically young system (8,000–10,000 yr BP), a lake that sustains high levels of phenotypic diversity of lake trout overall. The characterization of niche use among individuals, as done in this study, is necessary to understand the role that individual variation can play at the beginning of differentiation processes.     

Past, present and future of fishery management on one of the world’s last remaining pristine great lakes: Great Bear Lake, Northwest Territories, Canada

Great Bear Lake, Northwest Territories, Canada supports important sport and aboriginal-subsistence fisheries and is among the last remaining pristine great lakes of the world. The lake’s unique ecology is characterized by a harsh subarctic climate, low productivity and species diversity, and high intraspecific diversity of lake trout. These aspects in combination with geographical remoteness present special challenges to the management of two exceptionally different fisheries. The history of its management has not been well documented or reviewed; therefore, our objectives in this paper were to summarize the history and status of Great Bear Lake’s fisheries and their management, and to identify gaps in knowledge, future challenges, and actions required to meet those challenges. Prior to 1970, management goals for the lake had not been established formally, and harvest numbers and biological characteristics of fish were unknown. To reduce data gaps, creel surveys, gillnet assessments, fish tagging, and subsistence monitoring were implemented. During the 1980s, Canada established the management goal of conserving a high quality sport fishery, while protecting aboriginal access to the subsistence fishery. On the basis of assessment data, lodge harvest quotas, lodge guest capacity limits, individual angler harvest limits, and angler licensing were among the management actions taken to achieve that goal. Since 2005, decision making has been guided by the “water heart,” a management plan for Great Bear Lake. Management has since evolved into a complex co-management system among aboriginal, territorial, and federal governments. Changes in regulations, sport trophy and tourism industries, subsistence resource use, and social and cultural norms and practices contributed to changes in the Great Bear Lake fishery and its management. In the future, anthropogenic and climate change are the two main challenges facing co-management of the lake’s resources. We recommend the adoption of an ecosystem approach to management, establishment of a fishery technical committee, reformulation of the current plan, explicit commitment to evaluation, conducting community-based monitoring, and development and use of a joint strategic plan among co-managers to describe how to interact and implement the Great Bear Lake management plan.     

Comparison of brook trout reproductive success and recruitment in an acidic adirondack lake following whole lake liming and watershed liming

Limestone applications to the catchment of one tributary to Woods Lake were highly effective in reducing stream acidity and stabilizing seasonal fluctuations in pH. The resulting improvement in stream water quality also led to a dramatic shift in reproductive strategy of the Woods Lake brook trout population. Prior to catchment liming, brook trout in Woods Lake were restricted to spawning on poor quality near shore substrate with limited ground water seepage. Reproductive success was limited by high mortality of eggs and larvae and recruitment from in lake spawning was not successful. Spawning brook trout did not utilize the tributary for spawning prior to watershed liming. Mitigation of acidity in the tributary, by catchment liming, effectively extended the spawning habitat available to the Woods Lake brook trout population and one year following treatment brook trout spawned successfully in the tributary for the first time in 6 years of observation. Significant recruitment of young trout into the lake population occurred from 1991 through 1993, although the absolute number of fish captured was relatively small. In the fall of 1993, four year classes of naturally spawned brook trout were present in the lake. Although reproductive success was enhanced by improving tributary spawning habitat in the Woods Lake basin, self maintenance of the population may be limited by low recruitment rates of young trout, due to high levels of summer mortality resulting from predation. Mitigation of this constraint would require substantially higher levels of fry production than were observed in Woods Lake and/or enhanced refugia for young trout. The results of this experiment suggest that re-establishment of tributary spawning populations of brook trout may be possible, with future reductions in acidic deposition, in acidic Adirondack lakes with limited in-lake spawning habitat.     

Salvelinus namaycush spawning substratum attracts egg predators and opportunists through chemosensory cues

Two separate field experiments were conducted in a series of small boreal lakes to test for the attraction of egg predators to lake trout Salvelinus namaycush spawning shoals and subsequently to determine whether chemosensory cues attract egg predators to these sites. In the first experiment, minnow traps set on spawning sites captured significantly more egg predators than those set on structurally similar non‐spawning sites. Captures of slimy sculpin Cottus cognatus, common shiner Luxilus cornutus, blacknose shiner Notropis heterolepis and virile crayfish Orconectes virilis were more than double on spawning sites relative to non‐spawning sites for the two study lakes. To test whether chemosensory cues could attract egg predators to S. namaycush spawning sites, paired minnow traps were placed on eight to 10 sites in each of the three study lakes; one trap contained visually concealed S. namaycush spawning substratum and the other with visually concealed non‐spawning substratum. Traps containing spawning substratum consistently captured more fish and had higher mean daily catches than those that contained non‐spawning substratum. The combined results demonstrate a greater prevalence of egg predators on S. namaycush spawning shoals that appears to be the result of chemosensory attraction to spawning substratum.     

Differentiation of deep-water lake charr <Emphasis Type="Italic">Salvelinus namaycush</Emphasis> in North American lakes

Deep-water morphs of lake charr, Salvelinus namaycush, are found, with one exception, in four of the largest lakes in the world: lakes Superior and Mistassini (QC) and Great Bear and Slave lakes. This paper advances a hypothesis for resource polymorphisms involving two types of deep-water morph, one of which is characteristic of the humper and the other of the siscowet charrs of Lake Superior. My hypothesis states that, first, the humper, or a humper-like morph, diverged postglacially in sympatry from the ancestral common (shallow-water) lake charr and became a feeding specialist on Mysis relicta. Second, in at least two of the four lakes the siscowet, or a siscowet-like charr, diverged as a feeding specialist on postglacially derived forms of deep-water ciscoes. In Lake Superior a successional process may have resulted in dominance of the siscowet at the expense of the humper charr. I concur with a previous inference that the one occurrence of a deep-water charr in a small lake (the above exception) represents emigration from Lake Superior. I further infer that this event involved an early humper charr, which implies that this morphotype had differentiated in Lake Superior in less than 1,900 year. I suggest that innate differences in plasticity, breeding behavior and assortive mating, and philopatry account for why Arctic charr isolate readily in small lakes whereas lake charr do not. My hypothesis assumes divergence of deep-water morphs occurred postglacially, an idea consistent with genetic and biogeographical evidence.     

Reproductive ecology and early life history of a lacustrine sculpin, Cottus extensus (Teleostei, Cottidae)

Using a variety of sampling techniques and observations we describe aspects of the reproductive ecology and early life-history of Bear Lake sculpin, Cottus extensus, a species endemic to Bear Lake, Utah-Idaho. Adult sculpin spawned in shallow water (0.5–6.0 m depths) in cavities beneath large cobbles and boulders. During 1993, egg mass densities were highest (> 4.0 m²) at 1.0–2.0 m depths. Electivity indices verified substrate selection and also indicated an avoidance of s and- and gravel-embedded materials. During years of low water elevation, suitable spawning substrates were restricted to one or two limited areas of the lake and comprised < 0.004% of the total benthic area. Disturbance from turbulence, as would be produced by storm-induced waves, appeared to initiate hatching. After hatching, embryos passed through a pelagic interval that lasted for approximately one week. Laboratory experiments confirmed the short duration of this surface-swimming behavior. The pelagic behavior of free embryos appears to aid in dispersal of fish via currents from limited spawning habitats to areas throughout the 282 km² lake.     

Basic description and some notes on the evolution of seven sympatric morphs of Dolly Varden Salvelinus malma from the Lake Kronotskoe Basin

The study examines the basic morphological and ecological features of Dolly Varden from Lake Kronotskoe (Russia, Kamchatka). Seven valid morphs different in head proportions, feeding, timing, and place of spawning have been determined in this ecosystem. The basic morphometric characteristics clearly separate Lake Kronotskoe morphs from each other, as well as from its potential ancestor (Dolly Varden). According to CVA analysis, the most notable morphological characteristics determining the mouth position are the length of a lower jaw and rostrum. Furthermore, five of seven morphs inhabit different depth zones of the lake and feed on different food resources. Our data suggest that reproductive isolation may be maintained by temporal/spatial isolation for two morphs with lacustrine spawning, and by spatial isolation only for the rest of the morphs with riverine spawning. The sympatric diversity of the Lake Kronotskoe charrs is exceptionally wide, and there are no other examples for seven sympatric morphs of genus Salvelinus to coexist within a single ecosystem. This study puts forward a three‐step hypothetical model of charr divergence in Lake Kronotskoe as a potential ground for future studies.     

Integration of acoustic telemetry and GIS to identify potential spawning areas for lake trout (Salvelinus namaycush)

Locations of potential spawning areas for lake trout (Salvelinus namaycush) were predicted in Lake Opeongo, Ontario, Canada using information gained via acoustic telemetry and geographic information system (GIS) technologies. From 1998 to 2000, 18 adult lake trout (mean fork length 553 mm) implanted with acoustic transmitters (battery life 2 years) were manually tracked. For evening fall locations within the erosive zone of the lake (determined using an existing sedimentation model), habitat variables (slope, depth, and effective fetch) were summarised using GIS. Sites selected by lake trout during the spawning window were in areas of mean fetch equal to 1.5 km and mean slope of 10.6% (n = 50 fixes). We used GIS to identify areas that matched the mean habitat criteria and thus locate potential spawning areas. This model correctly identified 19 of 21 known spawning sites, as well as additional sites used by spawning females in an earlier telemetry study. Depths of traditional fall netting sites are shallow compared to areas in which telemetered lake trout were found during evenings of the spawning period (means 3.1 vs. 5.1 m, respectively). Through the use of information on spawning habitat selection gained through telemetry and knowledge of the physical characteristics of the lake, we provide an alternative means of identifying potential spawning habitat for lake trout.     

Field activity of lake trout during the reproductive period monitored by electromyogram radiotelemetry

Many lake trout Salvelinus namaycush spawn in shallow areas along windswept shores. However, precise determination of time and location of spawning is limited by its nocturnal occurrence, and possibly by some postulated, but unproven, spawning in deep water by certain populations. In White Pine Lake, Ontario, a well-studied experimental lake, lake trout spawn in two shallow locations where some direct observation is possible. Radiotelemetry apparatus was used in 1991 to obtain records of the electromyograms (EMGs) produced by muscle activity in one adult male and one adult female lake trout, captured from the lake before their reproductive period and returned to the lake at the beginning of the spawning period. Both fish survived, though only the male was detected as active over the main spawning site, where it showed evening EMG activity patterns indicating considerable activity. At other times of day, when not located over the spawning site, but present elsewhere in the lake, the male's EMG (i.e. muscle) values were considerably reduced. At the cessation of the spawning period, the transmitter-equipped male's EMG record showed no further pattern indicative of high activity during what had formerly been the daily spawning period. The female was tracked as she moved around the lake, but based on both her location and level of EMG activity, is thought not to have spawned. In 1992, a second male was captured, equipped with a transmitter and released again. This fish failed to show spawning activity but, as with the 1991 female, moved fairly considerable distances in the lake. In one instance, in a swim of short duration (6 min), well away from the spawning grounds, this fish displayed EMG activity levels resembling those of the high activity of spawning. The results obtained appear to demonstrate that EMG telemetry could be used to assess the level of muscular activity occurring in lake trout, especially in relation to reproductive behaviour, and when they cannot be directly observed.     

Morphological divergence and origin of sympatric populations of European whitefish (Coregonus lavaretus L.) in Lake Femund, Norway

Combining morphological and genetic analysis, we compared patterns of diversification within and between morphs among sympatric European whitefish (Coregonus lavaretus L.) populations in Lake Femund, Norway. Seven external populations, from potential colonization routes into Lake Femund were included. We found that deep-, shallow-, river- and bay spawning populations are distinct morphs in Lake Femund. Within morphs, populations range from being similar genetically (Fst=0-0.005) among deep-spawning populations to being highly differentiated (Fst=0.153) between bay-spawning populations. Between morphs, genetic differences ranged from a low (Fst=0.008-0.022) between deep- and shallow-spawning populations to high difference (Fst=0.125-0.143) between shallow- and bay-spawning populations. A higher proportion of molecular variance was seen among (3.9%) than within morphs (2.8%). The adaptive gene combinations behind the four morphs seem to have originated within the lake, although the lake could have been colonized from more than one source population.     

Sexual selection,reproductive behavior,and speciation in the mbuna species flock of Lake Malawi (Pisces: Cichlidae)

Synopsis The cichlid species flocks of the African Great Lakes represent the most extreme case of adaptive radiation among vertebrates. Recently, attention has focused on the potential for sexual selection to drive or accelerate speciation in these fishes. Cichlids as a whole are social in nature and display complex behavior, particularly during courtship and spawning; however, the extent to which changes in species recognition cues may account for species diversity among haplochromine lineages has remained speculative. Our investigations have indicated that oral incubating haplochromines show a reduction in diversity and extent of courtship relative to substrate brooding cichlids, and apparently retain aspects of a primitive specific mate recognition system. Laboratory observations of courtship in the Malawian endemicPseudotropheus zebra suggest that organization of the spawning bout is loose, and lacking in any well defined stimulus-response chain. Interspecific comparisons of in situ courtship behavior among male mbuna, lithophilous haplochromines of Lake Malawi, revealed only one potential example of species-specific behavior, and indicated that mate choice occurred prior to the onset of intense courtship. Courtship display by male mbuna does not appear critical to species recognition and may represent an evolutionary relict. Alternatively, male courtship display may have an indirect effect on successful reproduction or may be maintained through pleiotropic effects. The mbuna display no evidence of behavioral innovation and show limited interspecific differentiation in behavioral expression. More generally, there is no evidence at present to suggest that epigamic sexual selection, acting on courtship behavior, has been a major mechanism in the diversification of the haplochromine species flocks.     

No barriers to gene flow among sympatric polychromatic 'small'Telmatherina antoniae from Lake Matano, Indonesia

Genetic divergence, assortative courtship and intermale aggression were assessed between sympatric colour morphs of the sailfin silverside Telmatherina antoniae , endemic to Lake Matano, Indonesia. Genetic analysis using microsatellite markers showed no barriers to gene flow among T. antoniae primary colour morphs (blue and yellow) within sampling sites, sympatric populations or at the lake-wide level. Low but significant genetic differentiation was found between yellow morphs and mixed (blue–yellow) morphs. Behavioural surveys indicated assortative courtship does occur along primary colour lines; however, intermale aggression among paired and intruding male morphs appeared equal with respect to male colour. These observations support the hypothesis that males view other males as threats to their courtship regardless of their colour. This study supports recent work suggesting that assortative mating is present in T. antoniae despite a lack of reproductive isolation among colour morphs.     

Spatial and Temporal Relations between Fluvial and Allacustrine Yellowstone Cutthroat Trout, Oncorhynchus clarki bouvieri, Spawning in the Yellowstone River, Outlet Stream of Yellowstone Lake

Yellowstone cutthroat trout (YCT), Oncorhynchus clarki bouvieri, that spawn in the outlet of Yellowstone Lake show two potamodromous migration patterns, fluvial and allacustrine. The main purpose of this study was to determine whether those fluvial and allacustrine YCT represent reproductively isolated stocks. Redd surveys indicated spawning occurred during about 5 consecutive weeks between late May and mid-July 1993–1995. Lake fish (N=6), defined as radiotagged YCT that entered Yellowstone Lake after the spawning period (i.e. allacustrine pattern), were found in the river between the lake outlet (river kilometer [Rkm] 0) and Rkm 20.0 during spawning. Probable lake fish (N=28; tagged YCT that were last detected near the lake outlet) were found between Rkm 0 and Rkm 22.5 during spawning. River fish (N=4; tagged YCT that remained in the river when annual tracking concluded in fall, i.e. fluvial pattern) were found between Rkm 1.1 and Rkm 18.0 during spawning. Fidelity to spawning areas used between consecutive years was suggested by one of five lake fish and the single river fish for which data were available. Spatial overlap in spawning and a lack of temporal separation between the life-history types during spawning suggested that fluvial and allacustrine YCT were not reproductively isolated. Radiotagging, as well as visual observations made annually from boats during April and May, indicated fluvial YCT overwintered downstream from Rkm 14 and were few, probably on the order of 10% of all YCT that spawned in the Yellowstone River.     

Minimal gene flow from introduced brown trout (Salmo trutta L.) after 30 years of stocking

Lake Tinnsjø, Norway, has been heavily stocked over three decades with two different brown trout stocks. A population genetic survey based on 13 allozyme coding loci was conducted to investigate the genetic composition of the present trout population and the genetic impact of the two donors. Contrary to expectations, highly significant differences in allelic frequencies between Lake Tinnsjø trout and the two donor stocks were found at several loci, suggesting minimal gene flow from stocked trout to the wild populations. Pairwise genetic distance values were several times higher between donor stocks and Lake Tinnsjø samples than between the various spawning inlets to Lake Tinnsjø. F_st increased from 0.024 to 0.090, when samples from the donor stocks were included in the material. There were no deviations from expected Hardy–Weinberg distribution of genotypes in the spawning inlets. It was concluded that the donor stocks contributed little to the trout gene pool in the lake. A possible exception is the trout population below the lake outlet.