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Developmental instability and fitness in Periploca laevigata experiencing grazing disturbance

We investigated the sensitivity of developmental instability measurements (leaf fluctuating asymmetry, floral radial asymmetry, and shoot translational asymmetry) to a long‐standing natural stress (grazing) in a palatable tannin‐producing shrub ( Periploca laevigata Aiton). We also assessed the relationship between these measures of developmental instability and fitness components (growth and floral production). Developmental instability, measured by translational asymmetry, was the most accurate estimator of a plant’s condition and, consequently, environmental stress. Plants with less translational asymmetry grew more and produced more flowers. Plants from the medium‐grazed population were developmentally more stable, as estimated by translational and floral asymmetry, than either more heavily or more lightly grazed populations. Leaf fluctuating asymmetry was positively correlated with tannin concentration. The pattern of internode growth also responded to grazing impact. Plants under medium to heavy grazing pressure accelerated early growth and consequently escaped herbivory later in the season, i.e., at the beginning of the spring, when grazing activity was concentrated in herbaceous plants. Periploca laevigata accelerated growth and finished growing sooner than in the other grazing treatment. Thus, its annual growth was more mature and less palatable later in the season when grazers typically concentrate on shrubs. The reduction of developmental instability under medium grazing is interpreted as a direct effect of grazing and not as the release from competition.

International Journal of Plant Sciences↗

Higher nest predation favors rapid fledging at the cost of plumage quality in nestling birds

Life-history theory predicts that rapid growth comes at a cost to offspring quality and adult longevity. However, trade-offs have been examined primarily based on proximate variation within species rather than evolved differences across species. Evolved differences are important to examine because species may co-evolve mechanisms to reduce long-term costs of rapid growth. For example, trade-offs associated with fast growth and short development time might be expressed in ephemeral traits because costs associated with these traits are paid for only a short duration. We explored this idea using nestling body feathers, an ephemeral trait, and examined whether shorter development time yielded lower quality feathers. We found a strong trade-off whereby nestlings that spend more time in the nest produced higher quality plumage across 123 temperate and tropical species. Using a subset of these species (n=67), we found that plumage quality was lower in species with higher rates of nest predation, which favors rapid nestling growth and short development times. Our results suggest that evolution of more rapid growth may co-evolve with other mechanisms whereby long-term costs are reduced by shifting costs of allocational trade-offs to ephemeral traits that are short-lived.

The American Naturalist↗

Quantifying the contribution of habitats and pathways to a spatially structured population facing environmental change

The consequences of environmental disturbance and management are difficult to quantify for spatially structured populations because changes in one location carry through to other areas as a result of species movement. We develop a metric, G , for measuring the contribution of a habitat or pathway to network-wide population growth rate in the face of environmental change. This metric is different from other contribution metrics, as it quantifies effects of modifying vital rates for habitats and pathways in perturbation experiments. Perturbation treatments may range from small degradation or enhancement to complete habitat or pathway removal. We demonstrate the metric using a simple metapopulation example and a case study of eastern monarch butterflies. For the monarch case study, the magnitude of environmental change influences the ordering of node contribution. We find that habitats within which all individuals reside during one season are the most important to short-term network growth under complete removal scenarios, whereas the central breeding region contributes most to population growth over all but the strongest disturbances. The metric G provides for more efficient management interventions that proactively mitigate impacts of expected disturbances to spatially structured populations. use changes in one location carry through to other areas due to species movement. We develop a metric, G, for measuring the contribution of a habitat or pathway to network-wide population growth rate in the face of environmental change. This metric is different than other contribution metrics as it quantifies effects of modifying vital rates for habitats and pathways in perturbation experiments. Perturbation treatments may range from small degradation or enhancement to complete habitat or pathway removal. We demonstrate the metric using a simple metapopulation example and a case study of eastern monarch butterflies. For the monarch case study, the magnitude of environmental change influences ordering of node contribution. We find that habitats through which all migrants flow are the most important to short-term network growth under complete-removal scenarios. The metric G provides for more efficient management interventions that proactively mitigate impacts of expected disturbances to spatially structured populations.

American Naturalist↗

Arctic sea ice decline contributes to thinning lake ice trend in northern Alaska

Field measurements, satellite observations, and models document a thinning trend in seasonal Arctic lake ice growth, causing a shift from bedfast to floating ice conditions. September sea ice concentrations in the Arctic Ocean since 1991 correlate well ( r = +0.69, p < 0.001) to this lake regime shift. To understand how and to what extent sea ice affects lakes, we conducted model experiments to simulate winters with years of high (1991/92) and low (2007/08) sea ice extent for which we also had field measurements and satellite imagery characterizing lake ice conditions. A lake ice growth model forced with Weather Research and Forecasting model output produced a 7% decrease in lake ice growth when 2007/08 sea ice was imposed on 1991/92 climatology and a 9% increase in lake ice growth for the opposing experiment. Here, we clearly link early winter 'ocean-effect' snowfall and warming to reduced lake ice growth. Future reductions in sea ice extent will alter hydrological, biogeochemical, and habitat functioning of Arctic lakes and cause sub-lake permafrost thaw.

Alaska↗

Non-freezing cold event stresses can cause significant damage to mangrove seedlings: Assessing the role of warming and nitrogen enrichment in a mesocosm study

Mangroves are expanding poleward along coastlines globally as a response to rising temperatures and reduced incidence of freezing under climate change. Yet, knowledge of mangrove responses to infrequent cold events in the context of future global and regional environmental changes is limited. We initiated a mesocosm experiment in which the seedlings of two mangrove species were grown either at ambient temperature or under warming with and without nitrogen (N) loading. During a short winter period, an unusually severe cold event occurred with the lowest temperature of 2°C. We assessed the possible response of these two mangrove species to the cold stress. We found that the cold event caused various degrees of damage to the seedlings of both mangrove species, with the warming treatment seemingly protecting leaves and branches from the cold damage. However, warming did not buffer mangroves to mortality from those low temperatures in either species. The cold event resulted in a significant decrease in seedling growth rates and net ecosystem CO2 uptake in the post-cold period relative to the pre-cold period, though the cold event did not alter the effects of warming on these parameters of both mangrove species. The cold event differentially altered physiological responses of the two species growing under N loading, with A. marina growing in higher N concentrations having a reduced growth response after the cold event, whereas B. gymnorrhiza displayed no change in post-cold period versus pre-cold period growth. Our findings suggest the pivotal role of cold events, versus freeze events, in regulating mangrove survival and growth even under future warming scenarios. Two mangrove species exhibited differential survival and growth responses to the cold event at different N concentrations, which has implications for how we can restore and conserve mangroves among the world's eutrophied sub-tropical estuaries and with future warming.

Environmental Research↗

Effects of dietary selenium on the amphipod Hyalella azteca and the midge Chironomus dilutus

Chronic selenium (Se) water quality criteria are based primarily on dietary organoselenium exposure and subsequent reproductive effects in fish. Available chronic Se toxicity data suggests that invertebrates are less sensitive than fish, but chronic invertebrate studies are limited. We evaluated yeast-based diets for chronic toxicity studies with Hyalella azteca and Chironomus dilutus. Growth and survival were similar among C. dilutus fed a yeast-only diet, a mixture of yeast and fish flake food, and a mixture of yeast and diatoms. Survival and growth of H. azteca were best in the yeast + diatom diet, so this diet was used for subsequent tests. In rangefinder tests, H. azteca 28-day survival, growth, and biomass, and C. dilutus 11-day survival were all significantly reduced at a dietary Se concentration of 49 µg/g dry weight, whereas C. dilutus growth and biomass were only affected at 200 µg/g dry weight Se. H. azteca had similar sensitivity to dietary Se in a 28–42-day chronic test—survival, growth, and biomass were significantly reduced at 44.5 µg/g dry weight Se. Dietary selenium reduced H. azteca reproduction by 75%–100% relative to Controls, but differences were not significant. Tissue Se EC10s were lower for H. azteca (9.0–23 µg/g dry weight) than for C. dilutus (11–56 µg/g dry weight). However, model evaluations suggest that the C. dilutus 11-day survival and H. azteca 42-day EC10s were less reliable than other endpoints. When tissue EC10s were converted to equivalent Se concentrations in fish tissue with a food web model, H. azteca (18 µg/g dry weight) and C . dilutus (68 µg/g dry weight) ranked 6th and 12th of 13 freshwater genera, respectively. Overall, these results suggest that the current U.S. Environmental Protection Agency tissue-based Se water quality criteria would be protective for fish and invertebrates and could be used for management of fishless waters.

Environmental Toxicology and Chemistry↗

The potential for species distribution models to distinguish source populations from sinks

1. While species distribution models (SDM) are frequently used to predict species occurrences to help inform conservation management, there is limited evidence evaluating whether habitat suitability can reliably predict intrinsic growth rates or distinguish source from sink populations. Filling this knowledge gap is critical for conservation science, as applications of SDMs for management purposes ultimately depend on these typically unobserved population or metapopulation dynamics. 2. Using regression, we associate previously published population level estimates of intrinsic growth and abundance derived from a Bayesian analysis of mark-recapture data for 17 bird species found in the contiguous United States with SDM habitat suitability estimates fitted here to opportunistic data for these same species. We then use AUC to measure how well SDMs can distinguish populations categorized as sources and sinks, depending on their intrinsic growth rates estimated from the mark-recapture data. We built SDMs using two different approaches, boosted regression trees (BRT) and Generalized Linear Models (GLM), and compared their predictive performance. Each SDM was built with presence points obtained from eBird and 10 environmental variables previously selected to model intrinsic growth rates and abundance for these species. 3. We show that SDMs built with opportunistic data are poor predictors of species demography in general; both BRT and GLM explained very little spatial variation of intrinsic growth rate and population abundance (median R2 across 17 species was close to 0.1 for both SDM methods). SDMs do, however, estimate higher suitability for source populations as compared to sinks. Out of 13 species which had both source and sink populations, both BRT and GLM had AUC values greater than 0.7 for 7 species when discriminating between sources and sinks. 4. Habitat suitability have the potential to be a useful measure to indicate a population’s ability to sustain itself as a source population, however more research on a diverse set of taxa is essential to fully explore this potential. This interpretation of habitat suitability can be particularly useful for conservation practice, and identification of explicit cases of when and how SDMs fail to match population demography can be informative for advancing ecological theory.

Journal of Animal Ecology↗

Linking environmental variability to long-term demographic change of an endangered species using integrated population models

Understanding how species populations change with environmental conditions is important for implementing effective habitat management and conservation strategies. Challenges to evaluating population-level responses to environmental conditions arise when data are sparse or not spatiotemporally aligned, especially for at-risk species with small, declining numbers. We synthesized 30 years (1992–2021) of three partially aligned data sets to build a Bayesian integrated population model (IPM) and evaluate demographic and environmental drivers of growth rates for six separately managed ‘subpopulations’ (A–F) of the federally endangered Cape Sable seaside sparrow endemic to the Florida Everglades. We found that juvenile survival peaked at inundation periods (hydroperiods) around 100–220 days and dropped sharply outside those values, while adult survival increased with longer periods of water depth <20 cm, but not with longer periods of water depth >20 cm. Fecundity increased when water depths were more stable, more area was dry, intervals between fires were longer and less area was burned. Changes in population growth rates tended to occur in years that juvenile and adult survival were associated with hydroperiod, especially in the two largest subpopulations B and E. Population growth rates were also associated with hydrologic conditions during the breeding season and fire dynamics through changes in fecundity, most notably in the smaller subpopulations A, C/F and D. Synthesis and applications . Our IPM represents the first long-term population analysis of the Cape Sable seaside sparrow connecting demographic processes to environmental factors. Our results suggest that sustaining periods of shallow water year-round may enhance Cape Sable seaside sparrow survival and population growth. Also, limiting water depth variability and maintaining dry conditions during the breeding season and inhibiting fires in consecutive years may increase fecundity and population growth. Identifying the mechanistic links between environmental and population dynamics could inform how species are expected to respond to management decisions and anticipated ecosystem changes.

Florida↗

Modeling strategies and evaluating success during repatriations of elusive and endangered species

Wildlife repatriation is an important tool to decrease extinction risk for imperiled species, but successful repatriations require significant time, resources and planning. Because repatriations can be long and expensive processes, clear release strategies and monitoring programs are essential to efficiently use resources and evaluate success. However, monitoring can be challenging and surrounded by significant uncertainty, particularly for secretive species with extremely low detection probability. Here, we simulated how alternative repatriation strategies influence repatriation success for the eastern indigo snake Drymarchon couperi , a federally-Threatened species that is currently being repatriated in Alabama and Florida. Critically, we demonstrate how observed population growth can differ from true population growth when detection probabilities are low and mark-recapture analyses are not an option. Specifically, we built a stochastic stage-based population model to predict population growth and extinction risk under different release strategies and use information from ongoing repatriations to predict success and guide future releases. Because D. couperi is difficult to monitor, we modeled how detection probability influenced perceptions of abundance and population growth by monitoring programs. Simulated repatriation strategies releasing older, head-started snakes in greater abundance and frequency created wild populations with decreased extinction risk relative to scenarios releasing fewer and younger snakes less frequently. Ongoing repatriations currently have a 0.23 (Alabama) and 0.61 (Florida) probability of quasi-extinction, but extinction risk decreased to 0.07 and 0.10 at sites upon achieving the targeted number of releases. Abundances observed under realistic detection thresholds for D. couperi did not always predict true population growth; specifically, we demonstrate that monitoring programs during repatriations of secretive species may indicate that efforts have been unsuccessful when populations are actually growing. Overall, our modeling framework informs release strategies to maximize repatriation success while demonstrating the need to consider how detection processes influence assessment of success during conservation interventions.

Florida↗

Evaluation of ageing accuracy with complementary non‐lethal methods for slow‐growing, northern populations of shoal bass

In the upper Chattahoochee River basin, where some populations of shoal bass, Micropterus cataractae Williams & Burgess, are imperilled, age and growth data are lacking. Age and growth of shoal bass in this basin were assessed with non‐lethal means using scales and mark–recapture. Mark–recapture data allowed for estimation of accuracy and determination of effects of any scale‐based inaccuracies on growth models. Scale‐based age estimates were accurate for 57% of the samples, and errors of 1 to 3 years included equal numbers of over‐ and underestimates of age. von Bertalanffy growth models based on scale ages were similar to those based on mark–recapture ages for ages 3–8 but noticeably divergent for younger and older fish. Scales provided estimates of longevity up to 12 years of age, and growth models produced from mark–recapture suggest scale ages underestimated age, especially for older fish. These populations of shoal bass live longer and grow slower than other populations, suggesting regional management strategies may be needed.

Alabama, Florida, Gerogia↗

Temperature‐associated decreases in demographic rates of Afrotropical bird species over 30 years

Tropical mountains harbor globally significant levels of biodiversity and endemism. Climate change threatens many tropical montane species, yet little research has assessed the effects of climate change on the demographic rates of tropical species, particularly in the Afrotropics. Here, we report on the demographic rates of 21 Afrotropical bird species over 30 years in montane forests in Tanzania. We used mark–recapture analyses to model rates of population growth, recruitment, and apparent survival as functions of annual mean temperature and annual precipitation. For over one‐half of focal species, decreasing population growth rates were associated with increasing temperature. Due to the trend in temperature over time, we substituted a time covariate for the temperature covariate in top‐ranked population growth rate models. Temperature was a better explanatory covariate than time for 6 of the 12 species, or 29% of all focal species. Population growth rates were also lower for species found further below their elevational midpoint and for smaller‐bodied species. Changes in population growth rates were more closely tied to changes in recruitment than to changes in apparent survival. There were no consistent associations between demographic rates and precipitation. This study demonstrates temperature‐associated demographic impacts for 6 (29%) of 21 focal species in an Afrotropical understory bird community and highlights the need to incorporate the impacts of climate change on demographic rates into conservation planning across the tropics.

Global Change Biology↗

Pressure waves in a supersaturated bubbly magma

We study the interaction of acoustic pressure waves with an expanding bubbly magma. The expansion of magma is the result of bubble growth during or following magma decompression and leads to two competing processes that affect pressure waves. On the one hand, growth in vesicularity leads to increased damping and decreased wave amplitudes, and on the other hand, a decrease in the effective bulk modulus of the bubbly mixture reduces wave velocity, which in turn, reduces damping and may lead to wave amplification. The additional acoustic energy originates from the chemical energy released during bubble growth. We examine this phenomenon analytically to identify conditions under which amplification of pressure waves is possible. These conditions are further examined numerically to shed light on the frequency and phase dependencies in relation to the interaction of waves and growing bubbles. Amplification is possible at low frequencies and when the growth rate of bubbles reaches an optimum value for which the wave velocity decreases sufficiently to overcome the increased damping of the vesicular material. We examine two amplification phase-dependent effects: (1) a tensile-phase effect in which the inserted wave adds to the process of bubble growth, utilizing the energy associated with the gas overpressure in the bubble and therefore converting a large proportion of this energy into additional acoustic energy, and (2) a compressive-phase effect in which the pressure wave works against the growing bubbles and a large amount of its acoustic energy is dissipated during the first cycle, but later enough energy is gained to amplify the second cycle. These two effects provide additional new possible mechanisms for the amplification phase seen in Long-Period (LP) and Very-Long-Period (VLP) seismic signals originating in magma-filled cracks.

Geophysical Journal International↗

Invasive plants and their ecological strategies: Prediction and explanation of woody plant invasion in New England

Effective management of introduced species requires the early identification of species that pose a significant threat of becoming invasive. To better understand the invasive ecology of species in New England, USA, we compiled a character data set with which to compare non-native species that are known invaders to non-native species that are not currently known to be invasive. In contrast to previous biological trait-based models, we employed a Bayesian hierarchical analysis to identify sets of plant traits associated with invasiveness for each of three growth forms (vines, shrubs, and trees). The resulting models identify a suite of 'invasive traits' highlighting the ecology associated with invasiveness for each of three growth forms. The most effective predictors of invasiveness that emerged from our model were 'invasive elsewhere', 'fast growth rate', 'native latitudinal range', and 'growth form'. The contrast among growth forms was pronounced. For example, 'wind dispersal' was positively correlated with invasiveness in trees, but negatively correlated in shrubs and vines. The predictive model was able to correctly classify invasive plants 67% of the time (22/33), and non-invasive plants 95% of the time (204/215). A number of potential future invasive species in New England that deserve management consideration were identified. ?? 2007 The Authors.

Diversity and Distributions↗

Effects of carbon dioxide exposure on intensively cultured rainbow trout Oncorhynchus mykiss: Physiological responses and fillet attributes

Rainbow trout Oncorhynchus mykiss (261.6 × 24.7 g initial weight, mean × SEM) at 13.1 × 0.2 C were exposed for 94 d to one of three CO 2 treatments: control (22.1 × 2.8 mg/L), medium (34.5 × 3.8 mg/L), or high (48.7 × 4.4 mg/L). Trout were checked daily for survival, and fish were sampled at 0, 28, 56, and 84 d for physiological responses, growth, and fillet quality assessments. Trout were also challenged to a 15‐min crowding stress at 93 d to assess their ability to initiate a stress response during hypercapnia. Chronically exposed trout showed nearly 100% survival through 84 d exposure (1 of 1,500 fish died). Growth and physiological results showed that increasing elevated CO 2 , concentrations result in corresponding decreased growth rates and CO 2 specific physiological parameters: The medium and high CO 2 treatments had significantly slower growth and subsequently smaller fish by 84 d. Exposed trout also showed significantly ( P < 0.05) decreased plasma chloride for medium and high CO 2 treatments compared to the control from 28 through 84 d. Decreased growth and smaller fish in the medium and high CO 2 treatments resulted in correspondingly smaller fresh and smoked fillet weights. Chronic CO 2 exposure did not result in notable changes in ultimate muscle pH. Exposure to 15‐min crowding stress at 93 d resulted in significant changes in hematocrit, plasma cortisol, glucose, and chloride for all treatment groups. CO 2 ‐specific changes were detected in hematocrit, plasma cortisol, and plasma chloride responses following the 15‐min crowding stress.

Journal of the World Aquaculture Society↗

Plant-plant interactions in a subtropical mangrove-to-marsh transition zone: effects of environmental drivers

Questions Does the presence of herbaceous vegetation affect the establishment success of mangrove tree species in the transition zone between subtropical coastal mangrove forests and marshes? How do plant&ndash;plant interactions in this transition zone respond to variation in two primary coastal environmental drivers? Location Subtropical coastal region of the southern United States. Methods We conducted a greenhouse study to better understand how abiotic factors affect plant species interactions in the mangrove-to-marsh transition zone, or ecotone. We manipulated salinity (fresh, brackish or salt water) and hydrologic conditions (continuously saturated or 20-cm tidal range) to simulate ecotonal environments. Propagules of the mangroves Avicennia germinans and Laguncularia racemosa were introduced to mesocosms containing an established marsh community. Both mangrove species were also introduced to containers lacking other vegetation. We monitored mangrove establishment success and survival over 22 mo. Mangrove growth was measured as stem height and above-ground biomass. Stem height, stem density and above-ground biomass of the dominant marsh species were documented. Results Establishment success of A. germinans was reduced under saturated saltwater conditions, but establishment of L. racemosa was not affected by experimental treatments. There was complete mortality of A. germinans in mesocosms under freshwater conditions, and very low survival of L. racemosa . In contrast, survival of both species in monoculture under freshwater conditions exceeded 62%. The marsh species Distichlis spicata and Eleocharis cellulosa suppressed growth of both mangroves throughout the experiment, whereas the mangroves did not affect herbaceous species growth. The magnitude of growth suppression by marsh species varied with environmental conditions; suppression was often higher in saturated compared to tidal conditions, and higher in fresh and salt water compared to brackish water. Conclusions Our results indicate that herbaceous marsh species can suppress mangrove early seedling growth. Depending on species composition and density, marsh plants can slow mangrove landward migration under predicted climate change scenarios as salinity in freshwater and oligohaline wetlands increases with rising sea levels. Change in the relative coverage of mangrove forests and marshes will depend on both the ability of marsh species to migrate further inland as mangroves advance, and the ability of shoreline mangroves to adjust to rising sea level through accretionary processes.

Journal of Vegetation Science↗

Description of an estuarine methylotrophic methanogen, which grows on dimethyl sulfite

Characteristics of an obligately methylotrophic coccoid methanogen (strain GS-16) previously isolated from estuarine sediment are described. Growth was demonstrated on dimethyl sulfide (DMS) or trimethylamine (TMA), but not on methane thiol, methane thiol plus hydrogen, dimethyl disulfide, or methionine. DMS-grown cells were able to metabolize DMS and TMA simultaneously when inoculated into media containing substrate levels of these compounds. However, TMA-grown cells could not metabolize [ 14 C]DMS to 14 CH 4 , although they could convert [ 14 C]methanol to 14 CH 4 . These results suggest that metabolism of DMS proceeds along a somewhat different route than that of TMA and perhaps also that of methanol. The organism exhibited doubling times of 23 and 32 h for growth (25°C) in mineral media on TMA and DMS, respectively. Doubling times were more rapid (∼6 h) when the organisms were grown on TMA in complex broth. In mineral media, the fastest growth on DMS occurred between pH levels of 7.0 and 8.7, at 29°C, and with 0.2 to 0.4 M Na + and 0.04 M Mg 2+ . Somewhat different results occurred for growth on TMA in complex broth. Cells had a moles percent G+C value of 44.5% for their DNA. Growth on DMS, TMA, and methanol yielded stable carbon isotope fractionation factors of 1.044, 1.037, and 1.063, respectively. Fractionation factors for hydrogen were 1.203 (DMS) and 1.183 (TMA).

Applied and Environmental Microbiology↗

Vesiculation of basaltic magma during eruption

Vesicle size distributions in vent lavas from the Pu'u'O'o-Kupaianaha eruption of Kilauea volcano are used to estimate nucleation and growth rates of H 2 O-rich gas bubbles in basaltic magma nearing the earth's surface (≤120 m depth). By using well-constrained estimates for the depth of volatile exsolution and magma ascent rate, nucleation rates of 35.9 events ⋅ cm -3 ⋅ s -1 and growth rates of 3.2 x 10 -4 cm/s are determined directly from size-distribution data. The results are consistent with diffusion-controlled growth as predicted by a parabolic growth law. This empirical approach is not subject to the limitations inherent in classical nucleation and growth theory and provides the first direct measurement of vesiculation kinetics in natural settings. In addition, perturbations in the measured size distributions are used to examine bubble escape, accumulation, and coalescence prior to the eruption of magma.

Geology↗

Annulus formation on scales of four species of coregonids reared under artificial conditions

Scales from known-age coregonids reared in the laboratory were examined to determine when annuli formed and to learn possible factors of their formation. Scales were taken monthly from marked fish for periods up to 21 months. Scales were also examined from fish that died and from preserved specimens of young-of-the-year for each species. Two marks formed on almost all scales each calendar year. The stronger formed during March-April and the weaker in October-November. Both marks had all the usual characteristics of an annulus but the spring mark was considered the annulus and the fall mark an accessory check. The annulus formed during a period of constant temperatures and of little change in growth or increasing growth. The accessory check formed during a period of declining temperatures (1-5 degrees F, or 0.6-2.8 degrees C, per month) and of little change in growth or declining growth. Most fish grew throughout the winter; the only exceptions were one bloater ( Coregonus hoyi ) and several of the largest lake whitefish ( C. clupeaformis ). Fish were always given all the food they would eat to eliminate availability of food as a factor of mark formation. The temperature of the water during the winter (50 ±. 0.3 F; 10.0 ±. 0.2 C) did not arrest metabolic activity. The growth rate was related more closely to day length than to other variables examined.

Journal of the Fisheries Research Board of Canada↗