Limoilou-Amélie Renaud’s research while affiliated with Université du Québec en Abitibi-Témiscamingue and other places

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Publications (13)


A family history: Effects of parental age on offspring life-history traits in bighorn sheep
  • Article

May 2025

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47 Reads

Behavioral Ecology

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Limoilou-Amélie Renaud

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Long-term fitness effects of parental age could affect population dynamics, as age structure can vary considerably over time. Few studies of wild animals, however, have examined the possible long-term fitness consequences of parental age. Based upon 50 years of data on wild bighorn sheep, we investigated how parental age affected three offspring traits – mass at three years, longevity, and female lifetime reproductive success. We also tested for a survival filter which could mask or increase fitness effects of parental age. Our results showed a significant, quadratic negative association between maternal age, offspring longevity and female lifetime reproductive success. Offspring born to mothers aged 5-6 years lived about 2 years longer and weaned about 1.5 lambs more than siblings born to mothers aged 12-13 years. The effect of paternal age was not significant. There was also a positive association between the longevity of mothers and offspring. We did not detect any effects of parental age on mass at three years for offspring of either sex. These results demonstrate the presence of persistent maternal age effects in a long-lived species. The sex-specific effects support the importance of analyzing maternal and paternal age effects separately, as well as effects on female and male offspring. This study advances our understanding of evolutionary processes and population dynamics in wild long-lived mammals.


Figure 1: Portions of the St. Lawrence Estuary and Saguenay River (Quebec, Canada) included in this study and showing spatial age-and sex-segregation in beluga. Locations of Nunavik Inuit subsistence hunts in Hudson Bay and Hudson Strait, Canada, are in inset. Depth data are from Canadian Hydrographic Service.
Figure 2: Blubber progesterone concentration (ng per gram of tissue) in beluga samples from the a) SLE carcasses and b) combined NUN carcasses and SLE biopsies programs. In panel a), colours indicate reproductive status. In panel b), NUN carcasses and SLE biopsies are shown as light and dark grey bars, respectively. Raw progesterone concentrations were log-transformed.
Figure 3: Mixtures of Gaussian distributions used to assign a probability of being pregnant in beluga based on log-transformed blubber progesterone concentration. Panel a) shows distributions for necropsied beluga from the St. Lawrence Estuary, Canada (SLE carcasses) ; panel b) shows beluga harvested in Hudson Bay and Hudson Strait, Canada (NUN carcasses) ; panels c) and d) show free-ranging beluga biopsied in the St. Lawrence Estuary, Canada (SLE biopsies ) with samples respectively clustered into three and two groups. Means, standard deviations of each distribution and their respective 95% lower and upper credibility intervals, are given for each data set in Table A.1. The description of the different reproductive statuses observed in SLE carcasses is shown on Figure 2. Histograms were scaled between 0 and 1 to match the scale of the density plots.
Blubber progesterone concentrations for beluga of known and unknown reproductive status, coming from the St. Lawrence Estuary (SLE carcasses ), the harvest in Hudson Bay and Hudson Strait (NUN carcasses ) and free-ranging beluga from the St. Lawrence Estuary (SLE biopsies )
Estimating pregnancy rate from blubber progesterone levels of a blindly biopsied beluga population poses methodological, analytical and statistical challenges
  • Article
  • Full-text available

September 2023

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100 Reads

Conservation Physiology

L -A Renaud

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[...]

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Beluga (Delphinapterus leucas) from the St. Lawrence Estuary, Canada, have been declining since the early 2000s, suggesting recruitment issues as a result of low fecundity, abnormal abortion rates or poor calf or juvenile survival. Pregnancy is difficult to observe in cetaceans, making the ground truthing of pregnancy estimates in wild individuals challenging. Blubber progesterone concentrations were contrasted among 62 SLE beluga with a known reproductive state (i.e. pregnant, resting, parturient and lactating females), that were found dead in 1997 to 2019. The suitability of a threshold obtained from decaying carcasses to assess reproductive state and pregnancy rate of freshly-dead or free-ranging and blindly-sampled beluga was examined using three statistical approaches and two data sets (135 freshly harvested carcasses in Nunavik, and 65 biopsy-sampled SLE beluga). Progesterone concentrations in decaying carcasses were considerably higher in known-pregnant (mean ± sd: 365 ± 244 ng g−1 of tissue) than resting (3.1 ± 4.5 ng g−1 of tissue) or lactating (38.4 ± 100 ng g−1 of tissue) females. An approach based on statistical mixtures of distributions and a logistic regression were compared to the commonly-used, fixed threshold approach (here, 100 ng g−1) for discriminating pregnant from non-pregnant females. The error rate for classifying individuals of known reproductive status was the lowest for the fixed threshold and logistic regression approaches, but the mixture approach required limited a priori knowledge for clustering individuals of unknown pregnancy status. Mismatches in assignations occurred at lipid content < 10% of sample weight. Our results emphasize the importance of reporting lipid contents and progesterone concentrations in both units (ng g−1 of tissue and ng g−1 of lipid) when sample mass is low. By highlighting ways to circumvent potential biases in field sampling associated with capturability of different segments of a population, this study also enhances the usefulness of the technique for estimating pregnancy rate of free-ranging population.

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Study area for two colonies of harbour seals (Phoca vitulina) in the St. Lawrence Estuary, Québec, Canada. Water depth contours (in meters) are also shown. Coastline downloaded from “rnaturalearth” library version 0.3.2 (Massicotte and South 2023). Depth data are from Fisheries and Oceans Canada (© 2016, projection: NAD83, coordinate system: WGS84).
Preweaning survival of harbour seal (Phoca vitulina) pups in the St. Lawrence Estuary, 1998–2019, at Bic Island and Métis, Québec, Canada. Estimates are from a model with year and site effects on capture probability and preweaning survival. Bars represent 95% credible intervals around posterior means. Data were not available for years 2004–2007 and 2017–2018.
Harbour seal (Phoca vitulina) pup abundance in the St. Lawrence Estuary, 1998–2019, at Bic Island and Métis, Québec, Canada. Estimates are from an annual model with year and site effects on capture probability and preweaning survival. Bars represent 95% credible intervals around posterior means. Data were not available for years 2004–2007 and 2017–2018.
Cormack–Jolly–Seber models used to estimate harbour seal (Phoca vitulina) pup abundance and preweaning survival in the St. Lawrence Estuary, Quebec, Canada, 1998–2019.
Spatiotemporal variation in pup abundance and preweaning survival of harbour seals (Phoca vitulina) in the St. Lawrence Estuary, Canada

June 2023

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100 Reads

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5 Citations

Marine mammal populations worldwide greatly benefitted from conservation measures put in place since the 1970s following overexploitation, and many pinniped populations have recovered. However, threats due to bycatch, interspecific interactions or climate change remain, and detailed knowledge on vital rates, population dynamics, and their responses to environmental changes is essential for efficient management and conservation of wild populations. In this study, we quantified pup abundance and survival of individually marked harbour seal (Phoca vitulina Linnaeus, 1758) pups during the preweaning period at Bic Island and Métis sites in the St. Lawrence Estuary from 1998 to 2019. We used mark-recapture models to evaluate competing hypotheses regarding variation in daily preweaning survival rates and capture probability during the pups’ first 30 days of life. Pup abundance increased from 76 [95% CI: 59, 101] to 323 [95% CI: 233, 338] in the past two decades at Bic Island and from 66 [95% CI: 47, 91] to 285 [95% CI: 204, 318] at Métis. Preweaning survival was generally higher at Bic (0.73 [95% CI: 0.58, 0.82]) than at Métis (0.68 [95% CI: 0.52, 0.79]). We hypothesize that differences between habitats and human disturbance contribute to lower preweaning survival at Métis, but behavioural studies are needed to understand the impacts of disturbance on mother–pup interactions during the nursing period.



Effects of mismatch on bighorn ewe energy allocation

May 2022

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89 Reads

Phenological changes in response to climate change have been documented across taxa throughout the Northern Hemisphere. The rate of advancement often decreases with increasing trophic level, resulting in mismatch between consumers and their resources. The match-mismatch hypothesis has been investigated in bighorn sheep (Ovis canadensis) at Ram Mountain, Alberta (52ºN, 115ºW), a population which has been monitored since 1971. The gestation period of bighorn sheep is 172 ± 2 days, and our team previously showed that ewes have plastically responded to environmental changes in fall by advancing their conception date and giving birth on average 15 days earlier in recent years. However, spring conditions have changed at a much faster rate, resulting in lambs being born an average of 10 days later with respect to spring green-up. Ewes do not seem to pass on costs of mismatch onto lambs, leading us to hypothesize that mismatch may increase maternal reproductive costs. Our goal was to determine whether maternal summer mass gain is affected by increasing mismatch. We use parturition dates recorded since 1992, and spring green-up dates obtained from the Enhanced Vegetation Index available since 2000, to test the effects of phenological mismatch on bighorn ewe summer mass gain. A total of 152 females were measured, between 1-11 times each, with a total of 449 parturition dates recorded since 1992, 279 of which have been recorded since 2000. Mismatch between spring green-up and parturition, (parturition date - green-up date, in Julian days) ranges from -24 days (early parturition) to +120 days (late parturition), and population mismatch, (individual parturition date - population median parturition date, in Julian days), ranges from -18 to +95 days. Preliminary results indicate that ewes with a greater positive mismatch, both green-up and population, gain less mass over summer; those with the greatest mismatch gain up to 6kg less during summer than those whose parturition dates are matched with spring green-up or median parturition date. This suggests that ewes who are missing the highest quality forage have the greatest reproductive cost, using their fat reserves for lactation. Current knowledge of the effects of climate change on phenological plasticity are primarily from studies on birds, which are income breeders and thus more sensitive to daily changes in food abundance. This study improves our understanding of large mammal phenological plasticity, and the potential costs of phenological mismatch, which are expected to increase with climate change.


Temporal trends in autumn and spring phenologies, Ram Mountain, Alberta, Canada, 2000–2017. Spring temperature (a) showed marked interannual variation; thus, a linear trend was not significant from 2000 to 2017. Autumn (b) temperature increased linearly over time. Green‐up date (c) decreased linearly over years, but brown‐down date (d) showed no temporal trend. Together, variation in parturition date (e) and in green‐up date appear to drive interannual variation in mismatch (f), which is directly explained by spring temperature and to a lesser extent, autumn precipitation (Figure 2). In panels (b), (c), (e) and (f), the black line represents a prediction from a generalized additive mixed model. In panel (f), the green, dotted line is a prediction from a linear model with a quadratic effect of year. Dots represent individual values in (e) and (f). Shaded areas around model predictions are 95% CI. The Enhanced Vegetation Index was used to calculate green‐up and brown‐down dates [Colour figure can be viewed at wileyonlinelibrary.com]
Drivers of trophic mismatch in bighorn sheep, Ram Mountain, Alberta, Canada, 2000–2017. The effect of (a) spring temperature and (b) autumn precipitation on individual mismatch between parturition and spring green‐up is illustrated. Lines show predictions from a generalized additive mixed model, with female identity and year as random effects. Grey dots represent raw data. Shaded areas represent 95% CI around the prediction. Partial R² for spring temperature was 21%, and 9% for autumn precipitation once the model was refitted as a linear mixed‐effects model (total R²: 27%)
Individual and population effects of mismatch on lamb weaning mass in bighorn sheep, Alberta, Canada, 2000–2017. The individual‐ (a) and population‐level (b) components of fitness were tested in the same models but are presented separately. The annual population mismatch was defined as the mean of individual mismatch (the difference between the parturition date of each female and the green‐up date) values each year. The individual component of mismatch was calculated by standardizing individual mismatch within years by subtracting year‐specific population mismatch from individual mismatch values. Model estimates are in Table S4, Appendix 3. In (a), the line shows a significant within‐year effect of individual mismatch, accounting for between‐year effects of population mismatch. Shaded areas are 95% confidence intervals. Negative values for mismatch indicate early parturition relative to green‐up. The mismatch is calculated from the Enhanced Vegetation Index
Testing the match–mismatch hypothesis in bighorn sheep in the context of climate change

October 2021

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106 Reads

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13 Citations

In species with long gestation, females commit to reproduction several months before parturition. If cues driving conception date are uncoupled from spring conditions, parturition could be mistimed. Mismatch may increase with global change if the rate of temporal changes in autumn cues differs from the rate of change in spring conditions. Using 17 years of data on climate and vegetation phenology, we show that autumn temperature and precipitation, but not vegetation phenology, explain parturition date in bighorn sheep. Although autumn cues drive the timing of conception, they do not predict conditions at parturition in spring. We calculated the mismatch between individual parturition date and spring green‐up, assessed whether mismatch increased over time and investigated the consequences of mismatch on lamb neonatal survival, weaning mass and overwinter survival. Mismatch fluctuated over time but showed no temporal trend. Temporal changes in green‐up date did not lead to major fitness consequence of mismatch. Detailed data on individually marked animals revealed no effect of mismatch on neonatal or overwinter survival, but lamb weaning mass was negatively affected by mismatch. Capital breeders might be less sensitive to mismatch than income breeders because they are less dependent on daily food acquisition. Herbivores in seasonal environments may access sufficient forage to sustain lactation before or after the spring ‘peak’ green‐up, and partly mitigate the consequences of a mismatch. Thus, the effect of phenological mismatch on fitness may be affected by species life history, highlighting the complexity in quantifying trophic mismatches in the context of climate change.





Density distribution of green-up (on the right; in green in the online version) and snowmelt dates (on the left; in orange in the online version) as extracted from phenology time series, Ram Mountain, Alberta, Canada, 2000–2017. Julian day 150 is equivalent to May 30 in non-leap years
Relationship between maternal covariates and spring phenology for fatty acid concentrations in bighorn sheep, Ram Mountain, Alberta, Canada, 2011–2016 (n = 220). (a) Snowmelt date and (b) length of snow-free season (referred to as snow-free length) was tested as an explanatory variable in the multivariate model. Estimates are effect sizes of fixed effects for fatty acid concentration. Sampling year, mother’s identity and a sample-level factor were included as random effects. The horizontal line represents the 95% credibility intervals and the grey dot, the mean. The vertical grey line is drawn on zero for reference
Final path diagrams of causal effects between phenology, variation in milk composition, and bighorn lamb mass and survival at Ram Mountain, Alberta, Canada, 2011–2016 (n = 65). We first tested (a) the absence of effects on survival and (b) a direct effect of milk energy and maternal summer mass gain on lamb survival. Arrows indicate paths included in the models. Solid lines indicate statistically significant effects of a variable on another. Dashed arrows indicate not statistically significant paths included in a model. Standardized path coefficients and 95% confidence intervals are indicated directly on corresponding paths
Milk composition in a wild mammal: a physiological signature of phenological changes

June 2020

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159 Reads

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4 Citations

Oecologia

Understanding how spring phenology influences early life can provide important insights into drivers of future development and survival. We combined unique, long-term data from a bighorn sheep population and satellite-derived phenology indices to quantify the relative importance of maternal and environmental influences on milk composition and lamb overwinter survival. Based on 216 milk samples from 34 females monitored over 6 years, we found that longer snow-free and vegetation growing seasons increased milk fatty acid, iron and lactose concentrations. Structural equation modelling revealed no causality between milk energy content, lamb weaning mass and lamb overwinter survival. Our results suggest that spring conditions can affect milk energy content, but we did not detect any effect on lamb overwinter survival either directly or indirectly through lamb weaning mass. The effect of green-up date on milk composition and energy content suggests that herbivores living in seasonal environments, such as the bighorn sheep, might rely on a strategy intermediate between ‘capital’ and ‘income’ breeding when energy demands are high.


Citations (5)


... The study area consisted of 2 breeding colonies: Bic Island (40 km 2 ; 48°24' N, 68°51' W) and Métis area (10 km 2 ; 48°41' N, 68°01' W), located in the St. Lawrence Estuary, Canada (Dubé et al. 2003, Re naud et al. 2023. The population size of harbour seals in the St. Lawrence Estuary has been increasing since 2000 (Renaud et al. 2023). These 2 colonies have been monitored for 19 yr between 1998 and 2023. ...

Reference:

Neonatal growth patterns, sexual dimorphism, and compensatory growth in harbour seal pups
Spatiotemporal variation in pup abundance and preweaning survival of harbour seals (Phoca vitulina) in the St. Lawrence Estuary, Canada

... Failure to adjust birth phenology can potentially lead to a mismatch between the time offspring are born and the peak in food availability (Visser et al. 1998, Shipley et al. 2020, Renaud et al. 2022. Trophic mismatches are important to evaluate since they can have a critical impact on an individual's fitness as well as demography, with examples across a wide range of terrestrial and marine species (Miller-Rushing et al. 2010, Visser andGienapp 2019). ...

Testing the match–mismatch hypothesis in bighorn sheep in the context of climate change

... For instance, in alpine environments severe winter weather can affect adult and juvenile survival because cold and snowy winters increase thermoregulatory and traveling costs and reduce access to vegetation (Rupicapra rupicapra; Willisch et al. 2013). Spring conditions are also crucial because high-quality vegetation is accessible only over a brief period, which limits fetal growth and milk production (Renaud et al. 2020), influencing juvenile growth and survival (Portier et al. 1998;Côté and Festa-Bianchet 2001). Furthermore, interactions between intraspecific resource competition and environmental conditions are reported in several studies of large herbivore population dynamics (Portier et al. 1998;Bonenfant et al. 2009). ...

Milk composition in a wild mammal: a physiological signature of phenological changes

Oecologia

... In seasonal environments, animals time high energy-demanding events in their reproductive cycle, such as parturition and rearing, to the seasonal peak of abundance in high-quality food [8][9][10], using food availability, temperature and photoperiod as cues [11][12][13]. However, knowledge about adaptive responses is surprisingly elusive, especially for long-living mammals, such as ungulates [14,15]. Red deer, bighorn sheep and reindeer have shown trends towards earlier births in recent years, but the extent to which these changes result from reduced gestation length and earlier conception varies among species and is not yet fully understood [14][15][16][17][18][19]. ...

Phenotypic plasticity in bighorn sheep reproductive phenology: from individual to population

Behavioral Ecology and Sociobiology

... Since bite size and bite rate are negatively correlated (Denryter, 2020;Shipley, 2007), the decrease in bite rate with Julian date could also be partially explained by an increase in vegetation length and bite size. The absence of an interaction between lactation and Julian date also suggests that despite being partly capital breeders J€ onsson, 1997), bighorn sheep adjust their behaviour to use income resources for maternal expenditure (Renaud et al., 2019) and that individual differences in resource acquisition are linked to high resource allocation to lactation (Monteith et al., 2013;Moquin et al., 2010;Stephenson et al., 2020). We found that rumination effort showed strong variation in response to individual differences in yearly mass change. ...

Causes and short‐term consequences of variation in milk composition in wild sheep