ArticlePublisher preview available

Temperature and the maturation of fish: a simple sine-wave model for predicting accelerated spring spawning

Authors:
To read the full-text of this research, you can request a copy directly from the authors.

Abstract and Figures

Global warming affects the phenology of the Earth’s flora and fauna, notably by advancing the date at which many plants and animals tend to reproduce. We use fish, where this reproductive acceleration is well-documented, to present a simple approach based on sine curves to predict, in spring spawning fish, the minimum number of days (Δdmin) that spawning is advanced as the result of a given increase in water temperature (Δ°C). We show, via comparison with field estimates, that our simple model’s robust predictions correspond to observed values of Δdmin/Δ°C, and discuss both the potential uses and the limitations of the model.
This content is subject to copyright. Terms and conditions apply.
Vol.: (0123456789)
1 3
https://doi.org/10.1007/s10641-022-01212-0
Temperature andthematuration offish: asimple sine‑wave
model forpredicting accelerated spring spawning
DanielPauly · CuiLiang
Received: 28 April 2021 / Accepted: 7 January 2022
© The Author(s), under exclusive licence to Springer Nature B.V. 2022
Keywords Spring spawning in fish· Shifts in
phenology· Temperature differences· Size at first
maturity· Triggers for maturation· Amplitude of
seasonal oscillations
Introduction
Global change, or more precisely, global warming,
has modified much of the Earth’s flora and fauna’s
reproductive phenology and can be expected to con-
tinue doing so for decades. These modifications take
various forms, including earlier flowering in some
trees and earlier migrations in birds (see, e.g., Ahas
and Aasa 2006; Cohen etal. 2018), many due to sinu-
soid seasonal temperature oscillations with a higher
annual mean (Fig. 1). In the past few decades, the
temperatures of marine ecosystems have increased
(Belkin 2009), as they have in freshwater bodies (Car-
penter et al. 1992; Kangur et al. 2021). Increasing
temperatures, on the other hand, will alter the physi-
ology of fish, and particularly their reproduction, as
can be expected in ectotherms (Alix etal. 2020).
The fact that ocean and freshwater warming
is modifying the phenology of maturation and
spawning in fishes, i.e., the timing of their reach-
ing puberty, has been widely reported. A meta-
analysis of observed phenological shifts suggested
that seasonal events of marine species advanced by
an average of 4.4 days per decade during the late
twentieth century (Poloczanska etal. 2013). Also,
Abstract Global warming affects the phenology of
the Earth’s flora and fauna, notably by advancing the
date at which many plants and animals tend to repro-
duce. We use fish, where this reproductive accelera-
tion is well-documented, to present a simple approach
based on sine curves to predict, in spring spawn-
ing fish, the minimum number of days (Δdmin) that
spawning is advanced as the result of a given increase
in water temperature (Δ°C). We show, via comparison
with field estimates, that our simple model’s robust
predictions correspond to observed values of Δdmin/
Δ°C, and discuss both the potential uses and the limi-
tations of the model.
Supplementary information The online version
contains supplementary material available at https:// doi.
org/ 10. 1007/ s10641- 022- 01212-0.
D.Pauly(*)
Institute fortheOceans andFisheries, Sea Around Us,
University ofBritish Columbia, Vancouver, BC, Canada
e-mail: d.pauly@oceans.ubc.ca
C.Liang
CAS Key Laboratory ofMarine Ecology
andEnvironmental Sciences, Institute ofOceanology,
Chinese Academy ofSciences, Qingdao266071,
People’sRepublicofChina
e-mail: liangc@qdio.ac.cn
C.Liang
Laboratory forMarine Ecology andEnvironmental
Science, Qingdao National Laboratory forMarine
Science andTechnology, Qingdao266237,
People’sRepublicofChina
/ Published online: 8 February 2022
Environ Biol Fish (2022) 105:1481–1487
Content courtesy of Springer Nature, terms of use apply. Rights reserved.
... Changes in the oceanographic conditions suitable for spawning Areas that are most suitable for spawning can change along two margins. On the one hand, areas that have historically been suitable may become suitable for spawning earlier in the season 32,33 . On the other hand, areas that have historically not been suitable for spawning may become suitable for spawning with climate change 27,28 . ...
Article
Full-text available
Climate change is associated with altered oceanographic conditions that tend to shift the geographical distributions of fish. To assess the impact of climate change on fisheries, one must go beyond projections of catch potential and understand how fishers respond to moving target species. Many previous studies have explicitly or implicitly assumed that fishers follow fish that are displaced by climate warming. Here, we evaluate this assumption by analyzing a long-term, large-scale yet high-resolution dataset combined with a detailed oceanographic model. Our study case is the Atlantic cod ( Gadhus morhua ) fishery in Norway, one of the largest whitefish fisheries in the world, with little technological or judicial constraints on the potential spatial response of fishers. An oceanographic model is used to predict the areas that have been suitable for Atlantic cod spawning over the two last decades. We compare whether these areas overlap with actually observed fishing locations. While the areas that are suitable for spawning clearly predict how much fish are caught per trip, the suitability of an area does not predict how many vessels fish in a given area at a given point in time. In contrast, the number of vessels in the previous week and the previous year explain the current number of vessels in that area. Hence, future projections of climate change effects should account for the rich and nuanced behavioral responses of humans to project climate change effects on fisheries.
... Notable was the predicted increase for pelagic and demersal species that benefit from the permanent spatial fishing bans in purse and boat seining and trawling, as well as the seasonal ban of small-scale coastal vessels. The proposed ban of small-scale coastal fishing in MPA 5 is in line with the 2-4 month spawning period of most Mediterranean fishes that spans from April to August (Tsikliras et al., 2010) and that is expected to shift earlier due to ocean warming and resulting elevated water temperatures (Pauly and Liang 2022). The total boat seining ban suggested in scenario MPA 5 is based on research indicating it is a less sustainable, non-selective gear causing fish abundance reductions over Posidonia beds (Kalogirou et al., 2010;Vieira et al., 2020). ...
Article
Spatiotemporal simulation modeling is used in the context of ecosystem-based fisheries management to investigate different management options, including the size and allocation of marine protected and fisheries restricted areas. Here, we used ECOSPACE to assess the effectiveness of existing and potential future spatiotemporal fishing restrictions in the heavily exploited Thermaikos Gulf, Greece for the years 2000-2025 (calibration period 2000-2016; projection period 2017-2025). ECOSPACE combines temporal biomass and commercial catch data with spatial habitat and other environmental data, as well as species ecological preferences, feeding, and dispersal rates to depict changes in trophic interactions, biomasses, and commercial catches in time and space. ECOSPACE simulations supported the empirical data demonstrating that fisheries restricted areas are effective tools for rebuilding the biomass of exploited stocks, with their size and location playing a significant role in the way that different organisms respond to protection. Nevertheless, our results suggested that in order to achieve the highest benefits of protection, fisheries restricted areas would need to be accompanied by a parallel reduction in total fishing effort, rather than a redistribution of fishing activities. Such redistribution would just move the pressure on the boundaries of protected areas, causing a local increase of commercial catches owing to the beneficial spillover effects of protection. One of the tested spatiotemporal restriction scenarios (MPA 5) suggests certain additional management measures on top of the existing restrictions for all four fishing fleets operating in the area. This scenario predicted a considerable increase in the biomass of key commercial and vulnerable species groups, including hake, flatfishes, anglerfish, sharks, and rays and skates, by the end of the simulation period in 2025.
... The climatic changes observed in recent years have increased interest in monitoring studies, which make it possible to identify the peculiarities of the development of marine organisms and predict trends in their change. There are publications on the said problem (Akyol et al. 2000;Özbilgin et al. 2004;Güroy et al. 2007;Cherif et al. 2007; Karadurmuş 2012, 2013;Arslan and İşmen 2014;Kutsyn 2021, Pauly andLiang 2022), but the study of changes in GSI over a period of more than five years in warm-water fish species -Mediterranean horse mackerel, M. barbatus ponticus living in the seas of the Mediterranean basin, taking into account the trend of increasing sea surface temperature, has not been studied enough. ...
Article
Full-text available
The influence of the increase in Sea Surface Temperature (SST) over the period 2016–2021 on the reproductive system of warm-water fishes: Blunt-snouted mullet and Mediterranean horse mackerel are considered. It is shown that in the period from 2016 to 2021 in the Mediterranean basin in the Black Sea and the Marmara Sea, the tendency to increase the annual average temperature is higher than in the Aegean Sea and the Mediterranean Sea. It was found that in the North-Western part of the Black Sea, the temperature increase for a year and for five years is the highest compared to other considered areas of the Mediterranean basin. The analysis showed an influence of the trend of increasing the water temperature of the seas of the Mediterranean basin on the reproductive system of warm-water fish species. Thus, in the Aegean Sea and Mediterranean Sea, during the study period, there was a shift in the timing of spawning from the summer months to the spring months, with the highest GSI value in April-May. In the Marmara Sea and Black Sea, spawning period remains the same from April to September, but the highest GSI values have shifted from July-August to June.
... ideas on protein denaturation, efficiency of assimilation, etc.), particularly those that have come to light as it has evolved since the 1980s-that remain to be evaluated (Pauly, 1981(Pauly, , 2010(Pauly, , 2021. For example, the GOLT was recently expanded to encompass predictions surrounding spawning and maturation (Morbey & Pauly, 2022;Pauly, 2022;Pauly & Liang, 2022a, 2022b. In addition, there remains much to explore regarding oxygen limitation more broadly, as well as the role of oxygen in shaping body size and other life history traits (Atkinson, 1995;Audzijonyte et al., 2019;Pauly, 1981Pauly, , 2010Pauly, , 2021. ...
Article
Full-text available
Life history theory suggests that maximum size and growth evolve to maximize fitness. In contrast, the Gill Oxygen Limitation Theory (GOLT) suggests that growth and maximum size in fishes and other aquatic, water‐breathing organisms is constrained by the body mass‐scaling of gill surface area. Here, we use new data and a novel phylogenetic Bayesian multilevel modelling framework to test this idea by asking the three questions posed by the GOLT regarding maximum size, growth and gills. Across fishes, we ask whether the body mass‐scaling of gill surface area explains (1) variation in the von Bertalanffy growth coefficient ( k ) above and beyond that explained by asymptomatic size ( W ∞ ), (2) variation in growth performance (a trait that integrates the tradeoff between k and W ∞ ) and (3) more variation in growth performance compared to activity (as approximated by caudal fin aspect ratio). Overall, we find that there is only a weak relationship among maximum size, growth and gill surface area across species. Indeed, the body mass‐scaling of gill surface area does not explain much variation in k (especially for those species that reach the same W ∞ ) or growth performance. Activity explained three to five times more variation in growth performance compared to gill surface area. Our results suggest that in fishes, gill surface area is not the only factor that explains variation in maximum size and growth, and that other covariates (e.g. activity) are likely important in understanding how growth, maximum size and other life history traits vary across species.
... maximum size, timing of maturation and reproduction, spawning phenology, geographic distributions, activity level) and those based more on physiological processes (e.g. food consumption and conversion efficiency, the balance of oxidative versus glycolytic enzymes; Pauly, 1981Pauly, , 2010Pauly, , 2021Pauly & Liang, 2022). Although originally proposed in the early 1980s, interest in the GOLT has experienced a recent resurgence in light of research that aims to predict how species will respond to continued environmental change, particularly warming temperatures and shifting oxygen availability (Cheung et al., 2013;Lefevre et al., 2017Lefevre et al., , 2018Seibel & Deutsch, 2020). ...
Article
Full-text available
The Gill Oxygen Limitation Theory (GOLT) posits that a mismatch in oxygen supply and demand stemming from geometric constraints on gill surface area limits metabolic rate and energy available for biological processes. This theory has been suggested to explain numerous phenomena observed with warming yet is based upon a relationship among maximum size, growth, and gill surface area established over 40 years ago. However, the metric used in this relationship to characterize gill surface area, gill area index, fails to capture the known variability in the scaling of gill surface area and is biased by the sizes at which gills were measured. Here, we revisit a central prediction of the GOLT, asking four key questions that examine limitations in the original relationship. We find that gill area index does indeed explain variation in growth performance across 132 species of fish and this relationship is strikingly similar to the original relationship across 42 species. Yet, we argue that gill area index is not an adequate measure of gill surface area because (1) it has a non‐linear relationship with size and, thus, changes ontogenetically as an individual grows over time and (2) because it is based on mean estimates of both gill surface area and body mass. Indeed, we show that the value of gill area index for a given species is variable depending on how it is calculated. We therefore suggest a pathway forward for assessing whether gill surface area is an important factor in explaining variation in growth performance.
... Other crustaceans, particularly in warm-temperate or tropical areas, spawn throughout the year and in some cases repeatedly during a spawning season (Restrepo & Watson, 1991). Repeated in-season spawning in contrast to one spawning season per year also occur in fishes and are consistent with predictions of the GOLT (see Pauly, 2019;Pauly & Liang, 2022) and those arguments can potentially be extended to crustaceans. Limitations imposed by the GOLT may also explain the remarkable feature that female crustaceans, ranging from the smallest copepods to the largest lobsters, carry fertilized eggs outside their bodies. ...
Article
Gill surface area (S) and respiration (R) in juvenile and adult crustaceans scale with their body weight (W) such that S ∝ R ∝ Wd, with d ranging mostly between 0.6 and 0.9, but always <1, as in other water-breathing ectotherms (WBE). The growth of adult crustaceans therefore approaches an asymptote, whether or not seasonal growth oscillations are explicitly considered in the model used to describe that growth (e.g., von Bertalanffy growth function). On the other hand, the variation in asymptotic size (L∞ or W∞) among crustaceans is primarily determined by water temperature, which impacts on the oxygen requirements of WBE. Through multiple examples, this and related aspects of the Gill-Oxygen Limitation Theory (GOLT), first developed for fishes and later extended to other WBE, are shown to also apply to the growth of a wide range of crustacean taxa. The GOLT also explains certain aspects of crustacean reproduction, such as the relationship between size at first maturity and maximum size, and, possibly, the feature that female crustaceans hold their eggs outside of their bodies instead of internally.
Article
Full-text available
The annual entrainment of early life history stages of fishes (ELH) into the cooling system of the largest power plant in the Vistula River basin was assessed using passive capture techniques and DNA metabarcoding. Hydrological and thermal conditions during the 2022 breeding season were also analyzed. A total of 5,011 ELH individuals identified to four families and 23 species were recorded. ELH densities upstream of the power plant were an order of magnitude higher than they were downstream. Along with the phenology of occurrence and species composition of drift, this indicated very high (probably close to 100%) ELH mortality during passage through the cooling system. Taking into account low water discharge at constantly high water intake during the season, the absolute estimate of ELH individuals entrained into the cooling system was > 103 × 106. The power plant entrains a notable proportion of one-quarter to one-half of the total ELH drifting by, especially during low-flow periods. Together with the very high abundance of two invasive Ponto-Caspian gobies, the magnitude of the entrainment of ELH into the Kozienice Power Plant presents a truly alarming picture.
Article
Full-text available
Among fishery biologists and even ichthyologists, maturation and spawning of fish are viewed as processes that use “energy” that would otherwise be applied to somatic growth, which is supposed to explain why post-maturity growth in length tends to decline. This widespread conceptualization may be called the “reproductive drain hypothesis” (RDH). However, when growth is correctly viewed as involving body mass, and is thus expressed in weight, post-maturity turns out (in iteroparous bony fish whose maximum length exceeds 10 cm) to accelerate after first maturity, despite its energy cost. This, and other common observations flatly contradict the RDH, and the time has come to withdraw this hypothesis. As a contribution towards this task, we propose an alternative reconceptualization of fish spawning consistent with what is known about fish biology.
Article
Full-text available
This account serves as the introduction to a Special Issue of the journal Environmental Biology of Fishes. It includes 18 contributions, 12 of which deal predominantly with warming, four explicitly with deoxygenation, one providing the framework, and one addressing how warming and deoxygenation—which all contributions view, implicitly or explicitly, as leading to “oxythermic” stress—are used as a pretext to cover up overfishing. The “Mean Temperature of the Catch” (MTC) concept of Cheung et al. (2013, Nature 497:365–368) inspired five of the studies involving mainly temperature, including its first applications to fresh water and to the past, i.e., to the past 130, 7–8 thousand, and 2 million years. Four contributions, jointly representing 4200 + populations and 1100 + species, deal with the effect of temperature on the maximum and/or the asymptotic length of fish behaving as predicted by the Gill-Oxygen Limitation Theory (GOLT). This theory is also evoked in one of these studies to explain how cold denaturation causes fish to grow to a smaller size when temperatures decline below 4 °C. These contributions, which are here summarized and whose conceptual affinities are also presented in graphic form as a tree-like structure, provide a basis for understanding the changes in fish community composition and size structure resulting from marine or freshwater warming. Jointly, they explain some of the changes in fish behavior and position in the water column resulting from the deoxygenation of their habitats.
Article
Full-text available
This contribution applies the “mean tem-perature of the catch” (MTC) concept of Cheung et al. (Nature 497:365–368, 2013) to fish catch data for Lake Peipsi, Estonia/Russia, covering the years 1931 to 2019. The preferred temperature of each of the ten target fish species was obtained from the literature, and combined with the species-specific catch data to obtain MTC values for each year. The analysis of the MTC time series thus obtained with a segmented regression yielded two trend lines, one horizontal at 14.5 °C (1931–1986), and the other (1987–2019) ascending with a slope 0.85 °C·decade−1. Overall, the segmented regression model explains over half of the variance of the MTC data set (multiple R2 = 0.53; adjusted R2 = 0.51). Lake surface water temperatures correlate with MTC, even though weakly (r = 0.30), when considering a 2-year time lag. The fish community of the shallow Lake Peipsi reacts more strongly to temperature changes than marine ecosystems so far studied using the MTC.
Article
Full-text available
Fish generally mature of a smaller fraction of their maximum sizes than birds and mammals. In farmed tilapia (Family Cichlidae), which can tolerate very adverse conditions, the stunting caused by these conditions also cause them to spawn at very small sizes. Such spawning at small sizes (or 'early spawning') is usually perceived as a unique feature of tilapia. There are many reports in the literature that stressful environmental conditions, as occur in small freshwater reservoirs compared with larger ones, and in smaller tilapia aquaculture ponds, reduce both the maximal size that cichlid fish can reach and their size at first maturity. The mechanism that explains how stressful environmental conditions tend to reduce the maximum size that fish can reach, is very general and should apply to all fish. However, not all fish species are equally hardy, and most fish do not survive in stunted or dwarf form under stressful environmental conditions. Tilapia, and other cichids, on the other hand, can handle stressful conditions, if by remaining stunted. The present study shows that tilapia and other cichlids do not spawn 'earlier' than other teleosts. Rather, they are uncommonly tolerant of stressful environmental conditions that however, elevate their metabolism. By reducing their growth and the maximum size they can reach ('stunting'), also reduce the sizes at which their maturity is initiated ('early spawning'). This corroborates the Gill Oxygen Limitation Theory (GOLT), which identifies spawning as an event, rather than a determinant, of fish growth.
Article
Full-text available
The gill-oxygen limitation theory (GOLT) provides mechanisms for key aspects of the biology (food conversion efficiency, growth and its response to temperature, the timing of maturation, and others) of water-breathing ectotherms (WBEs). The GOLT’s basic tenet is that the surface area of the gills or other respiratory surfaces of WBE cannot, as two-dimensional structures, supply them with sufficient oxygen to keep up with the growth of their three-dimensional bodies. Thus, a lower relative oxygen supply induces sexual maturation, and later a slowing and cessation of growth, along with an increase of physiological processes relying on glycolytic enzymes and a declining role of oxidative enzymes. Because the “dimensional tension” underlying this argument is widely misunderstood, emphasis is given to a detailed refutation of objections to the GOLT. This theory still needs to be put on a solid quantitative basis, which will occur after the misconceptions surrounding it are put to rest.
Article
Full-text available
Substantial interannual variability in marine fish recruitment (i.e., the number of young fish entering a fishery each year) has been hypothesized to be related to whether the timing of fish spawning matches that of seasonal plankton blooms. Environmental processes that control the phenology of blooms, such as stratification, may differ from those that influence fish spawning, such as temperature-linked reproductive maturation. These different controlling mechanisms could cause the timing of these events to diverge under climate change with negative consequences for fisheries. We use an earth system model to examine the impact of a high-emissions, climate-warming scenario (RCP8.5) on the future spawning time of two classes of temperate, epipelagic fishes: "geographic spawners" whose spawning grounds are defined by fixed geographic features (e.g., rivers, estuaries, reefs) and "environmental spawners" whose spawning grounds move responding to variations in environmental properties , such as temperature. By the century's end, our results indicate that projections of increased stratification cause spring and summer phytoplankton blooms to start 16 days earlier on average (±0.05 days SE) at latitudes >40°N. The temperature-linked phenology of geographic spawners changes at a rate twice as fast as phyto-plankton, causing these fishes to spawn before the bloom starts across >85% of this region. "Extreme events," defined here as seasonal mismatches >30 days that could lead to fish recruitment failure, increase 10-fold for geographic spawners in many areas under the RCP8.5 scenario. Mismatches between environmental spawners and phytoplankton were smaller and less widespread, although sizable mismatches still emerged in some regions. This indicates that range shifts undertaken by environmental spawners may increase the resiliency of fishes to climate change impacts associated with phenological mismatches, potentially buffering against declines in larval fish survival, recruitment, and fisheries. Our model results are supported by empirical evidence from ecosystems with multidecadal observations of both fish and phytoplankton phenology. K E Y W O R D S earth system model, marine fish reproduction, phenology, phytoplankton blooms, species range shifts, trophic mismatches
Article
Full-text available
Shifts in phenology are already resulting in disruptions to the timing of migration and breeding, and asynchronies between interacting species1–5. Recent syntheses have concluded that trophic level¹, latitude⁶ and how phenological responses are measured⁷ are key to determining the strength of phenological responses to climate change. However, researchers still lack a comprehensive framework that can predict responses to climate change globally and across diverse taxa. Here, we synthesize hundreds of published time series of animal phenology from across the planet to show that temperature primarily drives phenological responses at mid-latitudes, with precipitation becoming important at lower latitudes, probably reflecting factors that drive seasonality in each region. Phylogeny and body size are associated with the strength of phenological shifts, suggesting emerging asynchronies between interacting species that differ in body size, such as hosts and parasites and predators and prey. Finally, although there are many compelling biological explanations for spring phenological delays, some examples of delays are associated with short annual records that are prone to sampling error. Our findings arm biologists with predictions concerning which climatic variables and organismal traits drive phenological shifts.
Article
The standard response to the question “why do fish reach first maturity when they do” is that, at some point (or age), they perceive environmental stimuli, which are converted via the pituitary and the hypothalamus into triggers for a hormonal cascade leading to gonadal maturation and the release of gametes. Yet, the question rarely asked is why fully formed young fish do not respond to the environmental stimuli that the adults react to by maturing and spawning. This question requires an answer, from ichthyologists and/or physiologists, e.g., in the form of a heuristic that individual fish can use, even if the explanation provided here (elaborating on a causal mechanism for the juvenile-to-adult proposed by the author in 1984) should be considered inadequate. This article is protected by copyright. All rights reserved.
Article
The slowing of growth as fish age has long been believed to be related to energy expenditure for maturation, and this rationalization has been used to explain why, across nearly all fish species, the relationship between size at first maturity (Lm ) and maximum (Lmax ) or asymptotic length (L∞ ) is relatively constant. In contrast, the Gill-Oxygen Limitation Theory (GOLT) postulates that (1) fish growth slows because as they grow, their two dimensional ability to extract oxygen from the water diminishes relative to their three dimensional weight gain, and (2) they can only invest energy for maturation if oxygen supply at their size at first maturity (Qm ) exceeds that needed for maintenance metabolism (Q∞ ). It has been reported previously across dozens of marine fish species that the relationship between Qm vs. Q∞ is linear and further, it can be mathematically converted to Lm vs. L∞ by raising both terms to the power of D (the gill surface factor), resulting in a slope of 1.36. If the GOLT is universal, a similar slope should exist for LmD vs. L∞D relationships for freshwater species across multiple individual populations that reside in disparate habitats, although to our knowledge this has never been evaluated. For analysis, we used existing data from previous studies conducted on 51 stream-dwelling populations of redband trout Oncorhynchus mykiss gairdneri, Yellowstone cutthroat trout O. clarkii bouvieri, and mountain whitefish Prosopium williamsoni. Resulting LmD vs. L∞D slopes combining all data points (1.35) or for all species considered separately (range = 1.29-1.40) were indeed equivalent to the slope originally produced for the marine species from which the GOLT-derived relationship was first reported. We briefly discuss select papers both supporting and resisting various aspects of the GOLT, note that it could potentially explain shrinking sizes of marine fish, and call for more concerted research efforts combining laboratory and field expertise in fish growth research. This article is protected by copyright. All rights reserved.
Article
Shifts in phenology are a well‐documented ecological response to changes in climate, which may or may not be adaptive for a species depending on the climate sensitivity of other ecosystem processes. Furthermore, phenology may be affected by factors in addition to climate, which may accentuate or dampen climate‐driven phenological responses. In this study, we investigate how climate and population demographic structure jointly affect spawning phenology of a fish species of major commercial importance: walleye pollock (Gadus chalcogrammus). We use 32 years of data from ichthyoplankton surveys to reconstruct timing of pollock reproduction in the Gulf of Alaska and find that the mean date of spawning has varied by over 3 weeks throughout the last >3 decades. Climate clearly drives variation in spawn timing, with warmer temperatures leading to an earlier and more protracted spawning period, consistent with expectations of advanced spring phenology under warming. However, the effects of temperature were nonlinear, such that additional warming above a threshold value had no additional effect on phenology. Population demographics were equally as important as temperature: An older and more age‐diverse spawning stock tended to spawn earlier and over a longer duration than a younger stock. Our models suggest that demographic shifts associated with sustainable harvest rates could shift the mean spawning date 7 days later and shorten the spawning season by 9 days relative to an unfished population, independent of thermal conditions. Projections under climate change suggest that spawn timing will become more stable for walleye pollock in the future, but it is unknown what the consequences of this stabilization will be for the synchrony of first‐feeding larvae with production of zooplankton prey in spring. With ongoing warming in the world’s oceans, knowledge of the mechanisms underlying reproductive phenology can improve our ability to monitor and manage species under changing climate conditions.
Article
Warming temperatures caused by climate change have the potential to impact spawning phenology of temperate marine fish as some species have temperature-dependent gonadal development. Inter-annual variation in the timing of Atlantic cod (Gadus morhua) spawning in the northern North Sea, central North Sea and Irish Sea was estimated by calculating an annual peak roe month (PRM) from records of roe landings spanning the last three decades. A trend towards earlier PRM was found in all three regions, with estimates of shifts in PRM ranging from 0.9 to 2.4 weeks per decade. Temperatures experienced by cod during early vitellogenesis correlated negatively with PRM, suggesting that rising sea temperatures have contributed to a shift in spawning phenology. A concurrent reduction in the mean size of spawning females excluded the possibility that earlier spawning was due to a shift in size structure towards larger individuals, as large cod spawn earlier than smaller-sized individuals in the North Sea. Further research into the effects of climate change on the phenology of different trophic levels within the North Sea ecosystem should be undertaken to determine whether climate change-induced shifts in spawning phenology will result in a temporal mismatch between cod larvae and their planktonic prey. © International Council for the Exploration of the Sea 2017. All rights reserved.