Dependence of recruitment on parent stock of the spiny lobster,Panulirus argus,in Florida

Despite management practices to achieve sustainability, commercial landings for Florida spiny lobster (Panulirus argus) have experienced a drastic decline (57%) since 2000. This is cause for concern not only for economic reasons, but for issues of sustainability. An annual index of P. argus post‐larval (puerulus) abundance, estimated with a generalized linear model with significant mean sea‐level effects, shows a 36% decrease in annual puerulus supply since 1988. In addition, local Florida spawning stock biomass estimated from an age‐structured sequential population analysis decreased 57% since 1988. Puerulus abundance follows a highly correlated (R = 0.76) trend with a 12‐month delayed spawning stock abundance, which supports the contention that the Florida spawning population is a significant contributing factor to post‐larval recruitment in Florida. Residuals about the puerulus on spawning stock abundance function follow closely an interannual North Atlantic Oscillation Index signal. This residual effect is thought of as a secondary regional population effect on Florida puerulus recruitment. The Florida spiny lobster stock is exploited with no fishing mortality controls due to the Pan Caribbean recruitment concept adopted in Florida spiny lobster management. Therefore, the potential of recruitment overfishing exists if fishing mortality controls to protect local spawning stock abundance, such as catch quotas, are not introduced.     

Dual effects of reversed winter–spring temperatures on year‐to‐year variation in the recruitment of chub mackerel (Scomber japonicus)

To clarify the effects of temperature on the recruitment of chub mackerel (Scomber japonicus) in the North Pacific, we investigated the influence of winter surface temperature (WST) on spawners at the time of maturity around the spawning grounds and the influence of ambient spring temperature on larvae using estimated temperature (ET) obtained from particle tracking experiments. We found a significant positive correlation between ET approximately 10 days following hatching and the recruitment per spawning stock biomass (RPS) after 2000. The closer (more meandering) the Kuroshio Current (KC) axis was in relation to the spawning ground, the higher (lower) the spring surface temperature and the higher (lower) RPS was in the spawning ground. In contrast, WST inside KC near the maturity/spawning ground was significantly negatively correlated with RPS. A significant negative correlation between the temperatures in winter and spring was detected in the area after 2000, when the conditions of the Pacific decadal oscillation index and the stability of the Kuroshio Extension were synchronous, indicating that KC shifted northward during this time. The reversed temperature pattern was consistent with the winter–spring movement of KC axis in the offshore direction and was correlated with the winter–spring difference in the intensity of the Aleutian low. These results suggest that the annual variation in chub mackerel recruitment after 2000 was strongly affected by the combined effects of ambient temperature because of the reversal of conditions that occurred between winter and spring around the maturity/spawning ground, which was related to the KC path.     

Implications of using small meshed gillnets for the sustainability of fish populations: a theoretical exploration based on three case studies

The study explores the impacts of varying gillnet mesh size and fishing level on yield per recruit (Y/R), escapement spawning stock (ESS) and mega‐spawners (MS) of three widely distributed freshwater fishery target species. Y/R is maximised when the optimal length of capture (L_cap) is above the size of maturity (L₅₀). However, the unimodal shape of gillnet selectivity results in lower impacts to ESS and MS with both smaller and larger mesh sizes. Under conditions of moderate exploitation, the fraction of MS was significantly larger if small meshed gillnets were used. This is due to the relatively smaller cumulative vulnerability from small mesh sizes through time, as they target a smaller size range of fish, which also grow more quickly through the vulnerable window due to higher growth rates. Therefore, unlike trawls and beach seines, which select all size classes beyond the minimum length of capture (L_c), small meshed gillnets are not necessarily destructive and may rather promote sustained production by allowing a higher proportion of the spawning biomass to remain in the stock. The work also helps to explain the observation of sustained fish production in many developing countries despite the persistent use of gillnets of small mesh size that target small, under‐sized individuals.     

Relationship between recruitment of Japanese sardine (Sardinops melanostictus) and environment of larval habitat in the low‐stock period (1995–2010)

Stock level of Japanese sardine (Sardinops melanostictus) was high from 1980s to early 1990s and low from late 1990s to 2000s. The warm and cold water masses in the vicinity of the Kuroshio axis from winter to early spring used to be critical for the recruitment in the high‐stock period, because most of the larvae were distributed there. However, the environmental fluctuation might not affect the recruitment in the low‐stock period. Some studies reported that spawning location and spawning season, and hence the larval habitat, differ depending on the stock level. Three points were investigated in this study: (a) how spawning location and spawning season shifted from the late 1990s, (b) confirmation of the distribution area of larvae in the recent low‐stock period and (c) whether the water temperature in the vicinity of the Kuroshio axis was still related to the recruitment in the low‐stock period. The spawning location and spawning season clearly changed after 1995. Consequently, particle tracking experiments suggested that the larvae appeared in the vicinity of the Kuroshio axis from winter to early spring decreased. Nevertheless, only the ambient temperature of larvae that appeared in the vicinity of the Kuroshio axis from winter had a significant negative correlation with an index of the recruitment in the low‐stock period. It is suggested that the warm and cold masses in the vicinity of the Kuroshio axis are critical for the recruitment regardless of the stock level.     

Stock–environment–recruitment models for North Atlantic albacore (Thunnus alalunga)

Different stock–recruitment models were fitted to North Atlantic albacore (Thunnus alalunga) recruitment and spawning stock biomass data. A classical density dependence hypothesis, a recent environmental‐dependence hypothesis and a combination of both were considered. For the latter case, four stock–environment–recruitment models were used: Ricker, Beverton‐Holt, Deriso's General Model (modified to take into account environmental effects) and conditioned Neural Networks. Cross‐validation analysis showed that the modified Deriso model had the best predictive capability. It detected an inverse effect of the North Atlantic Oscillation (NAO) on recruitment, a Ricker‐type behaviour with density dependent overcompensation when environmental conditions are unfavourable and a Beverton–Holt‐type behaviour towards an asymptotic recruitment carrying capacity with favourable environmental conditions. The Neural Network model also detected that under favourable environmental conditions high spawning stock biomass does not necessarily have a depensatory effect on recruitment. Moreover, they suggest that under extremely favourable environmental conditions, albacore recruitment could increase well above the asymptotic carrying capacity predicted by Beverton–Holt‐type models. However, the general decrease in spawning stock biomass in recent years and increasing NAO trends suggest that there is low probability of exceptionally large recruitment in the future and instead there is a danger of recruitment overfishing.     

Multi‐decadal variations in spawning ground use in Northeast Arctic haddock (Melanogrammus aeglefinus)

Despite the importance both in an ecological and management context, much uncertainty remains about the underlying factors controlling spawning ground use in marine fish. Here, we investigate how spawning ground use of Northeast Arctic (NEA) haddock (Melanogrammus aeglefinus) has changed over time. By combining data from a Soviet‐Russian egg survey conducted over a 35 year period (1959–1993) with a biophysical drift model, we estimate annual median spawning latitudes of NEA haddock. Furthermore, we test for statistical association between the median latitude of spawning and covariates that are candidates for shaping the spawning distribution of haddock. The results indicate no association with temperature, as measured at the Kola transect, or demographic structure, as measured by the mean weight of spawners. However, we find a positive and significant association between spawning latitude and spawning stock biomass, indicating that density‐dependence may be more important in shaping spawning location than other factors such as temperature and demographic structure. A potential mechanism for the reported effect of spawning stock biomass may act through the distribution at the feeding grounds, which is reported to expand north‐ and eastwards at high population density.     

Did lack of spawners cause the collapse of the European eel,Anguilla anguilla?

Since the 1980s, a 90% decline in recruitment of European eel, Anguilla anguilla (L.), has occurred across most of Europe. Whether the continental stock has equally declined is uncertain. This study compiles available landings statistics since the beginning of the 20th century and identifies trends over time and space, using a statistical model that takes varying levels of reporting into account. Landings in the pre‐1940s reached over 40 000 tonnes yr^?1, declined during World War II, rose to a peak of 40 000 tonnes yr^?1 in the 1960s (coincident with a peak in re‐stocking) and dropped to an all time low of <20 000 tonnes yr^?1 in the 1990s. The decline in recruitment since the early 1980s was preceded by a decline in landings two or more decades earlier, indicating a decline of the continental stock. Considering the continental stock and the spawning stock must have declined in parallel, insufficient spawning stock biomass might have caused the recruitment collapse currently observed.     

Oceanographic drivers of sablefish recruitment in the California Current

Oceanographic processes and ecological interactions can strongly influence recruitment success in marine fishes. Here, we develop an environmental index of sablefish recruitment with the goal of elucidating recruitment‐environment relationships and informing stock assessment. We start with a conceptual life‐history model for sablefish Anoplopoma fimbria on the US west coast to generate stage‐ and spatio‐temporally‐specific hypotheses regarding the oceanographic and biological variables likely influencing sablefish recruitment. Our model includes seven stages from pre‐spawn female condition through benthic recruitment (age‐0 fish) for the northern portion of the west coast U.S. sablefish stock (40°N–50°N). We then fit linear models and use model comparison to select predictors. We use residuals from the stock‐recruitment relationship in the 2015 sablefish assessment as the dependent variable (thus removing the effect of spawning stock biomass). Predictor variables were drawn primarily from ROMS model outputs for the California Current Ecosystem. We also include indices of prey and predator abundance and freshwater input. Five variables explained 57% of the variation in recruitment not accounted for by the stock‐recruitment relationship in the sablefish assessment. Recruitment deviations were positively correlated with (i) colder conditions during the spawner preconditioning period, (ii) warmer water temperatures during the egg stage, (iii) stronger cross‐shelf transport to near‐shore nursery habitats during the egg stage, (iv) stronger long‐shore transport to the north during the yolk‐sac stage, and (v) cold surface water temperatures during the larval stage. This result suggests that multiple mechanisms likely affect sablefish recruitment at different points in their life history.     

Changes in spawning stock structure strengthen the link between climate and recruitment in a heavily fished cod (Gadus morhua) stock

Atlantic cod (Gadus morhua) is one of the commercially most important fish species in the North Atlantic and plays a central role in several ecosystems. Fishing pressure has been heavy over a prolonged period and the recent decades have shown dramatic decline in abundance of many stocks. The Arcto‐Norwegian (or North‐east Arctic) cod stock in the Barents Sea is now the largest stock of Atlantic cod. Recruitment to this stock has varied extensively during the last 60 yr. There is evidence for fluctuations in climate, particularly sea temperature, being a main cause for this variability, higher temperatures being favourable for survival throughout the critical early life stages. Our studies of time series present compelling evidence for a strengthening of the climate–cod recruitment link during the last decades. We suggest this is an effect of the age and length composition of the spawning stock having changed distinctly. The age of the average spawner has decreased by more than 3 yr from between 10 and 11 in the late 1940s to 7–8 in the 1990s, average length from just above 90 cm to around 80 cm. The number of age classes contributing to the spawning stock has also decreased, while the number of length groups present increased slightly. Significant decrease in age of spawners has frequently been described for other heavily fished stocks worldwide. We therefore find it likely that the proposed mechanism of increased influence of climate on recruitment through changes in the spawning stock age and size composition is of a general nature and might be found in other systems.     

Environmentally driven forecasts of northern rock sole (Lepidopsetta polyxystra) recruitment in the eastern Bering Sea

Northern rock sole recruitment in the eastern Bering Sea has been hypothesized to (a) depend on wind‐driven surface currents linking spawning and nursery areas, (b) be density‐dependent, and (c) be negatively impacted by cold bottom temperatures over a large nursery area during the first summer of life. A suite of models was developed to test these hypotheses. Data included 32 years of recruitment and spawning biomass estimates derived from a stock assessment model and wind and temperature indices customized to the environmental exposure of age‐0 northern rock sole in the eastern Bering Sea. The predictive ability of the models was evaluated, and the models were used to forecast recruitment to age‐4 for recent year classes which are poorly retained by the standard multi‐species bottom trawl survey gear. Models which included wind and temperature indices performed better than a naïve forecast based on the running mean. The best‐performing model was a categorical model with wind and temperature thresholds, which explained 49% of the variation in recruitment. Ricker models performed more poorly than models without a spawning biomass term, providing no evidence that recruitment is related to stock size. The models forecast higher recruitment for the most recent year classes (2015–2018) than for prior year classes with observed poor recruitment (2006–2013). These environment‐based recruitment forecasts may improve recruitment estimates for the most recent year classes and facilitate study of the effects of future climate change on northern rock sole population dynamics.     

Proposed harvest control rule for managing the snow crab stock in the southwestern Gulf of St. Lawrence,Canada

We developed limit and target harvest control rules based on an age-, sex- and stage-structured model for managing the southwestern Gulf of St. Lawrence snow crab (Chionoecetes opilio) stock. We determined an F_x% (F resulting in a spawning biomass-per-recruit equivalent to x% of virgin spawning biomass-per-recruit) as a proxy for F_MSY and a minimum spawning stock biomass to open the fishery to incorporate them into the control rules. We evaluated the selected limit and target control rule parameters under stochastic simulations by considering various performance statistics. Because of the complexity in determining an effective female spawning biomass that involves a mating ratio, we choose the total mature male biomass (MMB) regardless of crab size as the spawning index to develop the stock–recruitment relationship for stochastic simulations. The MMB based F_37% appears a reasonable proxy for limit control rule while F_45% appears a reasonable target control rule. The corresponding limit and target harvest rates of legal size crab when the standing stock MMB exceeds the proxy MSY level are approximately 36% and 25%, respectively. The difficulty in establishing an appropriate stock–recruitment relationship for this stock was recognized hence a precautionary F_45% target level was chosen. Scenarios under biomass independent random recruitment were also investigated and performances of F_37% and F_45% control rules under this hypothesis behaved similar to those observed under S–R model based simulations.     

Environmental variability controls recruitment but with different drivers among spawning components in Gulf of St. Lawrence herring stocks

The factors affecting herring recruitment are still poorly understood, complicating the prediction of stock dynamics and the choice of operational management strategies. We investigated effects of intrinsic (SSB) and extrinsic factors (physical and biological environments, including competition and predation) on recruitment of the spring and fall spawning components of each of the two herring stocks occurring in the Gulf of the St. Lawrence between 1971 and 2014. Effects of potential explanatory factors on recruit (age 2) abundance were tested using Generalized Additive Models. Model fit was significantly improved by incorporating both physical and biological environmental variability, but effects of herring SSB and predation were not significant. Indices of zooplankton abundance and phenology explained more variance in recruitment than physical indices. Our results emphasize the dominance of bottom‐up processes over SSB in the regulation of herring recruitment. Environmental variability did not seem to act uniformly on the recruitment of either stock or their respective spawning components. A long‐term trend of decreasing recruitment in spring spawners was associated with a long‐term decline in abundance of cold water copepods. In fall spawners, optimal recruitment was dependent on warmer environmental conditions combined with an adequate supply (species composition and phenology) of zooplankton. These results provide the first empirical evidence that spring and fall spawning herring are adapted to contrasting environmental conditions and shed light on the potential mechanisms linking herring recruitment to key zooplankton community characteristics and phenology. Management strategies can be improved by incorporating this new knowledge on environmental drivers of herring recruitment.     

Greenland cod (Gadus morhua): modeling recruitment variation during the second half of the 20th century

Two approaches were used to qualify observed variability in Greenland cod (Gadus morhua) recruitment. In the first analysis, we used the linear trend of the Greenland cod recruitment time series and climatic variables, such as air temperatures from the Denmark Strait and wind conditions off East Greenland and Southwest Greenland, to explain the interannual variation in cod recruitment off Greenland. The model resulting from this ‘trend/environmental approach’, explained 79% of the interannual variation in cod recruitment off Greenland. In the second, analytical approach, the ‘regime approach’, multiple linear regression models were used, with the input data being the time series of cod recruitment and spawning stock biomass (SSB) from Iceland and Greenland, sea surface and air temperatures around Greenland, and zonal wind components between Iceland and Greenland. Model results indicated that, during the decades between 1950 and 1990, there were three different cause–effect regimes which significantly influenced the variability of cod recruitment. The three regimes included: (a) the 1950s and 1960s, a regime with favorable sea surface temperatures and a self‐sustaining cod stock off Greenland with high SSB that produced a series of above‐average, strong year classes; (b) the 1970s and 1980s, a regime of declining SSB and recruitment, with recruitment dependent on advection from Iceland; and (c) the 1990s, when the advective potential for recruitment from the Icelandic cod stock was the only available source for replenishment of the Greenland cod stocks, because cod recruitment in Greenland waters was negligible. The three models explained 76–77% of the observed interannual variation in cod recruitment off Greenland. Both approaches suggested that advective factors were the dominant influences for cod recruitment in the ‘Iceland–Greenland System’.     

Estimating fisheries reference points from catch and resilience

This study presents a Monte Carlo method (CMSY) for estimating fisheries reference points from catch, resilience and qualitative stock status information on data‐limited stocks. It also presents a Bayesian state‐space implementation of the Schaefer production model (BSM), fitted to catch and biomass or catch‐per‐unit‐of‐effort (CPUE) data. Special emphasis was given to derive informative priors for productivity, unexploited stock size, catchability and biomass from population dynamics theory. Both models gave good predictions of the maximum intrinsic rate of population increase r, unexploited stock size k and maximum sustainable yield MSY when validated against simulated data with known parameter values. CMSY provided, in addition, reasonable predictions of relative biomass and exploitation rate. Both models were evaluated against 128 real stocks, where estimates of biomass were available from full stock assessments. BSM estimates of r, k and MSY were used as benchmarks for the respective CMSY estimates and were not significantly different in 76% of the stocks. A similar test against 28 data‐limited stocks, where CPUE instead of biomass was available, showed that BSM and CMSY estimates of r, k and MSY were not significantly different in 89% of the stocks. Both CMSY and BSM combine the production model with a simple stock–recruitment model, accounting for reduced recruitment at severely depleted stock sizes.     

Impact of spring upwelling variability off southern‐central Chile on common sardine (Strangomera bentincki) recruitment

Off southern‐central Chile, the impact of spring upwelling variability on common sardine (Strangomera bentincki) recruitment was examined by analyzing satellite and coastal station winds, satellite chlorophyll, and common sardine recruitment from a stock assessment model. In austral spring, the intensity of wind‐driven upwelling is related to sea surface temperature (SST) from the Niño 3.4 region, being weak during warm periods (El Niño) and strong during cold periods (La Niña). Interannual changes in both spring upwelling intensity and SST from the Niño 3.4 region are related to changes in remotely sensed chlorophyll over the continental shelf. In turn, year‐to‐year changes in coastal chlorophyll are tightly coupled to common sardine recruitment. We propose that, in the period 1991–2004, interannual changes in the intensity of spring upwelling affected the abundance and availability of planktonic food for common sardine, and consequently determined pre‐recruit survival and recruitment strength. However, the importance of density‐dependent factors on the reproductive dynamic cannot be neglected, as a negative association exists between spawning biomass and recruitment‐per‐spawning biomass. Coastal chlorophyll, upwelling intensity, and SST anomalies from the Niño 3.4 region could potentially help to predict common sardine recruitment scenarios under strong spring upwelling and El Niño Southern Oscillation (ENSO)‐related anomalies.     

The influence of environment and spawning distribution on the survival of anchovy (Engraulis encrasicolus) larvae in the Bay of Biscay (NE Atlantic) investigated by biophysical simulations

A growth and survival model of the early life stages was run along virtual drift trajectories tracked in a hydrodynamic model to simulate the annual recruitment process of anchovy (Engraulis encrasicolus) in the Bay of Biscay (NE Atlantic). These biophysical simulations concerning three different years were analysed in order to investigate the influence of environment and spawning dynamics on the survival of larvae and juveniles. The location of space–time survival windows suggested major environmental mechanisms involved in simulated recruitment variability at the different scales – retention of larvae and juveniles in favourable habitats over the shelf margins and turbulence effects. These small‐scale and meso‐scale mechanisms were related to the variations in wind direction and intensity during spring and summer. Survival was also variable according to the origin of the drift trajectories, that is spawning distribution in space and time. The observed spawning distribution (according to field surveys) was compared with the spawning distribution that would maximize survival (according to the biophysical model) on a seasonal scale, which revealed factors not considered in the biophysical model (e.g. spawning behaviour of the different age classes). The variation of simulated survival according to spawning distribution was examined on a multi‐annual scale and showed a coherent pattern with past and present stock structures. The interaction processes between the population (influence on spawning) and its environment (influence on survival) and its implications on recruitment and stock dynamics are discussed.     

Factors influencing recruitment and growth of age‐0 yellow perch in eastern South Dakota glacial lakes

Variation in recruitment and growth of age‐0 yellow perch, Perca flavescens (Mitchill), was modelled across a range of nine eastern South Dakota glacial lakes to: (i) estimate factors influencing recruitment and growth dynamics during early ontogeny; and (ii) determine the relative importance of biotic versus abiotic processes in regulating recruitment and growth dynamics. Results provide a framework for future investigations and suggest that abiotic factors were more important in regulating recruitment of age‐0 yellow perch, whereas biotic factors were more important in regulating growth. Recruitment was positively related to springtime water levels and temperature and negatively related to spawning stock biomass and springtime wind conditions. By contrast, growth was negatively related to abundance of conspecifics and potential competitors (i.e. bluegill, Lepomis macrochirus Rafinesque) and positively related to abundance of potential predators [i.e. walleye, Sander vitreus (Mitchill) and northern pike, Esox lucius Linnaeus].     

Examining common assumptions about recruitment: a meta‐analysis of recruitment dynamics for worldwide marine fisheries

Assumptions about the future productivity of a stock are necessary to calculate sustainable catches in fisheries management. Fisheries scientists often assume the number of young fish entering a population (recruitment) is related to the biomass of spawning adults and that recruitment dynamics do not change over time. Thus, managers often use a target biomass based on spawning biomass as the basis for calculating sustainable catches. However, we show recruitment and spawning biomass are not positively related over the observed range of stock sizes for 61% of 224 stocks in the RAM Legacy Stock Assessment Database. Furthermore, 85% of stocks for which spawning biomass may not drive recruitment dynamics over the observed ranges exhibit shifts in average recruitment, which is often used in proxies for target biomasses. Our results suggest that the environment more strongly influences recruitment than spawning biomass over the observed stock sizes for many stocks. Management often endeavours to maintain stock sizes within the observed ranges, so methods for setting management targets that include changes within an ecosystem may better define the status of some stocks, particularly as climate changes.     

Variation in abundance of Norwegian spring-spawning herring (Clupea harengus, Clupeidae) throughout the 20th century and the influence of climatic fluctuations

A long-term (1907–98) virtual population analysis (VPA) was made for Norwegian spring-spawning herring (NSSH), which is a huge pelagic fish stock in the north-east Atlantic. It shows that this herring stock has had large fluctuations during the last century; these fluctuations have mainly been determined by variations in the temperature of the inflowing water masses to the region. The spawning stock biomass (SSB) increased from a rather low level in the early years of this century and reached a high level of around 14 million tons by 1930. The spawning stock biomass then decreased to a level of around 10 million tons by 1940, but increased again to a record high level of 16 million tons by 1945. The stock then started to decrease and during the next 20-year period fell to a level of less than 50 000 tons by the late 1960s. Through the 1970s and 1980s, the stock slowly recovered and after the recruitment of strong year classes in 1983 and 1990–1992 the stock recovered to a spawning stock biomass of about 10 million tons. The long-term fluctuation in spawning stock biomass is caused by variations in the survival of recruits. It is found that the long-term changes in spawning stock abundance are highly correlated with the long-term variations in the mean annual temperature of the inflowing Atlantic water masses (through the Kola section) into the north-east Atlantic region. The recruitment is positively correlated with the average temperature in the Kola section in the winter months, January–April, which indicates that environmental factors govern the large-scale fluctuations in production for this herring stock.     

Recruitment of burbot (Lota lota L.) in Lake Erie: an empirical modelling approach

Stapanian MA, Witzel LD, Cook A. Recruitment of burbot (Lota lota L.) in Lake Erie: an empirical modelling approach.
Ecology of Freshwater Fish 2010: 19: 326–337. Published 2010. This article is a US Government work and is in the public domain in the USA Abstract – World‐wide, many burbot Lota lota (L.) populations have been extirpated or are otherwise in need of conservation measures. By contrast, burbot made a dramatic recovery in Lake Erie during 1993–2001 but declined during 2002–2007, due in part to a sharp decrease in recruitment. We used Akaike’s Information Criterion to evaluate 129 linear regression models that included all combinations of one to seven ecological indices as predictors of burbot recruitment. Two models were substantially supported by the data: (i) the number of days in which water temperatures were within optimal ranges for burbot spawning and development combined with biomass of yearling and older (YAO) yellow perch Perca flavescens (Mitchill); and (ii) biomass of YAO yellow perch. Warmer winter water temperatures and increases in yellow perch biomass were associated with decreases in burbot recruitment. Continued warm winter water temperatures could result in declines in burbot recruitment, particularly in the southern part of the species’ range.