1. Introduction
Abundance indices are the bedrock of global fishery stock assessments and fishery conservation [
1]. The catch-per-unit-effort (CPUE) serves as a commonly utilized metric for revealing relative fluctuations in stock abundance [
2]. Currently, the stock assessment of most fisheries in the world relies heavily on fisheries-dependent data [
3]. However, the nominal CPUE from fisheries-dependent data is likely to be biased and limited, which may be influenced by factors such as catchability, fishing effort, spatial heterogeneity, and environmental changes [
4]. For instance, fishers may increase their efforts in response to declining fish populations, leading to higher CPUE values which may not accurately reflect the true fish abundance. Therefore, the assumption of a proportional relationship between the CPUE and stock abundance is often criticized in fishery stock assessment [
5]. CPUE standardization is a critical step in fisheries’ stock assessment that involves removing or adjusting for factors that may influence catchability [
6]. By standardizing the CPUE, fisheries’ managers can obtain more accurate information on fish stock status, which has significant implications for scientific stock assessment and management.
In recent years, many authors have employed various statistical models other than the traditional General Linear Model (GLM) and Generalized Additive Model (GAM) for CPUE standardization research [
7]. For example, two recruitment indices were constructed by Hashimoto et al. [
8] utilizing pelagic trawl survey data for the Chub mackerel (
Scomber japonicus) employing a delta-GLM model. Hazin et al. [
9] standardized the CPUE of swordfish in the equatorial and southwestern Atlantic Ocean using a Generalized Linear Mixed Model (GLMM). Thorson et al. [
10] introduced a spatio-temporal GLMM (referred to as VAST) for the purpose of estimating abundance indices for West Coast groundfish species and found that the model could improve the accuracy of CPUE standardization. The GLM, GLMM, and VAST models were commonly used methods for CPUE standardization in the past and have been successfully applied to various fish species [
11]. However, these models have their own advantages and limitations, and there is relatively little research on the comparative estimation performance between the models [
12]. Therefore, selecting CPUE standardization models with a higher estimation accuracy by comparing their evaluation performance is crucial for fishery stock assessment and management.
The Pacific sardine (
Sardinops sagax) is one of the essential target species for commercial fishing in the Northwest Pacific Ocean (NPO) [
13,
14]. Pacific sardines are a short-lived species, typically living for 6–7 years, and they reach sexual maturity at around one year of age and spawn in large schools near the surface of the water [
15,
16]. The Pacific sardine is a small pelagic fish, feeding primarily on zooplankton such as copepods, krill, and small fish larvae [
17], and it is also a critical prey for larger fish, seabirds, and marine mammals and plays a vital ecological role in the marine ecosystem. Therefore, sustainable management and conservation of this species can positively impact the entire marine food web. Currently, the primary harvesters of the Pacific sardine population in the NPO are China (including Chinese Taipei), Japan, and Russia. In 2021, China reported an annual catch of approximately 237,301 tons of Pacific sardines, representing 22.20% of the global production [
18,
19]. In addition, the catch percentage of Pacific sardines for China in the NPO has been increasing year by year [
20]. With increasing attention from global researchers, the North Pacific Fisheries Commission (NPFC) has officially recognized the Pacific sardine as a priority species, and preliminary fishery stock assessment and management have been conducted [
21].
Recently, the potential impact of climate and ocean environmental changes on Pacific sardine populations has been gathered increasing attention. Several researchers have suggested that the distribution and population size of Pacific sardines is highly impacted by environmental conditions. Shi et al. [
22] utilized the ensemble distribution model to examine the population variation for Pacific sardine in the NPO, and they pointed out that sea surface height (SSH) and sea surface temperature (SST) were critical environmental variables. Takasuka et al. [
23] studied the suitable temperature for the Pacific sardine and found that 16.2 °C is the optimal growth temperature for Pacific sardines. Ito [
24] stated that the spawning grounds’ temperature of Pacific sardines in the northern and southern Pacific coast of Japan were 14–17 °C and 17–19 °C, respectively. Wada et al. [
25] found that the population size of Pacific sardine has experienced drastic fluctuations, closely related to the climate and the oceanic environment Therefore, it is necessary to consider environmental factors when standardizing the CPUE for Pacific sardines. However, currently, research on Pacific sardines mainly focuses on biology [
26,
27] and potential habitat distribution [
28,
29], and there are few reports on the CPUE standardization for Pacific sardine fishery. Furthermore, it is crucial to accurately grasp the impact of explanatory variables in a model on the standardized CPUE in CPUE standardization studies [
30]. However, previous research has rarely taken this into account.
Due to the urgent need for stock assessment and management of Pacific sardine fishery, we used the GLM, GLMM, and VAST models to analyze the CPUE for the Pacific sardine, gauging the effectiveness of each model to select the most suitable model for standardizing the CPUE for Pacific sardine fishery. This can obtain an accurate abundance index of Pacific sardine resources. Additionally, we conducted an influence analysis to appraise the effect of each variable on standardized indices. Finally, simulation tests were used to assess the effectiveness of the various models in CPUE standardization. To our knowledge, this study marks the inaugural endeavor in this field to standardize Pacific sardine CPUE within the NPO of China, employing an array of standardization models. Our study aims to achieve the following three primary goals: (1) to assess and compare the performance of three models in CPUE standardization; (2) to derive accurate CPUE data and provide support for stock assessment of the Pacific sardine fishery; and (3) to examine how the inclusion of each explanatory variable affects the standardized CPUE and determine its main drivers. The results of this investigation can provide technical support for the stock assessment and management of Pacific sardines in the NPO region.
4. Discussion
Pacific sardine fishery is a traditional fishery in the NPO [
49]. Since the 1990s, the catch of Pacific sardines has experienced significant fluctuations. However, after 2010, the population of Pacific sardines gradually recovered, leading to an increase in the catch. In 2021, the catch of Pacific sardines exceeded one million tons [
50]. With the increase in fishing scale and economic worth, there has been a growing focus on the stock assessment and governance of Pacific sardine resources, and the NPFC has prioritized its stock assessment and management [
21]. Therefore, obtaining reliable relative abundance indices is of great significance for the management of Pacific sardines. The Pacific sardine, known for its short life cycle and highly migratory behavior, demonstrates continuous fluctuations in abundance and availability that are influenced by biotic and abiotic environmental factors, exhibiting a spatial structure [
51]. Hence, including spatial autocorrelation as a continuous covariate in the CPUE standardization model for Pacific sardines appears fitting. This will improve the model’s capacity to precisely grasp the spatial distinctions in the fish occurrence patterns, thus reflecting the variations more reasonably [
52].
This study assessed the effectiveness of three models—the GLM, the GLMM, and the VAST—using Chinese Pacific sardine fishery data and marine environmental data including the SST, the SSTG, and the SSH. The reason for selecting the SST, the SSTG, and the SSH is that previous research has indicated that the habitat distribution and abundance of small pelagic fish species, such as the Pacific sardine, are highly sensitive to marine environmental factors, particularly the SST, the SSTG, and the SSH [
53,
54]. The SST has long been considered the most important influencing factor for Pacific sardine resources, and the CAIC results revealed that the best-performing GLM, GLMM, and VAST models all included the SST, confirming its significance. Nevertheless, the findings from the influence analysis revealed that the annual influence values of the SST were distributed around one (
Figure 6D), suggesting that the factors affecting resource distribution might not always make a substantial contribution to the disparities between the nominal and standardized CPUE, aligning with the conclusions of Hsu et al. [
34].
Based on the CAIC and the Conditional
R2, the VAST demonstrates a better fitting performance compared to the GLM and the GLMM (
Table 5), which was consistent with the results of Kai [
55]. Meanwhile, according to the simulation tests, the VAST model showed smaller RMSE and model bias, indicating that the bias values were closer to one. This suggests that the VAST model can better accommodate the spatial variations in the CPUE data and has a higher assessment accuracy. Ducharme-Barth et al. [
47] conducted simulation tests on the VAST model for CPUE standardization and reached conclusions consistent with our study. This can be ascribed to three primary factors. To begin with, the VAST model encompasses both spatial and temporal data variations, accounting for spatial correlations between sampling locations and temporal correlations across different time intervals, and allows for a more accurate representation of the complex spatio-temporal patterns in the data [
56]. Additionally, by introducing spatial and temporal random effects, the VAST can effectively handle the inherent correlations and disparities within the CPUE data [
57]. Lastly, the VAST provides the flexibility to incorporate supplementary covariates that elucidate the fluctuations in CPUE. This helps in capturing the impacts of pertinent environmental factors and other significant variables on the standardized CPUE, facilitating a thorough and precise analysis [
58].
From
Figure 5, it can be observed that the values of the standardized CPUE surpassed the nominal CPUE between 2014 and 2019. However, in 2020 and 2021, the opposite trend was observed. This may be due to the fact that the values of the influence of the year × spatial or spatio-temporal variables were mostly below one from 2014 to 2019, whereas, in the following two years, their influence values exceeded one. This also demonstrates the importance of this explanatory variable in influencing the disparities between the nominal and standardized CPUE [
30]. The yearly relative standardized CPUE shows a progressive upward trend between 2014 and 2021 (
Figure 5), indicating the gradual recovery of Pacific sardine resources. This pattern corresponds to the noted surge in Pacific sardine catches in China during recent years. Additionally, these research findings align with those of Yang et al. [
14] and can offer valuable scientific support for the establishment or adjustment of management regulations in Pacific sardine fisheries. The objective is to strike a balance between ecological preservation and the concerns of fishers.
CPUE standardization models are applied to mitigate the confounding influences of external factors and derive an indicator that accurately reflects fish biomass [
59]. Therefore, it is imperative to thoroughly scrutinize the results, instead of merely adopting the CPUE data generated using the standardization model, in order to grasp the impact of including each explanatory factor in the standardized CPUE. We conducted an influence analysis for each explanatory variable of the three models, and the results indicated that, in contrast to other explanatory variables, the influence of the SST on the disparities among the standardized and nominal CPUE is relatively minor (
Figure 6). The overall influence of the year × spatial or spatio-temporal variable is the highest (
Table 6), which is consistent with the results of Hsu et al. [
34]. From
Figure 6, it can be observed that the influence values of the explanatory variables in the VAST model fluctuate less over time, while, in the GLM and GLMM models, the influence values of explanatory variables exhibit larger fluctuations over time. This also demonstrates the robustness of CPUE standardization in the VAST model.
In this study, the spatial stratification method “Spatial 1” was employed to stratify the fishery data of Pacific sardines in the NPO [
34]. The influence values of the year × spatial random effect or spatio-temporal random effect in the GLMM and VAST models were consistent with the influence criteria of the relationship between the nominal and standardized CPUE (
Figure 8 and
Figure 9), whereas the year × spatial effect of the GLM did not exhibit this trend (
Figure 7). This discrepancy highlights the differences in the data handling capabilities among the models [
60]. We also discovered that the distribution of data had a significant impact on the standardized CPUE. In instances where the data clustered in regions with high coefficients, the influence value for the respective year was relatively elevated, while it was diminished when the data dispersed across areas with low coefficients (
Figure 7,
Figure 8 and
Figure 9). Based on the existing literature, several approaches, such as the ad hoc approach [
61] and the binary recursive partitioning approach [
62], can be used to determine area stratification in CPUE standardization. Therefore, it is advisable to undertake future studies to examine the influence of various fishery data stratification approaches on CPUE standardization. This will impart valuable insights, delivering scientific management recommendations to a wide audience which includes fishers, managers, and stakeholders.
Our study has provided crucial insights for the stock assessment of Pacific sardine. It has also established a spatio-temporal model framework for the CPUE standardization for other small pelagic fish species worldwide, thereby supporting the conservation and sustainable use of other fish stocks. Importantly, our study once again demonstrated the impact of the SST on the distribution and abundance of Pacific sardines. However, we also discovered that explanatory variables with a high explanatory power in CPUE standardization models may not necessarily have a significant influence on the disparities between the nominal CPUE and the standardized CPUE. Therefore, this aspect should be clarified in future research. Our study revealed that the variables of year × spatial or spatio-temporal exerted the most significant overall impact concerning the standardized CPUE, indicating their significant role in explaining the disparities between the nominal and standardized CPUE. Furthermore, it highlighted the advantages of the VAST in standardizing the CPUE for highly migratory small pelagic fish species, suggesting its incorporation as a routine CPUE standardization tool. This research will facilitate the application of accurate biomass indices in stock assessment and ultimately promote the scientific management and conservation of Pacific sardines.