Jiyang Dong1，，a, Xiumiao Song2，b, Qigen Liu3，*，c, Zhongjun Hu4，*，b
1Centre for Research on Environmental Ecology and Fish Nutrition of the Ministry of Agriculture, Shanghai Ocean University, Shanghai201306, China
2Centre for Research on Environmental Ecology and Fish Nutrition of the Ministry of Agriculture,Shanghai Ocean University, Shanghai201306, China
3Centre for Research on Environmental Ecology and Fish Nutrition of the Ministry of Agriculture, Shanghai Ocean University, Shanghai201306, China
4Centre for Research on Environmental Ecology and Fish Nutrition of the Ministry of Agriculture, Shanghai Ocean University, Shanghai201306, China
aEmail: jiyangdong0527@163.com
bEmail: No
cEmail: qgliu@shou.edu.cn
dEmail:zjhu@shou.edu.cn
*Corresponding Author
Abstract: The assessment of fish resources is crucial for fishery management and often relies on gill nets due to their operational simplicity and cost-effectiveness compared to other fishing gears. Despite their widespread use in inland water fish surveys, gill nets’ catch and composition can be influenced by fish activity, as they represent a passive capture method. The study investigated diel catch variations of primary fish species in Gehu Lake using monofilament trammel nets from April to November of 2016. Fish sampling occurred monthly, with nets set and fish caught at four-hour intervals in each month. The results showed that significant diel effects and diel × month interactions were found on Chinese silver carp (SC), Chinese bighead carp (BC), Mongolian redfin (MR), common carp (CC), and Japanese grenadier anchovy (JGA). Topmouth, humpback, and Wuchang bream (WB) displayeded significantly higher number per unit effort during twilight and daytime than at night, and no diel × month interactions were detected. For goldfish, no diel effect and diel × month interaction was observed. The study suggested that most activities occurring in daytime and at twilight were caused by visual orientation to prey for topmouth and humpback, and by the herbivorous feeding habitat of WB. Food competition with BC led JGA to increase its nocturnal activity. Overnight gillnet fishing could underestimate the population sizes of herbivores such as WB and visually oriented predators, for example topmouth and humpback, and might not influence the estimation for goldfish. However, its effects on the stock estimation of SC, BC, CC, MR, and JGA would vary with months.
Keyword: Gehu Lake; monofilament net; catch per unit effort; diel effects
- Introduction
Assessment of fish resources serves as the cornerstone for the sustainable management of fishery resources. The most commonly used gear in fishery resource assessment is the gill net (Jackson and Harvey 1997; Zitek, Schmutz and Ploner 2004; Ye 2012). Compared to other gears such as trawls, purse seines, and beach seines, gill nets are easy to operate and cost-effective (Vašek et al. 2009; Rudolfsen et al. 2021). This dual benefit has driven their extensive utilization in fish surveys across inland waters (Li et al. 2015a). However, their catch and species composition are affected by fish activity since they represent a passive capture method. (Tušer et al. 2022). There are significant differences that exist in the diel rhythms of fish activities due to various reasons such as seasonal changes, feeding behaviour, predator-prey dynamics, temperature and light conditions, reproductive activities, and human disturbances (Hohausová, Copp, and Jankovský 2003; Mei et al. 2023). The accuracy of fish resource assessment can be affected by significant changes in gill net catches, caused by the diurnal changes in fish activity (Ford et al. 2020).
Several studies have been conducted to determine the effects of diel rhythms of fish catch in Europe and the United States of America. However, these investigations have predominantly focused on marine ecosystems, with limited attention given to the assessment of such effects in freshwater lakes (Prchalová et al. 2010; Kärgenberg et al. 2022; Sajdlová et al. 2018). In addition, the research conducted in China has also been skewed towards marine fish, neglecting the specific context of freshwater lakes and drinking water reservoirs (Kadye and Booth 2014; Yan 2008; Lian et al. 2017). Despite the existence of many reports on the circadian rhythms of feeding behavior in cultured freshwater fish, such as silver carp (Hypophthalmichthys molitrix) and bighead carp (Aristichthys nobilis), it is essential to note that the activity rhythm observed in these studies may not consistently align with the feeding rhythm of other fish in different ecosystems (Zhao 2016; Cooke 2016; Powell 2012). This emphasizes the need for additional research to improve our understanding of diel rhythms, particularly for specific freshwater fish species (Monterroso, Alves, and Ferreras 2014; Wang et al. 2015; Sun 2015; Lian et al. 2015).
Silver carp (SC) and bighead carp (BC) are commonly used as tool species in non-classical biomanipulation techniques to control cyanobacterial blooms. Meanwhile, they are stocked into freshwater ecosystems of China for economic benefits (Zhang et al. 2019). This study investigated the monthly variations of diel activitys of the above two carps in Gehu Lake of China, as well as the main bycatch species that are also caught in the process of fishing.
- Materials and Methods
2.1 Study area
Gehu Lake (119°44′15″E119°52′56″E, 31°43′04″N31°28′19″N) is loacated in the southern part of Jiangsu Province and the Taihu Lake Basin in the southeast China. It is a shallow and eutrophic lake, with an average water depth of 1.25 m and a total surface area of 164 km2. It fulfills a variety of functions, such as supplying drinking water, providing irrigation, developing tourism, facilitating aquaculture, and conserving genetic resources. (Kong 2017). With the rapid economic development and the resulting increase in pollution, the lake has undergone a complete transition from being dominated by macrophytes to being dominated by algae in the early years of this century (Ge, Zhang, and Yang 2018).
Figure 1: Map showing Gehu Lake
2.2 Fish sampling
Fish sampling was conducted monthly with a monofilament trammel net from April to November 2016 at the National Aquatic Germplasm Resource Reserve, located in the central part of the lake. Sampling was, however, not carried out in June because of severe weather conditions. This gap may affect our understanding of early summer dynamics, but the multi-month dataset still provides robust insights.
The trammel net is a specially designed gill net and was constructed by joining three parallel sheets of netting. Each trammel net had a length of 67 m and a height of 1.5 m, with a mesh size of 9.38 cm for the inner panel and 39.3 cm for the outer panel. It is commonly used by local fishermen to fish for silver and bighead carp. Each month, three sets of trammel nets, each consisting of 5 nets, were set at approximately 2:00, 6:00, 10:00, and 14:00, 18:00, and 22:00 time intervals at the sampling site depending on the weather at that time, and lifted after 1 h (Erzini et al. 2006). Each period was defined as daytime, nighttime, or twilight for the analysis of the diel rhythm of the catch. The 1 h sampling intervals encompassing sunrise and sunset were designated as twilight. The 1 h sampling intervals between sunrise and sunset (excluding the designated twilight) were referred to as daytime, while the 1-h intervals between sunset and sunrise (excluding the designated twilight) were defined as nighttime (Vašek et al. 2009). Fishes were collected according to each set of nets. All captured fish were classified and the body length and weight measured to the nearest centimeter and gram, respectively. All fishes caught were identified according to previous taxonomic descriptions (Huang, Wu, and Li 2013; Matsuura 2015).
2.3 Data analysis
Number per unit effort (NPUE, ind·1000 m-2·h-1) and biomass per unit effort (BPUE, g·1000 m-2·h-1) were calculated using the total number of fish captured and their weight. Each period was defined as daytime, nighttime, or twilight for the analysis of the diel rhythm of catch. The 1 h sampling intervals encompassing sunrise and sunset were designated as twilight. The 1 h sampling intervals between sunrise and sunset (excluding the designated twilight) were referred to as daytime, while the 1-h intervals between sunset and sunrise (excluding the designated twilight) were defined as nighttime (Vašek et al., 2009).
Data were transformed using the log to satisfy the normal distribution or homogeneity of variance before analysis. If they still did not meet the requirements of normal distribution and homogeneity of variance, a two-way analysis of variance (ANOVA) was run to test for the monthly and diel effects on NPUE of different fish species after nonparametric aligned rank transformation was conducted on the data. This data transformation could guarantee the integrity of the interaction (Mansouri, Paige, and Surles 2004). If there was an interaction, the diurnal differences in NPUE for each fish were tested in each month (Hu et al. 2015).
3 Results
3.1 Diel variations of monthly catches of silver carp and bighead carp
There was a substantial monthly variation in the diel pattern of NPUE of silver carp and bighead carp. The NPUE of silver carp was the highest at the time interval of 6:00–7:00 in April, October, and November. The highest NPUE for May and August, however, occurred at the time intervals of 18:00–19:00 and 22:00–23:00, respectively. Conversely, July and September had their highest NPUE occurring at the time interval of 2:00–3:00 (Fig. 2).
Bighead carp had its peak NPUE occurring between 10:00–11:00 in April with a sub-peak occurring between 6:00–7:00. The peak NPUE recorded for the months of May, July, and August occurred at the intervals of 18:00–19:00, 14:00–15:00, and 22:00–23:00, respectively, whereas that of September and November occurred from 6:00 to 7:00. During September, two sub-peaks also appeared at the intervals of 6:00–7:00 and 14:00–15:00, respectively. October had its peaks at the intervals of 6:00–7:00 and 18:00–19:00 (Fig.3).
Fig.2 Diel pattern of NPUE of Chinese silver carp caught by trammel net in different months
Fig.3 Diel pattern of NPUE of Chinese bighead carp caught by trammel net in different months
3.2 Diurnal differences in catches for the two carps and by-catch species
There were significant differences in the diurnal changes in the NPUE of silver carp, bighead carp, Mongolian redfin (Chanodichthys mongolicus), common carp (Cyprinus carpio), and Japanese grenadier anchovy (Coilia nasus) (P<0.05). However, there was a diel × month interaction for all five species (Table 1 and Table 2), indicating that the diurnal rhythms of these species changed according to month. The NPUE varied significantly with the time of day for silver carp in May, October and November, being higher at twilight and/or at night than in daytime (Fig. 4). For bighead carp, it varied significantly in November, with higher NPUE at twilight than at night as shown in Fig. 5. Mongolian redfin was captured only in April and November. Only in April did its NPUE fluctuate significantly with the time of day, with higher NPUE at twilight than throughout the day or at night (Figure 6). There was a significant diel difference in the NPUE of common carp in the months of April, May and September. In April, the NPUE was significantly higher in daytime than at night, whereas in the other two months the NPUE at twilight was significantly higher than in daytime and/or at night (Fig. 7).
There was a significant diurnal difference in NPUE for Japanese grenadier anchovy in July and October (P < 0.05). The nighttime NPUE recorded in July was significantly higher than that of twilight. In October, the NPUE recorded at night was significantly higher than that of twilight, which was also significantly higher than that of daytime (Fig.8).
The NPUE of topmouth (Culter alburnus), humpback (Chanodichthys dabryi), and Wuchang bream (Megalobrama amblycephala) varied with time of day, and no diel × month interactions were found as shown in Table 1. The NPUEs recorded at twilight and in the daytime were both significantly higher than at night (Table 2).The NPUE of goldfish (Carassius auratus) did not vary with time of day, and no interaction between the diel effect and monthly effect was observed (Table 1 and Table 2).
Table 1: Results of two-way analysis of variance performed on NPUE of different fish species showing the effects of the factors diel time, month and their interactions
| Species | Diel effect | Month effect | Interactions |
| Silver carp | 19.317 (0.001) | 4.835 (<0.001) | 3.855 (<0.001) |
| Bighead carp | 4.734 (0.011) | 3.026 (0.009) | 2.106 (0.022) |
| Topmouth | 10.813 (<0.001) | 2.346 (0.036) | 1.606 (0.101) |
| Humpback | 15.342 (<0.001) | 5.963 (<0.001) | 0.764 (0.686) |
| Mongolian redfin | 5.490 (0.009) | 14.076 (0.001) | 6.865 (0.004) |
| Common carp | 11.310 (<0.001) | 5.727 (<0.001) | 4.544 (<0.001) |
| Goldfish | 3.097 (0.051) | 2.393 (0.033) | 0.806 (0.644) |
| Wuchang bream | 5.186 (0.008) | 2.704 (0.053) | 0.726 (0.631) |
| Japanese grenadier anchovy | 7.704 (<0.001) | 12.645 (<0.001) | 3.840 (<0.001) |
Table 2: Significance test on NPUE of different fish species among daytime, twilight and nighttime
| Species | NPUE( ind. /1000 m2/ h-1) | ||
| Daytime | Twilight | Nighttime | |
| Silver carp | 43.34±6.22c | 119.47±21.04a | 85.89±9.54b |
| Bighead carp | 8.85±1.55ab | 11.58±1.96a | 6.18±1.02b |
| Topmouth | 1.22±0.44a | 2.33±0.71a | 0.33±0.19b |
| Humpback | 6.19±3.05a | 4.09±1.08a | 2.08±0.67b |
| Mongolian redfin | 0.66±0.45b | 2.31±0.94a | 0.17±0.17b |
| Common carp | 2.19±3.47a | 3.19±0.98a | 1.11±0.24b |
| Goldfish | 4.95±1.06 | 5.37±1.02 | 2.87±0.56 |
| Wuchang bream | 3.59±0.70a | 2.98±1.17a | 1.19±0.47b |
| Japanese grenadier anchovy | 2.76±0.57b | 5.92±1.95a | 5.34±1.07a |
Note: The different superscript letters in each column indicate significant differences in NPUE at the level of P < 0.05.
Fig.4 Change of NPUE of Chinese silver carp between daytime, twilight,and nighttime
Note: The different letters in each month denote significant differences at the level of P< 0.05, and the same applies to the figures below.
Fig.5 Change of NPUE of Chinese bighead carp between daytime, twilight, and nighttime
Fig. 6 Change of NPUE of Mongolian redfin between daytime, twilight,and nighttime
Fig.7 The diurnal change of NPUE of common carp (Cyprinus carpio)
Fig.8 The diurnal change of NPUE of Japanese grenadier anchovy (Coilia nasus)
3.3 NPUE correlation between fishes
The results showed a significant negative correlation in NPUE between bighead carp and Japanese grenadier anchovy. The NPUE of bighead carp, however, was positively correlated with those of silver carp, topmouth, and common carp. Significant positive correlations were also found between silver carp and topmouth, between common carp and topmouth, as well as between humpback and goldfish. No correlations between NPUEs of the other fishes were found (Table 3).
Table 3 Correlations between NPUE of different fish species (df=41)
| Species | Silver carp | Bighead carp | Topmouth | Humpback | Mongolian redfin | Goldfish | Common carp | Wuchang bream |
| Silver carp | 1 | |||||||
| Bighead carp | 0.529*** | 1 | ||||||
| Topmouth | 0.395* | 0.467** | 1 | |||||
| Humpback | -0.152 | -0.072 | 0.027 | 1 | ||||
| Mongolian redfin | -0.048 | 0.047 | 0.051 | 0.091 | 1 | |||
| Goldfish | -0.022 | -0.112 | 0.124 | 0.350* | 0.107 | 1 | ||
| Common carp | 0.117 | 0.408** | 0.412** | -0.045 | 0.202 | 0.104 | 1 | |
| Wuchang bream | -0.221 | -0.268 | -0.009 | -0.056 | -0.157 | 0.037 | -0.157 | 1 |
| Japanese grenadier anchovy | -0.262 | -0.453** | -0.229 | -0.011 | -0.193 | 0.102 | -0.103 | 0.173 |
Note: Superscript symbol of “*” / “**” or “***” denotes significant correlation between NPUE of different fish species at the level of P <0.05. 4. Discussion
The diel rhythm of fish activity, which varied according to species, could be classified into diurnal, nocturnal, crepuscular, and aphasic (arrhythmic) patterns(Schwartz 2016). The diel rhythm of a single fish species also has strong plasticity and changes with season, stage of life history, social interactions, and environmental conditions (Arndt and Evans 2022). The plasticity of a specific species even can be observed in specific times and places (Janáč and Jurajda 2013). White suckers (Catostomus commersonii) are reported by different authors to be either diurnal, nocturnal, or crepuscular (Grabowski and Isely 2006; Shuai et al. 2016; Peng et al. 2023), and in present study, the diel catch efficiencies of silver carp, bighead carp, Mongolian redfin, common carp, and Japanese grenadier anchovy changed with the months, indicating the plasticity in their diel activity. Despite the plasticity, the diel rhythms of some species remain relatively constant. For instance, whitefin gudgeon (Romanogobio albipinnatus) appeared to be nocturnal under all circumstances (Roche et al. 2021). Similarly, diel rhythms of topmouth, humpback, Wuchang bream, and goldfish were independent of month. Concretely, in all months the former three species were mainly captured during daytime and twilight and goldfish displayed no significant difference in diel catch. This may be because their activity patterns help them avoid predators and have enough energy since primary productivity is high during the daytime and water temperatures are within their preferred range (Bone and Moore 2008; Helfman et al. 2009).
Foraging success and predation risk are the two strong candidates to explain why freshwater fishes are active when they are most likely to encounter optimal feeding conditions, and why they sometimes change their activity patterns (Janáč and Jurajda 2013). Being active when predators are inactive can reduce the predation risk of prey species. This kind of predator avoidance is a mechanism allowing long-term survival of prey species. For instance, the diurnal inactivity of the three Anchoa species might be related to predator avoidance (Mehner 2012; Roman et al. 2019). In this study, no significant negative correlations were found in NPUE between the three predatory fishes and the other six fish species. Fishes may adjust their activity patterns to avoid direct competition with other species, thereby maximizing their foraging success (Beverton and Holt 2012). This could be attributed to the fewer juvenile samples resulting from the relatively large mesh size of the trammel net used in this study.
Foraging success is modulated by food availability, competition, and ease of food detection, depending on the lighting conditions. According to Reebs (2002), interspecific and intraspecific competition could be reduced by different diel rhythms. The Japanese grenadier anchovy is zooplanktivorous, while bighead carp are planktivorous with a preference for zooplankton (Sass et al. 2014). A significant negative correlation in NPUE was observed between the two species, likely driven by zooplankton resource competition, prompting the anchovy to shift toward nocturnal activity. Silver carp reached its activity peak at twilight or night and bighead carp did so at the intervals other than that of 2:00–3:00. Significant diel variations in their catches only occurred in a few months (Figure 5 and Figure 6). Similar results were also reported by previous studies which suggested that they feed actively in daytime, at twilight, or at night (Myers et al. 2016; Esmaeili and Johal 2015). The two carps are both filter feeders and had high nighttime catch rates in some months, which may be related to filter-feeders lack of dependence on visual detection for food(Guo et al. 2009).
Visually-oriented predators are more active during twilight or daytime (Vašek et al. 2009). For example, the European perch (Perca fluviatilis) was mostly captured at twilight and in daytime and rarely at night (Hanson et al. 2007). Yousif (2003) found that the catch rates of Brushtooth lizardfish (Saurida undosquamis) and Japanese Threadfin Bream (Nemipterus japonicus) were significantly higher during daytime compared to night time, which further supports the idea that visually-oriented predators are more active when the visibility is better. In this studies, the results of topmouth and humpback supported the findings and conclusions of the above-mentioned literature. Additionally, catch rates for the two species and Mongolian redfin in daytime were more than 3.7, 3.0, and 4.0 times greater than their catch rates at night, respectively, and the ratios of twilight to night catches were 7.0, 2.0, and 14.0, respectively. Although no significant diel variation of the catch rates of Mongolian redfin was detected in October, the catch rates during daytime and twilight were higher than those at night (NPUE = 0). Small fish and shrimp are the main food sources of topmouth and humpback (Yang 2002). The adult Mongolian redfin has a trophic level reaching up to 3.9 (fishbase) and can be considered carnivores with a preference for decapods and fish (Kottelat 2006). The diel variations of catches of these three species may therefore be related to their feeding habits, depending on visual orientation for detection of their prey.
Most herbivorous and omnivorous fish are primarily active during the daytime (Higham, Stewart, and Wainwright 2015). Furthermore, it was argued that among five functional groups, herbivores and cleaners are inactive during the night (Li et al. 2015b). Herbivorous Wuchang bream was active mostly during the daytime and twilight, which was consistent during the study months. This result supported the findings of Li and Zhang (2012). Common carp and goldfish are omnivorous fish. Individuals of common carp cultured in ponds or aquariums mainly fed between 14:00 to 18:00 or displayed no obvious diel feeding variation (Mesquita, Borcato, and Huntingford 2015; Kolar et al. 2007). Common carp in Gehu Lake were mostly active in daytime or at twilight in most months, and during other months at night. Similar results were also reported by other researchers (Baumgartner, Stuart, and Zampatti 2008; Li et al. 2017), but contrary to those of Říha (2011). Goldfish have been reported to be active during daytime, night, or twilight, or to be aphasic (Reebs 2002). In the Murray River, catches of goldfish were significantly higher at night than in daytime (Baumgartner, Stuart, and Zampatti 2008). No significant diel variations of goldfish catches were observed in Gehu Lake, which may be attributed to its non-preference for light during the time of day.
Conclusion and recommendation
To the best of our knowledge, we report for the first time the diel variation of NPUE of silver carp, bighead carp, Wuchang bream, Japanese grenadier anchovy, and the three culters (topmouth, humpback, and Mongolian redfin). The diel variations in fish catches have an important influence on fishery resource assessments based on the fishing method of setting a gillnet over-night; catches can be influenced by various factors. It is therefore recommended that further studies be conducted to confirm the plasticity of diurnal activity of the nine species investigated in the present study.
Acknowledgments
This work was supported by the National Key Research and Development Program of China (2023YFD2400901), Three New Aquaculture Project of Jiangsu Province (Item Number Y2017-23), Aquaculture and Environmental Management Technology Research and Demonstration (Item Number2015BAD13B00).
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