Better Fitness of Sharpbelly, Hemiculter leucisculus (Basilewsky, 1855) in Invaded Habitats: Evidence from Body Length-Weight Relationships Across the World

Yuebing Zhou1, Tao Ju2*, Lei Gan1, Zhenlu Wang1, Chao Luo3, Weiwei Wang1, Tao Xiang4, Lei Shi5, Haoyu He1, Shuhai Zhang1, Miao An1, Haibo Jiang1 and Xianghong Dong1*

1College of Animal Science, Guizhou University, Guiyang, 550025, China

2Guangxi Academy of Marine Sciences, Guangxi Academy of Sciences, Nanning, 530007, China

3College of Forestry, Guizhou University, Guiyang, 550025, China

4Nanjing Institute of Geography and Limnology, Chinese Academy of Sciences, Nanjing, 210008, China

5Institute for Ecological Research and Pollution Control of Plateau Lakes, School of Ecology and Environmental Science, Yunnan University, Kunming, 650500, China

ABSTRACT

As one of the main driving factors of biodiversity loss, biological invasion has attracted wide attention. Among the many scientific issues in this field, the most noteworthy one is to figure out the ecological mechanisms behind successful invasions since relevant knowledge is extremely important for conservation and management, but we still know very little about that to some species. In view of this, the notorious sharpbelly Hemiculter leucisculus (Basilewsky, 1855) was selected as the research object, by analyzing the difference between its fitness in the native habitats and the invaded habitats at global scale through data integration technology and body length-weight relationships, this study attempted to reveal the ecological mechanisms underlying the global rapid successful invasion of this invasive species. The results showed that the fitness of the invasive populations was higher than the native populations (t = 3.85; P < 0.05). Based on the above, this study put forward a series of measures and suggestions for prevention and control so as to mitigate the potential harm of its further dispersal in the near future.

Article Information

Received 20 April 2024

Revised 10 June 2024

Accepted 24 June 2024

Available online 08 January 2025

(early access)

Published 19 December 2025

Authors’ Contribution

YZ: Data curation, formal analysis, writing-original draft; TJ and XD: Conceptualization, methodology, writing- review and editing; LG, ZW, CL, WW, TX, LS, HH, SZ, MA, HJ: Data curation and language editing.

Key words

Hemiculter leucisculus, Body length-weight relationships, Difference analysis, Fitness, Growth patterns

DOI: https://dx.doi.org/10.17582/journal.pjz/20240420084908

* Corresponding author: [email protected], [email protected]

0030-9923/2026/0001-0167 $ 9.00/0

Copyright 2026 by the authors. Licensee Zoological Society of Pakistan.

This article is an open access article distributed under the terms and conditions of the Creative Commons Attribution (CC BY) license (https://creativecommons.org/licenses/by/4.0/).

Introduction

With the continuation of global climate change, international trade and transportation industry development, biological invasion has become the second factor leading to global biodiversity loss (Duenas et al., 2021; Early et al., 2016; Leuven et al., 2017). Generally, biological invasion means the process that a species was transferred from its original habitat to a new habitat which is outside the biogeographical barrier by natural or anthropogenic assistances, and formed breeding populations in the new habitat, and afterward caused adverse impacts on the local biodiversity, agricultural, and forestry production as well as human health, or ecological disasters (Richardson et al., 2011). Compared with terrestrial ecosystems, aquatic ecosystems are more vulnerable to alien species invasion (Pyšek et al., 2020).

The sharpbelly Hemiculter leucisculus (Basilewsky, 1855) is a small freshwater fish belonging to Cypriniformes, Cyprinidae, Culterinae (Chen et al., 1998; Zhang et al., 2016), and usually dwells in the middle and upper layers of lakes, rivers, and reservoirs. It mainly feeds on invertebrates and phytoplankton. It can be sexually mature at the age of 1 year, spawns from April to October, with peak period of May and June, and its body length is usually 10–14 cm (https://fishbase.mnhn.fr/search.php; Huang et al., 2022). This cyprinid is native to China, Mongolia, Russia, Vietnam, and the Korean Peninsula (Khurshut, 2016). However, due to improper management during introduction, it has spread beyond its native range to many regions including Georgia, Afghanistan, Iran, Iraq, Kazakhstan, Uzbekistan, Turkmenistan, and Azerbaijan (Epitashvili et al., 2023; Khurshut, 2016; Mustafayev et al., 2015). In addition, this fish also possesses typical characteristics of invasive species (Wang et al., 2013, 2016). For example, on the one hand, its life history pattern is r type, i.e. it shows short life span, fast growth rate, early sexual maturity, long breeding period, high fertility, and batch spawning. On the other hand, it also shows a certain degree of plasticity, e.g. with the change of climate or geographical environment, it can not only produce eggs with different ecological types, but also make its own brain grow with different patterns (Huang et al., 2022; Liu et al., 2022; Sun, 1987; Wang et al., 2016; Wu and Yi, 1959). Therefore, some studies even made a bold prediction that, with the continuation of global climate change and increase of human activities, the areas with high invasion risk by this fish will spread all over the world except Antarctica (Dong et al., 2020). In light of this species’ strong invasive potential, it is reasonable to speculate that it may have higher fitness in the invaded habitats than the native habitats.

However, after a systematic review and anatomizing of the existing works on this fish all over the world, it is not hard to find that they still just focused on its basic biology (Wang et al., 2016), ecology (Guo et al., 2021; Hung et al., 2015), physiology (Huang and Zeng, 2016), behavior (Lu, 2015), molecular biology (Luo et al., 2022), phylogeny and adaptive evolution (Liu et al., 2022; Vasileva et al., 2022), phylogeography or zoogeography (Sun et al., 2022; Vasileva et al., 2022), environmental toxicology (Hung et al., 2015; Zhang, 2007) as well as morphology and population genetics (Cho et al., 2012; Sun et al., 2022; Wang, 2021). By far, comparative research on the difference between its fitness in the native habitats and the invaded habitats has not been reported.

In view of this, by integrating the body length-weight data of H. leucisculus around the world and comparatively analyzing that data in both the native habitats and the invaded habitats, we attempted to uncover the ecological mechanisms behind the global rapid successful invasion of this species, and provide basic data or science and technology support for the scientific management of H. leucisculus and other similar invasive species.

Materials and Methods

Data collection

The body length-weight relationships data of H. leucisculus were collected by retrieving with the key words Hemiculter leucisculus, body length-weight relations, total length, and body length, etc. in both English and Chinese databases including the Web of Science (https://www.webofscience.com/wos), Google Scholar (https://scholar.google.com), and China National Knowledge Infrastructure (https://www.cnki.net), and data types include journal articles, dissertations, government reports and materials. By statistics, 71 valid records were initially gleaned, of which 45 and 26 were in the native areas and the invaded areas, respectively (Supplementary materials). The spatial distribution of relevant research sites is shown in Figure 1.

 

Data cleansing

Before taking next steps, this study cleansed the data obtained previously in order to get more reliable information or results. To be specific, firstly, we removed the records that were duplicative, obviously incorrect, and incomplete with key information (e.g. sampling location and time, sample sizes, body length-weight relationship, as well as length types and units); Secondly, using the equations a`cm= amm 10b and aTL= aLS (TL/LS)-b (if the relationship between total length and standard length or fork length is a proportional function) or aTL = aLS (2f/Lmax + g)-b (if that relationship follows the linear function TL = f + gLS), we standardized the a in the filtered body length-weight relationship into a’, so as to make the results of different studies with different body length types or units comparable to each other (i.e. unifying the body length in the body length-weight relationship into total length, and standardizing the units of body length and body weight into centimeter and gram, respectively). In the above formulas, LS is the standard length (SL) or fork length (FL) in the body length-weight relationship in the original records; TL is total length; aLS is the parameter corresponding to the length type before conversion; aTL is the parameter corresponding to the total length. Lmax is the maximum of standard length or fork length in the original studies. f and g are the regression intercept and slope, respectively, of the linear function describing the relationship between the total length and the standard length or the fork length (Froese, 2006). According to FishBase (https://fishbase.mnhn.fr/search.php) and relevant information (Qin et al., 2017), the conversion formulas of TL and FL, and, TL and SL for H. leucisculus in this study are TL/FL = 1.15 and TL = 0.02 + 1.19SL, respectively. Thirdly, based on the theoretical relationship between lg(a’) and b (Fig. 2), outliers (i.e. points outside the 95% confidence interval of the fitting line) are removed. Finally, a total of 48 data (36 and 12 from the native areas and the invaded areas, respectively) were obtained for following analysis.

 

Statistical analysis

Shapiro-Wilk and Levene tests were applied to evaluate normality and variance homogeneity of the b values of the body length-weight relationship of H. leucisculus populations, respectively. If the normal distribution and the variance homogeneity were both verified, an independent (single) sample t test was adopted; otherwise, non-parametric Mann-Whitney (Wilcoxon) tests were used to analyze whether there were significant differences between the b values of the native populations and the invasive populations (that two sets of b values and the theoretical value 3, respectively). All statistical analyses and graphics in this study were performed in ArcGIS 10.2 (Esri Co., USA) and OriginPro 2022 (OriginLab Corporation, USA). Unless otherwise specified, all statistics are expressed as mean ± standard error. The significance level (α) was set to 0.05 according to the past practice.

Results

Value ranges of b

The results showed that b values of H. leucisculus populations in the native habitats and the invaded habitats were 2.52–3.75 (3.07±0.05) and 3.00–3.54 (3.25±0.05), respectively (Table I). Both manifest normal distribution (W = 0.98; P > 0.05) and variance homogeneity (W = 2.53; P > 0.05).

 

Table I. b values of both the invasive populations and the native populations of H. leucisculus (CI represents confidence interval for the median of b; SE represents standard error).

	n	Min	Max	Mean	Median	SE	95%CI
Native	36	2.52	3.75	3.07	3.03	0.05	2.98–3.16
Invasive	12	3.00	3.54	3.25	3.23	0.05	3.15–3.35

 

 

Difference analysis on the invasive populations and the native populations

The results showed that there was significant difference in b values between the invasive populations and the native populations (t = 3.85; P < 0.05; Fig. 3); the b values of the invasive populations were greater than 3 with significance (t = 5.35; P < 0.05), while the b values of the native populations were greater than 3 with no significance (t = 1.53; P > 0.05).

Discussion

Value range of b

The b value of fish’s body length-weight relationship should theoretically be within the range from 2.50 to 3.50 (Carlander, 1977), with which our results are roughly consistent. However, we noticed that the b values in the Erlongshan reservoir of Jilin Province (3.62) and the Beijiang river of Guangdong Province (3.75) deviated far from the upper limit of the theoretical range. Looking up the primary data, it is not difficult to find that: due to the lack of food-competing fishes (e.g. silver carp Hypophthalmichthys molitrix (Valenciennes, 1844) and bighead carp Aristichthys nobilis (Richardson, 1845), H. leucisculus population in the Erlongshan reservoir is rich in food resources, which means better growth conditions, i.e. a larger b value. By contrast, because of sampling bias (the samples were all females with gonadal development stages of from IV to V), the b value in the Beijiang river had a large deviation from the theoretical value as well. This demonstrates that it is correct to cleanse the data after data collection or before further analysis, and hence our conclusions are reliable.

Fitness comparison between the native habitats and the invaded habitats

As mentioned above, the invasive populations’ b values (3.25 ± 0.05) were significantly greater than 3, and thus manifest positive allometry; the native populations’ b values (3.07 ± 0.05) were insignificantly greater than 3, and thus manifest isometry (Fig. 4). For this reason, it can be inferred that our previous conjecture is highly probably correct, i.e. H. leucisculus population may have better fitness in the invaded habitats than the native habitats. However, unlike previous studies, Lin et al. (2023) found that Pseudorasbora parva (Temminck and Schlegel, 1846) showed good fitness at both habitats. In contrast, Sun et al. (2023) discovered that Oreochromis niloticus (Linnaeus, 1758) owned poor fitness at both the native regions and the invaded regions, although the fitness in the latter is still higher than the former.

Management recommendations

Based on the results of this paper, we find that H. leucisculus has a strong invasion potential, which further supports the speculation that this fish may spread to new habitats in the near future. Hence, this study attempts to propose a series of adaptive management recommendations: (1) adopting new technologies (e.g. eDNA and eRNA) to conduct continuous monitoring on its global population dynamics; (2) attempting to analyze the mechanisms behind this fish’s rapid successful global invasion from other perspectives, e.g. the molecular level; (3) increasing input in the basic research on this fish so as to comprehensively understand its ecological habits; (4) enhancing transnational cooperation to achieve more coordinated and effective prevention and control.

 

Declarations

Acknowledgments

We would like to express our sincere appreciations to those anonymous reviewers who made contributions to this work for their valuable observations, comments, and recommendations, leading the huge improvement of the earlier draft of this paper. This study was founded by the Program Foundation for Talents of Guizhou University (No. [2021]65 and [2021]15), the Guizhou Provincial Science and Technology Projects (No. Yiban 104 2023 Qiankehe Jichu-ZK), the National Natural Science Foundation of China (No. 32102808), and the Basic Program of Guizhou University (No. [2023]14).

Supplementary material

There is supplementary material associated with this article. Access the material online at: https://dx.doi.org/10.17582/journal.pjz/20240420084908

Statement of conflict of interest

The authors declare that there is no conflict of interests regarding the publication of this article.

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