1. Introduction
Schizothorax pseudaksaiensis (Herzenstein, 1889) is a rare and typical cold-water native fish and one of the main economic fish, with large individuals. According to relevant data, the largest individual was 92 cm long and weighed 5 kg, the quantity of this fish is low, the yield is not high, and it is widely distributed in the waters of the Ili River-Balkhash Lake system, Issyk Lake, and the Chu River [
1]. At present, the distribution area has been greatly reduced, and only a small number of
S. pseudaksaiensis are concentrated in the Ili River and its main stream [
2]. The distribution of
S. pseudaksaiensis in Yamadu River is more concentrated, and there is a certain amount of resources, causing it to be one of the main fish caught. However, in the Yamadu River, the number of
S. pseudaksaiensis has been decreasing, causing the fish to be difficult to collect [
1]. As an ecologically sensitive species of lotus in the aquatic ecosystem of this basin,
S. pseudaksaiensis was listed as a second-level key protected aquatic species in Xinjiang in 2019 [
3]. Therefore, there is an urgent need to protect and manage
S. pseudaksaiensis resources.
Population characteristics are an important basis for fishery resources, among which the age, growth, and reproduction of fish are important indicators [
4,
5]. At present, descriptions of the geographical distribution and population genetic characteristics of
S. pseudaksaiensis are limited [
6], systematic studies on its biology and other aspects are scarce, and most studies focus on age identification [
7]. Artificial domestication and breeding technology of
S. pseudaksaiensis were initially successful only in 2011 [
8], and in May 2021, artificial breeding was successfully carried out to supplement population resources through breeding and release. Knowledge of the age structure and growth of
S. pseudaksaiensis is restricted to the study conducted by Wang et al. [
9] from 2018 to 2020. In their study, the female population ranged from 2 to 16 years, the male population ranged from 1 to 13 years, and the overall growth turning point age was 18.40 years old.
With the intensification of human activities such as fishing and construction of water conservancy projects [
10,
11,
12], the populations of
S. pseudaksaiensis have declined and the survival of this species has been greatly threatened. In addition,
S. pseudaksaiensis lives in high-altitude and limited areas [
2], and due to the unique geographical location and fragile environment, the region is facing serious ecological problems. The population structure of
S. pseudaksaiensis is unstable, sexual maturity is late, and the growth rate and fecundity are low. Because of its important characteristic factors [
13,
14], it is very sensitive to habitat changes.
S. pseudaksaiensis distributed in the upper reaches of the Turks River system was selected to study its age, growth, and reproductive characteristics in order to enhance relevant biological data, further understand its population dynamics, maintain the sustainable development of fisheries, and provide a basis for the assessment and protection of fish resources [
15].
4. Discussion
This is the first time the age, growth, and reproduction of S. pseudaksaiensis in the Ili River system have been studied. Our results showed that S. pseudaksaiensis prefers to live in high-altitude cold water, its uniform growth pattern is closely related to aquatic organisms in the Ili River, and it is a slow-growing fish with a long life span. The formation of its reproductive characteristics reflects its adaptability to the habitat environment. Therefore, to enhance and sustain the reproduction and survival of S. pseudaksaiensis, strengthening water environment management and the multiplication and discharge of the Ili River is necessary.
The water replenishment sources of the Turks River are mostly glaciers and mountain snow. The altitude of the river basin is high (800–4600 m), the water temperature is low, there are many tributaries and abundant water energy resources. Water physical and chemical indexes such as water temperature, pH, conductivity, dissolved oxygen, major ions (Na
+, K
+, Cl
−), and water chemical types are suitable for plateau fish survival [
1,
29,
30]. In fish maturation, size, age, and growth are closely related to water temperature and food supply. Borwn et al. [
31] found that in
W = a
Lb, the power exponent
b is used to judge fish growth. The total
b value was 3.1095, with 3.0436 for females and 2.9759 for males, so it was determined to be an isometrically growing fish. In the Turks River, the seasonal variation in phytoplankton was obvious and consistent, and diatom biomass dominated in all seasons; the seasonal variation in zooplankton was not obvious, and spring rotifer biomass was dominant [
1]. There are many organic debris types and benthic species in Turks River water, especially hookshrimp and Chironomidae [
1,
2], which provide effective food sources for indigenous fish and cause the growth trend of
S. pseudaksaiensis to be stable and uniform. In this study,
S. pseudaksaiensis preferred to live in cold water at high altitudes, and its adaptability varied with water temperature, resulting in slow growth, which was closely related to aquatic organisms in the Turks River, etc. These factors may be the major reasons for its uniform growth. The overall plumpness coefficient of
S. pseudaksaiensis in the female group was slightly higher than in the male group. The plumpness of female fish was higher than that of male fish at 1–5 years of age, and the growth of female fish was faster than that of male fish before sexual maturity. The plumpness of female fish decreased at 6–10 years of age, increased again at 11–15 years of age, and then dropped to a stable level. This change may be closely related to sexual maturity age, reproductive energy consumption, water temperature, and food levels [
1,
7,
32].
In fish age identification, the otolith radius is often used to infer the fish age change, with some researchers believing that otolith size and fish growth are controlled by the same metabolic process [
33], and its strong correlation can be reflected by
R2. Some scholars found that the
R2 values of 16 Mediterranean deep-sea fish and European bass (
Perca fluviatilis) were all greater than 0.8 [
34,
35]. In this study, the correlation of
S. pseudaksaiensis was high (
R2 > 0.8). Although the otolith had been growing, the growth on different axes was not uniform. Renones et al. [
36] believe that with increasing fish age, the continuous deposition of otoliths to the distal surface results in unequal growth of otoliths and fish standard length. Therefore, for older fish or slow-growing fish, the accuracy of using otoliths to calculate the length of the fish body is not high.
During fish ontogeny, reproductive parameters are fitted with information from the animal phenotypic index measured in different growth periods using mathematical models [
37], and their changes follow different strategies [
38]. The female population of
S. pseudaksaiensis showed
L∞ = 737.7,
W∞ = 6853.3,
k = 0.05,
t0 = −0.59, and
ti = 22.28; and the male population showed
L∞ = 667.0,
W∞ = 4447.2,
k = 0.05,
t0 = −0.36, and
ti = 19.55. At the turning point age, the male population was obviously smaller than the female population. Compared with
S. oconnori [
24] and
S. waltoni [
39], the
ti of
S. pseudaksaiensis in this study was higher, which was related to the difference in habitat, such as the habitat altitude of the fish species [
40]. The growth coefficient
k is a key parameter in the assessment of fish stock potential [
41,
42,
43], and fish with a value of
k between 0.05 and 0.10 are slow-growing fish.
S. pseudaksaiensis had a low
k value (0.05) and a high asymptotic standard length (
L∞), indicating that it is a fish with slow growth and a long life. Comparing this with the previous study on
S. pseudaksaiensis growth, it was found that the
L∞ obtained in this study was 654.80 mm higher than that in the previous study, and the growth coefficient
k (total 0.05, female group 0.05, male group 0.05) was lower than that in the previous study on
S. pseudaksaiensis (total 0.06). The reasons may lie in the different distribution of standard length or the different composition of age.
Reproductive characteristics are not affected by only internal factors such as genetics but are also closely related to external factors such as the habitat water environment and nutrition, including spawning type and fish fecundity [
44,
45]. The sex ratio is usually related to the reproductive habits of fish, and a change in the sex ratio is always conducive to the maintenance of a certain population [
46]. Having more females than males helps to increase the population [
20]. The gonad index (
GSI) reflects the variation in gonad development and the proportion of fish energy distribution [
47,
48]. The fecundity of fish reflects its investment in reproduction and its reproductive strategy [
49,
50]. The male/female ratio of the
S. pseudaksaiensis population in this study was 0.89:1, which was not significantly different from the theoretical ratio (1:1) (
p > 0.05), but the male population was larger than the female population, indicating that males were dominant in this reproductive population, which was similar to the
S. yunnanensis [
51] population. The gonadal index of
S. pseudaksaiensis reached its peak in June, the egg diameter in June was close to the maximum, and the egg diameter distribution was “unimodal,” indicating that
S. pseudaksaiensis belongs to the synchronous spawning type, breeds once a year, and the breeding period is short, concentrated in May to July. During this period, the Turks River has optimal external conditions (high water temperature, sufficient light, etc.), which are conducive to the development and normal growth of embryos and larvae [
52]. However,
S. pseudaksaiensis lays its eggs on the bottom, on gravel and other substrates during the breeding period, and the eggs sink [
20]. Therefore, the biggest potential danger is hypoxia, as the eggs may be covered by silt and infected by microorganisms, etc. However,
Schizothorax eggs are toxic and rarely eaten by predators [
53]. Many Schizothoracinae species have the same spawning type as
S. pseudaksaiensis, such as
S. waltoni [
39],
S. oconnori [
24], and
Oxygymnocypris stewartii [
54]. The fecundity of the
S. pseudaksaiensis population was significantly higher than that of other Schizothoracinae species [
24,
39,
54], and the relationships between absolute fecundity and the standard length and body weight were consistent with the results of studies on
Oxygymnocypris stewartii [
54]. The relationship between fecundity and body length of
S. pseudaksaiensis was linear, which was similar to that of
Oxygymnocypris stewartii and
S. oonnori. The differences in nutrient intake and energy distribution of fish living in the plateau water environment with scarce food resources may lead to differences in their fertility. Therefore, comparative studies on fertility should be conducted on the basis of eliminating the effects of individual fish or weight [
55].
Growth and reproduction constitute the processes of population growth and replenishment. The growth of fish tends to ensure that the species has the longest time to reproduce, and reproduction is linked to other life events to ensure reproduction of the population [
20]. The growth of fish is divided into three stages: the rapid growth stage before sexual maturity, the stable growth stage after sexual maturity, and the growth aging stage [
56]. The larger inflection point age of
S. pseudaksaiensis indicates that the sexual maturity is older, growth is rapid before sexual maturity, and material and energy accumulation can ensure high fertility and maintain the population number. This study will be essential for the protection and reproduction of
S. pseudaksaiensis resources in natural waters and their artificial reproduction.