Genetic analysis of floating Enteromorpha prolifera in the Yellow Sea with AFLP marker

Extremely large accumulation of green algae Enteromorpha prolifera floated along China’coastal region of the Yellow Sea ever since the summer of 2008.Amplified Fragment Length Polymorphism (AFLP) analysis was applied to assess the genetic diversity and relationships among E. prolifera samples collected from 9 affected areas of the Yellow Sea.Two hundred reproducible fragments were generated with 8 AFLP primer combinations,of which 194 (97%) were polymorphic. The average Nei’s genetic diversity, the coefficiency of genetic differentiation (Gst), and the average gene flow estimated from Gst in the 9 populations were 0.4018, 0.6404 and 0.2807 respectively. Cluster analysis based on the unweighed pair group method with arithmetic averages (UPGMA) showed that the genetic relationships within one population or among different populations were all related to their collecting locations and sampling time. Large genetic differentiation was detected among the populations.The E. prolifera originated from different areas and were undergoing a course of mixing.     

Population genetic structure of the mantis shrimp Oratosquilla oratoria(Crustacea: Squillidae) in the Yellow Sea and East China Sea

The mantis shrimp O ratosquilla oratoria is an ecologically and economically important species in the Western Pacific. In present study, the population genetic structure of O ratosquilla oratoria from the Yellow Sea and East China Sea was examined with mitochondrial DNA control region sequences. In total, 394 samples were collected from 18 locations and 102 haplotypes were obtained. For the Yellow Sea, the overall nucleotide diversity and haplotype diversity were 0.006 9 and 0.946 8, respectively; while across all the East China Sea locations, the overall nucleotide diversity and haplotype diversity were 0.027 94 and 0.979 0, respectively. The results of AMOVA and pairwise F_(ST)(0.145 2, P 0.001) revealed moderate differentiation between the Yellow Sea and East China Sea populations of O. oratoria. However, neither the neighbor-joining tree nor haplotype network showed clades with geographic pattern, which indicated considerable gene flow was existed between the Yellow Sea and East China Sea, and supporting the high larval dispersal ability in this species. Mismatch distribution analysis and neutrality tests suggested that O. oratoria has undergone population expansion event, and the Pleistocene glacial cycles might have an impact on the historical demography of O. oratoria. The genetic information obtained in this study can provide useful information for sustainable improvements for capture fisheries management strategies.     

Genetic diversity in two Japanese flounder populations from China seas inferred using microsatellite markers and COI sequences

Japanese flounder is one of the most important commercial species in China; however, information on the genetic background of natural populations in China seas is scarce. The lack of genetic data has hampered fishery management and aquaculture development programs for this species. In the present study, we have analyzed the genetic diversity in natural populations of Japanese flounder sampled from the Yellow Sea (Qingdao population, QD) and East China Sea (Zhoushan population, ZS) using 10 polymorphic microsatellite loci and cytochrome c oxidase subunit I (COI) sequencing data. A total of 68 different alleles were observed over 10 microsatellite loci. The total number of alleles per locus ranged from 2 to 9, and the number of genotypes per locus ranged from 3 to 45. The observed heterozygosity and expected heterozygosity in QD were 0.733 and 0.779, respectively, and in ZS the heterozygosity values were 0.708 and 0.783, respectively. Significant departures from Hardy-Weinberg equilibrium were observed in 7 of the 10 microsatellite loci in each of the two populations. The COI sequencing analysis revealed 25 polymorphic sites and 15 haplotypes in the two populations. The haplotype diversity and nucleotide diversity in the QD population were 0.746±0.072 8 and 0.003 34±0.001 03 respectively, and in ZS population the genetic diversity values were 0.712±0.047 0 and 0.003 18±0.000 49, respectively. The microsatellite data (F_st =0.048 7, P<0.001) and mitochondrial DNA data (F_st =0.128, P<0.001) both revealed significant genetic differentiation between the two populations. The information on the genetic variation and differentiation in Japanese flounder obtained in this study could be used to set up suitable guidelines for the management and conservation of this species, as well as for managing artificial selection programs. In future studies, more geographically diverse stocks should be used to obtain a deeper understanding of the population structure of Japanese flounder in the China seas and adjacent regions.     

The loss of genetic diversity during captive breeding of the endangered sculpin, Trachidermus fasciatus, based on ISSR markers: implications for its conservation

Inter-simple sequence repeat (ISSR) markers were used to determine the genetic variation and genetic differentiation of cultured and wild populations of Trachidermus fasciatus, an endangered catadromous fish species in China. Six selected primers were used to amplify DNA samples from 85 individuals, and 353 loci were detected. Relatively low genetic diversity was detected in the cultured population (the percentage of polymorphic loci PPL = 73.80%, Nei’s gene diversity h = 0.178 2, Shannon information index ...     

Genetic differentiation of Japanese sardinella (Sardinella zunasi) populations in the Northwest Pacific revealed by ISSR analysis

Knowledge of population genetic structure plays an important role in fisheries management.In this research,In-ter-Simple-Sequence-Repeat(ISSR)markers were employed to evaluate the genetic structure of Japanese sardinella(Sardinella zun-asi)populations in the Northwest Pacific.Eighty seven individuals from 5 locations were screened using 4 highly polymorphic prim-ers.A total of 173 polymorphic loci were detected out of 191 loci amplified.Small but significant genetic differentiation was de-tected between the Chinese and Japanese populations by both AMOVA and pairwise FST analyses,which was further supported by cluster analysis.We consider that climate change during glaciations should be responsible for the genetic differentiation.Isolation by geographic distance among populations was observed,indicating that the distance might also lead to the genetic differentiation.However,no genetic structure was found within the populations off both the Chinese and Japanese coasts,indicating a high-level along-coast gene flow,which might result from ocean current transport and common ground for over-wintering.     

Distribution of heavy metals and nutrients in rainwater in coastal regions between the southern Yellow Sea and East China Sea

Rainwater samples were collected in series in Qianliyan Island (southern Yellow Sea) and Shengsi Archipelago (East China Sea) between May 2000 and May 2002, chemical analysis for pH values,concentrations of heavy metals (Cu, Pb, Zn and Cd) and nutrients (NH4^-, NO3^-, PO43^-, SiO32^-) were performed.Results indicate that concentrations of most of the heavy metals and nutrients in rainwater show clear seasonal variation, i.e. high level in winter and low level in summer. Regionally, concentrations are higher in the southern Yellow Sea than in the East China Sea, but the annual input of heavy metals into oceans by wet deposition is similar in both stations. However, the input of nutrients by wet deposition in the East China Sea is 2-3 times higher than that in the southern Yellow Sea. In individual, Pb and PO4^3- are input to the sea mainly by dry deposition; whereas Cu, Zn, Cd and N compounds are input dominantly by wet deposition, the N/P ratios in the rainwater from two stations are much higher than those in seawater, showing a significant impact of atmospheric wet deposition on marine production and biogeochemical circulation of nutrients in these sea regions.     

Molecular identification and population differentiation of Aurelia spp. ephyrae in sea cucumber aquaculture ponds of northern China

Aurelia spp. ephyrae have been reported to form blooms in sea cucumber aquaculture ponds in the Bohai and Yellow Seas. To identify the species, we carried out a genetic analysis of Aurelia spp. ephyrae and medusae based on mitochondrial 16 S rRNA gene. Samples offour Aurelia sp. ephyrae populations were collected in sea cucumber aquaculture ponds and samples offour Aurelia sp. medusae populations were collected in coastal waters. Using a BLASTn search, we found that both the ephyrae collected in the aquaculture ponds and medusae collected in coastal waters belong to Aurelia coerulea. Seventeen haplotypes were recovered from the 16 S rRNA gene. The overall haplotype diversity and nucleotide diversity of the 166 A. coerulea individuals were 0.686% and 0.329%, respectively, indicating high haplotype diversity and low nucleotide diversity. Moreover, the haplotype diversity of ephyrae populations were generally lower than that of medusae populations with close sampling points. The genetic differentiation between ephyrae populations collected in the sea cucumber aquaculture ponds and A. coerulea medusae collected in coastal waters was not significant, suggesting the ephyrae populations in the sea cucumber culture ponds were part of the same genetic group as the medusae populations in the coastal waters. Phylogeographic analysis of the 16 S rRNA region revealed that there was no significant correlation between the haplotypes and the geographic distribution of populations. Pairwise fixation index values showed significant genetic differentiation and limited gene flow between A. coerulea population of Weifang and other locations.     

Biomarker evidence for paleoenvironmental changes in the southern Yellow Sea over the last 8200 years

Previous studies have indicated that the Yellow Sea underwent significant environmental changes during the Holocene, but many questions remain concerning the timing of the establishment of the modern circulation system, which would have major implications for the Yellow Sea ecosystem and carbon cycle. In this study, marine and terrestrial biomarkers were analyzed in Core YE-2 from a muddy area in the southern Yellow Sea to reconstruct Holocene environmental and phytoplankton community change. The content of three individual marine phytoplankton biomarkers (alkenones, brassicasterol and dinosterol) all display a similar trend, and their total contents during the early Holocene (362 ng/g) were lower than those during the mid-late Holocene (991 ng/g). On the other hand, the contents of terrestrial biomarkers (C27+C29+C31n-alkanes) during the early Holocene (1 661 ng/g) were about three times higher than those during the mid-late Holocene (499 ng/g). Our biomarker results suggest that the modern circulation system of the Yellow Sea was established by 5-6 ka, and resulted in higher marine productivity and lower terrestrial organic matter inputs. Biomarker ratios were used to estimate shifts in phytoplankton community structure in response to mid-Holocene (5-6 ka) environmental changes in the Yellow Sea, revealing a transition from a dinoflagellate-diatom dominant community structure during the early Holocene to a coccolithophore-dominant community structure during the mid-late Holocene.     

Ulva and Enteromorpha (Ulvaceae, Chlorophyta) from two sides of the Yellow Sea: analysis of nuclear rDNA ITS and plastid rbcL sequence data

Ulvacean green seaweeds are common worldwide; they formed massive green tides in the Yellow Sea in recent years, which caused marine ecological problems as well as a social issue. We investigated two major genera of the Ulvaceae, Ulva and Enteromorpha, and collected the plastid rbcL and nuclear ITS sequences of specimens of the genera in two sides of the Yellow Sea and analyzed them. Phylogenetic trees of rbcL data show the occurrence of five species of Enteromorpha (E. compressa, E. flexuosa, E. intestinalis, E. linza and E. prolifera) and three species of Ulva (U. pertusa, U. rigida and U. ohnoi). However, we found U. ohnoi, which is known as a subtropical to tropical species, at two sites on Jeju Island, Korea. Four ribotypes in partial sequences of 5.8S rDNA and ITS2 from E. compressa were also found. Ribotype network analysis revealed that the common ribotype, occurring in China, Korea and Europe, is connected with ribotypes from Europe and China/Japan. Although samples of the same species were collected from both sides of the Yellow Sea, intraspecific genetic polymorphism of each species was low among samples collected worldwide.     

东、黄海鲐鱼群体遗传差异的RAPD分析

利用随机扩增多态DNA(RAPD)技术分别分析了五岛西部日本鲐、东海西部日本鲐和澳洲鲐三群体的样本,共产生226个分子标记,其中143个为多态性标记,多态百分率达到63.3%,每条引物产生的平均标记数为7.29。五岛西部日本鲐、东海西部日本鲐和澳洲鲐群体内遗传相似度分别为0.949、0.877和0.936,而群体间遗传距离依次为:五岛西部日本鲐-东海西部日本鲐(0.2357)<东海西部日本鲐-澳洲鲐(0.3030)<五岛西部日本鲐-澳洲鲐(0.3528),五岛西部日本鲐与东海西部日本鲐个体在UPGMA系统树独立聚类为两个类群,初步支持东、黄海的日本鲐存在东海西部和五岛西部的两个种群的观点。Shannon多样性指数表明,其遗传多样性大小依次为东海西部日本鲐、澳洲鲐、五岛西部日本鲐。     

Distribution of diatoms and silicoflagellates in surface sediments of the Yellow Sea and offshore from the Changjiang River,China

The spatial distribution of siliceous microfossils (diatoms and silicoflagellates) in the surface sediments was mapped at 113 sites in the Yellow Sea and sea areas adjacent to the Changjiang (Yangtze) River, China. In total, 267 diatom taxa and two silicoflagellate species were identified from the sediments. The spatial variations in abundance and diversity were classified into three distinct geographic patterns using Q mode clustering: a south-north geographic pattern, a coastal-offshore pattern and a unique pattern in the Changjiang River mouth. The south-north geographic pattern was related to the spatial variations in sea temperature. Coscinodiscus oculatus, a warm-water species, indicated these variations by a gradual decrease in abundance from the south to the north. The coastal-offshore pattern was in response to the spatial variations in salinity. Cyclotella stylorum, Actinocyclus ehrenbergii and Dictyocha messanensis, the dominant brackish species in coastal waters, significantly decreased at the isobaths of approximately 30 m, where the salinity was higher than 31. Paralia sulcata and Podosira stelliger indicated the impact of the Yellow Sea Warm Current in the central Yellow Sea. The unique pattern in the Changjiang River mouth showed the highest species diversity but lower abundance, apparently because: freshwater input can significantly increase the proportion of brackish species; nutrients can supply the growth of phytoplankton; and high sedimentation rates can dilute the microfossil abundance in the sediments. Our results show that an integration of environmental factors (e.g., nutrient levels, sedimentation rate, sea temperature, salinity and water depth) determined the spatial characteristics of the siliceous microfossils in the surface sediments.     

Spatiotemporal variation in community structure of marine benthic ciliates in the Yellow Sea during and after macroalgal and giant jellyfish blooms

The annual bloom of the green macroalgal Ulva prolifera from May through July since 2008 and another of giant jellyfish Nemopilema nomurai from June through September have been frequent events in the Yellow Sea. However, the patterns of benthic ciliate communities during and after the blooms are still not known. In combination with analyses of benthic environmental factors, we investigated the distribution and community composition of benthic ciliates in the Yellow Sea in July and November 2011. In July, ciliates had high standing crops and diversity in the northern Yellow Sea, and in the inshore area off the southern Shandong Peninsula, where large numbers of green macroalgae accumulated. In November, the abundance, biomass and diversity of ciliates were high in the sea areas off the Shandong Peninsula and Changjiang estuary, where a large quantity of jellyfish occurred in August. Neither the abundance nor the biomass had significant difference between seasons, or between different compartments of the Yellow Sea. The species number, and both Margalef and Shannon-Wiener indices of ciliates were all significantly higher in November than in July. In both seasons, prostomateans and karyorelicteans consistently constituted the first and second most important ciliate groups in biomass; and carnivorous ciliates constituted the primary feeding type in terms of biomass as well as species richness, followed by bacterivores, algivores and omnivores. Compared with that in June 2007 when no macroalgae occurred, the percentage of small-sized bacterivores (e.g. Metacystis spp., Euplotes spp. and scuticociliates) increased in July 2011. The proportion of carnivorous ciliates increased in November, and this increased dominance of carnivorous ciliates may be a response to the increase in predominance of heterotrophic nanoflagellates, which might in turn be ascribed to an effect of green macroalgal and giant jellyfish blooms in the Yellow Sea.     

Relocation of the Yellow River estuary in 1855 AD recorded in the sediment core from the northern Yellow Sea

Relocation of the Yellow River estuary has significant impacts on not only terrestrial environment and human activities, but also sedimentary and ecological environments in coastal seas. The responses of regional geochemical characteristics to the relocation event, however, have not been well studied. In the present study, we performed detailed geochemical elemental analyses of a sediment core from the northern Yellow Sea and studied their geochemical responses to the 1855 AD relocation of the Yellow River estuary. The results show that TOC/TN, Co/Al₂O₃, Cr/Al₂O₃, Ni/Al₂O₃ and Se/Al₂O₃ ratios all decreased abruptly after 1855 AD, and similar decreases are observed in the sediments of the mud area southwest off the Cheju Island. These abrupt changes are very likely caused by the changes in source materials due to the relocation of the Yellow River estuary from the southern Yellow Sea to the Bohai Sea, which the corresponding decreasing trends caused by the changes in main source materials from those transported by the Liaohe River, the Haihe River and the Luanhe River to those by the Yellow River. Because the events have precise ages recorded in historical archives, these obvious changes in elemental geochemistry of sediments can be used to calibrate age models of related coastal sea sediments.     

Analysis of the genetic diversity and differentiation of <Emphasis Type="Italic">Fenneropenaeus penicillatus</Emphasis> populations using AFLP technology

Fenneropenaeus penicillatus (redtail shrimp) is an important marine commercial animal in China. Recently, its resources have been depleted rapidly as a result of, for example, over-exploitation and environmental degradation of spawning grounds. Therefore, we analyzed the genetic diversity and differentiation of nine wild populations of F. penicillatus of China (Ningde, Lianjiang, Putian, Xiamen, Quanzhou, Zhangpu, Dongshan, Nanao, and Shenzhen populations) by amplified fragment length polymorphism (AFLP) technology, to provide genetic information necessary for resource protection, rejuvenation, artificial breeding, and sustainable use of the resource. Eight AFLP primer pairs were used for amplification, and 508 bands were detected among the populations. The results show that the percentage of polymorphic loci (P) ranged from 41.34% to 63.58%; the Nei’s gene diversity (H) of the populations was 0.119 4-0.230 5; and Shannon’s Information Index (I) was 0.184 1-0.342 5. These genetic data indicate that the genetic diversity of F. penicillatus was high. The genetic differentiation coefficient (GST=0.216 2) and gene flow (Nm=1.812 4) show that there was a high level of genetic differentiation and a moderate level of gene flow among populations. More studies on the genetic differentiation mechanism of F. penicillatus along the south-eastern coast of China need to be conducted to find more effective scientific protection strategies for the conservation of F. penicillatus genetic resources.     

Microsatellite variation in the oyster <Emphasis Type="Italic">Crassostrea plicatula</Emphasis> along the coast of China

Genetic diversity and differentiation of the oyster Crassostrea plicatula populations from China’s coast were studied based on seven microsatellite loci. All loci showed high polymorphism for all five C. plicatula populations, with an average number of allele per locus of 19.3–27.9 and an average expected heterozygosity of 0.889–0.952. Significant departures from Hardy-Weinberg equilibrium and deficits of heterozygotes were observed over most populations at each locus, which were fully explained by null alleles. Microsatellite analysis revealed significant subdivision in the C. plicatula populations. According to the neighbor-joining tree constructed on the basis of the D _A distance, the five populations fell into three regional groups, showing a relatively homogeneous genetic structure in geographically close populations. Assignation tests correctly assigned high percentages of individuals to their original populations and groups, and also confirmed the existence of genetic differentiation among C. plicatula populations. The results obtained in this study will facilitate the formulation of appropriate fisheries management programs, stock identification and conservation of biodiversity for the species.     

Heterosis and combining ability: a diallel cross of three geographically isolated populations of Pacific abalone Haliotis discus hannai Ino

We conducted a complete diallel cross among three geographically isolated populations of Pacific abalone Haliotis discus hannai Ino to determine the heterosis and the combining ability of growth traits at the spat stage. The three populations were collected from Qingdao (Q) and Dalian (D) in China, and Miyagi (M) in Japan. We measured the shell length, shell width, and total weight. The magnitude of the general combining ability (GCA) variance was more pronounced than the specific combining ability (SCA) variance, which is evidenced by both the ratio of the genetic component in total variation and the GCA/SCA values. The component variances of GCA and SCA were significant for all three traits (P<0.05), indicating the importance of additive and non-additive genetic effects in determining the expression of these traits. The reciprocal maternal effects (RE) were also significant for these traits (P<0.05). Our results suggest that population D was the best general combiner in breeding programs to improve growth traits. The DM cross had the highest heterosis values for all three traits.     

Phase evolution of Holocene paleoenvironmental changes in the southern Yellow Sea: Benthic foraminiferal evidence from core C02

Assemblages of benthic foraminifera in a sediment core (C02) near the western margin of the southern Yellow Sea Mud were studied to decipher the phase evolution of Holocene paleoenvironmental changes associated with the Holocene marine transgression. It appears that during the early Holocene (11.2–10.1 kyr BP), the faunal was dominated by low salinity and shallow water species Cribrononion subincertum, Buccella frigida and Ammonia beccarii, reflecting a near coast depositional environment. A rapid increase of the relative abundance of Ammonia compressiuscula between 10.1–9.3 kyr BP indicates that the sea level rose rapidly during that time period. From 9.3–7.7 kyr BP, the benthic foraminiferal assemblage was dominated by high percentage of A. compressiscula, suggesting that the sea level was relatively stable. An obvious transition of benthic foraminifera, from the A. compressiuscula-dominated assemblage to an Ammonia ketienziensis-dominated assemblage, occurred between 7.7–6.2 kyr BP, possibly corresponding to a second sea level rapid rise period in the Yellow Sea during the Holocene. This transition may correspond to the gradually strengthened Yellow Sea warm current (YSWC) and finally is established the modern-type circulation in the Yellow Sea. It may also mark the formation of the Yellow Sea cold bottom water (YSCBW) during that period. Since then, the benthic foraminiferal assemblage based on core C02 was dominated by typical YSCBW species, A. ketienziensis, Astrononion italicum and Hanzawaia nipponica, at 6.2–4 kyr BP. A non-deposition period occurred since ～4 kyr BP, which possibly related to the hydrology changes caused by the East Asia monsoon. The two obvious benthic foraminiferal transitions recorded in core C02 during the early and middle Holocene provide evidence that the Yellow Sea has undergone a two-phase rapid sea level rise during the Holocene marine transgression.     

Significant population genetic structure detected in the rock bream <Emphasis Type="Italic">Oplegnathus fasciatus</Emphasis> (Temminck &amp; Schlegel, 1844) inferred from fluorescent-AFLP analysis

Oplegnathus fasciatus (rock bream) is a commercial rocky reef fish species in East Asia that has been considered for aquaculture. We estimated the population genetic diversity and population structure of the species along the coastal waters of China using fluorescent-amplified fragment length polymorphisms technology. Using 53 individuals from three populations and four pairs of selective primers, we amplified 1 264 bands, 98.73% of which were polymorphic. The Zhoushan population showed the highest Nei’s genetic diversity and Shannon genetic diversity. The results of analysis of molecular variance (AMOVA) showed that 59.55% of genetic variation existed among populations and 40.45% occurred within populations, which indicated that a significant population genetic structure existed in the species. The pairwise fixation index F _st ranged from 0.20 to 0.63 and were significant after sequential Bonferroni correction. The topology of an unweighted pair group method with arithmetic mean tree showed two significant genealogical branches corresponding to the sampling locations of North and South China. The AMOVA and STRUCTURE analyses suggested that the O. fasciatus populations examined should comprise two stocks.     

Influence of the physical environment on the migration and distribution of <Emphasis Type="Italic">Nibea albiflora</Emphasis> in the Yellow Sea

This study investigates the migration and distribution of the warm-temperate fish Nibea albiflora. Their spawning migration and wintering migratory routes within in the Yellow Sea are described in detail. Considering the main physical features and environment of the Yellow Sea, it appears to be have one wintering ground and three migratory routes from the wintering ground to the spawning grounds. The fish begin to migrate from the wintering ground to the spawning grounds in the northwest region of the Yellow Sea in late March. The Yellow Sea has three spawning grounds. The first is located near the Yalu River on the Liaodong Peninsula and the second one is located in Rushan Bay of Shandong Peninsula. The third spawning ground is located in Haizhou Bay in the southern region of the Yellow Sea. This study found that the temperature of the Yellow Sea influences the migration of N. albiflora, and that the migratory routes coincide with the thermal fronts in the sea. Nutrients for juvenile fish are taken from the coastal upwelling area. Chlorophyll is a good environmental indicator of phytoplankton biomass and thereby provides the status of biological resources. Different types of sediment in near-shore zones are also of practical significance for the growth of fish. The study of the effects of marine environments on the migration of various fishes is not only significant to the fishing industry, but can also provide a scientific basis for the understanding of the ecological implications of the relevant physical processes.     

Genetic analysis of four populations of redlip mullet (Chelon haematocheilus) collected in China seas

Horizontal starch gel electrophoresis was used to investigate genetic structures of four populations (Dandong, Rizhao, Ningbo and Guangzhou) of redlip mullet (Chelon haematocheilus). Fourteen loci of ten enzymes (G3PDH, IDHP, LAP, LDH, MDH, MPI, PGDH, PGM, SDH, and SOD) were identified. The proportions of polymorphic loci varied from 0.07 to 0.14. The average observed and expected heterozygosities ranged from 0.02 to 0.05 and from 0.02 to 0.04 respectively among the 4 populations. The average number of efficient alleles of each locus varied from 1.02 to 1.06. The genetic distance among populations was from 0.0004 to 0.0021. The genetic distances between Dandong, Rizhao and Ningbo populations were small, while those between Guangzhou population and other three populations were relatively high. The result indicated the possible divergence of redlip mullet between the South China Sea and the other sea areas of China due to geographic isolation.