Osteology of Zacco pachycephalus, sensu Jordan & Evermann (1903), with special reference to its systematic position

The skeletal system of Zacco pachycephalus, sensu Jordan & Evermann (1903), is described and compared with that of other related barilines to clarify its systematic position. Osteological characters fall generally into two categories in terms of apomorphic features. The first category is exemplified by autapomorphies such as lower jaw bone shape and the number of branchiostegal rays and hypurals. The latter characters are highly unusual among Cyprinidae. The second category consists of synapomorphies linking Z. pachycephalus and Opsariichthys uncirostris. Specialization of jaw bones for a piscivorous habit is also notable. The quadrate-pterygoid fenestra of Z. pachycephalus and O. uncirostris is considered derived. We conclude that Zacco pachycephalus should be transferred from Zacco to Opsariichthys based on the present osteological analysis.     

Phylogenetic structure of Zacco platypus (Teleostei, Cyprinidae) populations on the upper and middle Chang Jiang (=Yangtze) drainage inferred from cytochrome b sequences

We examined the genetic structure and phylogenetic relationships of some Chinese populations from the Chang Jiang (=Yangtze) drainage of the cyprinid Zacco platypus. We sequenced the complete mitochondrial cytochrome b gene of 64 individuals from 6 upper and middle tributaries of the Sichuan and Hunan Provinces to assess their population structure and systematics. The combined analyses of the phylogenetic information and the population structure suggested that Chinese Z. platypus consist of four distinct mtDNA lineages which exhibit high genetic variation and haplotypic diversity (Zacco A-D). The high molecular divergence observed among Zacco A-D mtDNA lineages (TrN+I (0.76) distance, mean 8.9%+/-1.7%) and their phylogeographic structure indicate that all four lineages have evolved independently. Analysis of molecular variance (AMOVA) indicates that most of the genetic variation observed is found among the four Zacco mtDNA lineages (thetaCT = 0.94) suggesting restricted gene flow among the Chang Jiang populations. Long-term interruption of gene flow was also evidenced by thetaST values higher than 0.9 that could be favoured by the discontinuous distributions of the lineages inhabiting upper (Sichuan Province) and middle (Hunan Province) Chang Jiang tributaries. The significant correlation between the geographic and genetic distances provide support for the importance of geographic discontinuity in shaping the Zacco genetic structure. Nested clade analysis (NCA) results were congruent with phylogenetic relationships recovered and confirm the genetic distinctiveness of four independent Zacco groups. These four groups correspond to the four Zacco A-D mtDNA lineages recovered in the phylogeny and were defined by nucleotide synapomorphies permitting bootstrapped and Bayesian confidence of 95% or greater. The high level of mitochondrial sequence divergence separating all Zacco mtDNA lineages suggested that the Z. platypus populations from the Chang Jiang drainage probably correspond to four different species.     

丹江口水库三种鲤科鱼类寄生木村小棘吻虫的季节动态

作者于2004年2月至2005年2月调查了丹江口水库木村小棘吻虫Micracanthornhynchina motomurai(Harada,1935)感染三种小型经济鲤科鱼类--马口鱼(Opsariichthys uncirostris)、宽鳍(鱼鼠)(Zacco platypus)和油(粲鱼)(Hemiculter bleekeri bleekeri)的季节动态情况.在感染率和感染丰度方面,寄生于马口鱼和油(鱼鼠)的木村小棘吻虫呈现出显著的季节变化:秋冬季节感染率和感染丰度都比较高,而春夏季节比较低;与此不同的是,寄生于宽鳍(鱼鼠)中的木村小棘吻虫则全年都维持着较稳定的感染水平;木村小棘吻虫种群在三种宿主体内的总体感染丰度差异显著,但都呈聚集分布,三者的方均比与感染率和感染丰度之间无显著相关关系.     

Influence of glaciation on divergence patterns of the endemic minnow, Zacco pachycephalus, in Taiwan

The genetic subdivision of the endemic minnow, Zacco pachycephalus, in 12 rivers of Taiwan was studied by allozyme electrophoresis. Among 26 loci surveyed, six were significantly differentiated among sites. Allozyme genotype frequencies within samples accord with Hardy-Weinberg expectations. A highly divergent genetic structure among sites (FST = 0.497) with low genetic variability within samples (H = 0.006-0.062, mean = 0.03) suggests that local populations originated from a small number of founders. UPGMA and Fitch trees derived from Nei's genetic distance and hierarchical analysis of FST values reveal that samples of Z. pachycephalus can be divided into northern, middle, and southern subgroups and that most genetic variation (87%) is distributed among rather than within groups. It is proposed that there were two isolated refugia during the most recent glacial period, and that populations of the southern group (Group I) originated from a southern refugium in Taiwan below the boundary of the Kaoping River, and other groups (Group II), including northern and middle subgroups, originated from a refugium located on the extinct land-bridge between Taiwan and mainland China. The postglacial division between northern and middle subgroups may be due to hydrological differences between these two regions.     

Study on Centrocestus armatus in Korea. I. Infection status of Zacco platypus and Z. temminckii with the metacercariae of C. armatus

This study was performed to observe the infection rate and infection intensity of fresh water fish such as pale chub (Z. platypus) and dark chub (Z. temminckii) with the metacercariae of C. armatus. The fish were caught in several rivers and streams from June to September, 1988 and examined by artificial digestion technique. Total 370 fish were caught at 19 rivers. The metacercarial (C. armatus) infection rate and average burden in Z. platypus were 86.8% and 224, and those in Z. temminckii were 78.5% and 131. The infection rate and burden per fish were 73.4% and 32 in the fish caught from the Han river, and 88.0% and 44 from the Youngsan river. All fish caught from the Keum and the Tamjin river were infected and their mean metacercarial density was 45 and 59 respectively. The infection rate and density in fish caught from the Seomjin and Nagdong rivers were 100% and 119, and 81.0% and 348 respectively. From this study, it is confirmed that C. armatus is widely distributed along 5 major rivers in Korea.     

The fish community of a high mountain stream in Taiwan and its relation to dam design

Synopsis The fish community of a high mountain stream in Taiwan was studied from 1988 to 1990 in a 16 km section of Ta-chia river where a new dam is planned for construction. There were 16 species of fish belonging to 7 families; 15 genera were recorded. Six species are endemic to Taiwan. The dominant species wereZacco pachycephalus, Acrosscheileus paradoxus andRhinogobius brunneus. The composition of fish fauna was different between the eight study sites. Adjacent sites did not show a higher similarity than remote ones. AdultZ. pachycephalus andA. paradoxus showed seasonal aggregation behavior in the stream. At two distinct sites, Kukan and Tungmau, these two species had similar breeding and stock composition patterns with an intensified production period of juveniles from late autumn to early winter and a low, but continuous production period of juveniles in summer months. This information suggest that fishes in the river have special adaptations to the typhoon season in summer and benthic algal blooming in winter. The similarity of fish community and the independent breeding sites of endemic fishes in Kukan and Tungmau also suggest that these are important factors to consider for conservation when the flooding area of a high mountain dam is designed.     

华南长江和西江流域鲤科宽鳍鱲和马口鱼不同线粒体DNA谱系之间的体形分化

本文研究了两种鲤科鱼类,即分布于华南长江与西江的宽鳍鱲和马口鱼的不同线粒体DNA谱系之间的体形差异。近期基于线粒体DNA的这两个物种的系统发育分析认为这两个类元内包含多个谱系(宽鳍鱲A-D和马口鱼1-5)。本文采用几何形态度量学的方法研究了这些不同线粒体DNA谱系间是否存在体形方面的差异。在宽鳍鱲的4个不同的线粒体DNA谱系中,有3个被纳入本研究内容。结果显示:在宽鳍鱲的3个线粒体DNA谱系中,2个具有体形的差异。在马口鱼的5个线粒体DNA谱系中,3个被纳入本文的研究。结果显示所有3个谱系均存在体形差异。因此,线粒体DNA的谱系差异很大程度上表现在谱系间的体形差异方面,表明线粒体DNA谱系可能对应于不同的物种。     

Infection status of pond smelts, Hypomesus olidus, and other freshwater fishes with trematode metacercariae in 6 large lakes

In order to determine the current infection status of pond smelts, Hypomesus olidus, and other freshwater fishes with trematode metacercariae, a total of 4,861 pond smelts and 18 other freshwater fishes, collected from the Soyang and Uiam Lakes in Gangwon-do, the Unam Lake in Jeollabuk-do, the Jangseong Lake in Jeollanam-do, the Uirim-ji (lake) in Chungcheongbuk-do, and the Andong Lake in Gyeongsangbuk-do, were individually digested with 1% pepsin-HCl and examined under a dissecting microscope. In all pond smelts caught from the 6 lakes, we were unable to detect any known human infectious trematode metacercariae in Korea. However, in other freshwater fishes, such as, Squalidus japonicus coreanus (Unam Lake), and Zacco platypus (Jangseong Lake) and Opsariichthys uncirostris amurensis (Jangseong Lake), metacercariae of human-infecting trematodes, i.e., Clonorchis sinensis and Metagonimus sp. were detected, respectively.     

Developmental morphology of the Indian cyprinid fish Barilius canarensis

Embryonic and larval development of an Indian cyprinid fish, Barilius canarensis, is described from laboratory-reared specimens. The eggs, measuring 2.1–2.4 mm in diameter, were demersal, almost spherical in shape, transparent and unpigmented, with a pale yellow yolk without oil globules. Hatching occurred 39–45 h after fertilization at 26.8°–27.4°C. The newly hatched larvae, measuring 4.8–5.1 mm in body length (BL) with 22 + 17 = 39 myomeres, were characterized by melanophores already deposited on the eyes. The eggs of B. canarensis resembled those of the related danionin species Candidia barbatus, Opsariichthys uncirostris uncirostris, Zacco platypus, Z. sieboldii, and Z. temminckii. Although the larvae of B. canarensis were also similar to those of the foregoing species in general morphology, they differed in having a straight notochord tip and pigmentation on the eyes at hatching and the almost entire absence of melanophores on the ventral body surface from the yolk sac to postflexion larval stages. Conversely, melanophores occurred on the anterior abdominal and pericardial cavities from the preflexion to postflexion larval stages.     

Negative effect of ayu on the growth of omnivorous pale chub in experimental pools

The growth rate of pale chub Zacco platypus was negatively correlated with the number and biomass of ayu Plecoglossus altivelis in experiment pools regardless of their body weight. Both species were omnivorous, feeding upon both benthic algae and invertebrates; ayu fed principally upon algae whereas pale chub consumed invertebrates. Pale chub shifted their moving tactics from bottom moving to surface moving or sit-and-wait when they coexisted with numerous ayu, presumably due to occasional attack by ayu. It seems likely that the reduced pale chub growth rate was due to interspecific aggression and exploitation of benthic algae and invertebrates by ayu.     

AFLPs congruent with morphological differentiation of Asian common minnow Zacco (Pisces: Cyprinidae) in Taiwan

Morphological and amplified fragment length polymorphism (AFLP) analyses were used to examine the evolutionary relationships among Asian common minnows ( Zacco spp.) in Taiwan, where four morphotypes were recently reported. Congruent results from the two sets of analyses indicate that the four morphotypes represent four separate morphometric clades with distinct AFLPs. Consequently, we conclude that four species exist in Taiwan and the systematics of Zacco should be revised accordingly. We also discuss conservation implications and offer a key to the four proposed Zacco species.     

北京地区宽鳍鱲的早期发育

作者对分布于北京市怀柔区怀九河的宽鳍鱲(Zacco platypus)的胚胎发育及仔鱼前期发育进行了研究,过程中采用人工干法授精获取受精卵, 观察并描述了宽鳍鱲早期发育过程及其特点。结果显示, 宽鳍鱲成熟卵呈圆球状, 平均卵径1.04 mm, 为沉性卵。在平均23.0℃(17.1-28.0℃)水温条件下, 从受精卵到孵化经历了73h1min, 积温为1682.3 度时; 孵化后6.5d 进入弯曲期仔鱼。仔鱼前期发育速度与出膜前相比明显减慢;弯曲期仔鱼出现大量死亡可能与有限的人工培育条件、混合营养期能量供给不足等原因有关。通过比较发现, 宽鳍鱲与鲤科中其他21 个种相比, 早期发育时间比其中7 个种均长。宽鳍鱲早期发育时间比同域分布、相同发育水温的马口鱼略长, 明显长于同属的纵纹鱲的发育时间。宽鳍鱲南、北方种群仔鱼发育速度存在差异, 北方种群出膜前发育速度比南方种群快, 但出膜后发育速度减慢。       

Mitochondrial phylogeny reveals the artificial introduction of the pale chub Zacco platypus (Cyprinidae) in Taiwan

The presence of the pale chub Zacco platypus (Japanese name, Oikawa) in Taiwan has been suggested to be a result of its inadvertent introduction from Lake Biwa in Japan in the 1980s in conjunction with the Japanese Ayu, Plecoglossus altivelis altivelis, which was released several times into the Tamsui River to restock the extinct Ayu population of Taiwan. However, it is also possible that Z. platypus is native to Taiwan and has not been previously described for reasons of its narrow range. Knowledge of the colonizing history of Z. platypus is of considerable importance because it provides insight into the evolutionary process and, hence, impacts management decisions regarding this species in Taiwan. A portion of the mitochondrial D-loop was sequenced for 77 specimens from five populations of Z. platypus from Japan and Taiwan. A total of 22 haplotypes were identified, and nucleotide divergence among haplotypes ranged from 0.20% to 2.82%. Haplotype diversity was high in all populations examined, with a range from 0.718 in the Tagiri River population to 0.909 in the Lake Biwa population. Phylogenetic and statistical parsimony analyses of the molecular data revealed a close genetic relationship between Taiwanese and Japanese Z. platypus and supported the previous report that the Taiwanese Z. platypus originated in Lake Biwa in Japan.     

Isolation and cross‐species amplification of microsatellite loci in the freshwater minnow Zacco pachycephalus (Teleostei: Cyprinidae) for diversity and conservation genetic analysis

Asian minnows of the genus Zacco are dominant fish in various freshwater ecosystems. Two species, Zacco pachycephalus and Z. platypus, occur in Taiwan and are favourite targets for local sport anglers. The introduction of Zacco fish into nonindigenous habitats in Taiwan has become a conservation issue. We developed eight polymorphic microsatellites for Z. pachycephalus (average H_E = 0.779) and these microsatellites were applicable to Z. platypus, which showed a comparable polymorphic level (average H_E = 0.784). These loci can be used as genetic markers for identifying conservation units and studying population differentiation for both Zacco species.     

Molecular characterization of Calocedrus rupestris Aver., H.T. Nguyen & L.K. Phan, 2008 (Cupressaceae) based on ITS1 partial sequence

Calocedrus rupestris Aver., H.T. Nguyen & L.K. Phan was described in 2008 based on some morphological characters that were not sufficiently significant to discriminate it as a species distinct from C. macrolepis Kurz. We applied a new approach to resolve these conflicting views by using sequence data from DNA (ITS) to elucidate phylogenetic relationships between the two species. Analyses of a partial ITS1 sequence in 5 individuals of 2 subpopulations of C. macrolepis and 18 individuals of 8 subpopulations of C. rupestris collected in Vietnam were done. Molecular characterization of the two species showed its low divergence with the lack of autapomorphic characters. In addition, the ITS1 partial sequences of some C. rupestris individuals were identical with C. macrolepis. Due to the less distinctive morphology between C. rupestris and C. macrolepis, the divergence between them does not exceed the interspecific levels, and therefore, C. rupestris could not be regarded as an independent species in relation to C. macrolepis but only as one of its varieties, C. macrolepis var. rupestris (Aver., H.T. Nguyen & L.K. Phan) L.K. Phan, Long K. Phan & Aver.     

Clonorchis sinensis metacercarial infection in the pond smelt Hypomesus olidus and the minnow Zacco platypus collected from the Soyang and Daechung Lakes

The pond smelt Hypomesus olidus and minnow Zacco platypus were collected from the Soyang and Daechung Lakes in January 2003, and their metacercarial infections was examined by the muscle compression and artificial digestion techniques. In the Soyang Lake, 161 metacercariae of Clonorchis sinensis (0.35 per fish) were harvested from 459 pond smelts examined. Also, 13 metacercariae of C. sinensis (0.43 per fish), 1 of Metagonimus sp., 4 of Echinostoma sp., 148 of Centrocestus armatus and 44 unidentified species were collected from 30 minnows. In the Daechung Lake, 369 metacercariae of C. sinensis (3.69 per fish) and 51 unidentified species were recovered from 100 pond smelts. The metacercariae of C. sinensis were fed to experimental rats, in which the adult flukes were identified. The pond smelts and minnows collected from the Soyang and Daechung Lakes were verified to be the second intermediate hosts and the sources of human C. sinensis infection.     

徽水河宽鳍的年龄、生长和繁殖

认识鱼类的生活史特征及其对生态环境变化的响应,是鱼类物种保护与资源合理利用的基础。于2009年5月至2010年4月研究了黄山地区徽水河中宽鳍鱲(Zacco platypus)的年龄、生长和繁殖。共采集标本352尾,雌雄性比为0.49∶1,与1∶1差异显著。以鳞片为年龄鉴定材料,雌、雄个体的最大寿命均为3龄;年轮主要形成于3月份。全长和体重呈幂函数关系,两性间无显著性差异,关系式表达为W=6×10–6L3.10;全长和鳞径呈线性关系,且两性间差异显著,关系式分别表达为L♀=29.58R+38.84和L♂=33.17R+34.99;2龄个体的退算全长在雌、雄两性间无显著性差异。繁殖时间为4—7月份;2龄时达50%初次性成熟,其个体全长为98.22(雌)和105.69 mm(雄);绝对繁殖力为(758±362)卵粒,相对繁殖力为(77.38±22.15)卵粒/g。同已有的少量研究资料相比较,徽水河宽鳍鱲的年龄结构较北京地区种群的相对简单,个体生长较同为黄山地区的浦溪河种群相对快速,这种生活史特征差异可能是生态环境空间异质性的作用结果,但有关宽鳍鱲各生活史特征之间的权衡及其对生态环境的响应还有待于进一步研究。     

Population structure and phylogeography of Aphyocypris kikuchii (Oshima) based on mitochondrial DNA variation

Aphyocypris kikuchii is a cyprinid species endemic to northern and eastern Taiwan and is the only primary freshwater fish native east of the Coastal Mountain Range. In total, 92 individuals of A. kikuchii from seven populations in three regions of the island were surveyed for mitochondrial DNA (mtDNA) variation. High haplotype diversity ( h = 0·989) and low nucleotide diversity ( π = 0·009) of mtDNA were detected. Negative values of Tajima's D and unimodal mismatch distributions probably reflect a history of recent demographic expansions from small populations. Three major haplotype clusters displayed geographically non-overlapping distributions, indicating a long-term isolation between regions. Hierarchical analysis of molecular variance showed significant genetic structuring among populations ( Φ _ST= 0·66). Significant haplotype heterogeneity was also detected among populations within regions ( Φ _SC= 0·41, P < 0·001) and among regions ( Φ _CT= 0·43, P < 0·05). Molecular clock estimates of coalescence in the three major mtDNA lineages indicated coalescence in the most recent common ancestor c. 0·11–0·39 million years ago. Haplotypes of cluster B nested as interior nodes in the haplotype network, indicating that migrations from Shueilian (SL) populations to the northern region (cluster A) and to the eastern region (cluster C) may have occurred independently. Lineages A and B + C should be managed as two distinct evolutionarily significant units, while the northern, SL and southern groups should be managed as separate management units.     

Differentiation of golden-ringed dragonfly Anotogaster sieboldii (Selys, 1854) (Cordulegastridae: Odonata) in the insular East Asia revealed by the mitochondrial gene genealogy with taxonomic implications

Molecular phylogeographical analyses of Anotogaster sieboldii (Selys, 1854) were conducted to reveal the differentiation process of insular populations. The gene genealogy based on 845 bp of the mitochondrial genes (cytochrome oxidase subunit I and subunit II) indicated that A. sieboldii includes two deeply separated lineages. These two major lineages seem to have differentiated in Miocene before the formation of the insular East Asia. One lineage includes three inner clades that correspond to the populations of northern area (the Japanese main islands, Korean Peninsula, Yakushima), Amamioshima and Okinawajima. Populations of Central Ryukyu, including Amamioshima and Okinawajima, might have been divided from the northern populations in early Pleistocene. The other major lineage includes populations of the Yaeyama Group, Taiwan and East China. The former two populations were reconstructed as a reciprocal monophyletic group. Populations of Taiwan and Yaeyama Groups would have been separated from the continental ones in Pleistocene. These two highly divergent lineages should be recognized as distinct species. Furthermore, the mitochondrial lineages revealed six genetically distinct and geographically isolated assemblages: (1) northern populations, (2) Amamioshima, (3) Okinawajima, (4) Yaeyama Group, (5) Taiwan and (6) East China.