Comparative phylogeography of the Yellow River schizothoracine fishes (Cyprinidae): Vicariance, expansion, and recent coalescence in response to the Quaternary environmental upheaval in the Tibetan Plateau

National Natural Science Foundation of China (NSFC) [30225008, 30300036, 30530120]; Key Innovation Plan [KSCX2-SW-106]; National Basic Research Project in China [2005cb422005]; National Natural Science Foundation of China [30600062]

Otolith microstructure of Oxygymnocypris stewartii (Cypriniformes, Cyprinidae, Schizothoracinae) in the Lhasa River in Tibet, China

Otolith microstructure of Oxygymnocypris stewartii collected from the Lhasa River was examined and described with regards to the early life history events. The monthly changes in the number of microincrements on the margin of the otolith were examined to validate the approximately daily periodicity of otolith increment formation. The microstructure of otoliths was used to detect changes in microincrement deposition patterns corresponding with events during early life. The annuli, microincrements and checks including the hatch check, yolk absorption check and several recurrent patterns in the otolith were described. Periodicity of the recurrent patterns was weekly, fortnightly and monthly. Through counting the number of the microincrements, it was confirmed that the primary growth of O. stewartii was in a period of 7 or 8 months from late March to October; it was estimated that O. stewartii might hatch between April and May.

Otolith characteristics and age determination of an endemic Ptychobarbus dipogon (Regan, 1905) (Cyprinidae: Schizothoracinae) in the Yarlung Tsangpo River, Tibet

We describe the microincrements, checks and annuli in the lapilli of the schizothoracine Ptychobarbus dipogon, an endemic species of the Tibetan plateau. We collected samples in the Yarlung Tsangpo River and its tributaries on a monthly basis (from April 2004 to August 2006). We describe the shape features of the three pairs of otoliths and document the full trajectory of lapillus development. We found that five to seven checks were clearly visible in the opaque zone of the first annulus. The pattern of 21-23 daily growth increments within each check might be explained as a lunar-induced deposition. We counted between 137 and 154 increments within the first annulus. Annuli appeared as a sequence of gradually declining increment widths, whereas false rings were characterized by abrupt checks. Our oldest estimates were 23(+)years for males and 44(+) for females. The time of annulus completion was clearly between March and April each year using monthly marginal increments analysis. We consider the factors responsible for daily increment formation as an endogenous circadian rhythm. Environmental information, such as strong sunlight and cold water temperatures in the Tibetan Plateau, could reinforce the endogenous daily cycle. Our results provided important data addressing the ecology and population dynamics of P. dipogon.

The reproductive traits of brown trout (Salmo trutta fario L.) from the Yadong River, Tibet

Some key aspects of the reproductive strategy of the brown trout (Salmo trutta fario L.) in the Yadong River, Tibet, including spawning season, age at sexual maturity, fecundity and egg size, have been studied. The majority of the samples were less than 215 mm and age ranged from 1 to 4 in both sexes, indicating that the majority of the fish were younger and the pressure by overfishing was high. The spawning periodicity was determined to be between the end of October and January, mainly in November and December. The ratio of male to female brown trout population (1.29:1 with P > 0.05) suggested no sex significant differences, although males were significantly more abundant than females in October (P < 0.0001) on monthly basis. Age and size of males and females at maturity was different and males matured earlier than females. Fecundity was markedly correlated with their body weight (P < 0.001, r = 0.9255), standard length (P < 0.01, r = 0.8879), and gonad weight (P < 0.001, r = 0.9366). The mean size of mature eggs in the spawning season was: 4.0 +/- 0.45 mm and tended to increase along with the female spawners size (P < 0.001, r = 0.9641). Further researches about the brown trout population in the Yadong River should be conducted on issues such as artificial reproduction, culture, conservation, management, and restocking.

Age structure, growth and mortality estimates of an endemic Ptychobarbus dipogon (Regan, 1905) (Cyprinidae: Schizothoracinae) in the Lhasa River, Tibet

Ptychobarbus dipogon is an endemic fish in the Yarlung Tsangpo River, but its biology is poorly known. We sampled 582 specimens (total length, TL, between 70.6 and 593.0 mm) from April 2004 to August 2006 in the Lhasa River, Tibet. We estimated ages based on the counts of alternating opaque and translucent zones (annuli) in thin transverse sections of lapilli otoliths. Ages ranged from 1(+) to 23(+) years for males and 1(+) to 44(+) for females. The observed 44(+) years was the oldest reported for schizothoracine fishes. Females attained a larger size than males. The TL weight relationship was W=7.12 x 10(-6)TL(3.006) for combined sexes. The growth parameters fitted von Bertalanffy growth functions were L-infinity = 598.66 mm, k=0.0898 year(-1), t(0)=-0.7261 year and W-infinity = 1585.38 g for females and L-infinity = 494.23mm, k=0.1197 year(-1), t(0)=-0.7296 year and W-infinity = 904.88g for males. The longevities of 32.7 year for females and 24.3 year for males were similar to the observed ages. Using an empirical model we estimated the instantaneous rate of total mortality (Z) at 0.28 per year in the lower reaches. Z in the upper and middle stocks was close to the M because of unexploited or lightly exploited stock. Protracted longevity, slow growth, low natural mortality and large body size were typical characteristics of P. dipogon. The current declining trend of P. dipogon could be prevented by altering fishing regulations.

Age and growth of Schizopygopsis younghusbandi younghusbandi in the Yarlung Zangbo River in Tibet, China

Schizopygopsis younghusbandi younghusbandi is an endemic species whose distribution is restricted to the middle reaches of the Yarlung Zangbo River, being one of the most important commercial fishes in this area. Age and growth of 606 specimens captured between October 2002 and April 2005 were studied. The range in standard length (L) was 65.7-387.3 mm and total weight (W) was 3.3-772.0 g. The relationship between L and W was W=0.000909L(2.2493) for males and W=0.000259L(2.4781) for females. Age, determined from anal scales and lapillus otoliths, ranged from 3 to 18 years. The parameters of von Bertalanffy growth functions, estimated by back-calculated length, were L-infinity = 442.7mm L, k=0.0738 year(-1) and t(0)=-1.4 year for males, and L-infinity = 471.4mm L, k=0.0789 year(-1) and t(0)=0.2 year for females. Males and females exhibited statistically significant differences in growth. chi(2)-test indicated that von Bertalanffy growth functions could well describe the growth of S. y. younghusbandi. The longevities were 39.2 and 38.2 years for males and females, respectively. Growth inflexion points were 10.2 and 12.0 years for males and females, respectively, but 84.8% of the captures were at the smaller ages. So conservation and management schemes for this population should be considered urgently. In addition, we found that populations from the upstream of the Lhasa River, the downstream of the Lhasa River and the middle reaches of the Yarlung Zangbo River showed statistically significant differences in growth patterns.

Phylogeography of Schizothorax o'connori (Cyprinidae: Schizothoracinae) in the Yarlung Tsangpo River, Tibet

Schizothorax o'connori is endemic to the Yarlung Tsangpo River on the Tibetan Plateau. We assessed the relative impacts of historical and contemporary factors in organizing genetic variation in S. o'connori populations using mitochondrial cytochrome b sequences. We analyzed 191 samples from 11 populations and identified 78 haplotypes. The phylogenetic analyses and analysis of molecular variance all supported the same conclusions of two well-differentiated east-west phylogroups, separated by the Tsangpo Great Gorge. The split between the two clades accounted for 58% of the genetic variance observed among the examined samples. Waterfalls as effective barriers played an important role in shaping the phylogeographical structure of this species. Analyses of migration rates revealed that upstream dispersal was limited crossing waterfalls. Our study revealed substantial spatial and temporal variation in the influence of landscape features on contemporary patterns of genetic structure in S. o'connori. Interglacial range expansions clearly left their mark on contemporary populations above the Tsangpo Great Gorge.

Using drift nets to capture early life stages and monitor spawning of the Yangtze River Chinese sturgeon (Acipenser sinensis)

P>A sampling system for capturing sturgeon eggs using a D-shaped bottom anchored drift net was used to capture early life stages (ELS) of Chinese sturgeon, Acipenser sinensis, and monitor annual spawning success at Yichang on the Yangtze River, 1996-2004, before and just after the Three Gorges Dam began operation. Captured were 96 875 ELS (early life stages: eggs, yolk-sac larvae = eleuthero embryos, and larvae); most were eggs and only 2477 were yolk-sac larvae. Most ELS were captured in the main river channel and inside the bend at the Yichang spawning reach. Yolk-sac larvae were captured for a maximum of 3 days after hatching began, indicating quick dispersal downstream. The back-calculated day of egg fertilization over the eight years indicated a maximum spawning window of 23 days (20 October-10 November). Spawning in all years was restricted temporally, occurred mostly at night and during one or two spawning periods, each lasting several days. The brief temporal spawning window may reduce egg predation by opportunistic predators by flooding the river bottom with millions of eggs. During 1996-2002, the percentage of fertilized eggs in an annual 20-egg sample was between 63.5 to 94.1%; however, in 2003 the percentage fertilized was only 23.8%. This sudden decline may be related to the altered environmental conditions at Yichang caused by operation of the Three Gorges Dam. Further studies are needed to monitor spawning and changes in egg fertilization in this threatened population.

The influence of topography and land use on water quality of Xiangxi River in Three Gorges Reservoir region

A self-organizing map (SOM) was used to cluster the water quality data of Xiangxi River in the Three Gorges Reservoir region. The results showed that 81 sampling sites could be divided into several groups representing different land use types. The forest dominated region had low concentrations of most nutrient variables except COD, whereas the agricultural region had high concentrations of NO3N, TN, Alkalinity, and Hardness. The sites downstream of an urban area were high in NH3N, NO2N, PO4P and TP. Redundancy analysis was used to identify the individual effects of topography and land use on river water quality. The results revealed that the watershed factors accounted for 61.7% variations of water quality in the Xiangxi River. Specifically, topographical characteristics explained 26.0% variations of water quality, land use explained 10.2%, and topography and land use together explained 25.5%. More than 50% of the variation in most water quality variables was explained by watershed characteristics. However, water quality variables which are strongly influenced by urban and industrial point source pollution (NH3N, NO2N, PO4P and TP) were not as well correlated with watershed characteristics.

Age and growth of Schizothorax waltoni in the Yarlung Tsangpo River in Tibet, China

Age and growth were studied for Schizothorax waltoni in the middle reaches of the Yarlung Tsangpo River in Tibet, southwest of China, from April 2004 to September 2006. A total of 201 specimens were collected ranging from 110 to 580 mm in standard length (SL). In contrast to other otoliths, sectioned lapillus showed a clear pattern of alternating opaque and hyaline zones. Marginal increment analyses showed that the increments, each composed of one opaque and one hyaline zone, are deposited annually. Opaque edges were prevalent from May to August. The von Bertalanffy growth parameters based on sectioned otolith data were L (t) = 689.8{1 - exp[0.051 (t + 3.275)]} for males, and L (t) = 691.1{1 - exp[0.056 (t + 2.466)]} for females. The slow growth and long life indicate that S. waltoni is vulnerable to overfishing and that harvesting strategies for the species should be conservative.

Communities of gastrointestinal helminths of fish in historically connected habitats: habitat fragmentation effect in a carnivorous catfish Pelteobagrus fulvidraco from seven lakes in flood plain of the Yangtze River, China

Background: Habitat fragmentation may result in the reduction of diversity of parasite communities by affecting population size and dispersal pattern of species. In the flood plain of the Yangtze River in China, many lakes, which were once connected with the river, have become isolated since the 1950s from the river by the construction of dams and sluices, with many larger lakes subdivided into smaller ones by road embankments. These artificial barriers have inevitably obstructed the migration of fish between the river and lakes and also among lakes. In this study, the gastrointestinal helminth communities were investigated in a carnivorous fish, the yellowhead catfish Pelteobagrus fulvidraco, from two connected and five isolated lakes in the flood plain in order to detect the effect of lake fragmentation on the parasite communities. Results: A total of 11 species of helminths were recorded in the stomach and intestine of P. fulvidraco from seven lakes, including two lakes connected with the Yangtze River, i.e. Poyang and Dongting lakes, and five isolated lakes, i.e. Honghu, Liangzi, Tangxun, Niushan and Baoan lakes. Mean helminth individuals and diversity of helminth communities in Honghu and Dongting lakes was lower than in the other five lakes. The nematode Procamallanus fulvidraconis was the dominant species of communities in all the seven lakes. No significant difference in the Shannon-Wiener index was detected between connected lakes (0.48) and isolated lakes (0.50). The similarity of helminth communities between Niushan and Baoan lakes was the highest (0.6708), and the lowest was between Tangxun and Dongting lakes (0.1807). The similarity was low between Dongting and the other lakes, and the similarity decreased with the geographic distance among these lakes. The helminth community in one connected lake, Poyang Lake was clustered with isolated lakes, but the community in Dongting Lake was separated in the tree. Conclusion: The similarity in the helminth communities of this fish in the flood-plain lakes may be attributed to the historical connection of these habitats and to the completion of the life-cycles of this fish as well as the helminth species within the investigated habitats. The diversity and the digenean majority in the helminth communities can be related to the diet of this fish, and to the lacustrine and macrophytic characters of the habitats. The lake isolation from the river had little detectable effect on the helminth communities of the catfish in flood-plain lakes of the Yangtze River. The low similarities in helminth communities between the Dongting Lake and others may just be a reflection of its unique water environment and anthropogenic alterations or fragmentation in this lake.

Acoustic Observation of the Spawning Aggregation of Megalobrama hoffmanni in the Pearl River

Using the Simrad EY60 split-beam echosounder, the spawning aggregation of Megalobrama hoffmanni was observed at the Luopang spawning grounds in the Pearl River, China, from April 19 to 22 2006. With the boat anchored, the transducer was stationary and was aimed horizontally to monitor the migration of the fish. Using fishery information, the echoes of M. hoffmanni were identified. The results showed that the spawning aggregation of M. hoffmanni at Luopang was obvious and easy to discriminate. The target strength of M hoffmanni in situ ranged from -33.8 dB to -52.3 dB (average 42.2dB). The aggregation of M. hoffmanni was obviously affected by light. With a speed of -0.31 m/s, 88.9% of the spawning stocks migrated upstream. Most M hoffmanni were recorded moving near the bottom. Their distinctive acoustic signature demonstrated the suitability of the stationary acoustic observation for M. hoffmanni identification and discrimination.

Age and Growth of an Endemic Tibetan Fish, Schizothorax o'connori, in the Yarlung Tsangpo River

We investigated the age and growth of Schizothorax o'connori in the Yarlung Tsangpo River by examination of annuli from otoliths. The von Bertalanffy model was the most acceptable statistical growth model. Its parameters were as follows: SL infinity = 492.4 mm, K = 0.1133, t(0) = -0.5432 year and W-infinity = 1748.9 g for females; SL infinity = 449.0 mm, K = 0.1260, t(0) = -0.4746 year and W-infinity = 1287.0 g for males. Theoretical longevity was 25.9 years for the female and 23.3 years for the male. Moreover, females had larger asymptotic length and weight compared with males.

Pseudobagrus fui Miao, a valid bagrid species from the Yangtze River drainage, South China (Teleostei: Bagridae)

Pseudobagrus fui Miao is a valid bagrid species that can be separated from all its congeners in having 27-33 anal-fin rays. Among the species of Pseudobagrus, it forms part of a group of approximately 20 species putatively characterized by having a smooth anterior margin of the pectoral-fin spine and short maxillary barbels not extending to the base of the pectoral-fin spine. Pseudobagrus fui, together with P. nitidus and P. vachelli, can be differentiated from all other fork-tailed species of this group by having no fewer than 20 branched anal-fin rays, the posterior end of the anal-fin base posterior to the vertical through the posterior end of the adipose-fin base, and anal-fin base longer than the adipose-fin base. It further differs from P. nitidus in having a gas bladder without beaded lateral and posterior margins, and from P. vachelli in having short maxillary barbels not extending to the base of the pectoral-fin spine. Pseudobagrus fui is currently known from the main stream of the upper Yangtze River and its tributaries, the Min River, Jialing River, Tuo River, Wu River and Chishui River in Sichuan Province, Guizhou Province, and the Chongqing City. The identity and nomenclature of Pseudobagrus nitidus is also discussed.

Trends of Superoxide Dismutase and Soluble Protein of Aquatic Plants in Lakes of Different Trophic Levels in the Middle and Lower Reaches of the Yangtze River, China

A limnological study was carried out to determine the responses of superoxide dismutase (SOD) activities and soluble protein (SP) contents of 11 common aquatic plants to eutrophication stress. Field investigation in 12 lakes in the middle and lower reaches of the Yangtze River was carried out from March to September 2004. Our results indicated that non-submersed (emergent and floating-leafed) plants and submersed plants showed different responses to eutrophication stress. Both SOD activities of the non-submersed and submersed plants were negatively correlated with their SP contents (P < 0.000 1). SP contents of non-submersed plants were significantly correlated with all nitrogen variables in the water (P < 0.05), whereas SP contents of submersed plants were only significantly correlated with carbon variables as well as ammonium and Secchi depth (SD) in water (P < 0.05). Only SOD activities of submersed plants were decreased with decline of SD in water (P < 0.001). Our results indicate that the decline of SOD activities of submersed plants were mainly caused by light limitation, this showed a coincidence with the decline of macrophytes in eutrophic lakes, which might imply that the antioxidant system of the submersed plants were impaired under eutrophication stress.