Origin, radiation, dispersion and allopatric hybridization in the chub Leuciscus cephalus

The phylogenetic relationships of 492 chub (Leuciscus cephalus) belonging to 89 populations across the species' range were assessed using 600 base pairs of cytochrome b. Furthermore, nine species belonging to the L. cephalus complex were also analysed (over the whole cytochrome b) in order to test potential allopatric hybridization with L. cephalus sensu stricto (i.e. the chub). Our results show that the chub includes four highly divergent lineages descending from a quick radiation that took place three million years ago. The geographical distribution of these lineages and results of the nested clade analysis indicated that the chub may have originated from Mesopotamia. Chub radiation probably occurred during an important vicariant event such as the isolation of numerous Turkish river systems, a consequence of the uplift of the Anatolian Plateau (formerly covered by a broad inland lake). Dispersion of these lineages arose from the changes in the European hydrographic network and, thus, the chub and endemic species of the L. cephalus complex met by secondary contacts. Our results show several patterns of introgression, from Leuciscus lepidus fully introgressed by chub mitochondrial DNA to Leuciscus borysthenicus where no introgression at all was detected. We assume that these hybridization events might constitute an important evolutionary process for the settlement of the chub in new environments in the Mediterranean area.     

Hybrids of bleak, Alburnus alburnus, and chub, Leuciscus cephalus in English rivers

The occurrence of hybrids between bleak, Alburnus alburnus , and chub, Leuciscus cephalus is reported from four discrete river systems in England. The hybrids are described and illustrated, and means of identification from the parent species are given. Bleak × chub hybrids appear to be moderately common and on superficial examination resemble the bleak parent species more than the chub.     

Hybrids of chub and roach (Leuciscus cephalus and Rutilus rutilus) in English rivers

The occurrence of hybrids between chub, Leuciscus cephalus , and roach, Rutilus rutilus , from three rivers in England is reported. The hybrids are described and illustrated, and means of identification are indicated. Chub X roach hybrids, like those between other leuciscine cyprinid species, can be positively identified by the number and form of the pharyngeal teeth.     

Proposed standard mass equations for European chub Leuciscus cephalus in Italy

Total length (L(T)) and mass measurements of 28,596 specimens of European chub Leuciscus cephalus, collected from a variety of waterways across Italy, were used to compute standard mass (W(s)) equations by both empirical percentile (EmP) and regression line percentile (RLP) methods. The use of the EmP W(s) equation [log(10) W(s) = -4·79 + 2·68log(10) L(T) + 0·10(log(10) L(T))(2)] to compute relative mass (W(r)) of L. cephalus in Italy is suggested, as it was not influenced by length-related bias (L(T) range of application = 70-470 mm).     

Observations on the growth and production of chub Leuciscus cephalus and dace Leuciscus leuciscus in a small lowland river in southeast England

SUMMARY. The River Eden (Kent, England) holds a mixed coarse fish population in which minnows ( Phoxinus phoxinus ), gudgeon ( Gobio gobio ), chub ( Leuciscus cephalus ) and dace ( Leuciscus leuciscus ) are numerically predominant. Chub and dace provide the major interest to anglers and their growth and production were studied. Observed growth rates of both species were marginally below recorded averages from other British habitats. Back-calculations showed that year-class strength and relative growth rates varied between years. Instantaneous mortality rate Z was 0.147 for chub aged 1–10, 0.438 for chub aged 10 and over, and 0.172 for dace. Exclusive of the 0-group, fish numbers, biomass (wet wt) and production were found to be 0.1305 m^-2, 19.12 gm^-2 and 5.38 gm^-2 year^-1, respectively, for chub and 0.0968 m^-2, 4.33 gm^-2 and 1.69 gm^-2 year^-1 for dace. Smoothing of data produced theoretical production figures of 6.69 gm^-2 year^-1 for chub and 1.15 gm^-2 year^-1 for dace.     

Lifetime consequences of variable 0 year group length in riverine populations of chub Leuciscus cephalus (L.)

The fork length ( L _F) of individual chub Leuciscus cephalus in English riverine populations at the end of their first growth season varied considerably, ranging from <25 to >70 mm. This had a significant influence on the subsequent growth of individuals over their lifetime. Chub of small L _F at age 1 year generally produced smaller annual growth increments throughout life than those of longer L _F at age 1 year, although they had the potential to attain greater ultimate L _F. This variability in L _F at age 1 year resulted, at least in part, from multiple spawning events over a protracted period, that caused variation in the growing seasons for 0 year group chub of the same year class. Since the adult population contained individuals that only attained L _F of <25 mm at the end of their first growing season, it is possible that L _F at age 1 year may not be such a major precursor to strong chub recruitment as previously thought.     

Seasonal and Diel Utilisation of Inshore Microhabitats by Larvae and Juveniles of Leuciscus Cephalus and Leuciscus Leuciscus

From July 1995 to January 1996, we examined the seasonal variations of the diel dynamics of habitat use by young-of-the-year cyprinid fishes (chub Leuciscus cephalus and dace L. leuciscus) in a lotic stream (River Ourthe, Southern Belgium). Inshore bays and neighbouring habitats (riparian shelters, entrance of the bay and shallow riffles) were sampled every three hours from 6:00 to 22:00 h, using DC electrofishing with prepositioned frames. In early summer, chub larvae moved exclusively in between the middle of the bay and riparian shelters inside the bay. Juvenile dace and, later in the season, juvenile chub showed diel dynamics of which the amplitude was dependent on temperature and illumination: they moved into the bay in the morning, gathered in greater numbers at mid-day (up to 586 chub and 387 dace m^-2), then progressively left the bay and entered neighbouring riffles in the late afternoon or evening. Small fish immigrated earlier into the bay and emigrated later than fish of larger size. By late September, most dace had left the bays, but returned there when water temperatures were <7°C. During autumn and winter, juvenile dace and chub of all sizes occupied exclusively inshore shelters with submerged riparian macrophytes or fallen tree leaves (corresponding densities of 0.7, 9.5 and 15.8 dace m^-2, and of 6.8, 20.2 and 94.2 chub m^-2). These results support the idea that young-of-the-year dace and chub shift from a restricted use of inshore shallow bays to a diel dynamics with alternate inshore-offshore movements at the time when they become juveniles, although the precise timings of these movements are still influenced by fish size and water temperature afterwards. The significance of these dynamics is discussed within a context of trade-off between the use of food resources and avoidance of predators.     

Growth and morphological development of chub, Leuciscus cephalus (L.), during the first year of life

The growth and the morphological development of larvae and juveniles of chub, Leuciscus cephalus (L.), from a small stream of western Greece is described, based on field samples and laboratory-raised fish. In the laboratory, size hierarchy was developed and morphological differentiation occurred at a younger age in the fastest growing larvae. Growth was faster in the field and differentiation of fins and scales began at a larger size, but apparently at a younger age, than in the laboratory.     

Hybridization between Alburnus albidus (C.) and Leuciscus cephalus cabeda R. in Italy

Thirty-six specimens of hybrid between the cyprinid species Alburnus albidus and Leuciscus cephalus cabeda were recognized from samples collected from five rivers in southern Italy. In the Alento river hybridization was marked and in a single sample of 66 specimens, 23 were found to be hybrids. The identification of hybrids is easy due to the combination of intermediate characters such as body pigmentation, gill rakers, branched rays of anal and pelvic fins, and some proportional measurements. The growth rate of hybrids is intermediate between the parental forms. Males are more numerous than females.     

Ligula intestinalis infection as a potential source of bias in the bioindication of endocrine disruption in the European chub Leuciscus cephalus

European chub Leuciscus cephalus collected from five localities in the lowland and subalpine regions of Austria were analysed for oestrogenic effects of endocrine-disrupting chemicals and the presence of the plerocercoid of the tapeworm Ligula intestinalis. Of 1494 chub analysed, only seven (six males, one female) were found to be infected with single, but large plerocercoids up to 15 cm in length. Ligula-infected fish showed comparatively immature gonads, as demonstrated by the gonadosomatic index and gamete developmental stages. Plasma levels of the egg precursor protein vitellogenin also showed concentrations ranging below the detection limit. The present results indicate that chub infected with L. intestinalis and exposed to exogenous oestrogenic compounds can result in reduced gonadal maturation and produce false oestrogen-positive diagnoses in male fish. For plasma vitellogenin levels, L. intestinalis infections can result in false oestrogen-negative diagnoses in male and female fish.     

Karyotypes of Cyprinus carpio and Leuciscus cephalus

Karyological investigations on the common carp (Cyprinus carpio) and chub (Leuciscus cephalus) are described. The diploid chromosome complement in the carp was found to be 98, with 50 meta- to submetacentric chromosomes, 48 subtelo- to telocentric chromosomes, and an arm number of 148. Chromosomal studies on the chub revealed the diploid chromosome complement to be 50, with 34 meta- to submetacentric chromosomes, 16 subtelo- to telocentric chromosomes, and an arm number of 84.     

Local and global multivariate analysis of geographical mitochondrial DNA variation in Leuciscus cephalus L. 1758 (Pisces: Cyprinidae) in the Balkan Peninsula

Geographical variation of cytochrome b mitochondrial DNA (mtDNA) in chub (Leuciscus cephalus L.) was analysed in 31 samples from 20 different river basins in the Balkan Peninsula and Danube catchment. Multivariate methods of ordination were used to analyse variation of the data sets. The results were interpreted in the context of the proposed ichthyogcographic districts separating the Balkan Peninsula into two main ichthyogeographic divisions (Eastern Greece/Ponto-Acgcan and Western Greece/South Adriatic-Ionian). Boundary detection supported these two ichthyogeographic districts for L. cephalus , revealing a boundary that ran from north to south through the Balkan Peninsula and the middle of Greece. The results also revealed the existence of a third division in Central Greece. The results of ordination techniques on homogeneous zones and analysis of the molecular variance confirmed the results obtained in studying local variability (boundaries). They also allowed us to test the existence of possible subdivisions proposed by different authors inside the two main ichthyogeographic districts. These subdivisions were not supported. The multivariate methods used in this study allowed us to propose a coherent picture of chub ichthyogeographic districts in terms of boundary detection and maximal autocorrelation between populations and to explain the patterns of chub mtDNA variation. A complete interpretation of results concerning L. cephalus requires careful consideration of both boundary analysis and autocorrelative approach. Results from an the autocorrclativc approach alone could lead to substantial misinterpretations.     

Amplitude of ecological potential: chub Leuciscus cephalus (L.) spawning in an artificial lowland canal

Chub Leuciscus cephalus (L.) is a rheophilic cyprinid which prefers lotic habitat conditions, shallow water (0.1–0.3 m) and gravel banks with moderate to high water flow (0.15–0.75m s^–1) for spawning. In contrast to these spawning requirements, a self-sustaining chub population was detected in a German lowland canal, the Oder–Havel Kanal. This canal is monotonously embanked with rip rap and almost completely lacks water flow and habitats that are suitable for riverine species. In 1999 chub spawned in the absence of water flow, in depths up to 1.28 m on allochthonous gravel with a mean diameter of 39 ± 16 mm (SD). These findings indicate a substantially higher environmental plasticity and tolerance of chub than previously reported.     

Variability in the growth rate of chub Leuciscus cephalus along a longitudinal river gradient

An increase in individual growth rate of chub Leuciscus cephalus along a longitudinal river gradient of the Rhône River basin was observed and supported by significant differences between mean growth rates of successive river size groups. The potential implications of the results for studies on species traits variability along large-scale environmental gradients are discussed.     

Use of the chromosomal co-location of the minor 5S and the major 28S rDNA as a cytogenetic marker within the genus Leuciscus (Pisces, Cyprinidae)

The chromosomes of three species from the genus Leuciscus (the ide L. idus, the European chub L. cephalus and the common dace L. leuciscus) were examined with the FISH technique for 5S and 28S rDNA probes. The analysis showed that among the three examined species, 5S rDNA signals were located on two large and four small subtelocentric chromosomes in L. leuciscus, on one large and five small subtelocentric chromosomes in L. idus, while in L. cephalus the probe signals were found on two metacentric chromosomes and one large and one small subtelocentric chromosome pairs. In all analysed species, the 28S rDNA probe signals were placed on only one chromosome pair, subtelocentric in the common dace and the European chub, and submetacentric in the ide. The three species differed in the number of sites in which both probe signals were present. In conclusion, the co-location of the 5S and 28S rDNA proved to be a useful cytogenetic marker among the studied fishes. Moreover, this marker could be adapted to other cyprinids.     

Genetic and morphological variation in a common European cyprinid, Leuciscus cephalus within and across Central European drainages

Genetic variability and differentiation of the common European cyprinid Leusciscus cephalus was investigated within and across the drainages of the rivers Rhine, Elbe and Danube using starch gel electrophoresis and morphological characters. Eleven of 24 presumptive enzyme loci showed some allozymic variation. Compared to other freshwater fish, genetic variability within all sites was high ( H _e=0·074–0·113). This was explained by the wide geographical and ecological range of the chub. In contrast, genetic and morphological differentiation between sites and drainages was low (F_ST=0·15), which may be due to the high dispersal ability of the chub and translocations of spawn by waterfowl. The phylogeographic structure of Leuciscus cephalus in north-eastern Bavaria showed low congruence with Cottus gobio populations sampled in the same area.     

Pomphorhynchus laevis: the intestinal acanthocephalan as a lead sink for its fish host, chub (Leuciscus cephalus).

Aqueous lead exposure of chub (Leuciscus cephalus) experimentally infected with the parasite Pomphorhynchus laevis resulted in a rapid accumulation of this metal in the intestinal acanthocephalans, reaching concentrations which were significantly greater than in the host muscle, liver, and intestine and approximately 1000 times higher than the exposure con centration. Parasitized chub accumulated significantly less lead in their intestinal wall than their uninfected conspecifics (Mann-Whitney U test, P 相似文献   

Comparison between lead accumulation of Pomphorhynchus laevis (Palaeacanthocephala) in the intestine of chub (Leuciscus cephalus) and in the body cavity of goldfish (Carassius auratus auratus)

This experimental study assessed the role of the microhabitat in the uptake of metals by adult acanthocephalans. We examined the accumulation of lead by adult Pomphorhynchus laevis in the intestine of chub (Leuciscus cephalus) and compared it with that in goldfish, Carassius auratus auratus, in which the parasites penetrate the intestinal wall and enter the body cavity. Chub and goldfish experimentally infected with adult Pomphorhynchus laevis were exposed to 0.01 mg l(-1) Pb(2+) over 3 weeks. Lead was rapidly accumulated in the intestinal acanthocephalans reaching a mean concentration of 7.3 microg g(-1). This concentration was significantly greater than in the host muscle, liver and intestine and more than 730 times higher than the exposure concentration. Intraperitoneal P. laevis in goldfish exposed to lead did not accumulate the metal. Thus, it was conclusively shown that metal accumulation in acanthocephalans is associated with the intestinal location and does not occur in the body cavity.     

Metazoan parasites of freshwater cyprinid fish ( Leuciscus cephalus): testing biogeographical hypotheses of species diversity

The diversity and similarity of parasite communities is a result of many determinants widely considered in parasite ecology. In this study, the metazoan parasite communities of 15 chub populations (Leuciscus cephalus) were sampled across a wide geographical range. Three hypotheses of biogeographical gradients in species diversity were tested: (1) latitudinal gradient, (2) a 'favourable centre' versus 'local oasis' model, and (3) decay of similarity with distance. We found that the localities in marginal zones of chub distribution showed lower parasite species richness and diversity. A latitudinal gradient, with increasing abundance of larvae of Diplostomum species, was observed. There was a general trend for a negative relationship between relative prevalence or abundance and the distance from the locality with maximum prevalence or abundance for the majority of parasite species. However, statistical support for a 'favourable centre' model was found only for total abundance of Monogenea and for larvae of Diplostomum species. The phylogenetic relatedness of host populations inferred an important role when the 'favourable centre' model was tested. Testing of the hypothesis of 'decay of similarity with geographical distance' showed that phylogenetic distance was more important as a determinant of similarity in parasite communities than geographical distance between host populations.     

Genetic divergence in Greek populations of the genus Leuciscus and its evolutionary and biogeographical implications

In 23 populations of Greek Leuciscus ( Squalius ), the percentage of polymorphic allozyme loci ranged between 0·034 and 0·379 (P=0·19) and expected heterozygosity was 0·011–0·166 (H_e=0·067). Current taxonomy is confusing and does not correspond to genetic data that support the presence in Greece of at least seven different species: L. cephalus , L. peloponnensis , L. prespensis , L. moreoticus , L. borysthenicus , L. keadicus and Leuciscus sp. from Euboea Island. The maximum Nei modified genetic distance was found among L. keadicus and the rest of subgenus Squalius populations D * Nei=0·446–0·705). Accepting the molecular clock hypothesis, speciation for the genus Leuciscus in Greece must have occurred during the Cenozoic period (between the Middle Miocene and the Later Pliocene). The two main biogeographical events causing speciation on mainland Greece were the Uplift of the Pindic cordillera and the isolation of the southern part of Peloponnesus. The faunistic composition of the lakes studied, in which new taxa are reconsidered, suggests the same faunistic origin in the Early Pliocene for Lakes Prespa and Stymphalia and a younger one in the Late Pliocene for Lake Trichonis. Euboea Island was not a zoogeographical unit during the Cenozoic. The isolation of all the freshwater fish fauna of Euboea has occurred since the Pliocene. The biogeographical model proposed here differs from classical hypotheses in considering of lesser importance the dispersion of L. cephalus on the Greek mainland during the Neogene and Quaternary.