Phylogeography and postglacial dispersion of the chub (Leuciscus cephalus) in Europe

A phylogeographic analysis of mitochondrial DNA variation was performed in order to test the hypothesis of a postglacial recolonization of mid- and north-European rivers from a Danubian refuge. Over 345 chub specimens from European rivers covering most of the species' native range were investigated using 600 bp of the cytochrome b gene. Chub in European rivers belong to four highly divergent mitochondrial groups (lineages) differing by mean divergence estimates from 5.2% to 7.89%. These four lineages have a largely allopatric distribution, implying four geographical sets: two Mediterranean, and two north-European sets. This pattern provided strong evidence for: (i) the eradication of this species from most of Europe during maximum ice extent; (ii) its survival in four refugia (Adriatic side of the Balkans, eastern Greece (Aegean Rivers), southern tributaries of the Danube, and periphery of Black and Caspian Seas); (iii) a differential postglacial recolonization of mid- and northern Europe from the last two refugia only; (iv) the occurrence of this recolonization in two steps for the Danubian (western) lineage that entered western Europe (Rhine-Rhone-Loire drainages) during the Riss-Würm interglacial period and survived the last glaciation there before colonizing Garonne, UK and German drainages up to the Elbe during the Holocene; and (v) the occurrence of this recolonization in a single step for the Ponto-Caspian (eastern) lineage that entered the Baltic area as far as the Oder in the Holocene. Both lineages came into contact in the River Elbe without evident mixing.     

Isolation and characterization of microsatellite loci from chub Leuciscus cephalus (Pisces: Cyprinidae)

Twelve microsatellite markers were isolated from chub Leuciscus cephalus Linné, 1766 (Cyprinidae), a freshwater fish widely distributed in Europe. We assessed the level of genetic diversity for these loci in 24 individuals sampled in the Rhone river watershed. Nine loci were polymorphic, displaying from two to 13 alleles per locus with expected heterozygosity ranging from 0.160 to 0.887. Simultaneous migration of different polymerase chain reaction products was developed for routine analysis of L. cephalus populations.     

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).     

Morphological development and growth of chub, Leuciscus cephalus (L.), larvae

This study describes the morphological development and early growth of laboratory-raised chub, Leuciscus cephalus (L. 1758), larvae. Larvae hatched with relatively large yolk sacs, stayed motionless at the tank bottom and exhibited short and sudden bursts from time to time. They were olive in colour and with complete absence of melanophores on the body. Larvae were transparent but showed brownish eye pigment. Intense body pigmentation first appeared on day 4 after hatching. By day 8 the yolk sac was fully absorbed; only 30% of the initial population of larvae successfully established exogenous feeding. Fin rays first appeared on the dorsal and anal fins of larvae around day 12. Growth during the yolk sac stage was initially fast but slowed down with the increasing size of larvae at the time of yolk absorption. The specific growth rate (SGR) of larvae declined with time, although the total observation period was relatively short (12 days). SGR values ranged from 5.18 (day 2) to 2.97 (day 12). Only a negligible egg mortality was observed during the period of early endogenous feeding (between days 1 and 6), and about 45% towards the end of endogenous feeding and immediately after the yolk sac phase (between days 7 and 9). During the exogenous feeding period (between days 10 and 12) deaths were negligible.     

Goussia chalupskyi n. sp. (Apicomplexa: Eimeriidae) from the chub Leuciscus cephalus (Cyprinidae)

Oöcysts of Goussia chalupskyi n. sp. are described from the faeces of the chub Leuciscus cephalus (Cyprinidae) from Southern Bohemia, Czech Republic. The oöcysts are spherical, 11.0 (10.5–11.5) m in diameter. The sporocysts are broadly oval to subspherical, 7.7 (7.0–8.5) × 5.7 (5.0–6.5) m, with a sporocyst residuum composed of 2–4 granules. Merogonial and gamogonial stages were found in the posterior part of the intestine.     

Pomphorhynchus laevis (Palaeacanthocephala) in the intestine of chub (Leuciscus cephalus) as an indicator of metal pollution

Chub experimentally infected with the acanthocephalan parasite Pomphorhynchus laevis were exposed to 0.01 mg l(-1) Pb(2+) over 5 weeks. Lead was rapidly accumulated in the intestinal worms reaching a steady-state concentration after 4 weeks. This concentration was significantly greater than in the host muscle, liver and intestine and more than 9,000 times higher than the exposure concentration. Lead accumulation in P. laevis increased at a higher exposure concentration but was not correlated with either parasite intensity or with pooled or individual worm weight. The highest lead concentrations were recorded in those specimen of P. laevis that were attached in the posterior intestine. Interestingly, parasitised chub accumulated less lead in their own tissues than uninfected conspecifics. A mechanism of lead uptake by P. laevis and the application of acanthocephalans as accumulation indicators of metal pollution is discussed.     

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.     

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.     

Structure of the seminal pathway in the European chub, Leuciscus cephalus (Cyprinidae); Teleostei

The testicular efferent duct system of Leuciscus cephalus (Cyprinidae), is described for three phases of testicular development. Testicular main ducts were analyzed by means of conventional histology and transmission electron microscopy. Additional techniques were applied for lectin histochemistry to determine secretory activity, as well as immunohistochemistry for cell proliferation activity and for muscle actin to demonstrate the distribution and amount of contractile cells. The contribution of the main ducts' epithelia and of degenerating spermatocytes to seminal fluid composition was confirmed, with the former being a source of carbohydrates and the latter that of phospholipids. The apical glycocalyx of epithelial cells, which is important in cell recognition and potentially involved in sperm storage, was marked by RCA I, LCA, and WGA lectin. Higher numbers of proliferating epithelial cells were ascertained during spawning phase compared to pre- and postspawning phases. In the ducts' stroma, a large number of cells expressed muscle actin and tropomyosin, indicating the ducts' contractile potential for the transport of seminal fluid towards release. Adjacent to these contractile cells, numerous nerves were found, indicating neuronal control of sperm fluid flow.     

Larval rearing of chub, Leuciscus cephalus (L.), using decapsulated Artemia as direct food

Little is known about the larviculture of the chub, Leuciscus cephalus (L.), an endangered cyprinid species endemic to European flowing waters. The use of decapsulated Artemia cysts as food for chub larviculture was investigated. After 3‐day feeding with the rotifer Brachionus calyciflorus, the larvae were fed on different diets: (i) dried decapsulated Artemia cysts, (ii) Artemia nauplii, (iii) rotifers for seven more days and then Daphnia collected from a pond, and (iv) an artificial diet. After a 24‐day rearing period, the highest survival rate was obtained with the larvae receiving decapsulated Artemia cysts. Feeding of the larvae with an artificial diet resulted in a significantly lower survival rate compared with the other groups. At the end of the experiment, the larvae fed on Artemia nauplii yielded a significantly higher mean length compared with the other groups. Feeding an artificial diet resulted in a significantly lower average weight and mean length gain compared with the other groups.     

Myxobolus cycloides on the swimbladder of chub Leuciscus cephalus: a controlled,host-specific localisation

Of 150 wild stock chub, Leuciscus cephalus L. captured in Lower Austrian watercourses, 112 revealed disc like plasmodia of Myxobolus cycloides Gurley, 1893 on the caudal chamber of the swim bladder. Other cyprinid species from the same waters lacked M. cycloides or other myxosporeans in this specific localisation. In chub, the intensity of infection (number of discs on the swim bladder) showed a logarithmic, age-dependent increase. The plasmodia of M. cycloides were situated in the connective tissue--mainly along blood vessels--and exhibited a delicate envelope of host tissue, thus forming a characteristic myxosporean cyst. Occasionally single trophozoites seemed to merge. A general process of fibroblast proliferation leading to encapsulation and degradation of the parasite was observed. This process was initiated by the formation of small multiple encapsulations within the spore containing trophozoid, before thickening of the outer cyst wall occurred. The general non-inflammatory course of the M. cycloides infection, and the obvious good health of the investigated chub suggest that this myxosporean in its host specific localisation cannot be regarded as a serious pathogen--on the contrary: parasite multiplication and degradation seemed to occur in a well-defined equilibrium controlled by the host fish.     

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.     

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.     

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.     

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.     

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.     

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.     

Interactions between immunocompetence,somatic condition and parasitism in the chub Leuciscus cephalus in early spring

Relationships between immunocompetence, somatic condition, parasitism and water temperature in a wild population of chub Leuciscus cephalus were investigated. The effects of a rapid temperature increase in early spring were studied for both sexes as water temperature affects immunocompetence. Investment in gonads and activity of mucus lysozyme were negatively correlated; lysozyme activity decreased as temperature increased. No correlations were found between lysozyme activity and parasitism or intensity of infection by monogeneans, the most abundant metazoan parasite group in L. cephalus. There was a positive correlation, however, between respiratory burst intensity and parasitism. Indices of investment in gonads and spleen were correlated, showing that energetic reserves allowed either investment in gonads and spleen, or that spleen investment, even if often used in other studies in immunoecology, was not always a significant indicator of immunocompetence during this period. This last proposition is supported by the lack of correlation between spleen investment and other factors linked to immunocompetence.     

The intestinal parasite Pomphorhynchus laevis (Acanthocephala) interferes with the uptake and accumulation of lead (210Pb) in its fish host chub (Leuciscus cephalus)

Uninfected chub as well as fish experimentally infected with the acanthocephalan parasite Pomphorhynchus laevis were exposed to ²¹⁰Pb²⁺ for up to 38 days and the uptake and distribution of lead within different fish organs and the parasites was determined at various time points. Highest metal concentrations were detected in the acanthocephalans, followed by intestine, bile, liver, gill and muscle of the fish host. Infected chub had significantly lower ²¹⁰Pb levels in the gills on day 17 (P≤0.01), in the bile on day 24 (P≤0.05) and in the gills as well as in the intestine on day 38 compared with uninfected fish. A subsequent polynomial regression revealed that lead levels for the infected fish ranged below the levels determined for uninfected fish during most of the exposure period. This is the first proof that P. laevis reduces lead levels in the bile thereby diminishing or even impeding the hepatic intestinal cycling of lead, which may reduce the amount of metals available for the fish organs. This is especially important for ecotoxicological research. For example, organisms used as accumulation indicators may erroneously indicate low levels of pollution if they are infected with parasites which alter their pollutant uptake mechanisms. Additionally, the results gave further experimental evidence for acanthocephalans as accumulation indicators for metals.