The autecology of the chub, Squalius cephalus (L.), of the River Lugg and the Afon Llynfi

(1) The pattern of absolute atid relative gonad growth during the life-span and the seasonal gotiad cycle were determined from monthly samples of chub taken from two tributaries of the Herefordshire Wye. (2) A marked change in the pattern of absolute gonad growth was thought to indicate the attainment of sexual maturity. This was found to occur synchronously with corresponding changes in the pattern of body growth described previously and was in agreement with the age at which a seasonal gonad cycle was detectable. (3) Sexual maturity was attained during the eighth year of life (7+) in female chub and in males from the Afon Llynfi, but most males from the River Lugg matured a year earlier. (4) The chub gonads continued to increase in relative size after the attainment of sexual maturity. This was more evident in female chub. (5) Seasonal changes in the gonad condition and weight were followed. Gonad maturation occurred during spring and the gonad mass increased rapidly just prior to spawning. (6) Spawning occurred during June.     

The autecology of the chub, Squalius cephalus (L.), of the River Lugg and the Afon Llynfi

(1) Seasonal variation in the intensity of feeding activity was demonstrated in two chub populations. Feeding activity was higher in summer and more pronounced in the young age-classes. The intensity of feeding activity was significantly correlated with temperature. (2) The diet of the total chub populations embraced a wide range of food items but vegetable material predominated. Substrate material was frequently encountered which suggested considerable feeding on the river bed. Fish were the most important animal food, by volume, in the River Lugg but Trichoptera larvae were most important in the Afon Llynfi. No remains of salmon parr or trout were found in chub stomachs but grayling were eaten. The most important prey fish, by volume, was the eel but cyprinids, especially minnows, were most important by frequency of occurrence. (3) Seasonal variation in the composition of the diet could be related to availability of food organisms but some selection was shown. Vegetable foods were most important in summer; animal foods, Trichoptera larvae in particular, were important in winter. (4) Analysis of the variation in the composition of the diet with age revealed several trends, notably a decrease in the importance of plant material, aquatic and aerial insects with age and a corresponding increase in the importance of larger prey foods such as fish, frogs and crayfish. (5) The commonly held view that chub are deleterious to salmonid stocks was examined. The absence of direct predation and the small likelihood of direct competition suggests that this view is incorrect.     

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.     

Observations on the age, growth, reproduction and food of the chub Squalius cephalus (L.) in the River Stour, Dorset

Opercular bones from 399 chub from the River Stour, Dorset were used for age and backcalculated growth measurements. Scales were only used to aid the interpretation of difficult operculars. Annuli were laid down through the period mid-April to mid-June. Growth in length was minimal between October and March. Growth rates were similar to those published for chub in other European waters, but the Stour chub were longer-lived and attained a greater ultimate size. Female chub grew faster than the males. Spawning occurred from late May into June and elaboration of the gonads took place between September and May. Immature chub had an annual cycle of condition; the 0 group having a maximum in August and older immature fish reaching their maximum in June. Both categories had a minimum condition in early spring. The cycle of gonad development affected the condition of mature fish. The numbers of eggs in chub of lengths 359–467 mm ranged from 27 000–65 000. Some females attained sexual maturity at age V or VI, but most by age VII. The majority of males matured at age V, though some at ages III or IV. Growth rates and year-class strengths varied from year to year but independently of one another. Thirty-one per cent of chub aged II and over belonged to the 1959 year class. Young chub ate insect larvae and small crustacea, but the occurrence of fish and macrophytes was greater in the diet of older fish.     

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.     

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.     

The growth, reproduction and food of the roach Rutilus rutilus (L.), of the River Lugg, Herefordshire

Checks were formed on the scales in June and July. A dominant 1959 year class accounted for half the population present in 1964. Females grew faster than males and attained greater ultimate size. Growth in the River Lugg was good in comparison with rates in other British waters. The sex ratio was found to change with age; a predominance of males in the youngest age classes changed to a preponderance of females in the older classes. The youngest fishes caught (3+) were already sexually mature. The gonads were found to be a constant proportion of the body weight for fishes of all sizes at a given season. Spawning was completed by June; males were ripe in May. The ovaries recovered more rapidly than the testes after spawning. Seasonal variation in feeding activity, with higher activity in summer, was observed and could be correlated with temperature. Seasonal changes in the diet could be related to availability. The diet was predominantly vegetable; the most important animal component, aquatic insect larvae, accounted for 11 % of the total diet. Aerial insects were rarely eaten. Diet varied with age: molluscs being particularly important to older roach; younger roach consumed large amounts of substrate material. There is some evidence that young roach indiscriminately ingest substrate material but older roach are more selective and reject inert material.     

The fine structure of muscle fibres of roach, Rutilus rutilus (L.), and chub, Leuciscus cephalus (L.), Cyprinidae, Teleostei: interspecific differences and effects of habitat and season

Red and white axial muscle fibres from roach and chub were investigated by electron microscopy. Fish from three different localities were compared. Qualitative and quantitative analysis of myofibrils, mitochondria, lipid and subsarcolemmal cytoplasm with regard to muscle fibre type, species, season and habitat were made. Muscle fibre types differ significantly with the exception of the subsarcolemmal cytoplasm in roach. Within-species lipid content of red fibres differs between seasons. However, the most marked effect on red muscle fibres within species and season as regards volume density of lipid and mitochondria can be attributed to the different localities. The results are discussed in relation to mode of life and differences in habitat.     

The food of the grayling Thymallus thymallus (L.) of the River Lugg, Herefordshire

The examination of the stomach contents of 253 grayling from the River Lugg has provided some evidence of a possible depression of feeding activity during the summer months. The diet was varied but bottom food predominated, especially insect larvae, Crustacea and molluscs. Some surface foods were taken but plant material was rarely eaten. Substrate material, detritus and debris and even large pebbles were eaten, mainly in summer. Most seasonal variations in the diet could be related to variations in availability. Fish were not eaten but cyprinid and salmonid eggs were eaten at the appropriate season. In general, increasing amounts of larger food items were eaten by the older grayling.     

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.     

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.     

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.     

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.     

Age, growth and reproductive characteristics of chub, Leuciscus cephalus (L., 1758) in the İkizcetepeler dam lake (Balikesir), Turkey

In this study, the population structure, growth and reproduction characteristics of 414 chub (Leuciscus cephalus L., 1758) from the ?kizcetepeler dam lake were investigated monthly between January and December 2000. Age groups ranged between I and VI for this species in the reservoir, with the second and third year‐classes dominating. Sex ratio was 1 : 1.4 (M : F), corresponding to 58.4% males and 41.6% females. Females attained greater size and age than males. The largest female captured was 24.8 cm FL, the largest male was 24.1 cm FL, both age VI. The von Bertalanffy growth equations and length–weight relationships were found as: L_t = 28.89[1 ?e^{?0.224(t+1.55)}] for females, L_t = 26.71[1 ? e^{?0.259(t+1.55)}] for males; W_t = 347.386[1?e^{?0.224h (t+1.55)}]^2.86 for females, W_t =286.48[1?e^{?0.259 (t+1.55)}]^2.92 for males; W = 0.0227 × L^2.87 for females and W = 0.0194 × L^2.92 for males. Significant statistical differences in condition factors between age classes and sexes were not found (P > 0.05, t‐test). Spawning period of this species in the lake was between April and May.     

Water and ion balance in hydrating oocytes of the grey mullet, Mugil cephalus (L.), during hormone-induced final maturation

Sequential changes in water content and inorganic ion concentrations occurring in intra-ovarian oocytes of the grey mullet, Mugil cephalus L., during hormone-induced final maturation are described. Post-vitellogenic oocytes have a water content of 59.4%. During final maturation, oocyte water increases rapidly to a maximum of 84.8% at oviposition. Hydration is accompanied by a relatively small increase in dry matter due to the concurrent uptake of inorganic ions and an apparent uptake of organic matter. Net uptake of Ca²⁺ during hydration is negligible. Net uptake of Mg^{2 +} and Na⁺ occurs in less than equimolar proportions. Net uptake of K⁺ and Cl^? occurs in greater than equimolar proportions prior to ovulation, returning to equimolarity at oviposition. Hence, hydration results in a dilution of intracellular Ca²⁺, Mg²⁺ and Na⁺ to minimum concentrations at oviposition. In contrast, K⁺ and Cl^? concentrations are slightly elevated prior to ovulation, returning to initial levels at oviposition. A 1:1 stoichiometric relationship in net uptake of K ⁺ andCl^? is evident. Intracellular osmotic pressure in post-vitellogenic oocytes is 203.4 ± 18.0 mOsmolkg^?1. Osmotic pressure rises to a peak of 421.3+ 17.3 mOsmol kg^?1 prior to ovulation, then declines to 313.3 ± 15.0 mOsmol kg^?1 at oviposition. This rise in osmotic pressure prior to ovulation cannot be accounted for solely on the basis of inorganic ion uptake and must be attributed to changes in intracellular concentrations of organic solutes as well. An apparent increase in non-ash dry matter during hydration suggests that a net uptake of organic solute is involved. A relatively small organic molecule with high osmotic activity (e.g., amino acids) is implicated.     

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.     

Myxosporidian Parasites from the Fish of the River Lugg

SYNOPSIS. Six different species of Myxosporidia were recorded from fish of the River Lugg. Myxobolus muelleri, M. artus, M. macrocapsularis, Henneguya zschokkei, H. psorospermica and H. oviperda .
Leuciscus cephalus, L. leuciscus and Rutilus rutilus infected with M. muelleri were common; of these 3 hosts, L. cephalus had the highest intensity of infection and the greatest number of infected organs and tissues. M. muelleri was compared from all the different infection sites each month where possible, from the same host ( i.e., L. cephalus ) and from the same site on different hosts (i.e. from the gills of all 3 hosts). No seasonal variation in occurrence, spore shape or size was found for M. muelleri , but spore size varied depending on the host and site of infection.
Tailed and tailless forms of H. zschokkei were found on L. leucicus and H. oviperda on E. lucius . Only tailless H. psorospermica were found.
Variations in spore form and size are discussed in relation to their use as taxonomic characters.