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

(1) Analysis of data from samples of chub taken at monthly intervals from two tributaries of the Herefordshire Wye revealed an annual cycle of scale growth with the formation ofa check during May or June. The frequency distributions of ages deter-mined from scale checks and the length-frequency distributions showed close simi-larities. It was concluded that age determinations based on an examination of the scales were valid, provided care was taken in the identification ofthe first scale cheek. The maximum age observed was 15 years. (2) Examination ofthe population structure revealed an overall sex ratio close to unity but there was a significant increase in the proportion of females in the 11 + and older age groups. Dominant year-classes were observed. The strongest was 1959, which was attributed to exceptionally warm, dry, sunny weather during the period May-September. (3) Female chub grew more rapidly and attained greater ultimate size than the males. Changes in the pattern of growth during the life-span were attributed to the attainment of sexual maturity, especially in the females. The length-weight relation-ships of the sexes diflFered slightly but were almost identical for the same sex in the two rivers studied. Seasonal variations in the growth-rate were followed. Most rapid growth occurred after check formation in May or June and continued until September.     

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.     

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.     

Otolith shape analysis and daily increment validation during ontogeny of larval and juvenile European chub Squalius cephalus

This assesses features of otoliths from laboratory-reared embryos, larvae and juvenile European chub Squalius cephalus from hatching to 180 days post-hatching (dph). We observed the development of the three pairs of otoliths (lapilli, sagittae and asterisci) and more precisely shape changes, as well as timing and deposition rate of increments of the lapilli. The lapilli and the sagittae were present at hatching, whereas the asterisci formed between 20 and 30 dph. The lapillus and sagitta shapes were round until 20 dph. From 60 dph the anterior and the posterior rostra of the sagittae were well developed, but very thin, making this otolith too fragile to manipulate for further studies of shape and validation of otolith increment deposition rate. The lapilli provided reliable age estimates for free embryos, larvae and juveniles up to 120 dph. However, caution should be taken when ageing fish older than 150 dph as an underestimation was noticeable. The regression of the number of otolith increments on age showed a slope and an intercept not significantly different from 1 and 0, respectively, which indicated that otolith growth increments were deposited on a daily basis, with the first microincrement occurring at hatching. Increment counts were consistent between three interpreters, indicating a consistent and reliable age estimate. This study validates that the otolith increment deposition rate can be used to assess hatching dates and daily growth of wild S. cephalus under 150 dph and in environments similar to the conditions used in this study.     

Diet composition of chub,Squalius cephalus (Teleostei: Cyprinidae), in Lake Tödürge,Sivas, Turkey

Digestive tract contents of 241 specimens captured between March 1998 and November 1999 were analysed using both qualitative and quantitative methods to determine diet composition and feeding habits of Squalius cephalus (L., 1758) inhabiting Lake Tödürge, Turkey. Ages ranged from I to VI years; 60.6% of the fish were females, 39.4% males. The chub population appeared to consume a wide range of food items such as phytoplankton, zooplankton, nematodes, insects, fish, macrophytes, plant and animal detritus, whereby zooplankton (69.2%) predominated. Although individuals in the age I group preferred plankton (phytoplankton frequently; zooplankton infrequently), the older specimens (age groups IV, V and VI) behaved mostly as carnivores but also consumed some macrophytes. While there were no significant differences between diet compositions of males and females, seasonal changes were observed in frequency of occurrence, quality and quantity of consumption, and feeding activity. The chub specimens fed most heavily during spring (the pre‐spawning period: March, April and May). Highest food volume (13.0 cm³) consumed was in May and the lowest in September (2.0 cm³) while also being correlated with temperature. All of the food items were found in the digestive tract (82.1% of the specimens examined over all seasons). Highest fullness (84.9%) was reached in autumn and the lowest during summer (79.8%).     

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.     

Squalius semae,a new species of chub from the Euphrates River,Eastern Anatolia (Teleostei: Cyprinidae)

In a study of the fishes of the Euphrates River (Persian Gulf basin) three species of Squalius belonging to the cephalus-group were found: The Tohma population was identified as S. seyhanensis and the Merzimen and Hilvan populations as S. berak. The comparison of 28 metric and 5 meristic parameters and morphological characters showed that the populations of the northern Euphrates River drainages are distinct and belong to a hitherto unnamed species. We describe it here as Squalius semae sp. n.

http://www.zoobank.org/urn:lsid:zoobank.org:pub:D195B69C-C6CB-4DEB-B85D-5B695F917CE5     

Multiple Pleistocene refugia and post‐glacial colonization in the European chub (Squalius cephalus) revealed by combined use of nuclear and mitochondrial markers

Aim To analyse patterns of nuclear and mitochondrial genetic variation in the European chub, Squalius cephalus (Linnaeus, 1758), in order to understand the evolutionary history of this species and to test biogeographical hypotheses for the existence of co‐distributed European freshwater fish species. Location Rivers in Europe (Finland, Poland, Czech Republic, France, Bulgaria, Spain, Italy). Methods We genotyped 12 polymorphic microsatellite markers derived from 310 individuals collected from across the distribution of S. cephalus in Europe (including a total of 15 populations) and sequenced mitochondrial DNA (mtDNA) from a subset of 75 individuals. Sequences of mtDNA cytochrome b were analysed using both phylogenetic (median‐joining networks) and population genetic methods (tests for demographic history, mismatch distributions, Bayesian coalescent analysis). Geographical structure in microsatellite loci was examined using a distance method (F_ST), factorial correspondence analysis (FCA) and a Bayesian clustering method (structure ). Results The mtDNA network showed a clear split into four different haplogroup lineages: Western (separated into Atlantic and Danubian sublineages), Eastern, Aegean (occurring in two distinct sublineages in the Balkans and in Spain) and Adriatic. Our results indicate recent population expansion in the Eastern and Western Atlantic lineages and the admixture of two previously separate sublineages (Atlantic and Danubian) in the Western lineage. Bayesian structure analysis as well as FCA results roughly corresponded to the mtDNA‐based structure, separating the sampled individuals into almost non‐overlapping groups. Main conclusions Our results support hypotheses suggesting origins of extant lineages of freshwater fishes in multiple refugia and the subsequent post‐glacial colonization of Europe via different routes. We confirmed the previously proposed two‐step expansion scenario from the Danube refuge, the existence of a secondary (Atlantic) refuge during the last glaciation (probably in the Rhone River) and population expansion of this lineage. Conspicuous divergences among Mediterranean populations reflect their different origin, as well as their low contribution to the recent genetic pool of chub in central Europe.     

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.     

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.