Massive invasion of exotic Barbus barbus and introgressive hybridization with endemic Barbus plebejus in Northern Italy: where,how and why?

Biological invasions and introgressive hybridization are major drivers for the decline of native freshwater fish. However, the magnitude of the problem across a native species range, the mechanisms shaping introgression as well as invader's dispersal and the relative role of biological invasions in the light of multiple environmental stressors are rarely described. Here, we report extensive (N = 665) mtDNA sequence and (N = 692) microsatellite genotypic data of 32 Northern Adriatic sites aimed to unravel the invasion of the European Barbus barbus in Italy and the hybridization and decline of the endemic B. plebejus. We highlight an exceptionally fast breakthrough of B. barbus within the Po River basin, leading to widespread introgressive hybridization with the endemic B. plebejus within few generations. In contrast, adjacent drainage systems are still unaffected from B. barbus invasion. We show that barriers to migration are inefficient to halt the invasion process and that propagule pressure, and not environmental quality, is the major driver responsible for B. barbus success. Both introgressive hybridization and invader's dispersal are facilitated by ongoing fisheries management practices. Therefore, immediate changes in fisheries management (i.e. stocking and translocation measures) and a detailed conservation plan, focussed on remnant purebred B. plebejus populations, are urgently needed.     

Morphologic and genetic variability in the Barbus fishes (Teleostei,Cyprinidae) of Central Italy

Italian freshwaters are highly biodiverse, with species present including the native fishes Barbus plebejus and Barbus tyberinus that are threatened by habitat alteration, fish stocking and invasive fishes, especially European barbel Barbus barbus. In central Italy, native fluvio‐lacustrine barbels are mainly allopatric and so provide an excellent natural system to evaluate the permeability of the Apennine Mountains. Here, the morphologic and genetic distinctiveness was determined for 611 Barbus fishes collected along the Padany–Venetian (Adriatic basins; PV) and Tuscany–Latium (Tyrrhenian basins; TL) districts. Analyses of morphological traits and mitochondrial DNA sequence data explored the natural and anthropogenic factors that have shaped their distribution ranges. Over 100 alien B. barbus were recorded in the Tiber basin (TL district) and Metauro basin (PV district). Comparisons of genetic and morphometric data revealed that morphometric data could identify alien B. barbus from native Barbus, but could not differentiate between B. tyberinus and B. plebejus. Genetic analyses revealed ~50 D‐loop mtDNA haplotypes and identified a distinct Barbus lineage present only in the Vomano River at the southern boundary of PV district. Demographic expansion and molecular variance analyses revealed a lack of geographic structuring across the sampling regions. While the contemporary B. plebejus distribution has been driven primarily by anthropogenic fish translocations, the dispersal of B. tyberinus has been via natural dispersion, including their crossing of the Apennine Mountains via temporary river connectivity. The results also revealed that the Barbus fishes of the mid‐Adriatic region of Europe have a complex pattern of local endemism. To conserve these patterns of genetic uniqueness, especially in the mid‐Adriatic basins, Barbus fishes should be managed by treating them as unique evolutionary units and ceasing translocations of all Barbus fishes between river basins.     

Combining geometric morphometrics with molecular genetics to investigate a putative hybrid complex: a case study with barbels Barbus spp. (Teleostei: Cyprinidae)

This integrative study examined the morphological and genetic affinities of three endemic barbel species from Italy (brook barbel Barbus caninus, Italian barbel Barbus plebejus and horse barbel Barbus tyberinus) and of putative hybrid specimens to their species of origin. Two of the species frequently occur together with the non‐native barbel Barbus barbus. DNA barcoding indicates that mitochondrial (mt) haplotypes often do not match the species expected from morphology. Linear distance measurements and meristics are not informative for discrimination of the species and putative hybrids, but a discriminant analysis of principal components (DAPC) of geometric landmark data produces reassignments largely in congruence with mt and nuclear genetic data. Cyto‐nuclear conflicts confirm the presence of hybridization in B. plebejus and B. tyberinus and identify additional introgressed specimens. A comparison between mixed genotypes and their morphology‐based assignment reveals no predictable pattern. The finding that most individuals of the morphologically similar B. plebejus and B. tyberinus have very high assignment probabilities to their respective species suggests that the presented approach may serve as a valuable tool to distinguish morphologically very similar taxa.     

Utility of fish scales from stock assessment surveys in stable isotope analysis for initial assessments of trophic relationships in riverine fish communities

The utility of using fish scales collected during stock assessment exercises to assess the trophic relationships of riverine fishes using their stable isotopes of d¹³C and d¹⁵N was tested using three riverine fish communities in England (Rivers Great Ouse, Ivel and Goyt). In each river, European barbel Barbus barbus was an important species, with other cyprinid species, including chub Squalius cephalus, present. Stable isotope analyses was completed using relatively small sample sizes per species (<11) from fish samples collected in 2001, 2005 and 2006 when up to 5 scales were collected from each fish. The calculation of standard ellipse areas (as a measure of trophic niche size) revealed that relative to other fishes, B. barbus occupied high trophic positions with minimal overlap in their trophic niche with other species, especially S. cephalus. As the analysed fish samples comprised species of various length ranges and as length has strong ontogenetic consequences for fish diet composition, generalized linear models were developed in which length was the covariate; model outputs included length‐adjusted mean δ¹³C and δ¹⁵N for each species. In each fish community, significant differences in δ¹³C and δ¹⁵N were apparent between B. barbus and S. cephalus, but were less apparent between B. barbus and other fishes. Thus, whilst the utility of using fish scales from stock assessments in stable isotope analyses are limited due to the differing length ranges of the sampled fishes, they can be useful in identifying trophic differences between species when methods such as stomach content analyses are unavailable.     

Compensatory growth,condition and food utilization in barbel Barbus barbus juveniles reared at different feeding periodicities with a dry diet

In a 40 day experiment, barbel Barbus barbus juveniles were fed a dry diet continuously, or food was withheld for 2, 5 or 10 days, and then resumed feeding. Improved feed utilization efficiency allowed fish deprived of food for 2 or 5 days to fully compensate for body mass loss. Short‐term food deprivation as a method of preventing negative effects of overfeeding in intensive rearing of juvenile B. barbus is proposed.     

Movements of adult fish in a hybrid zone revealed by microsatellite genetic analysis and capture-recapture data

• 1 Barbus barbus and Barbus meridionalis (Cyprinidae) form a hybrid zone in the Lergue river (southern France). We attempted to characterise the pattern of adult dispersal with the aim of understanding the evolutionary interactions maintaining this hybrid zone. In a previous capture/recapture study, movements between localities at a distance of 2.5 km or more appeared extremely rare. We therefore decided to investigate adult movement using a different method, based on genetic markers.
• 2 As this hybrid zone has produced allele frequency clines, massive population movements would produce temporal variations in genetic composition at a given locality. In order to determine the relationship between gene frequency and position, we surveyed four diagnostic or semidiagnostic microsatellite loci over the hybrid zone and estimated the cline produced by introgression. Then, we focused on a single locality at the centre of the hybrid zone and established its introgression index over 11 periods during two years.
• 3 The introgression index varied between periods producing significant Fst. A synthetic hybrid index, based on principal component analysis of the logit frequencies, was used for regression analyses and appeared significantly correlated with the river flow. This may be explained by displacement of adult fishes in response to flow increase. Using the information from the cline, we estimate that the most important gene frequency changes among dates, if created exclusively by population movement, correspond to a distance of 1500 m.
• 4 Additionally, we performed recaptures on a finer geographic scale than previously, around the central locality. No recaptured fishes were observed at 1875 m or more downstream, and at 875 m or more upstream, from the central locality. A high proportion of recaptured fishes (20% and 12%) was found in the nearest points (312 m downstream and 437 m upstream, respectively).
• 5 Thus, we established that (i) individual movements appear limited to a few hundred metres and (ii) individuals in a location tend to move in the same direction at the same time, probably in response to the same environmental factor.

     

Trophic consequences for riverine cyprinid fishes of angler subsidies based on marine‐derived nutrients

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Interstitial movement and emergence of barbel Barbus barbus free embryos and larvae

The movement of barbel Barbus barbus free embryos in the interstitial zone and the timing of larval emergence were studied for 12 days in an artificial stream. After egg fertilization (day 1), hatching occurred on day 6, downstream interstitial movements of free embryos on day 9 and emergence and downstream movement of all larvae on day 12, with density of free embryos decreasing as day of emergence approached. The results suggest that a memory‐based response of B. barbus to environmental conditions, previously observed in 0+ year juveniles, begins at an early age.     

Mortality induced by electrofishing and handling in five young‐of‐the‐year cyprinids: effect of the fish size,species and anode size

Immediate mortality induced by electrofishing and handling of young‐of the‐year (YOY) fish of five cyprinid species (bitterling Rhodeus amarus, chub Leuciscus cephalus, bleak Alburnus alburnus, barbel Barbus barbus, roach Rutilus rutilus) was quantified during routine YOY fish monitoring (point abundance sampling with backpack electrofisher, in situ identification and measurement). Sampled, identified and measured fish were held in screened containers immersed in the river for 5 h after electrofishing. Bleak exhibited significantly higher mortality (17%) than all other species (1–3%). Mortality decreased significantly with fish length; fish <35 mm exhibited the highest mortality. Anode size (10, 20, and 30 cm diameter) had no direct effect on mortality. Point abundance electrofishing with in situ identification and measurement is a safe method for sampling riverine YOY cyprinids.     

Comparison of diatom community structure from epilithon and fish guts: implications for inferring past changes in water quality

The sampling skills of three common European fish species (Barbus barbus, Chondrostoma nasus and Squalius cephalus) were tested to assess their potential as biomonitors of past changes in river water quality through the analysis of diatoms in fish guts. The study was performed on three rivers with different chemical and physical conditions. Comparison of similarity indices revealed low diatom assemblage resemblance between epilithic samples and samples collected from fish guts at the same locality. In contrast, a mixed-effect linear model identified significant differences between locations through comparison of diatom-based water quality indices. Among fish species, diatom indices calculated from gut samples of B. barbus and C. nasus were in most cases not significantly different from those from epilithic samples while those from S. cephalus were often significantly different. The results of the study demonstrate that diatom analysis of fish guts provides a clear distinction between eutrophic and hypereutrophic rivers. Finer nutrient variations within hypereutrophic conditions were also found to be significant for some diatom indices. In this context, choice of appropriate diatom water quality index proved to be crucial in maximizing the sensitivity of the method.     

Spawning strategies in cypriniform fishes in a lowland river invaded by non-indigenous European barbel Barbus barbus

Spawning strategies of lowland river fishes include single spawning, where reproduction generally occurs in early spring to provide 0+ fish with an extended growth season through the summer, but with a high risk of stochastic mortality events occurring, such as early summer floods. This risk can be reduced by multiple or protracted spawning strategies, where 0+ fish are produced over an extended period, often into mid-summer, but with the trade-off being a shorter growth season. The spawning strategies of cypriniform fish were explored in the River Teme, a spate river in Western England, which has non-indigenous European barbel Barbus barbus present. Sampling 0+ fish in spring and summer and across three spawning periods, B. barbus, chub Squalius cephalus and minnow Phoxinus phoxinus always revealed multiple spawning events, with 0+ fish of < 20 mm present in samples collected from June to August. Fish below 20 mm in August remained relatively small by the end of their growth season (October). For dace Leuciscus leuciscus, only single spawning events were evident, but with 0+ dace always being relatively large. Therefore, multiple spawning appears to be a common strategy that provides resilience in 0+ fish against stochastic mortality events in lowland rivers.

     

Characterizing the trophic niches of stocked and resident cyprinid fishes: consistency in partitioning over time,space and body sizes

Hatchery‐reared fish are commonly stocked into freshwaters to enhance recreational angling. As these fishes are often of high trophic position and attain relatively large sizes, they potentially interact with functionally similar resident fishes and modify food‐web structure. Hatchery‐reared barbel Barbus barbus are frequently stocked to enhance riverine cyprinid fish communities in Europe; these fish can survive for over 20 years and exceed 8 kg. Here, their trophic consequences for resident fish communities were tested using cohabitation studies, mainly involving chub Squalius cephalus, a similarly large‐bodied, omnivorous and long‐lived species. These studies were completed over three spatial scales: pond mesocosms, two streams and three lowland rivers, and used stable isotope analysis. Experiments in mesocosms over 100 days revealed rapid formation of dietary specializations and discrete trophic niches in juvenile B. barbus and S. cephalus. This niche partitioning between the species was also apparent in the streams over 2 years. In the lowland rivers, where fish were mature individuals within established populations, this pattern was also generally apparent in fishes of much larger body sizes. Thus, the stocking of these hatchery‐reared fish only incurred minor consequences for the trophic ecology of resident fish, with strong patterns of trophic niche partitioning and diet specialization. Application of these results to decision‐making frameworks should enable managers to make objective decisions on whether cyprinid fish should be stocked into lowland rivers according to ecological risk.     

Number and Distribution of Superficial Neuromasts in Twelve Common European Cypriniform Fishes and Their Relationship to Habitat Occurrence

This paper gives the first detailed data on the number and body part related distribution of superficial neuromasts in twelve common European Cypriniform species and examines whether such anatomical variables can be related to rough scale habitat occurrence. The fishes (Barbatula barbatula, Barbus barbus, Chondrostoma nasus, Cobitis taenia, Leuciscus cephalus, Leuciscus leuciscus, Phoxinus phoxinus, Rutilus rutilus, Rhodeus sericeus, Scardinius erythrophthalmus, Tinca tinca, Vimba vimba) were classified in two generalized ‘ecological guilds’, 1) rheophilic and 2) limnophilic or indifferent, based on literature data. The total number of superficial neuromasts was consistent within each species, but differed considerably between species. Lowest numbers of superficial neuromasts were found in Barbatula barbatula (21 ± 4.9 superficial neuromasts per cm body length) (mean ± SD), highest numbers in Vimba vimba (233 ± 36.1). Both species can be classified as rheophilic. Over all no relationship was found between the total number of superficial neuromasts and large scale habitat occurrence. (© 2010 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Physical changes in specimens of five species of Cyprinidae preserved in ethanol and frozen

Preservation in 30% ethanol and freezing to a temperature of ?20 ± 2° C is an appropriate method for measurement of fish eggs, larvae and juveniles. Egg diameter of the common carp Cyprinus carpio increased insignificantly by 1·32% after preservation compared with live size. The total length (L_T) of 1 day post‐hatching (dph) larvae as well as the standard length (LS) of 16 dph larvae of C. carpio increased significantly (2·95 and 1·50%, respectively) after preservation. Egg diameter as well as the L_T of 1 dph larvae of barbel Barbus barbus increased significantly after preservation, by 1·74 and 1·96%, respectively over their original size. The standard length (L_S) of 14 dph larvae of B. barbus as well as juveniles of B. barbus, crucian carp Carassius carassius, common nase Chondrostoma nasus and tench Tinca tinca decreased significantly after preservation (?0·56 to ?5·54%), whereas their body mass increased significantly (11·46–18·57%). Preserved eggs of C. carpio and B. barbus were hard, round and transparent. The larvae and juveniles of examined fishes, preserved in frozen ethanol, were straight, flexible and easily measurable after 60 days. Integrity of body surface and fins, as well as preservation of colours were much better in larvae or juveniles frozen and thawed only once than in specimens frozen and thawed thrice. Cooling in 30% ethanol to a temperature of 6 ± 2° C and freezing in water to a temperature of ?20 ± 2° C are not appropriate preservation methods for eggs and larvae of C. carpio (1 and 16 dph).     

Length–weight relationships of fish species in the freshwater rift valley lakes of Kenya

This study reports length–weight relationships for 10 species of fish encountered in the only two freshwater lakes wholly in the Rift Valley of Kenya, Naivasha and Baringo. In Lake Naivasha, none of the species analysed was native to the lake. Two were introduced species non‐native to East Africa (Micropterus salmoides, Cyprinus carpio), with three native to East Africa but not to the lake (Tilapia zillii, Oreochromis leucostictus, Barbus paludinosus). In Lake Baringo, all species were native (Oreochromis niloctictus baringoensis, Labeo cylindricus, Barbus intermedius autralis, Barbus lineomaculatus, Clarias gariepinus). Specimens were collected by gill netting in September 2004 and October 2005. With the exception of B. lineomaculatus, the b values in the relationship W = aL^b were between 2.90 and 3.22.     

Differential Parasitism of Native and Introduced Snails: Replacement of a Parasite Fauna

The role of parasites in a marine invasion was assessed by first examining regional patterns of trematode parasitism in the introduced Japanese mud snail, Batillaria cumingi (= B. attramentaria), in nearly all of its introduced range along the Pacific Coast of North America. Only one parasite species, which was itself a non-native species, Cercaria batillariae was recovered. Its prevalence ranged from 3 to 86%. Trematode diversity and prevalence in B. cumingi and a native sympatric mud snail, Cerithidea californica, were also compared in Bolinas Lagoon, California. Prevalence of larval trematodes infecting snails as first intermediate hosts was not significantly different (14% in B. cumingi vs 15% in C. californica). However, while the non-native snail was parasitized only by one introduced trematode species, the native snail was parasitized by 10 native trematode species. Furthermore, only the native, C. californica, was infected as a second intermediate host, by Acanthoparyphium spinulosum(78% prevalence). Given the high host specificity of trematodes for first intermediate hosts, in marshes where B. cumingi is competitively excluding C. californica, 10 or more native trematodes will also become locally extinct.     

New and known species of Dactylogyrus Diesing, 1850 (Monogenea,Dactylogyridae) from cyprinid fishes of the River Tigris,Iraq

Four new Dactylogyrus species are described and two redescribed from cyprinids of the River Tigris, Iraq. These are as follows: Dactylogyrus barbioides n. sp. from Barbus grypus; D. orbus n. sp. from Barbus lacerta; D. barbuli n. sp. from Barbus barbulus; D. macrostomi n. sp. from Cyprinion macrostomi; D. pavlovskyi Bychowsky, 1949 from Barbus grypus and Barbus sharpeyi; and D. inutilis Bychowsky, 1949 from Barbus xanthopterus. A phylogenetic and zoogeographical analysis is presented.     

Solitary terminal cells of Aphanizomenon gracile (Cyanobacteria,Nostocales) can divide and renew trichomes

An attempt was made to find evidence that morphologically distinct terminal cells of filamentous cyanobacterium Aphanizomenon gracile strain CCALA 8 are capable of dividing and forming trichomes. Based on our current knowledge, the division of morphologically diversified terminal cells is possible in nostocalean cyanobacteria. However, this process has been observed only in a few species. Terminal cells of A. gracile differ morphologically from other vegetative cells of a trichome, as they are not hyaline and can sometimes be found as solitary cells in cultures. Hence, it was reasonable for us to suspect that these cells are capable of dividing and forming trichomes. We observed terminal cells under a light and transmission electron microscope. Microscopic observations revealed that the septum formed in both solitary terminal cells and in terminal cells attached to trichomes. Our study is the first to demonstrate division and renewal of trichomes in terminal cells of A. gracile. Previously, such mode of reproduction was described only for another nostocalean cyanobacterium Raphidiopsis mediterranea. Moreover, our findings further emphasize the variability among members that belong to the genus Aphanizomenon , in which a type species (A. flos‐aquae) has hyaline cells incapable of dividing and renewing trichomes, while A. gracile can additionally propagate by solitary terminal cells division. This additional feature distinguishing A. gracile from typical species of Aphanizomenon, such as A. flos‐aquae, might be valuable for resolving taxonomic position of the species considering ambiguous genetic relationship between A. gracile and A. flos‐aquae.     

Competition for hosts between two hyperparasitoids of aphids,Dendrocerus laticeps andDendrocerus carpenteri (Hymenoptera: Megaspilidae): The benefit of interspecific host discrimination

The two hyperparasitoids of aphids, Dendrocerus carpenteri(Curtis) and Dendrocerus laticeps(Hedicke), differ in their interspecific host discrimination behavior: D. carpenterireadily accepts hosts for oviposition previously parasitized by D. laticeps,whereas D. laticepsoften refrains from ovipositing on hosts previously parasitized by D. carpenteri.The two species also differ in their competitive abilities: D. carpenteriis competitively superior to D. laticepswhen the time between ovipositions is 1 h. D. carpenteri's superiority probably arises from (a) the action of the venom produced by the female to paralyze the host, as evidenced by the comparison of emergence patterns from naturally vs artificially parasitized hosts, and (b) its more rapid egg development. Because of its competitive abilities, D. carpenterigains only limited benefit from discriminating interspecifically, at least when encountering a host parasitized by a competitively inferior species, such as D. laticeps.In contrast, D. laticepsgains considerable benefit from recognizing hosts previously parasitized by D. carpenteri,as the probability to produce offspring from such hosts is greatly reduced.     

Anglers' records as a tool for assessing changes in fish populations

Published anglers’ records from Polish rivers between 1966–1991 were used to show shifts in body weight of two obligatory riverine species: barbel [Barbus barbus(L.)], and chub [Leuciscus cephalus (L.)]. The body weight of barbel significantly decreased while that of chub did not. In 1966–89, the quality of inland waters continuously decreased, a result mainly from nutrient element input (domestic and agricultural). Hence, we consider two factors which were mainly responsible for reduction in fish size: overfishing and, perhaps, eutrophication.