Population genetic structure in the Iberian cyprinid fish Iberochondrostoma lemmingii (Steindachner, 1866): disentangling species fragmentation and colonization processes

Interplay between the complex geography, hydrogeomorphological history, past climatic changes, and anthropogenic pressures is likely responsible for the current diversity and species' distribution of freshwater fishes in the Iberian Peninsula. To further disentangle the evolutionary processes promoting the diversification of endemic Iberian Cyprinids through time and space, we explored the patterns of genetic diversity of the Iberian arched‐mouth nase, Iberochondrostoma lemmingii (Steindachner, 1866), using molecular markers rendering at different timescales: the mitochondrial gene cytochrome b and seven microsatellite loci. Both markers showed significant differentiation of populations though the relative genetic distances among populations were different between markers. Mitochondrial DNA results indicate the isolation of hydrographic basins as the main driver of population differentiation, with Tejo as the centre of diversification. The results also support connections between Tejo, Guadiana, and Guadalquivir, with levels of divergence suggesting an earlier severance of Guadalquivir, whereas Guadiana and Tejo maintained connections until a more recent past. Establishment of more peripherial populations in small southern basins (Quarteira and Almargem) could have been ruled by founder events. However, the analysis of present‐day genetic configuration suggested by microsatellite data implies, for the first time, the involvement of other factors in the evolution of arched‐mouth Iberian nase populations. Relative low genetic distances between inter‐basin populations (Tejo and Guadiana) and the lack of concordance between differentiation and geography suggest a possible influence of human‐mediated translocations in the population genetic patterns of I. lemmingii. High intra‐basin differentiation levels were found within Tejo and Guadiana and may be associated with factors intrinsic to the species (e.g. low dispersal capability) or natural and/or artificial barriers to gene flow. The low vagility of the species appears to be an important factor influencing the evolutionary processes shaping the phylogeographical patterns of I. lemmingii, which could be relevant for the conservation of this threatened species. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 559–572.     

First record of a population of the exotic mummichog Fundulus heteroclitus (L., 1766) in the Mediterranean Sea basin (Ebro River delta)

Until now, mummichog Fundulus heteroclitus have been recorded only from the estuaries of the southernmost river basins of the Iberian Peninsula (Guadiana and Guadalquivir Rivers). Data reported here indicate the first record of F. heteroclitus in the Ebro River delta (north-east Spain), and hence also the first citation for F. heteroclitus as an invasive species present in any estuary of the Mediterranean Sea basin.     

Phylogeographic patterns in Hypochaeris section Hypochaeris (Asteraceae, Lactuceae) of the western Mediterranean

Aim  To analyse phylogeographic patterns in the four species of Hypochaeris sect. Hypochaeris , evaluating possible areas of origin and the microevolutionary processes that have shaped their morphology, genetics and distribution.
Location  Western Mediterranean area.
Methods  We applied amplified fragment length polymorphism (AFLP) markers to a total of 494 individuals belonging to 82 populations of Hypochaeris arachnoidea , H. glabra , H. radicata and H. salzmanniana to determine population structure.
Results  Populations with the largest proportion of private and rare AFLP fragments were found in Morocco. This region was consequently inferred to be the ancestral area for H. arachnoidea , H. glabra , H. radicata and H. salzmanniana . The Guadalquivir River (southern Spain) was inferred to be an effective dispersal barrier for H. glabra and H. radicata. The Strait of Gibraltar was inferred to be a somewhat weaker barrier than the Guadalquivir River for H. radicata and a much weaker barrier for H. glabra . The main barrier for H. salzmanniana coincides with the extension of the Rif Mountains to the Atlantic coast in Morocco, and the Strait of Gibraltar is a much weaker barrier for this species. Hypochaeris arachnoidea appears to have originated in the Atlas Mountains.
Main conclusions  The highest levels of genetic variation in La Mamora forest ( H. glabra and H. salzmanniana ) or the adjacent central Middle Atlas ( H. arachnoidea and H. radicata ) in Morocco suggest that these areas were a centre of origin of Hypochaeris sect. Hypochaeris . All three potential barriers – the Guadalquivir River, the Strait of Gibraltar, and the Rif Mountains – have been important in shaping genetic diversity in species of section Hypochaeris .     

The situation of penduline tit (Remiz pendulins) in southern Europe: A new stage of its expansion

Summary The present status ofRemiz pendulinus in the Iberian Peninsula is updated after a new expansive stage, so that we can distinguish four expansive waves. The recent spreading resulted in an extension by ca. 200 km of the breeding area following the river Tajo and by ca. 300 km following the Guadiana. Likewise,Remiz pendulinus has colonized the only remaining great river in the Iberian Peninsula, the Guadalquivir, at least along 170 km. This range extention is considerable and compares with a similar development in Central Europe. A factor possibly supporting the expansion may have been the artificial creation of wetlands (reservoirs), improving existing and creating new habitats suitable for the species. At present,Remiz pendulinus breeds discontinuously from the southern coast of Finland to the southern coast of Spain.

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Zusammenfassung Die heutige Verbreitung der Beutelmeise (Remiz pendulinus) auf der Iberischen Halbinsel wird nach einer neuerlichen Ausbreitungsphase in 4 Karten für die Zeiträume 1960–1970, 1971 und 1972, 1973–1978 und 1979–1989 dargestellt. Die jüngste Ausbreitungswelle führte zu einer Erweiterung des Brutgebietes um ca. 200 km entlang des Tajo und um ca. 300 km entlang des Guadiana. Außerdem besiedelt die Beutelmeise jetzt auch den letzten übriggebliebenen Fluß der Iberischen Halbinsel, den Guadalquivir, auf einer Länge von mindestens 170 km. Diese Ausbreitung ist von ihrer Ausdehnung her noch weitreichender als die in jüngster Zeit in Mitteleuropa festgestellte. Möglicherweise wird diese letzte Ausbreitungswelle durch den künstlichen Bau von neuen Feuchtgebieten (Stauseen) begünstigt, die für die Art neue Lebensräume darstellen. Die Beutelmeise ist heute von der Südküste Finnlands bis zum südlichen Spanien als Brutvogel verbreitet.

     

Life‐history variability of non‐native centrarchids in regulated river systems of the lower River Guadiana drainage (south‐west Iberian Peninsula)

Life‐history variability of two non‐native centrarchids, pumpkinseed Lepomis gibbosus and largemouth bass Micropterus salmoides, was evaluated in three stream stretches of the lower River Guadiana drainage (south‐west Iberian Peninsula) with different degrees of regulated flows. Abundance, condition and population structure differed among populations for both species, but invasion success was lower in the least regulated river. Lepomis gibbosus were abundant and had multiple age classes in the three river sites, whereas M. salmoides were less abundant and mainly represented by young‐of‐the‐year fish. Juvenile growth in L. gibbosus was similar in all three populations, though longevity was slightly greater in the population from the River Guadiana mainstream. Lepomis gibbosus exhibited a long reproductive season, but the duration of season, size at maturity and reproductive effort varied among populations. The life‐history differences found demonstrate the importance of species adaptation to local conditions which might favour their invasion success. Lepomis gibbosus were more adaptable and resilient to local conditions, whereas M. salmoides seemed dependent on reservoirs and large rivers for maintenance of riverine populations.     

Description of the Nymphs and Eggs of Acentrella almohades sp. n. from Morocco and Southern Spain (Ephemeroptera: Baetidae)

The nymph and egg of Acentrella almohades sp. n. are described and illustrated on the basis of material collected in southern Spain (in the upper reaches of the Guadalquivir River Basin) and Morocco (Rif Mountains and Atlas Mountains). Features distinguishing the new species from other European species are discussed and keyed.     

First record of intraspecific cleaning behavior by the threatened Leuciscidae Squalius alburnoides (Steindachner, 1866) at the Guadiana River basin (Portugal)

Squalius alburnoides (Steindachner, 1866) is an endemic threatened species from the Iberian Peninsula. Here, we report the first observations of intraspecific cleaning behavior in isolated summer pools in the Guadiana River Basin (Portugal). We found that focal S. alburnoides solicited cleaning by adopting an immobile tail-stand position known as “posing,” which immediately signaled a response to a few conspecifics that approached and inspect them. Our study expands the list of cleanerfish species in freshwaters, giving emphasis to the importance of mutual positive behavior within an endangered species, particularly when facing seasonal disturbance.     

Identifying priority sites for the conservation of freshwater fish biodiversity in a Mediterranean basin with a high degree of threatened endemics

The Guadiana River basin’s freshwater fish species richness, endemicity and threatened status (92% of native species are threatened) highlight the need for a large-scale study to identify priority areas for their conservation. One of the most common problems in conservation planning is the assessment of a site’s relative value for the conservation of regional biodiversity. Here we used a two-tiered approach, which integrates an assessment of biodiversity loss and the evaluation of conservation value through site-specific measures. These measures based on the reference condition approach introduce the ability to make objective comparisons throughout the Guadiana River basin, thus avoiding a priori target areas. We identified a set of biodiversity priority areas of special conservation significance—which contain rare taxa as well as intact fish communities—because of their outstanding contribution to the basin’s biodiversity. The inclusion of complete sub-basins in these priority areas might guarantee an optimal solution in terms of spatial aggregation and connectivity. However, the high spatial fragmentation to which the Guadiana River basin is submitted due to river regulation highlights the necessity of a systematic approach to evaluate the capability of the identified priority areas to maintain the Guadiana’s freshwater fish biodiversity. Handling editor: R. H. Norris     

Seasonal changes in fish community structure of intermittent streams in the middle reaches of the Guadiana basin, Portugal

Seven sites on three tributaries with intermittent flow regimes in the middle reaches of the Guadiana, i.e. Xévora, Caia and Degebe, were sampled approximately every 3 months from April 1995 to April 1997. The density and biomass of fish were greater in downstream rivers and sites. Along the Xévora River, a series of point abundance samples showed evidence of fish zonation along the river. Canonical correspondence analysis (CCA) suggested a simple relationship between the different species and their preferred habitat. The small-sized Rutilus alburnoides and Leuciscus pyrenaicus used most or all of the habitats with some cover and flowing water. Greater concentrations of larger fish were found in downstream reaches which generally had greater depths as stream width increased: large Barbus microcephalus , B. steindachneri and B. comiza preferred such sites on the Degebe River. Droughts in seasonal Mediterranean streams such as the Guadiana River are a predictable natural disturbance. In the summer, flow ceases and some stretches of river consist of isolated pools. This results in intense aggregations of fish and possible competition for food and/or space. Although some species are well adapted to natural droughts, habitat degradation and possibly the introduction of exotic species contribute to marked variability in species composition.     

Genetic Identification of Western Mediterranean Sturgeons and its Implication for Conservation

To date, the only native sturgeon species in Western Europe was believed to be Acipenser sturio. However, this species is currently restricted to the Gironde River (Southern France), and it poses serious difficulties for rearing in captivity and for using in recovery programme. Furthermore, it has been questioned whether A. sturio is in fact the only species within the rivers of Western Europe, as A. naccarii, a species previously considered endemic of the Adriatic region, has been reported from the Iberian Peninsula in recent years. Here, we have used forensic techniques to obtain DNA from several museum specimens of sturgeons caught in the Spanish Guadalquivir River and in other European rivers. We analysed DNA sequences from these specimens for five genetic markers (three nuclear and two mitochondrial regions), which were subsequently compared with sequences obtained from A. sturioand A. naccarii. Our study demonstrates that A. naccarii coexists withA. sturio, from the Adriatic Sea to the Iberian Peninsula, a finding that could be taken into account in future sturgeon recovery programmes in Western Europe.     

Population genetic structure of the sidespot barb,Barbus neefi,from the north-eastern escarpment,South Africa

Allozyme analysis was used to determine patterns of genetic variation within and between populations of Barbus neefi. The products of 29 loci were analysed, with 17 loci being monomorphic in all populations. The genetic variability estimates compared well with values reported in the literature. The percentage of polymorphic loci (0.95 criterion) ranged from 0–20.69% and average expected heterozygosity from 0.004–0.069. Headstream populations generally revealed lower genetic variability than populations in low lying areas, which emphasizes the importance of conserving headstream populations. Fixed allele mobility differences were observed at the MPI-1 protein coding locus between the Selons River population and the other populations. The measures of genetic differentiation as assessed by F-statistics, the effective number of migrants per generation and Nei's unbiased genetic distance consistently separated the Selons River population from the rest. Nevertheless, these estimates all fell within the range considered for conspecific populations. There was also a clear genetic division between populations from north and south of the Olifants River. The three populations from the Dorps, Spekboom and Blyde Rivers (Limpopo River system) and the population from the Crocodile River (Incomati River system) showed negligible genetic differentiation. Human-assisted transfer of populations may be responsible for this close similarity.     

First records of the North American catfish Ameiurus melas in Portugal

The North American catfish Ameiurus melas is recorded for the first time in Portugal. Its presence in the Tagus River Basin is probably due to natural spread of individuals from Spanish populations, while episodic translocation could explain its occurence in the previously non-invaded Guadiana and Sado River basins.     

Ecology of some Species of Thiobacillus and Sulfate-reducing Bacteria in the Middle Course of the River Guadalquivir (Spain)

During 1985, a study was made of the physical, chemical and biological characteristics of the River Guadalquivir. At the same time, the density of certain microorganisms involved in the transformation of sulfur compounds in the water was studied. The results obtained give the following numerical order of density: T. thioparus >sulfate reducers > T. denitrificans > T. thiooxidans. Urban waste was found to increase density levels of these microorganisms. Their relationship with certain environmental parameters is discussed in this study. Regression equations have been obtained of bacteria numbers for three of the four groups studied, using water temperature and permanganate oxidability as dependent variables.     

Age, diet, growth and reproduction of a Population of Cobitis paludica (de Buen, 1930) in the Palancar Stream (southwest of Europe, Spain) (Pisces: Cobitidae)

A total sample of 99 specimens of the vulnerable fish species Cobitis paludica(de Buen, 1930), was collected on a monthly basis from a tributary of the River Guadalquivir between September 1990 and September 1991. Female longevity was 1 year longer than that of males, the former reaching age-class 5+. The diet of C. paludicaconsisted mainly of Ostracoda and chironomid larvae, benthic prey of small size and occasionally other resources, such as Macrobiotid eggs spawned during winter. Detritus in gastrointestinal content was scarce, which differs from other populations. Back-calculations of length were performed from scale annuli measurements. The relationship between standard length and scale radius was different for males and females. Females were larger than males of the same age. Both back-calculated length and theoretical length, calculated by applying von Bertalanffy's model, were larger for females. The relationship between female standard length (SL) and fecundity (F) was log F= –7.465 + 5.5753 log SL.     

First isolation of Prototheca species in Spain

One-hundred forty samples of waste water were studied in search of algae of the genus Prototheca, that are potentially pathogenic to man. The samples were taken from collector outlets opening onto the Guadalquivir River.Nineteen cultures of Prototheca were isolated and identified as P. wickerhamii 9, P. zopffi 8 and P. stagnore 2.This is the first report of these algae in Spain and it is to be anticipated that these opportunist human pathogens will eventually be found to cause disease in the general population.     

Recent,small beginnings: genetic analysis suggests Catostomus rimiculus (Klamath smallscale sucker) in the Smith River,California, are introduced

Identification of introduced species can be important to understanding ecological systems and meeting conservation and management goals, but the process can be surprisingly challenging. The Klamath smallscale sucker Catostomus rimiculus seems likely to be native to the Smith River because the drainage separates two basins believed to be within the fish's native range, the Rogue and Klamath rivers. Further, C. rimiculus is broadly distributed in the Smith River, and the indigenous Dee-ni’ People of the Smith River have a unique word for sucker. Nonetheless, a historical survey of fishes that described C. rimiculus from the Rogue and Klamath rivers did not include C. rimiculus among the fishes of the Smith River. To determine whether the genetic structure of the Smith River C. rimiculus reflects expectations for a native sucker population, the authors of this study examined variation in microsatellite and mitochondrial genetic markers from the Smith River and surrounding drainages. The genetic analyses revealed a pattern consistent with extreme founder effects in Smith River C. rimiculus, as would be expected from a single introduction of six or fewer effective individuals. The sharing of a high-frequency haplotype between the Smith River and Klamath River that is not detected in the Rogue River suggests the Klamath River as the likely source for the introduction. The findings highlight that local-scale introductions can be easily overlooked because the newly established populations can appear to be parts of contiguous natural distributions.     

Variability in diet and foraging behaviour between sexes and ploidy forms of the hybridogenetic Squalius alburnoides complex (Cyprinidae) in the Guadiana River basin, Portugal

Samples of the Squalius alburnoides complex, a hybridogenetic Iberian minnow composed of diploid, triploid and, less commonly, tetraploid forms, were collected from a tributary of the Ardila River (Guadiana River basin) between February 1999 and January 2001. Seasonal variation in diet was evident, and was probably linked to prey availability. Distinct foraging behaviours between ploidy forms were found towards several prey items, suggesting that diploid adult males fed mostly near the surface, whereas diploid adult females tended to feed near the bottom and submerged vegetation. Triploid females exhibited an intermediate foraging behaviour, although there was greater affinity towards diploid male feeding behaviour. Diploid males which, in contrast to diploid and triploid females, have non‐hybrid genomes in the Guadiana drainage, exhibited a higher specialization for food. Despite considerable dietary overlap, there appeared to be spatial segregation of feeding niches between the three forms, especially during dry periods when prey availability was lower, which may be a strategy for diminishing competition for food. Therefore, considering asexual generalist and specialist hypotheses, it appears that the different ploidy levels are generalist, opportunistic feeders that partition the resources when limited.     

First record of Barbonymus schwanenfeldii (Bleeker) in the Iberian Peninsula

The South-east Asian cyprinid Barbonymus schwanenfeldii is recorded for the first time in the Iberian Peninsula from two specimens collected in the Guadiana River Basin in 2005. Their presence is probably due to release from aquaria.     

Variability,genetic structure and phylogeography of the dolomitophilous species Convolvulus boissieri (Convolvulaceae) in the Baetic ranges,inferred from AFLPs,plastid DNA and ITS sequences

Convolvulus boissieri is an edaphic endemic plant which grows in the Baetic ranges always in association with high mountain xeric dolomitic outcrops. As these dolomitic areas appear in a ‘soil‐island’ pattern, the distribution of this species is disjunct. Populations of this species frequently include a low number of individuals, which could have an important impact on their genetic diversity and viability. Convolvulus boissieri provides an excellent opportunity to study the genetic and phylogeographical aspects of species linked to dolomites. We used amplified fragment length polymorphism markers and nuclear (internal transcribed spacer region of the nuclear ribosomal cistron) and plastid sequences (trnL‐trnF, rpl32‐trnL and trnQ‐5′rps16). Data were generated from 15 populations, representing the distribution area of the species. For sequence analysis and estimation of divergence times we also used sequences from other Convolvulus species. Results revealed low intrapopulational genetic diversity and a strong interpopulational structure. Furthermore, we found clear‐cut differentiation caused by the existence of two large population groups separated by the Guadiana Menor river basin. Estimation of divergence times indicated that divergence took place during the Pleistocene glaciations. Genetic diversity and differentiation are similar to those other species exhibiting naturally fragmented distribution with a sky islands pattern. In phylogeographical terms, the successive glaciation–interglaciation cycles caused the species to spread from the western sites to eastern sites, the latter being more exposed to the effects of glaciation. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176 , 506–523.     

Genetic diversity and differentiation of Daphnia galeata in the middle and lower reaches of the Yangtze River,China

Mitochondrial 16S rDNA and CO I gene were used as molecular markers for the analysis of the genetic diversity and differentiation of Daphnia galeata populations in nine water bodies in the middle and lower reaches of the Yangtze River. In the combined 16S rDNA and CO I gene sequences, 54 variable sites and 44 haplotypes were observed among 219 individuals belonging to nine D. galeata populations. Average haplotype diversity and nucleotide diversity were, respectively, 0.72% and 0.56%. The F‐statistics (F_ST) value of the D. galeata populations was 0.149. According to the results of the neutral test, D. galeata in the middle and lower reaches of the Yangtze River had experienced a bottleneck effect in the history. Molecular variance analysis indicated that the genetic differentiation of the D. galeata populations mainly occurred within populations (85.09%). Greater genetic differentiations of D. galeata among individuals within populations appeared in the populations from the Huaihe River basin, whereas smaller genetic differentiations occurred in the populations from the middle reaches of the Yangtze River. Strong gene flows were all observed between Group I (four populations from the middle reaches of the Yangtze River) and Group ΙΙ (three populations from the middle and lower reaches of the Yangtze River), and Group ΙΙΙ (two populations from the Huaihe River basin). The effective migration rates (M) were 851.49 from Group I to Group ΙΙ and 685.96 from Group I to Group ΙΙΙ, respectively. However, no significant relationship was observed between the genetic differentiation and geographical distance of the nine populations (r = .137, p > .05). Results suggested that the genetic differentiation of D. galeata in the water bodies in the middle and lower reaches of the Yangtze River resulted mainly from geographical isolation.