Taxonomy of the African giant pouched rats (Nesomyidae: Cricetomys): molecular and craniometric evidence support an unexpected high species diversity

Our study combined a mitochondrial cytochrome b phylogeny with cranial measurements from giant pouched rats collected across sub‐Saharan Africa. The mitochondrial phylogeny resolves two West African clades and a clade with east and central Africa representatives. This last clade can be further divided into four subclades. Altogether they represent six species (Cricetomys gambianus, Cricetomys ansorgei, Cricetomys emini, and three undescribed taxa) that can be distinguished on the basis of their mitochondrial DNA sequences and craniometry. In the absence of adequate craniometric data the existence of Cricetomys kivuensis cannot be confirmed by our data. Our combined molecular and craniometric data allowed us to broadly delineate the distribution ranges of the detected species. Cricetomys gambianus occurs in the savannah and forest clearings of West Africa. Cricetomys ansorgei is distributed in the savannah of East and southern Africa. Cricetomys emini, as currently recognized across the Guineo‐Congolian forest of Africa, is shown to be diphyletic. Cricetomys sp. 1, a separate operational taxonomic unit closely resembling C. emini, occurs in the forest zone of West Africa. An undescribed sister‐species of C. ansorgei, Cricetomys sp. 2, occurs in the forest of Central Africa along the left bank of the Congo River. Cricetomys sp. 3 occurs on the right bank of the Congo River from Cameroon to the Republic of Congo, whereas the true C. emini also occurs on the right bank of the Congo River but appears to be restricted to the Democratic Republic of Congo. Cranial phenotype within the genus tends to conform to ecological zonation (either forest or savannah) rather than to phylogenetic affiliation of the species concerned, suggesting that diversifying selection across environmental gradients could be responsible for biological diversification within the genus.     

A revision of the genus Sclerosperma (Arecaceae)

Summary  A taxonomic revision of the palm genus Sclerosperma (Arecaceae) is presented. Three species are recognised: S. mannii H. Wendl., which is relatively widespread from Liberia to the Democratic Republic of Congo; S. walkeri A. Chev., which is apparently confined to the interior of Gabon and a band along the Congo River; and S. profiziana, a new species previously considered conspecific to S. mannii that is found in southwest Ghana, Congo, the Democratic Republic of Congo and Angola. The taxonomic history, morphology, distribution and conservation status of the genus and each species are discussed.     

The impact of the Congo River and its tributaries on the rodent genus Praomys: speciation origin or range expansion limit?

We studied the taxonomy, distribution, and ecology of species within Praomys, a common rodent genus present in rainforests and montane forests in sub‐Saharan Africa. The taxonomy of the group is problematic, and for the sampled region of Kisangani (Democratic Republic of Congo) no prior genetic study has been published. We used a combination of molecular (cyt b sequencing) and craniometric techniques (canonical analyses of skull measurements) for the species identification of a total of 654 specimens. We confirm the presence of Praomys minor in the region, up to now only known from the type and paratype specimens. At least seven species are present in the Kisangani region, and two clades occur along both banks of the Congo River. The present‐day distribution of the genus seems to be influenced by large‐scale rainforest fragmentation related to drier periods in geological history. The Congo River could in this case constitute a modern barrier to gene flow when the climate enabled rainforest expansion. The tributaries of the Congo River play no role in limiting Praomys species distribution, apart from the Aruwimi River for Praomys jacksoni s.l. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 983–1002.     

Pan‐African phylogeography of a model organism,the African clawed frog ‘Xenopus laevis’

The African clawed frog Xenopus laevis has a large native distribution over much of sub‐Saharan Africa and is a model organism for research, a proposed disease vector, and an invasive species. Despite its prominent role in research and abundance in nature, surprisingly little is known about the phylogeography and evolutionary history of this group. Here, we report an analysis of molecular variation of this clade based on 17 loci (one mitochondrial, 16 nuclear) in up to 159 individuals sampled throughout its native distribution. Phylogenetic relationships among mitochondrial DNA haplotypes were incongruent with those among alleles of the putatively female‐specific sex‐determining gene DM‐W, in contrast to the expectation of strict matrilineal inheritance of both loci. Population structure and evolutionarily diverged lineages were evidenced by analyses of molecular variation in these data. These results further contextualize the chronology, and evolutionary relationships within this group, support the recognition of X. laevis sensu stricto, X. petersii, X. victorianus and herein revalidated X. poweri as separate species. We also propose that portions of the currently recognized distributions of X. laevis (north of the Congo Basin) and X. petersii (south of the Congo Basin) be reassigned to X. poweri.     

Tropical Monodominance: A Preliminary Test of the Ectomycorrhizal Hypothesis1

This study reports results from the first explicit test of the ectomycorrhizal hypothesis for tropical monodominance in the Ituri Forest of the Democratic Republic of Congo (formerly Zaire), where the canopy tree Gilbertiodendron dewevrei forms large, monospecific stands. To test the hypothesis that ectomycorrhizae are important to the success of dominant species, we surveyed the mycorrhizal status of dominant species, as well as other common, but not dominant, species in the forest. The survey reveals that two dominant species, Gilbertiodendron dewevrei and Julbernardia seretii, form ectomycorrhizae and vesicular-arbuscular mycorrhizae, while Cynometra alexandri, another dominant, forms only vesicular-arbuscular mycorrhizae. These results, along with those of other species in this and other forests, are discussed within the context of the ectomycorrhizal hypothesis for tropical mondominance. This study demonstrates that the relationship between EM and tropical monodominance is more complex than has been previously recognized.     

Phylogenetic affinities of evolutionarily enigmatic African galliforms: the Stone Partridge Ptilopachus petrosus and Nahan's Francolin Francolinus nahani,and support for their sister relationship with New World quails

The monotypic Stone Partridge Ptilopachus petrosus (Galliformes: Phasianidae), restricted to arid rocky areas of the northern savanna belt including the Sahel on the southern border of the Sahara Desert, is a taxonomic enigma. Historically, it has been grouped with Asian forest partridges (Galloperdix and Bambusicola spp.). However, recent DNA‐based phylogenetic research has suggested that its closest relative is Nahan's Francolin Francolinus nahani, another taxonomically enigmatic African galliform, and a globally threatened, narrow endemic species associated with the interior of remnant primary forests of the eastern equatorial lowlands of the Democratic Republic of the Congo and Uganda. This hypothesis is investigated in greater detail using additional DNA evidence and information on behaviour and vocalizations. Phylogenetic analyses of the combined sequences from three nuclear and four mitochondrial markers (5554 bases for 84 galliform taxa) overwhelmingly support the sister relationship between F. nahani and P. petrosus. They, in turn, are the distantly related sister taxon of the New World quails (Odontophoridae), and are not related to any other Old World galliform.     

The species identity and biogeography of Blanus (Amphisbaenia: Blanidae) in Lebanon

The genus Blanus Wagler, 1830 represents limbless, burrowing reptiles of the family Blanidae with disjunct circum-Mediterranean distribution. The recently described species Blanus alexandri Sindaco, Kornilios, Sacchi & Lymberakis, 2014 is known from south-eastern Turkey with a presumed occurrence in the Levant and Iraq. We provide here records from Lebanon and confirm the affiliation of Lebanese populations to this species by mitochondrial and nuclear DNA. Blanus alexandri comprises at least seven deeply evolved phylogenetic clades with up to 11% of uncorrected p-distances in their mitochondrial DNA. This suggests a probably older than the Miocene origin of some of these clades. Populations from Lebanon form a different clade with a genetic diversity that is close to populations from southern Turkey.     

Phylogeography of Mus (Nannomys) minutoides (Rodentia,Muridae) in West Central African savannahs: singular vicariance in neighbouring populations

We studied the phylogeography of the strict savannah pygmy mice Mus (Nannomys) minutoides in West Central Africa. A total of 846 base pairs of the cytochrome b sequence were obtained for 66 individuals collected in Gabon, Cameroon, Republic of Congo and Central African Republic. These sequences were compared to those of M. minutoides from other African countries and to eight other species of the genus Mus. We performed maximum likelihood, Bayesian and nested clade analyses, as well as neutrality tests and time estimates. We show that M. minutoides is a well‐differentiated monophyletic species that separated from other pygmy mice 1.17 Myr ago. A distinct West Central African M. minutoides clade diverged early from the other African populations of the species, with a more recent common ancestor dating 0.14 Myr. West Central African populations are globally homogeneous, despite the present fragmentation of savannahs by the rain forest. However, our analyses show an unexpected vicariance between geographically close savannahs, embedded in the rain forest in Central Gabon. One of these populations is genetically more similar to very distant peripheral populations than to three closely neighbouring populations situated on both sides of the Ogooué River. A non‐river geographical barrier probably persisted in this area, durably isolating these local populations. This hypothesis about the history of the savannah landscape should be testable through the biogeographical analysis of other strict savannah small mammal species.     

Conservation and genetics of the frosted flatwoods salamander (Ambystoma cingulatum) on the Atlantic coastal plain

The federally threatened frosted flatwoods salamander, Ambystoma cingulatum, occurs in isolated populations on the coastal plain of northern Florida, Georgia and South Carolina. An earlier phylogeographic study based primarily on mitochondrial DNA (mtDNA) and morphometrics demonstrated that the previously recognized species A. cingulatum contained two cryptic species, and that two distinct mtDNA clades were contained within the newly restricted A. cingulatum. However, salamanders from the northern extent of the species range in South Carolina were not previously available for analysis. Here, we used individuals from a newly discovered South Carolina breeding site to determine whether A. cingulatum from South Carolina are genetically distinct from their more southerly conspecifics. These analyses included the mitochondrial gene studied previously as well as broad geographic sampling of three rapidly evolving nuclear genes that allowed us to further investigate lineage diversification of flatwoods salamanders. The mitochondrial and nuclear results are largely congruent, yielding strong support for two distinct species of flatwoods salamanders and also two lineages within the eastern species. Further, the South Carolina specimens are closely related to other haplotypes found in eastern Georgia and Florida. Our summary of field surveys over the past 20 years indicates that this South Carolina population may be one of only three remaining in the entire Atlantic coastal plain distribution of this rare and declining amphibian species.     

A taxonomic revision and species delimitation of the genus Purpuraria Enderlein, 1929 (Orthoptera: Pamphagidae) using an integrative approach

Recent studies on the endemic Canarian genus Purpuraria have shown that the taxonomy of its only recognized species (P. erna) is probably erroneous. In this study, an integrative revision of the genus is performed, based on a large number of specimens and geographical sampling. As a result, (1) the diagnostic characters at the genus level are re‐described, (2) Purpuraria magna n. sp. based on morphological, morphometric and genetic data is described and (3) the taxonomic status of a formerly described subspecies is clarified. Intraspecific and interspecific morphometric differences have been found, indicating that the genus is undergoing a process of morphological diversification. Nevertheless, the possibility of interspecific mating between individuals of the two species is suggested, because no significant differences have been found between their respective calling songs. Genetic analyses using mitochondrial and nuclear DNA sequences suggest that P. erna and P. magna are recent species with evidences of secondary contact episodes in the past.     

Cytonuclear discordance among Southeast Asian black rats (Rattus rattus complex)

Black rats are major invasive vertebrate pests with severe ecological, economic and health impacts. Remarkably, their evolutionary history has received little attention, and there is no firm agreement on how many species should be recognized within the black rat complex. This species complex is native to India and Southeast Asia. According to current taxonomic classification, there are three taxa living in sympatry in several parts of Thailand, Cambodia and Lao People's Democratic Republic, where this study was conducted: two accepted species (Rattus tanezumi, Rattus sakeratensis) and an additional mitochondrial lineage of unclear taxonomic status referred to here as ‘Rattus R3’. We used extensive sampling, morphological data and diverse genetic markers differing in rates of evolution and parental inheritance (two mitochondrial DNA genes, one nuclear gene and eight microsatellite loci) to assess the reproductive isolation of these three taxa. Two close Asian relatives, Rattus argentiventer and Rattus exulans, were also included in the genetic analyses. Genetic analyses revealed discordance between the mitochondrial and nuclear data. Mitochondrial phylogeny studies identified three reciprocally monophyletic clades in the black rat complex. However, studies of the phylogeny of the nuclear exon interphotoreceptor retinoid‐binding protein gene and clustering and assignation analyses with eight microsatellites failed to separate R. tanezumi and R3. Morphometric analyses were consistent with nuclear data. The incongruence between mitochondrial and nuclear (and morphological) data rendered R. tanezumi/R3 paraphyletic for mitochondrial lineages with respect to R. sakeratensis. Various evolutionary processes, such as shared ancestral polymorphism and incomplete lineage sorting or hybridization with massive mitochondrial introgression between species, may account for this unusual genetic pattern in mammals.     

Length-weight relationships of 37 freshwater fish species from the Lefini river,middle Congo River basin (Republic of the Congo)

The present study reports on length-weight relationships (LWRs) for 37 of the most abundant fish species in the Lefini River basin, one of the largest right bank tributaries of the Congo River in the Republic of the Congo. The study area comprised 12 stations along 103 km of the river. Each station was sampled during March 2007 to August 2008 following standardized methods utilizing five monofilament gill nets of 10, 12, 15, 20 and 30 mm mesh size. Sample size, lengths, weights, LWRs, 95% confidence intervals of a and b, coefficient of correlation r² and growth type are provided for each species. Values of allometric coefficient b of the LWRs (W = aSL^b) ranged from 2.556 for Synodontis nigriventris to 3.363 for Brycinus poptae. These results represent the first study of length-weight relationships for fish species in a large affluent of the Congo basin as a whole.     

Incongruence between mtDNA and nuclear data in the freshwater mussel genus Cyprogenia (Bivalvia: Unionidae) and its impact on species delineation

Accurately identifying species is a crucial step for developing conservation strategies for freshwater mussels, one of the most imperiled faunas in North America. This study uses genetic data to re‐examine species delineation in the genus Cyprogenia. Historically, Cyprogenia found west of the Mississippi River have been ascribed to Cyprogenia aberti (Conrad 1850 ), and those east of the Mississippi River were classified as Cyprogenia stegaria (Rafinesque 1820). Previous studies using mitochondrial DNA sequences indicated that C. aberti and C. stegaria were not reciprocally monophyletic groups, suggesting the need for systematic revision. We generated a novel dataset consisting of 10 microsatellite loci and combined it with sequence data from the mitochondrial ND1 gene for 223 Cyprogenia specimens. Bayesian analysis of the ND1 nucleotide sequences identified two divergent clades that differ by 15.9%. Members of these two clades occur sympatrically across most sampling locations. In contrast, microsatellite genotypes support recognition of three allopatric clusters defined by major hydrologic basins. The divergent mitochondrial lineages are highly correlated with the color of the conglutinate lures used by mussels to attract and infest host fishes, and tests for selection at the ND1 locus were positive. We infer that the incongruence between mtDNA and microsatellite data in Cyprogenia may be the result of a combination of incomplete lineage sorting and balancing selection on lure color. Our results provide further evidence that mitochondrial markers are not always neutral with respect to selection, and highlight the potential problems of relying on a single‐locus‐marker for delineating species.     

Evolutionary systematics of the Australian Eocyzicus fauna (Crustacea: Branchiopoda: Spinicaudata) reveals hidden diversity and phylogeographic structure

Although ‘large branchiopods’ are an important faunal element of the temporary water bodies in Australia's vast (semi)arid regions, knowledge of their diversity, distribution and ecology is still poor. Here, on the basis of one mitochondrial [cytochrome oxidase subunit I (COI)] and three nuclear (EF1α, ITS2 and 28S) markers, we present new data relating to the diversity and phylogeography of eastern and central Australian Eocyzicus (Spinicaudata) fauna. Using a combination of phylogenetic, haplotype network and DNA barcoding analyses of COI, 312 individuals were grouped into eleven main lineages. To infer whether these lineages are reproductively isolated from each other (the prerequisite for species delineation according to the Biological or Hennigian Species Concepts), separate analyses of each nuclear marker were performed on a subset of specimens. Although some lineages are non‐monophyletic in the analysis of one nuclear marker, this is mostly attributed to processes such as incomplete lineage sorting rather than ongoing reproduction. The eleven lineages translate into at least seven species whose reproductive isolation is additionally indicated by sympatry, including both Australian Eocyzicus species previously described. Another three lineages may constitute further species, but their clear allopatric distribution rendered the test for reproductive isolation inapplicable. One lineage appears not to be reproductively isolated and is therefore considered a genetically distinct lineage within one of the other species, and one divergent lineage within E. argillaquus may constitute an additional species. Although sympatry is very common – six species occur in the central Paroo River catchment in eastern Australia, for instance – syntopic occurrence is rare. It is possible that a combination of differing habitat preferences and priority effects inhibits the presence of more than one Eocyzicus species per water body. There is little to no genetic differentiation between certain populations of the species found in eastern and central Australia (e.g. the Murray–Darling Basin, the Bulloo River catchment and the eastern and northern Lake Eyre Basin; LEB), suggesting high dispersal rates within this large area. Between the central Australian populations themselves, however (e.g. those inhabiting the central and western LEB), genetic differentiation is pronounced, probably as a result of the lack of abundance of important dispersal vectors (aquatic birds) and the lower diversity and density of suitable habitats in the area. The most prominent biogeographical break exists towards north‐eastern Australia (north‐east LEB), which does not share species with any other region studied.     

Patterns of diversification in a North American endemic fish,the Blackbanded Darter (Perciformes,Percidae)

The coastal plain of the south‐eastern United States shows multiple biogeographic patterns of plant and animal dispersal; however, few freshwater fish taxa span these biogeographic barriers. Percina nigrofasciata, the Blackbanded Darter (Teleostomi: Percidae), is a small, benthic, freshwater fish species with an extensive range in the south‐eastern United States. Recently, two species have been elevated from within P. nigrofasciata: P. crypta and P. westfalli, but their ranges have not been established. We broadly sampled across the south‐eastern United States, encompassing the range of P. nigrofasciata sensu lato. We reconstruct the phylogeny of Percina using both mitochondrial and nuclear markers. Eighty‐four specimens of Percina nigrofasciata were sampled for the mitochondrial gene cytochrome b (1,119 bp) to form a base phylogeny. The nuclear marker S7‐I1 was subsampled across populations to detect instances of hybridization. Phylogenetic relationships with other members of the genus Percina were assessed through Bayesian inference. Our results suggest that Percina nigrofasciata sensu stricto occurs from the Lake Pontchartrain Basin in Louisiana to the rivers of the Mobile Basin with little genetic structuring throughout its range. Percina westfalli occurs from the Apalachicola River drainages to the Atlantic Slope from the Savannah River to the St. Johns River. We find that P. crypta is not genetically distinct from P. westfalli in the Chattahoochee and Flint Rivers. Possible ancestral hybridization occurred between the P. nigrofasciata and P. westfalli in the panhandle of Florida between Mobile Bay and the Apalachicola River.     

Review of the cichlid genus Thysochromis (Teleostei: Ovalentaria) with the description of a new species from the Kouilou Province in the Republic of Congo,west–Central Africa

A new species of the chromidotilapiine genus Thysochromis, is described from the Noumbi and Kouilou River drainages in the Republic of Congo. Based on the current investigation, Thysochromis is resolved as containing two geographically disjunct species, T. ansorgii from localities in the upper Guinean ichthyofaunal province (Ivory Coast, Ghana, Benin and Nigeria) and Thysochromis emili sp. nov. restricted to coastal regions of the Republic of Congo in the lower Guinean province.     

Analysis of <Emphasis Type="Italic">Canis</Emphasis> mitochondrial DNA demonstrates high concordance between the control region and ATPase genes

Background  

Phylogenetic studies of wild Canis species have relied heavily on the mitochondrial DNA control region (mtDNA CR) to infer species relationships and evolutionary lineages. Previous analyses of the CR provided evidence for a North American evolved eastern wolf (C. lycaon), that is more closely related to red wolves (C. rufus) and coyotes (C. latrans) than grey wolves (C. lupus). Eastern wolf origins, however, continue to be questioned. Therefore, we analyzed mtDNA from 89 wolves and coyotes across North America and Eurasia at 347 base pairs (bp) of the CR and 1067 bp that included the ATPase6 and ATPase8 genes. Phylogenies and divergence estimates were used to clarify the evolutionary history of eastern wolves, and regional comparisons of nonsynonomous to synonomous substitutions (dN/dS) at the ATPase6 and ATPase8 genes were used to elucidate the potential role of selection in shaping mtDNA geographic distribution.     

Molecular phylogeny of African hinge‐back tortoises (Kinixys): implications for phylogeography and taxonomy (Testudines: Testudinidae)

We examine the phylogeography, phylogeny and taxonomy of hinge‐back tortoises using a comprehensive sampling of all currently recognized Kinixys species and subspecies and sequence data of three mitochondrial DNA fragments (2273 bp: 12S rRNA, ND4 + adjacent DNA coding for tRNAs, cytb) and three nuclear loci (2569 bp: C‐mos, ODC, R35). Combined and individual analyses of the two data sets using Bayesian and Maximum Likelihood methods suggest that the savannah species of Kinixys are paraphyletic with respect to the rainforest species K. homeana and K. erosa, and that the rainforest species may be derived from a savannah‐living ancestor. The previously recognized savannah species K. belliana was a conglomerate of three deeply divergent clades that we treat here as distinct species. We restrict the name K. belliana (Gray, 1830) to hinge‐back tortoises ranging from Angola to Burundi, while five‐clawed hinge‐back tortoises from the northernmost part of the formerly recognized range of K. belliana, together with four‐clawed tortoises from West Africa, are assigned to the species K. nogueyi (Lataste, 1886). These two species are allied to K. spekii, whereas Southeast African and Malagasy hinge‐back tortoises formerly lumped together with K. belliana represent the distinct species K. zombensis Hewitt, 1931, which is sister to K. lobatsiana. The latter two species together constitute the sister group of the rainforest species K. homeana and K. erosa. Mitochondrial data suggest that K. natalensis has a basal phylogenetic position in a clade embracing K. belliana sensu stricto, K. nogueyi and K. spekii, while nuclear data and the two data sets combined favour a sister group relationship of K. natalensis to all other hinge‐back tortoises. Phylogeographic structure is present in all wide‐ranging species and correlates in K. homeana and K. erosa with the Dahomey Gap and former rainforest refugia. The Malagasy population of K. zombensis is weakly differentiated from its South African conspecifics and further sampling is needed to determine whether there is support for the subspecific distinctness of Malagasy tortoises.     

Genetic isolation in an endemic African habitat specialist

The Chestnut‐banded Plover Charadrius pallidus is a Near‐Threatened shorebird species endemic to mainland Africa. We examined levels of genetic differentiation between its two morphologically and geographically distinct subspecies, C. p. pallidus in southern Africa (population size 11 000–16 000) and C. p. venustus in eastern Africa (population size 6500). In contrast to other plover species that maintain genetic connectivity over thousands of kilometres across continental Africa, we found profound genetic differences between remote sampling sites. Phylogenetic network analysis based on four nuclear and two mitochondrial gene regions, and population genetic structure analyses based on 11 microsatellite loci, indicated strong genetic divergence, with 2.36% mitochondrial sequence divergence between individuals sampled in Namibia (southern Africa) and those of Kenya and Tanzania (eastern Africa). This distinction between southern and eastern African populations was also supported by highly distinct genetic clusters based on microsatellite markers (global F_ST = 0.309,  = 0.510, D = 0.182). Behavioural factors that may promote genetic differentiation in this species include habitat specialization, monogamous mating behaviour and sedentariness. Reliance on an extremely small number of saline lakes for breeding and limited dispersal between populations are likely to promote reproductive and genetic isolation between eastern and southern Africa. We suggest that the two Chestnut‐banded Plover subspecies may warrant elevation to full species status. To assess this distinction fully, additional sample collection will be needed, with analysis of genetic and phenotypic traits from across the species’ entire breeding range.     

Identification of natural hybrids between Cottus poecilopus,Heckel, 1837, and Cottus gobio,Linnaeus, 1758, at a hybrid zone on the Svratka River (Czech Republic)

Sculpins occasionally form interspecific hybrids in areas of mutual contact. These can be verified genetically and, potentially, morphologically. At the study site (Svratka River, Czech Republic), Cottus gobio are common whereas C. poecilopus represent a geographically isolated alien population. Potential hybrids between these two species were compared with the parent populations to establish simple identification criteria. Analysis of nuclear and mitochondrial DNA confirmed the existence of a new European hybridisation zone. Hybrids were the result of unidirectional crossing, with female C. gobio taking the maternal position. With the exception of the lateral line position, morphometric characteristics and the epidermis structure were similar in many traits, reflecting the contribution of both species, as well as in the number of sacciform cells where hybrids were similar to C. poecilopus. The vertical position of the lateral line allows recognition of hybrids directly in the field, a useful feature for future studies. The occurrence of these hybrids confirms expansion of non‐native C. poecilopus at the expense of native C. gobio in rivers under anthropogenic pressure.