Northern refugia and recent expansion in the North Sea: the case of the wrasse Symphodus melops (Linnaeus, 1758)

Pleistocene climate changes have imposed extreme conditions to intertidal rocky marine communities, forcing many species to significant range shifts in their geographical distributions. Phylogeographic analyses based on both mitochondrial and nuclear genetic markers provide a useful approach to unravel phylogeographic patterns and processes of species after this time period, to gain general knowledge of how climatic changes affect shifts in species distributions. We analyzed these patterns on the corkwing wrasse (Symphodus melops, Labridae), a rocky shore species inhabiting North Sea waters and temperate northeastern Atlantic Ocean from Norway to Morocco including the Azores, using a fragment of the mitochondrial control region and the first intron of the nuclear S7 ribosomal protein gene. We found that S. melops shows a clear differentiation between the Atlantic and the Scandinavian populations and a sharp contrast in the genetic diversity, high in the south and low in the north. Within each of these main geographic areas there is little or no genetic differentiation. The species may have persisted throughout the last glacial maximum in the southern areas as paleotemperatures were not lower than they are today in North Scandinavia. The North Sea recolonization most likely took place during the current interglacial and is dominated by a haplotype absent from the south of the study area, but present in Plymouth and Belfast. The possibility of a glacial refugium in or near the English Channel is discussed.     

What is natural? The scale of cryptogenesis in the North Atlantic Ocean

Aim Cryptogenic species may include those taxa that were historically introduced and are now falsely viewed as native. Investigated here is the scale of cryptogenesis in the North Atlantic Ocean by examining disjunct distributions, defined as temperate species occurring only on both sides of the North Atlantic. Disjunct distributions can be explained by four scenarios: glacial relicts, taxonomic artefacts, natural trans‐oceanic dispersal and human‐mediated introduction. Location North Atlantic Ocean. Methods Model taxa included ascidians, bivalves and hydrozoans. Biogeographic status (native, introduced or cryptogenic) was assigned to all species exhibiting a disjunct distribution, based upon multiple criteria. Results Of 1030 species, 60 have a strictly disjunct distribution. Of these disjunct species, for five species there is no reason to doubt their native status, and 55 species are cryptogenic or introduced. Groups with high relative dispersal capacities do not have disjunct distributions more often. Infaunal bivalves have the lowest relative number of disjunct species; none are cryptogenic or naturally disjunct. This supports the concept that glaciations are unlikely to cause disjunct distributions: there are no studies that provide conclusive evidence for the glacial relict model. Hydrozoa have the highest relative number of disjunct species, which, while historically explained by undocumented rafting, may more likely be the result of dispersal by ships, which travel relatively fast, are independent of currents and provide greater surface area. Main conclusions This reanalysis of the historical biogeography of the North Atlantic marine biota reveals that far more species may have been introduced than previously recognized, potentially significantly altering our fundamental understanding of community evolution and ecology. Species that have been present for centuries and can be important ecological engineers who have shaped contemporary communities are possibly falsely viewed as native: they may in fact be the unrecognized introductions of historical times.     

A comparison of benthic macroinvertebrate assemblages among different types of alpine streams

1. Benthic macroinvertebrate assemblages were compared among a diverse array of first‐order alpine tundra streams of the Swiss Alps. 2. A principal components analysis separated sites into three main groups: rhithral streams, rhithral lake outlets, and kryal sites including outlets and streams. Rhithral streams contained the most diverse and taxon rich assemblages, being colonised by both non‐insect taxa and Ephemeroptera, Plecoptera, Trichoptera and Diptera. 3. Rhithral lake outlets supported high densities of non‐insect taxa such as Oligochaeta, Nemathelminthes and crustaceans. Despite low taxon richness, kryal sites had high Ephemeroptera and Plecoptera abundances. Chironomidae were most common at all sites. 4. Collector‐gatherers were dominant at all sites, whereas filter‐feeders were rare. Scrapers and shredders were more common in streams than lake outlets. 5. Water temperature and algal standing crops were higher at rhithral lake outlets than rhithral streams, perhaps providing more favourable habitat for non‐insect taxa. Glacial runoff was the dominant factor influencing macroinvertebrate assemblages of kryal streams and kryal lake outlets. Alpine lakes influenced the environmental conditions of their outlets and, consequently, their macroinvertebrate assemblages unless being constrained by a glacial influence.     

Phylogeography of the cold‐water barnacle Chthamalus challengeri in the north‐western Pacific: effect of past population expansion and contemporary gene flow

Aim Phylogeographical patterns of marine organisms in the north‐western Pacific are shaped by the interaction of past sea‐level fluctuations during glacial maxima and present‐day gene flow. This study examines whether observed population differentiation in the barnacle Chthamalus challengeri, which is endemic to the north‐western Pacific, can be explained by the interactions between historical glacial events and patterns of contemporary gene flow. Location Eleven locations in the north‐western Pacific. Methods Partial sequences of mitochondrial cytochrome c oxidase subunit I (COI), 12S, 16S and nuclear internal transcribed spacer 1 (ITS1) were obtained from 312 individuals. Parsimony haplotype networks and analysis of molecular variance (AMOVA) were used to determine whether the observed genetic structure corresponds to marine provinces (Kuroshio Current and China Sea Coastal Provinces), zoogeographical zones (oriental and Japan warm‐temperate zones) and/or potential refugial areas (Sea of Japan and East China Sea) in the north‐western Pacific. Neutrality tests, mismatch distribution analysis and Bayesian skyline plots were used to infer the demographic history of C. challengeri. Results In total, 312, 117, 182 and 250 sequences were obtained for COI, 12S, 16S and ITS1, respectively. A panmictic population was revealed, which did not conform to the ‘isolation by distance’ model. None of the a priori population groupings based on marine provinces, zoogeographical zones or potential refugial areas was associated with observed genetic patterns. Significant negative values from neutrality tests and the unimodal mismatch distribution and expansion patterns in Bayesian skyline plots for the COI and 16S data sets indicate a population expansion in the mid‐Pleistocene (c. 200 ka). Information from the fossil record suggests that there has been a northward range shift of this species from the East China Sea or the Palaeo‐Pacific coast of Japan to the Sea of Japan since the mid‐Pleistocene. Main conclusions Chthamalus challengeri has experienced a population expansion and range shift since the mid‐Pleistocene. The observed lack of population differentiation can be explained by this past population expansion and present‐day wide‐scale larval dispersal (owing to the long planktonic larval duration) across marine provinces, which have led to the successful establishment of the species in different zoogeographical zones and habitats.     

Testing a relict distributional pattern of fen plant and terrestrial snail species at the Holocene scale: a null model approach

Aim The term relict refers to a formerly widespread species currently occurring in refugia that provide a persistent combination of specific ecological conditions. In peatlands, direct palaeoecological evidence of relict status exists for some plant species and, in the case of calcareous sediments, for some snail species. We tested whether some species are significantly linked to old calcareous fens at the millennial scale independent of the effect of recent fen area. We focused on three organism groups – vascular plants, bryophytes and land snails – that differ in the degree of preservation of their remains in calcareous fen sediments and in their dispersal ability. Location Western Carpathians (Slovakia and the Czech Republic). Methods The sample sites comprised 47 well‐preserved calcareous fens, from which we compiled complete recent species lists, measured the area and analysed radiocarbon‐dated samples from the deepest sediment and from the beginning of complete deforestation, as indicated by plant and snail fossils. Using the species co‐occurrences in large data sets, we identified calcareous fen specialists and compared their recent distribution patterns against a null model that controlled for the effect of fen area. Results Two land snail species, eleven vascular plant species and no bryophyte species have statistically significant affinities with old fens, independent of the effect of recent fen area. For one bryophyte and one snail, the effects of age and area are not distinguishable. Main conclusions The results for land snails, being abundantly preserved and easily determinable in calcareous fen deposits, are in full accordance with the direct macrofossil evidence. This suggests that our approach indirectly revealed a relict distribution of the species. Identification of species that are significantly linked to ancient localities at the millennial scale has great potential in palaeoecology for the detection of stands with old sediments, and in nature conservation as a tool for the identification of long‐term‐persisting rare species that infrequently colonize young sites and thus deserve priority in the protection of their habitats. From a theoretical perspective, limited dispersal from old to new localities of the same habitat can contribute to spatial effects in biotic assemblages, even at relatively fine scales.     

Evolution of sexual size monomorphism: the influence of passive mate guarding

Some species have potential for intense mate competition yet exhibit little or no sexual size dimorphism, despite predictions from sexual selection theory. Using a conceptual model, we show the conditions for which passive mate guarding with copulatory plugs can be an alternative and more successful strategy to active (direct) guarding, reducing selection pressure on large male size. The model predicts that copulatory plugs in mammals should be favoured in species for which females have short sexual receptivity periods. Using data on 62 primate species and a phylogenetic regression approach, we show that, as predicted, copulatory plugs are negatively associated with degree of sexual dimorphism and females’ sexual receptivity length. Penile spines are also significantly associated with plug use and short receptivity periods suggesting a possible offensive role in sperm competition. Results highlight that life‐history characteristics, such as sexual receptivity lengths, may alter the costs and benefits of alternative male strategies and thus alter the strength of sexual selection.     

Austrian gudgeons of the genus Gobio (Teleostei: Gobionidae): A mixture of divergent lineages

Gudgeons of the genus Gobio are small Eurasian fishes whose systematics, taxonomy, and phylogenetic relationships have been the matter of a long-standing debate. Two species, Gobio gobio and G. obtusirostris have been reported for Austria, with a potential hybrid zone in the upper Danube. Phylogenetic and phylogeographic analysis of mitochondrial cytochrome oxidase subunit I and control region sequences, as well as nuclear ribosomal protein S7 sequences, however, shows that the proposed hybrid zone is not restricted to the upper Danube, but spans large parts of the Austrian Danube system (upper Danube, Rába & Mur systems). Moreover, our data show that also a third lineage, closely related to species from the southern Balkan, contributed to the gene pool of Austrian Gobio. Patterns of intra-lineage genetic diversity indicate that the distinct Gobio lineages expanded their distribution recently (most likely post-glacially) to come into secondary contact and hybridize in the Danube system.