Refugial races and postglacial colonization history of the freshwater amphipod Gammarus lacustris in Northern Europe

The systematic structure and postglacial population history of the freshwater amphipod Gammarus lacustris were explored in an allozyme survey of 65 populations across Northern Europe. A strong multilocus pattern of differentiation discriminated populations of the north‐east (north‐eastern Norway, northern Finland) from those in the west and the south (southern and central Scandinavia, Denmark, Poland). This principal division is attributed to postglacial colonization of the area by two main refugial races or lineages, one from the east (Russia), the other from the south (north‐western European continent). The strongly diverged Eastern and Western races (Nei's D= 0.3, from 22 loci) now meet in a secondary contact zone across a narrow sector of northernmost Norway. Genetic population compositions in this zone vary in a mosaic pattern, and show no evidence of reproductive incompatibility. Similar contacts of eastern and western lineages, far older than the latest glaciation, are now known from a number of taxa and they constitute a general pattern in Fennoscandian phylogeography. Within the Western Gammarus race, the populations through coastal north‐western Norway are further distinguished from those in southern Scandinavia and Denmark by a set of unique alleles at high frequencies (D = 0.12). This suggests an independent early colonization of the coastal region by another distinct stock, either along an early deglaciated coastal corridor from the south‐west, or directly from the ice‐free continental shelf off the Norwegian coast – a hypothesis that has also previously been presented for G. lacustris, and parallels controversial suggestions of local refugia for other taxa in Scandinavia. The coastal population type only later could come into contact with Gammarus invading over the mountains from the south; these two population types now smoothly intergrade. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 79, 523–542.     

Evidence of changing intrinsic water‐use efficiency under rising atmospheric CO2 concentrations in Boreal Fennoscandia from subfossil leaves and tree ring δ13C ratios

Investigating the many internal feedbacks within the climate system is a vital component of the effort to quantify the full effects of future anthropogenic climate change. The stomatal apertures of plants tend to close and decrease in number under elevated CO₂ concentrations, increasing water‐use efficiency (WUE) and reducing canopy evapotranspiration. Experimental and modelling studies reveal huge variations in these changes such that the warming associated with reduced evapotranspiration (known as physiological forcing) is neither well understood or constrained. Palaeo‐observations of changes in stomatal response and plant WUE under rising CO₂ might be used to better understand the processes underlying the physiological forcing feedback and to link measured changes in plant WUE to a specific physiological change in stomata. Here we use time series of tree ring (Pinus sylvestris L.) δ¹³C and subfossil leaf (Betula nana L.) measurements of stomatal density and geometry to derive records of changes in intrinsic water‐use efficiency (iWUE) and maximum stomatal conductance in the Boreal zone of northern Finland and Sweden. We investigate the rate of change in both proxies, over the recent past. The independent lines of evidence from these two different Boreal species indicate increased iWUE and reduced maximum stomatal conductance of similar magnitude from preindustrial times (ca. ad 1850) to around ad 1970. After this maximum stomatal conductance continues to decrease to ad 2000 in B. nana but iWUE in P. sylvestris reaches a plateau. We suggest that northern boreal P. sylvestris might have reached a threshold in its ability to increase WUE as CO₂ rises.     

The stream as a habitat templet for bryophytes: species' distributions along gradients in disturbance and substratum heterogeneity

1. The structure of bryophyte communities in streams in relation to habitat characteristics, especially disturbance, is described. Disturbance in rivers is quantified as movement of the stream bed, whereas in small streams water level fluctuation is used as an indicator of disturbance frequency. 2. Canonical correspondence analysis differentiated frequently disturbed sites from more stable ones. The existence of a disturbance gradient was confirmed in a subset of the study sites using long-term records of discharge variation. A parallel change was detected in the species composition of bryophyte communities with low-statured, potentially fast colonizers dominating the disturbed end and large perennial species the stable end of the gradient. 3. A consistent pattern of zonation of bryophyte species was found along the gradient from continually submersed to persistently dry conditions in small streams and lake outlets. An abrupt increase in species richness occurred at or just above the water line, where facultatively aquatic species tolerant of both conditions formed the bulk of the community. 4. The relationship between species richness and standing crop in stream bryophyte communities was consistent with the hump-backed model of Grime (1979), especially at the within-habitat scale. Quadrats of low and very high standing crop were characterized by low species richness, while peak richness was observed at intermediate standing crops. 5. A few perennial species (e.g. Fontinalis spp. and Rhynchostegium riparioides) capable of monopolizing space dominated the most stable habitats. Species composition in low biomass sites was more variable, yet only one basic growth-form (small-statured species with high allocation to spore production) seemed possible in these highly disturbed streams. In habitats of intermediate biomass, small-scale disturbances apparently allow the formation of a more varied bryophyte community. 6. A habitat templet for stream bryophyte life strategies and community structure is presented. Disturbance is proposed to be the factor filtering out traits unsuitable for a given environment. The potential of stream bryophytes for testing and developing general ecological theory is emphasized.     

Molecular and morphological analysis of secondary contact zones of Cottus gobio in Fennoscandia: geographical discordance of character transitions

Northern Europe was postglacially colonized from different directions by distinct phylogeographical lineages of the bullhead Cottus gobio L. (Pisces: Scorpaeniformes). These lineages have then come into contact in coastal habitats of the currently brackish Baltic Sea and in the freshwaters north of it. We studied the patterns of intergradation in the contact zones in four morphometric and six molecular characters. In the north, intergradation between the western (W) and eastern (E) bullhead lineages is found both among rivers (west‐to‐east) and along individual rivers (south‐to‐north). The locations of the transition zones probably relate to the timing of the initial contact, subsequent Baltic shoreline displacement (i.e. emergence of the lower river reaches), and dispersal barriers caused by variations of coastal salinity. The transitions (clines) in different characters are, however, not geographically coincident. Mitochondrial DNA clines are generally found upstream and to the east of the other transitions, and GPI‐1 allozyme clines are mostly shifted downstream in the rivers, and west of the other transitions on the broader scale of the Baltic Sea. The location of the mtDNA clines may best reflect the initial contact between lineages, and the displacement of the other clines could result from dispersal being overall asymmetric (predominantly downstream) and sex‐biased (stronger in males). Alternatively, the non‐coincidence might reflect selection against deleterious cytonuclear character combinations. No clear evidence of reproductive incompatibility between the lineages was seen in local population structures; no remaining genetic correlations were observed locally among traits. In another transition area, a coastal transect in southern Finland, clinal patterns similar to those in the northern contact zone were recorded, but the population compositions could not be explained by simple in situ mixing of any of the putatively pure, invading refugial lineages. Probably, the bullhead stocks that initially came into contact in this southern study area already represented mixtures of the invading lineages. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 81 , 535–552.     

Relationships between distribution and abundance vary with spatial scale and ecological group in stream bryophytes

1. We examined whether the local abundance of stream bryophytes in a boreal drainage basin (Koutajoki system in northeastern Finland) correlated with their: (i) regional occupancy; (ii) provincial distribution in northwestern Europe; and (iii) global range size. We specifically tested whether aquatic and semi‐aquatic species differ in their distribution–abundance relationships. We also analysed the frequency distributions of occupancy at two spatial scales: within the focal drainage system and across provinces of northwestern Europe. 2. Regional occupancy and mean local abundance of stream bryophytes were positively correlated, and the relationship was rather strong in aquatic species but very weak in semi‐aquatic species. Local abundance was related neither to provincial distribution nor global distribution. 3. Species frequency distributions differed between regional occupancy and provincial distribution. While most species were rare with regard to their regional occupancy within the focal drainage system, most of the same set of species were common and occurred in most provinces in northwestern Europe. 4. The results indicate the presence of dominants (core species) and transients/subordinates (satellite species) among stream bryophytes, highlighting marked differentiation in life‐history strategies and growth form. The observed abundance–occupancy relationships suggest that dispersal limitation and metapopulation processes may govern the dynamics of obligatory aquatic stream bryophytes. In semi‐aquatic species, however, habitat availability may be more important in contributing to regional occupancy.     

The enigma of the landlocked Baikal and Caspian seals addressed through phylogeny of phocine mitochondrial sequences

The endemic seals of Lake Baikal ( Phoca sibirica ) and of the Caspian Sea ( Phoca caspica ) inhabit ancient continental basins that have remained isolated from primary marine seal habitats for millions of years. The species have been united with the Arctic ringed seal, Phoca hispida , into (sub)genus Pusa , but the age and route of invasions to/from the continental basins remain controversial. A phylogenetic analysis of nine northern phocines based on three mitochondrial genes (Cyt b , COI, COII, total 3369 bp) provided no support for the monophyly of the Pusa group. The three species are involved in an apparent polytomy with the boreal harbour seal, Phoca vitulina , and grey seal, Halichoerus grypus . From the average estimated interspecies divergence (4.1%), the radiation of this group plausibly took place in the Late Pliocene 2–3 Mya. This dating does not fit the prevailing hypotheses on the origin of the landlocked taxa in association with Middle Pleistocene glacial events, or of the Caspian seal as a direct descendant of Miocene fossil phocines of the continental Paratethyan basin. The current phocine diversity more likely results from marine radiations, and the continental seals invaded their basins through Plio-Pleistocene (marine) connections from the north. The palaeohydrography that would have enabled the invasions at that time still remains an enigma.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 61–72.