Variation in offspring size towards the high elevational range edge of a montane annual

Understanding the ecological determinants of species’ distribution is a fundamental goal of ecology, and is increasingly important with changing limits to species’ range. Species often reach distributional limits on gradients of resource availability, but the extent to which offspring provisioning varies towards range limits is poorly understood. Selection is generally expected to favour higher provisioning of individual offspring in environments with short growing seasons and limited moisture, nutrients, or hosts for parasitism. However, individual provisioning may decline if parent size is limited by resources. This thesis focuses on three major questions: 1) does seed size vary over an elevational gradient? 2) does this variation respond adaptively towards the range limit? and 3) is potential elevational variation environmentally or genetically controlled? I tested variation in seed investment towards the upper elevational limit of the hemiparasitic annual herb Rhinanthus minor, sampled across an elevational range of 1,000m in the Rocky Mountains of Alberta, Canada. I also used a reciprocal transplant experiment to address the heritability of seed mass. Seed mass increased marginally towards higher elevations, while seed number and plant size declined. There was a strong elevational increase in seed mass scaled by overall plant size. Therefore, investment in individual seeds was higher towards the upper range edge, indicating potential adaptation of the reproductive strategy to allow for establishment in marginal environments. Genetic, environmental, and genotype-by-environment interactions were observed in transplanted populations, but the relative proportions of these effects on seed size were unclear.

Range-edge effects promote clonal growth in peripheral populations of the one-sided wintergreen Orthilia secunda

Existing in suboptimal conditions is a frequent occurrence for species inhabiting the cusp of their ecological range. In range-edge populations of plants, the scarcity of suitable habitat may be reflected in small population sizes which may result in increased self-pollination and/or inbreeding and an increase in the incidence of clonal reproduction. These factors may result in a decrease in levels of genetic diversity and a loss of potential adaptive variation that may compromise species' ability to cope with changes in their environment, an issue that is particularly relevant today with the current concern surrounding global climate change and its effect on species' distributional ranges. In the present study, we have compared the levels of clonal reproduction in the one-sided wintergreen Orthilia secunda (L.) House in (1) populations from its main continuous distribution range, (2) populations occurring on the limits of the continuous range, and (3) peripheral populations outwith the species' continuous distribution range. Range-edge populations in Scotland and Sweden displayed significantly lower genotypic richness and diversity than those from the main area of the species' distribution in these countries. Populations from Ireland, which occur in the temperate zone rather than the boreal conditions that are the preferred habitat for the species, and which represent relict populations left over from cooler periods in the Earth's history, displayed no within-population genetic diversity, suggesting a complete lack of sexual reproduction. Furthermore, the genetic distinctiveness of the Irish populations, which contained alleles not found in either the Scottish or the Swedish populations, highlights the value of 'trailing edge' populations and supports the concept of 'parochial conservation', namely the conservation of species that are locally rare but globally common.

Genetic structure at range edge: Low diversity and high inbreeding in Southeast Asian mangrove (Avicennia marina) populations

Understanding the genetic composition and mating systems of edge populations provides important insights into the environmental and demographic factors shaping species’ distribution ranges. We analysed samples of the mangrove Avicennia marina from Vietnam, northern Philippines and Australia, with microsatellite markers. We compared genetic diversity and structure in edge (Southeast Asia, and Southern Australia) and core (North and Eastern Australia) populations, and also compared our results with previously published data from core and southern edge populations. Comparisons highlighted significantly reduced gene diversity and higher genetic structure in both margins compared to core populations, which can be attributed to very low effective population size, pollinator scarcity and high environmental pressure at distribution margins. The estimated level of inbreeding was significantly higher in northeastern populations compared to core and southern populations. This suggests that despite the high genetic load usually associated with inbreeding, inbreeding or even selfing may be advantageous in margin habitats due to the possible advantages of reproductive assurance, or local adaptation. The very high level of genetic structure and inbreeding show that populations of A. marina are functioning as independent evolutionary units more than as components of a metapopulation system connected by gene flow. The combinations of those characteristics make these peripheral populations likely to develop local adaptations and therefore to be of particular interest for conservation strategies as well as for adaptation to possible future environmental changes.

Historical comparisons reveal multiple drivers of decadal change of an ecosystem engineer at the range edge

Biogenic reefs are important for habitat provision and coastal protection. Long-term datasets on the distribution and abundance of Sabellaria alveolata (L.) are available from Britain. The aim of this study was to combine historical records and contemporary data to (1) describe spatiotemporal variation in winter temperatures, (2) document short-term and long-term changes in the distribution and abundance of S. alveolata and discuss these changes in relation to extreme weather events and recent warming, and (3) assess the potential for artificial coastal defense structures to function as habitat for S. alveolata. A semi-quantitative abundance scale (ACFOR) was used to compare broadscale, long-term and interannual abundance of S. alveolata near its range edge in NW Britain. S. alveolata disappeared from the North Wales and Wirral coastlines where it had been abundant prior to the cold winter of 1962/1963. Population declines were also observed following the recent cold winters of 2009/2010 and 2010/2011. Extensive surveys in 2004 and 2012 revealed that S. alveolata had recolonized locations from which it had previously disappeared. Furthermore, it had increased in abundance at many locations, possibly in response to recent warming. S. alveolata was recorded on the majority of artificial coastal defense structures surveyed, suggesting that the proliferation of artificial coastal defense structures along this stretch of coastline may have enabled S. alveolata to spread across stretches of unsuitable natural habitat. Long-term and broadscale contextual monitoring is essential for monitoring responses of organisms to climate change. Historical data and gray literature can be invaluable sources of information. Our results support the theory that Lusitanian species are responding positively to climate warming but also that short-term extreme weather events can have potentially devastating widespread and lasting effects on organisms. Furthermore, the proliferation of coastal defense structures has implications for phylogeography, population genetics, and connectivity of coastal populations.

Historical comparisons reveal multiple drivers of decadal change of an ecosystem engineer at the range edge

Biogenic reefs are important for habitat provision and coastal protection. Long-term datasets on the distribution and abundance of Sabellaria alveolata (L.) are available from Britain. The aim of this study was to combine historical records and contemporary data to (1) describe spatiotemporal variation in winter temperatures, (2) document short-term and long-term changes in the distribution and abundance of S. alveolata and discuss these changes in relation to extreme weather events and recent warming, and (3) assess the potential for artificial coastal defense structures to function as habitat for S. alveolata. A semi-quantitative abundance scale (ACFOR) was used to compare broadscale, long-term and interannual abundance of S. alveolata near its range edge in NW Britain. S. alveolata disappeared from the North Wales and Wirral coastlines where it had been abundant prior to the cold winter of 1962/1963. Population declines were also observed following the recent cold winters of 2009/2010 and 2010/2011. Extensive surveys in 2004 and 2012 revealed that S. alveolata had recolonized locations from which it had previously disappeared. Furthermore, it had increased in abundance at many locations, possibly in response to recent warming. S. alveolata was recorded on the majority of artificial coastal defense structures surveyed, suggesting that the proliferation of artificial coastal defense structures along this stretch of coastline may have enabled S. alveolata to spread across stretches of unsuitable natural habitat. Long-term and broadscale contextual monitoring is essential for monitoring responses of organisms to climate change. Historical data and gray literature can be invaluable sources of information. Our results support the theory that Lusitanian species are responding positively to climate warming but also that short-term extreme weather events can have potentially devastating widespread and lasting effects on organisms. Furthermore, the proliferation of coastal defense structures has implications for phylogeography, population genetics, and connectivity of coastal populations.

Broad-scale patterns of sex ratios in Patella spp.: a comparison of range edge and central range populations in the British Isles and Portugal

Sex change, or sequential hermaphroditism, occurs in the plant and animal kingdoms and often determines a predominance of the first sex. Our aim was to explore changes in sex ratios within the range of the species studied: Patella vulgata and Patella depressa. The broad-scale survey of sex with size of limpets covered a range of latitudes from Zambujeira do Mar (southern Portugal) to the English Channel. Indirect evidence was found for the occurrence of protandry in P. vulgata populations from the south of England, with females predominating in larger size-classes; cumulative frequency distributions of males and females were different; sex ratios were biased towards males and smallest sizes of males were smaller than the smallest sizes of females. In contrast in Portugal females were found in most size-classes of P. vulgata. In P. depressa populations from the south coast of England and Portugal females were interspersed across most size-classes; size distributions of males and females and size at first maturity of males and females did not differ. P. depressa did, however, show some indications of the possibility of slight protandry occurring in Portugal. The test of sex ratio variation with latitude indicated that P. vulgata sex ratios might be involved in determining the species range limit, particularly at the equatorward limit since the likelihood of being male decreased from the south coast of England to southern Portugal. Thus at the southern range limit, sperm could be in short supply due to scarcity of males contributing to an Allee effect.

Broad-scale patterns of sex ratios in Patella spp.: a comparison of range edge and central range populations in the British Isles and Portugal

Sex change, or sequential hermaphroditism, occurs in the plant and animal kingdoms and often determines a predominance of the first sex. Our aim was to explore changes in sex ratios within the range of the species studied: Patella vulgata and Patella depressa. The broad-scale survey of sex with size of limpets covered a range of latitudes from Zambujeira do Mar (southern Portugal) to the English Channel. Indirect evidence was found for the occurrence of protandry in P. vulgata populations from the south of England, with females predominating in larger size-classes; cumulative frequency distributions of males and females were different; sex ratios were biased towards males and smallest sizes of males were smaller than the smallest sizes of females. In contrast in Portugal females were found in most size-classes of P. vulgata. In P. depressa populations from the south coast of England and Portugal females were interspersed across most size-classes; size distributions of males and females and size at first maturity of males and females did not differ. P. depressa did, however, show some indications of the possibility of slight protandry occurring in Portugal. The test of sex ratio variation with latitude indicated that P. vulgata sex ratios might be involved in determining the species range limit, particularly at the equatorward limit since the likelihood of being male decreased from the south coast of England to southern Portugal. Thus at the southern range limit, sperm could be in short supply due to scarcity of males contributing to an Allee effect.

Unidirectional hybridization at a species' range boundary: implications for habitat tracking

Aim Introgressive hybridization between a locally rare species and a more abundant congener can drive population extinction via genetic assimilation, or the replacement of the rare species gene pool with that of the common species. To date, however, few studies have assessed the effects of such processes at the limits of species' distribution ranges. In this study, we have examined the potential for hybridization between range-edge populations of the wintergreen Pyrola minor and sympatric populations of Pyrola grandiflora. Location Qeqertarsuaq, Greenland and Churchill, Manitoba, Canada. Methods Genetic analysis of samples from Greenland and Canada was carried out using a combination of nuclear and chloroplast single nucleotide polymorphisms (SNPs). Results Analysis of nuclear SNPs confirmed hybridization in populations of morphologically intermediate individuals, as well as revealing the existence of cryptic hybrids in ostensibly morphologically pure P. minor populations. Analysis of chloroplast SNPs revealed that this hybridization is unidirectional and suggests that hybrids originate via pollen swamping of P. minor by the more common P. grandiflora. Main conclusions Extensive unidirectional hybridization may lead to the extinction of peripheral populations of P. minor where the two species grow sympatrically. Extinction could occur as a result of genetic assimilation where F1s are fertile, or via the removal of unidirectionally pollinated sterile F1s, or by a combination of these processes. This could compromise the ability of species to respond to climate change via habitat tracking, although the final outcome of these processes may ultimately depend on the rate of global climate change and its effect on the species' distributions. © 2009 Blackwell Publishing Ltd.

Detecting range shifts among Australian fishes in response to climate change

One of the most obvious and expected impacts of climate change is a shift in the distributional range of organisms, which could have considerable ecological and economic consequences. Australian waters are hotspots for climate-induced environmental changes; here, we review these potential changes and their apparent and potential implications for freshwater, estuarine and marine fish. Our meta-analysis detected 300 papers globally on 'fish' and 'range shifts', with ∼7% being from Australia. Of the Australian papers, only one study exhibited definitive evidence of climate-induced range shifts, with most studies focussing instead on future predictions. There was little consensus in the literature regarding the definition of 'range', largely because of populations having distributions that fluctuate regularly. For example, many marine populations have broad dispersal of offspring (causing vagrancy). Similarly, in freshwater and estuarine systems, regular environmental changes (e.g. seasonal, ENSO cycles not related to climate change) cause expansion and contraction of populations, which confounds efforts to detect range 'shifts'. We found that increases in water temperature, reduced freshwater flows and changes in ocean currents are likely to be the key drivers of climate-induced range shifts in Australian fishes. Although large-scale frequent and rigorous direct surveys of fishes across their entire distributional ranges, especially at range edges, will be essential to detect range shifts of fishes in response to climate change, we suggest careful co-opting of fisheries, museum and other regional databases as a potential, but imperfect alternative. © 2011 CSIRO Open Access.

Oriented Percolation in One-dimensional 1/vertical bar x-y vertical bar(2) Percolation Models

We consider independent edge percolation models on Z, with edge occupation probabilities. We prove that oriented percolation occurs when beta > 1 provided p is chosen sufficiently close to 1, answering a question posed in Newman and Schulman (Commun. Math. Phys. 104: 547, 1986). The proof is based on multi-scale analysis.

Hydrophobic attraction may contribute to aqueous flocculation of clays

SEM observations of low solid content vitrified clay suspensions reveal that clay platelets build porous three-dimensional networks with platelets contacting each other mostly by their edges. To explain this behaviour, which must require long range edge-to-edge (EE) attractive forces, a hydrophobic-like interaction has been proposed. This interaction may be induced by the presence of nano-bubbles existing on the edges of clay crystals which may cause clay particles to flocculate. The following indirect evidence for such hydrophobic behaviour is presented. First, a clay platelet is shown attached to an oil drop by its edge; second, clay flocs were attracted by a vertically placed Teflon strip but not to the hydrophilic mica basal surface; third, a much thicker porous sediment occurred in CO2-saturated water solution compared with vacuum degassed water.

The structure of biogenic habitat and epibiotic assemblages associated with the global invasive kelp Undaria pinnatifida in comparison to native macroalgae

Kelp forests dominate temperate and polar rocky coastlines and represent critical marine habitats because they support elevated rates of primary and secondary production and high biodiversity. A major threat to the stability of these ecosystems is the proliferation of non-native species, such as the Japanese kelp Undariapinnatifida (‘Wakame’), which has recently colonised natural habitats in the UK. We quantified the abundance and biomass of U. pinnatifida on a natural rocky reef habitat over 10 months to make comparisons with three native canopy-forming brown algae (Laminaria ochroleuca, Saccharina latissima, and Saccorhiza polyschides). We also examined the biogenic habitat structure provided by, and epibiotic assemblages associated with, U. pinnatifida in comparison to native macroalgae. Surveys conducted within the Plymouth Sound Special Area of Conservation indicated that U. pinnatifida is now a dominant and conspicuous member of kelp-dominated communities on natural substrata. Crucially, U. pinnatifida supported a structurally dissimilar and less diverse epibiotic assemblage than the native perennial kelp species. However, U. pinnatifida-associated assemblages were similar to those associated with Saccorhiza polyschides, which has a similar life history and growth strategy. Our results suggest that a shift towards U. pinnatifida dominated reefs could result in impoverished epibiotic assemblages and lower local biodiversity, although this could be offset, to some extent, by the climate-driven proliferation of L. ochroleuca at the poleward range edge, which provides complex biogenic habitat and harbours relatively high biodiversity. Clearly, greater understanding of the long-term dynamics and competitive interactions between these habitat-forming species is needed to accurately predict future biodiversity patterns.

The structure of biogenic habitat and epibiotic assemblages associated with the global invasive kelp Undaria pinnatifida in comparison to native macroalgae

Kelp forests dominate temperate and polar rocky coastlines and represent critical marine habitats because they support elevated rates of primary and secondary production and high biodiversity. A major threat to the stability of these ecosystems is the proliferation of non-native species, such as the Japanese kelp Undariapinnatifida (‘Wakame’), which has recently colonised natural habitats in the UK. We quantified the abundance and biomass of U. pinnatifida on a natural rocky reef habitat over 10 months to make comparisons with three native canopy-forming brown algae (Laminaria ochroleuca, Saccharina latissima, and Saccorhiza polyschides). We also examined the biogenic habitat structure provided by, and epibiotic assemblages associated with, U. pinnatifida in comparison to native macroalgae. Surveys conducted within the Plymouth Sound Special Area of Conservation indicated that U. pinnatifida is now a dominant and conspicuous member of kelp-dominated communities on natural substrata. Crucially, U. pinnatifida supported a structurally dissimilar and less diverse epibiotic assemblage than the native perennial kelp species. However, U. pinnatifida-associated assemblages were similar to those associated with Saccorhiza polyschides, which has a similar life history and growth strategy. Our results suggest that a shift towards U. pinnatifida dominated reefs could result in impoverished epibiotic assemblages and lower local biodiversity, although this could be offset, to some extent, by the climate-driven proliferation of L. ochroleuca at the poleward range edge, which provides complex biogenic habitat and harbours relatively high biodiversity. Clearly, greater understanding of the long-term dynamics and competitive interactions between these habitat-forming species is needed to accurately predict future biodiversity patterns.