Biotic disturbance and benthic community dynamics in salmon-bearing streams

1. Organisms can impact ecosystems via multiple pathways, often with positive and negative impacts on inhabitants. Understanding the context dependency of these types of impacts remains challenging. For example, organisms may perform different functions at different densities. 2. Anadromous salmon accumulate > 99% of their lifetime growth in marine ecosystems, and then return to spawn, often at high densities, in relatively confined freshwaters. While previous research has focused on how salmon nutrients can fertilize benthic communities, we examined how an ecosystem engineer, sockeye salmon Oncorhynchus nerka, influences seasonal dynamics of stream benthic communities through their nest-digging activities in south-western Alaska, USA. Benthic invertebrate and algal abundance were quantified every 7-14 days during the open water seasons of 10 streams in riffle and run habitats across multiple years, leading to 25 different stream-year combinations that spanned a large gradient of salmon density. 3. In streams with few or no salmon, benthic algal and insect biomass were fairly constant throughout the season. However, in streams with more than 0.1 salmon m(-2), algal and insect biomass decreased by an average of 75-85% during salmon spawning. Algal biomass recovered quickly following salmon disturbance, occasionally reaching pre-salmon biomass. In contrast, in streams with more than 0.1 salmon m(-2), aquatic insect populations did not recover to pre-salmon levels within the same season. We observed no positive impacts of salmon on algae or insects via fertilization from carcass nutrients. 4. Salmon, when their populations exceed thresholds in spawning density, are an important component of stream disturbance regimes and influence seasonal dynamics of benthic communities. Human activities that drive salmon densities below threshold densities, as has likely happened in many streams, will lead to altered seasonal dynamics of stream communities. Human activities that alter animal populations that are sources of biogenic disturbance can result in shifts in community dynamics.     

Quantitative Links Between Pacific Salmon and Stream Periphyton

Species’ impacts on primary production can have strong ecological consequences. In freshwater ecosystems, Pacific salmon (Oncorhynchus spp.) may influence stream periphyton through substrate disturbance during spawning and nutrient subsidies from senescent adults. The shape of relationships between the abundance of spawning salmon and stream periphyton, as well as interactions with environmental variables, are incompletely understood and may differ across the geographic range of salmon. We examined these relationships across 24 sockeye salmon (Oncorhynchus nerka) spawning streams in north-central British Columbia, Canada. The influence of salmon abundance and environmental variables (temperature, light, dissolved nutrients, water velocity, watershed size, and invertebrate grazer abundance) on post-spawning periphyton abundance and nitrogen stable isotope signatures, which can indicate the uptake of salmon nitrogen, was evaluated using linear regression models and Akaike Information Criterion. Periphyton nitrogen stable isotope signatures were best described by a positive log-linear relationship with an upstream salmon abundance metric that includes salmon from earlier years. This suggests the presence of a nutrient legacy. In contrast, periphyton abundance was negatively related to the spawning-year salmon density, which likely results from substrate disturbance during spawning, and positively related to dissolved soluble reactive phosphorus prior to spawning, which may indicate phosphorus limitation in the streams. These results suggest that enrichment from salmon nutrients does not always translate into elevated periphyton abundance. This underscores the need to directly assess the outcome of salmon impacts on streams rather than extrapolating from stable isotope evidence for the incorporation of salmon nutrients into food webs.     

Movers and shakers: nutrient subsidies and benthic disturbance predict biofilm biomass and stable isotope signatures in coastal streams

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Separating physical disturbance and nutrient enrichment caused by Pacific salmon in stream ecosystems

1. Pacific salmon (Oncorhynchus spp.) deliver marine‐derived nutrients to the streams in which they spawn and die, and these resource subsidies can increase the abundance of stream biota. In strong contrast, physical disturbance from salmon spawning activity can reduce the abundance of benthic organisms. Previous experimental designs have not established the relative effects of these two contrasting processes on stream organisms during a salmon run. 2. We combined manipulative and observational field studies to assess the degree of nutrient enrichment, physical disturbance, and the net effect of salmon on the abundance of benthic periphyton. Related salmon‐mediated processes were also evaluated for benthic macroinvertebrates. Mesh exclosures (2 × 2 m plots) prevented salmon from disturbing areas of the stream channel, which were compared with areas to which salmon had access. Sampling was conducted both before and during the late‐summer spawning run of pink (O. gorbushca) and chum (O. keta) salmon. 3. Streamwater nitrogen and phosphorus concentrations increased sharply with the onset of the salmon run, and highly significant positive relationships were observed between the numbers of salmon present in the stream and these dissolved nutrients. Before the salmon run, periphyton biomass (as chlorophyll a) and total macroinvertebrate abundance were very similar between control and exclosure plots. During the salmon run, exclosures departed substantially from controls, suggesting significant disturbance imparted on benthic biota. 4. Comparing exclosures before and during the salmon run enabled us to estimate the effects of salmon in the absence of direct salmon disturbance. This ‘nutrient enrichment potential’ was significant for periphyton biomass, as was a related index for macroinvertebrate abundance (although enhanced invertebrate drift into exclosures during the salmon run could also have been important). Interestingly, however, the net effect of salmon, evaluated by comparing control plots before and during the salmon run, was relatively modest for both periphyton and macroinvertebrates, suggesting that nutrient enrichment effects were largely offset by disturbance. 5. Our results illustrate the importance of isolating the specific mechanisms via which organisms affect ecosystems, and indicate that the relative magnitude of salmon nutrient enrichment and benthic disturbance determines the net effect that these ecologically important fish have on stream ecosystems.     

Recent local adaptation of sockeye salmon to glacial spawning habitats

Salmonids spawn in highly diverse habitats, exhibit strong genetic population structuring, and can quickly colonize newly created habitats with few founders. Spawning traits often differ among populations, but it is largely unknown if these differences are adaptive or due to genetic drift. To test if sockeye salmon (Oncorhynchus nerka) populations are adapted to glacial, beach, and tributary spawning habitats, we examined variation in heritable phenotypic traits associated with spawning in 13 populations of wild sockeye salmon in Lake Clark, Alaska. These populations were commonly founded between 100 and 400 hundred sockeye salmon generations ago and exhibit low genetic divergence at 11 microsatellite loci (F _ST < 0.024) that is uncorrelated with spawning habitat type. We found that mean P _ST (phenotypic divergence among populations) exceeded neutral F _ST for most phenotypic traits measured, indicating that phenotypic differences among populations could not be explained by genetic drift alone. Phenotypic divergence among populations was associated with spawning habitat differences, but not with neutral genetic divergence. For example, female body color was lighter and egg color was darker in glacial than non-glacial habitats. This may be due to reduced sexual selection for red spawning color in glacial habitats and an apparent trade-off in carotenoid allocation to body and egg color in females. Phenotypic plasticity is an unlikely source of phenotypic differences because Lake Clark sockeye salmon spend nearly all their lives in a common environment. Our data suggest that Lake Clark sockeye salmon populations are adapted to spawning in glacial, beach and tributary habitats and provide the first evidence of a glacial spawning ecotype in salmonids. Glacial spawning habitats are often young (i.e., <200 years old) and ephemeral. Thus, local adaptation of sockeye salmon to glacial habitats appears to have occurred recently.     

Microsatellite DNA Data Indicate Distinct Native Populations of Kokanee, Oncorhynchus nerka, Persist in the Lake Sammamish Basin, Washington

Large-scale introductions of resident and anadromous salmonids from exogenous sources and urbanization have led to major changes in, and concern for the fate of, indigenous fish populations of the Lake Sammamish/Lake Washington Basin. Specifically, introductions of kokanee (the resident form of Oncorhynchus nerka) from the Lake Whatcom Hatchery and sockeye (the anadromous form of O. nerka) from Baker Lake have caused uncertainty about the ancestry of the kokanee that currently spawn in the basin. We used nine microsatellite loci to investigate the inter-relationships of kokanee populations that spawn in streams in the Sammamish sub-basin, sockeye salmon populations that share spawning areas with the kokanee, Lake Whatcom Hatchery kokanee and Baker Lake sockeye, and an outgroup, Meadow Creek kokanee, from Lake Kootenay which drains into the upper Columbia River. We observed high levels of genetic variation (5–49 alleles per locus). Explicit tests of population sub-division revealed that collections from most spawning aggregations differed from each other. Observed allele frequency distributions strongly suggest that natural spawning kokanee in the basin are not descended from recent Lake Whatcom stock introductions. We found no compelling evidence to suggest that the kokanee sampled from spawning areas within the Lake Sammamish sub-basin have resulted from, or been altered substantially by, past introductions of non-native kokanee or sockeye.     

Do fall additions of salmon carcasses benefit food webs in experimental streams?

Research showing that salmon carcasses support the productivity and biodiversity of aquatic and riparian ecosystems has been conducted over a variety of spatial and temporal scales. In some studies, carcasses were manipulated in a single pulse or loading rate or manipulations occurred during summer and early fall, rather than simulating the natural dynamic of an extended spawning period, a gradient of loading rates, or testing carcass effects in late fall-early winter when some salmon stocks in the US Pacific Northwest spawn. To address these discrepancies, we manipulated salmon carcass biomass in 16 experimental channels located in the sunlit floodplain of the Cedar River, WA, USA between mid-September and mid-December, 2006. Total carcass loads ranged from 0?C4.0 kg/m² (0, 0.001, 0.01, 0.1, 0.5, 1.0, 2.0 and 4.0 kg/m², n = 2 per treatment) and were added to mimic the temporal dynamic of an extended spawning period. We found little evidence that carcasses influenced primary producer biomass or fish growth; however, nutrients and some primary consumer populations increased with loading rate. These effects varied through time, however. We hypothesize that the variable effects of carcasses were a result of ambient abiotic condition, such as light, temperature and disturbance that constrained trophic response. There was some evidence to suggest peak responses for primary producers and consumers occurred at a loading rate of ~1.0?C2.0 kg/m², which was similar to other experimental studies conducted during summer.     

Critical swimming velocities of juvenile sockeye salmon and kokanee, the anadromous and non-anadromous forms of Oncorhynchus nerka (Walbaum)

In streams tributary to the North Pacific, anadromous sockeye salmon and non-anadromous kokanee, Oncorhynchus nerka (Walbaum), occasionally spawn sympatrically and male kokanee may act as 'sneaks’to spawn with the larger female sockeye. Despite this interbreeding, sockeye and kokanee exhibit persistent biochemical genetic differences at several enzyme loci. Genetic differences between forms may be maintained by selection against‘hybrids’due to the different life histories of sockeye and kokanee; sockeye make extensive smolt, oceanic, and spawning migrations while kokanee reside permanently in fresh water. We tested the sustained swimming abilities of juvenile sockeye, kokanee, and sockeye (female) × kokanee (male) hybrids to see if hybrids were inferior to sockeye in a trait that is probably under stronger selection in an anadromous life history. Sockeye had significantly greater mean critical swimming velocities (U_crit) than kokanee of the same size raised under identical conditions (8.3 v. 7.3 body lengths s^?1 respectively). When tested 1 month later the mean U_crit of sockeye was only marginally greater than that for sockeye × kokanee hybrids (both c. 6.6 body lengths s^?1). Sockeye swimming performance was also less variable than that of either kokanee or hybrids. Sockeye tended to have slimmer bodies and longer caudal regions than kokanee or sockeye × kokanee hybrids of the same size. Sockeye also had significantly more vertebrae than kokanee and hybrids, while hybrids had more vertebrae than kokanee. These morphological differences may have contributed to the differences in swimming performance. We concluded: (i) that juvenile sockeye and kokanee have diverged with respect to sustained swimming performance and that reduced performance by kokanee may be due to relaxed selection for sustained swimming performance associated with their non-anadromous life history, (ii) that sockeye × kokanee hybrids appear to have modestly lower swimming capabilities than pure sockeye, and (iii) if the variability in swimming performance is associated with differences in survival in nature, then such differences may promote divergence between sympatric sockeye and kokanee.     

Energy expenditures during spawning by chum salmon Oncorhynchus keta (Walbaum) in British Columbia

Physiological telemetry and proximate tissue analyses were used to assess energy expended by chum salmon Oncorhynchus keta on various behaviours during spawning in Kanaka Creek, British Columbia, Canada, and results were compared with published data on Fraser River sockeye salmon Oncorhynchus nerka , the only other species for which both types of measurements have been taken. Chum salmon arrived at the spawning grounds with body energy densities of 4·84 MJ kg⁻¹ in males and 4·62 MJ kg⁻¹ in females, lower than most sockeye salmon populations, and died with energy densities of c . 4 MJ kg⁻¹, similar to that observed in sockeye salmon and other salmonids. Moisture levels generally increased in body tissues over the spawning life, particularly in female gonads, and lipid levels decreased. Declines in protein observed over the spawning life of other Pacific salmon Oncorhynchus sp. were less evident in Kanaka Creek chum salmon. Holding behaviour constituted the dominant component of the activity schedule and energy budget of both sexes. After holding, the most expensive behaviours were nest digging in females and aggressive displays in males. Dominant males expended the most energy on behaviours each day, as indexed by oxygen consumption (3600 mgO₂ kg⁻¹), while satellite males expended nearly as much (3504 mgO₂ kg⁻¹) but females expended considerably less (2327 mgO₂ kg⁻¹). Kanaka chum salmon engaged more frequently in energetically expensive reproductive behaviours than Stuart River sockeye salmon.     

MOLECULAR GENETIC EVIDENCE FOR PARALLEL LIFE-HISTORY EVOLUTION WITHIN A PACIFIC SALMON (SOCKEYE SALMON AND KOKANEE,ONCORHYNCHUS NERKA)

The Pacific salmon Oncorhynchus nerka typically occurs as a sea-run form (sockeye salmon) or may reside permanently in lakes (kokanee) thoughout its native North Pacific. We tested whether such geographically extensive ecotypic variation resulted from parallel evolutionary divergence thoughout the North Pacific or whether the two forms are monophyletic groups by examining allelic variation between sockeye salmon and kokanee at two minisatellite DNA repeat loci and in mitochondrial DNA (mtDNA) Bgl II restriction sites. Our examination of over 750 fish from 24 populations, ranging from Kamchatka to the Columbia River, identified two major genetic groups of North Pacific O. nerka: a “northwestern” group consisting of fish from Kamchatka, western Alaska, and northwestern British Columbia, and a “southern” group consisting of sockeye salmon and kokanee populations from the Fraser and Columbia River systems. Maximum-likelihood analysis accompanied by bootstrapping provided strong support for these two genetic groups of O. nerka; the populations did not cluster by migratory form, but genetic affinities were organized more strongly by geographic proximity. The two major genetic groups resolved in our study probably stem from historical isolation and dispersal of O. nerka from two major Wisconsinan glacial refugia in the North Pacific. There were significant minisatellite DNA allele frequency differences between sockeye salmon and kokanee populations from different parts of the same watershed, between populations spawning in different tributaries of the same lake, and also between sympatric populations spawning in the same stream at the same time. MtDNA Bgl II restriction site variation was significant between sockeye salmon and kokanee spawning in different parts of the same major watershed but not between forms spawning in closer degrees of reproductive sympatry. Patterns of genetic affinity and allele sharing suggested that kokanee have arisen from sea-run sockeye salmon several times independently in the North Pacific. We conclude that sockeye salmon and kokanee are para- and polyphyletic, respectively, and that the present geographic distribution of the ecotypes results from parallel evolutionary origins of kokanee from sockeye (divergences between them) thoughout the North Pacific.     

Evaluating Relationships between Wild Skeena River Sockeye Salmon Productivity and the Abundance of Spawning Channel Enhanced Sockeye Smolts

The enhancement of salmon populations has long been used to increase the abundance of salmon returning to spawn and/or to be captured in fisheries. However, in some instances enhancement can have adverse impacts on adjacent non-enhanced populations. In Canada''s Skeena watershed, smolt-to-adult survival of Babine Lake sockeye from 1962–2002 was inversely related to the abundance of sockeye smolts leaving Babine Lake. This relationship has led to the concern that Babine Lake smolt production, which is primarily enhanced by spawning channels, may depress wild Skeena (Babine and non-Babine) sockeye populations as a result of increased competition between wild and enhanced sockeye smolts as they leave their natal lakes and co-migrate to sea. To test this hypothesis we used data on Skeena sockeye populations and oceanographic conditions to statistically examine the relationship between Skeena sockeye productivity (adult salmon produced per spawner) and an index of Babine Lake enhanced smolt abundance while accounting for the potential influence of early marine conditions. While we had relatively high power to detect large effects, we did not find support for the hypothesis that the productivity of wild Skeena sockeye is inversely related to the abundance of enhanced sockeye smolts leaving Babine Lake in a given year. Importantly, life-time productivity of Skeena sockeye is only partially explained by marine survival, and likely is an unreliable measure of the influence of smolt abundance. Limitations to our analyses, which include: (1) the reliance upon adult salmon produced per spawner (rather than per smolt) as an index of marine survival, and (2) incomplete age structure for most of the populations considered, highlight uncertainties that should be addressed if understanding relationships between wild and enhanced sockeye is a priority in the Skeena.     

Long‐term zooplankton responses to nutrient and consumer subsidies arising from migratory sockeye salmon (Oncorhynchus nerka)

Migratory animals, such as Pacific salmon, can significantly shape communities in recipient habitats both by altering the flux of resources, and changing community composition and subsequent trophic interactions. Here we mainly used paleoecological records from natural sockeye salmon nursery lakes to quantify the response of plankton communities to the influx of salmon‐derived nutrients and consumers (juvenile salmon). Our long‐term data show that increases in the density of spawning salmon often elevated influx of nutrients, and, in turn, zooplankton production over the past few centuries. In contrast, significant correlations were not detected in two lakes with extremely low or high average spawner densities (i.e. 1.5 and 34.7 × 10³ spawners km^?2 year^?1 respectively). With increasing spawner densities across lakes, analysis of the size structure of subfossils in sediments revealed a strong decrease in body size of a main juvenile salmon prey item (Eubosmina longispina; r²= 0.36, p < 0.001, n = 67), consistent with an overriding effect of predation in lakes with high salmon densities. These long‐term data not only highlight the key role of salmon‐derived nutrients in stimulating plankton communities, but also suggest that the relative effect of nutrient and consumer subsidies varies along gradients of lake production, despite a single ultimate causal mechanism (migrating fish).     

Recovery of Benthic Megafauna from Anthropogenic Disturbance at a Hydrocarbon Drilling Well (380 m Depth in the Norwegian Sea)

Recovery from disturbance in deep water is poorly understood, but as anthropogenic impacts increase in deeper water it is important to quantify the process. Exploratory hydrocarbon drilling causes physical disturbance, smothering the seabed near the well. Video transects obtained by remotely operated vehicles were used to assess the change in invertebrate megafaunal density and diversity caused by drilling a well at 380 m depth in the Norwegian Sea in 2006. Transects were carried out one day before drilling commenced and 27 days, 76 days, and three years later. A background survey, further from the well, was also carried out in 2009. Porifera (45% of observations) and Cnidaria (40%) dominated the megafauna. Porifera accounted for 94% of hard-substratum organisms and cnidarians (Pennatulacea) dominated on the soft sediment (78%). Twenty seven and 76 days after drilling commenced, drill cuttings were visible, extending over 100 m from the well. In this area there were low invertebrate megafaunal densities (0.08 and 0.10 individuals m⁻²) in comparison to pre-drill conditions (0.21 individuals m⁻²). Three years later the visible extent of the cuttings had reduced, reaching 60 m from the well. Within this area the megafaunal density (0.05 individuals m⁻²) was lower than pre-drill and reference transects (0.23 individuals m⁻²). There was a significant increase in total megafaunal invertebrate densities with both distance from drilling and time since drilling although no significant interaction. Beyond the visible disturbance there were similar megafaunal densities (0.14 individuals m⁻²) to pre-drilling and background surveys. Species richness, Shannon-Weiner diversity and multivariate techniques showed similar patterns to density. At this site the effects of exploratory drilling on megafaunal invertebrate density and diversity seem confined to the extent of the visible cuttings pile. However, elevated Barium concentration and reduced sediment grain size suggest persistence of disturbance for three years, with unclear consequences for other components of the benthic fauna.     

Effects of Salmon-Derived Nutrients and Habitat Characteristics on Population Densities of Stream-Resident Sculpins

Movement of nutrients across ecosystem boundaries can have important effects on food webs and population dynamics. An example from the North Pacific Rim is the connection between productive marine ecosystems and freshwaters driven by annual spawning migrations of Pacific salmon (Oncorhynchus spp). While a growing body of research has highlighted the importance of both pulsed nutrient subsidies and disturbance by spawning salmon, their effects on population densities of vertebrate consumers have rarely been tested, especially across streams spanning a wide range of natural variation in salmon densities and habitat characteristics. We studied resident freshwater prickly (Cottus asper), and coastrange sculpins (C. aleuticus) in coastal salmon spawning streams to test whether their population densities are affected by spawning densities of pink and chum salmon (O. gorbuscha and O. keta), as well as habitat characteristics. Coastrange sculpins occurred in the highest densities in streams with high densities of spawning pink and chum salmon. They also were more dense in streams with high pH, large watersheds, less area covered by pools, and lower gradients. In contrast, prickly sculpin densities were higher in streams with more large wood and pools, and less canopy cover, but their densities were not correlated with salmon. These results for coastrange sculpins provide evidence of a numerical population response by freshwater fish to increased availability of salmon subsidies in streams. These results demonstrate complex and context-dependent relationships between spawning Pacific salmon and coastal ecosystems and can inform an ecosystem-based approach to their management and conservation.     

Asynchrony in population dynamics of sockeye salmon in southwest Alaska

Ecologists have examined the synchronization of population dynamics across space as a means to understand how populations respond to climate variation. However, response diversity may reflect important variation among local population dynamics driven by population‐specific responses to regional environmental change. We used long‐term data on sockeye salmon Oncorhynchus nerka from pristine watersheds of southwestern Alaska to show that populations spawning in close proximity (<40 km) to one another have a limited degree of synchrony in their dynamics, even after accounting for density‐dependent processes. In fact, the dynamics of local populations of stream‐spawning sockeye salmon were no more coherent than those of stocks at a much coarser resolution across this region of Alaska. We examined four hypotheses to explain the observed patterns of asynchrony among stream‐spawning populations, and found that populations spawning in dissimilar habitats, and using different nursery lakes were less synchronized in their productivity. Similarity in the age structure of spawning adults was less correlated with synchrony in productivity. These results emphasize the importance of maintaining diverse spawning and rearing habitat for the conservation of Pacific salmon, and should guide conservation planning for Pacific salmon populations in regions where natural dynamics have been altered by habitat loss, hatchery practices, and over‐fishing.     

Long-Term Growth and Succession in Restored and Natural Mangrove Forests in Southwestern Florida

We compared colonization, growth and succession from 1989 to 2000 in a restored mangrove site and in gap and closed canopy sites in a natural mangrove forest. The restored site was created in 1982 and planted with Rhizophora mangle (≈2 m⁻²) propagules. By 1989, Laguncularia racemosa, with densities up to 12.9 tree m⁻², was a dominant in all plots, although densities were greater at edge plots relative to inner plots, and near open water (west plots) relative to further inland (east plots), and in tall mangrove plots relative to scrub plots. Rhizophora mangle (1989 tree densities about 2 m⁻²) was a codominant in inner and scrub plots, while Avicennia germinans had the lowest densities (<1 tree m⁻²) in all plots. From 1989 to 2000 L. racemosa experienced reduced recruitment and apparent density-dependent mortality of canopy individuals in plots with high initial densities. Scrub plots experienced high rates of colonization by R. mangle and L. racemosa, rapid growth in height of all species (1989–1996), followed by a dieoff of L. racemosa in later years (1997–2000) as the canopy came to resemble that of tall mangrove plots. Colonization and growth rates were lower in gap and closed canopy regions of the natural forest relative to rates in the restored site. After 11 years, densities of L. racemosa were 10–20× lower and R. mangle slightly less in the gap relative to densities in tall mangrove plots in the restored site at the same age. Although the restored stand had converged with the natural forest by 2000 in terms of some factors such as species richness, vegetation cover, litterfall, and light penetration, trees were still much smaller and stem densities much higher. Full development of mature structure and ecological function will likely require decades more development.     

Export of non-native gastropod shells to a coastal lagoon: Alteration of habitat structure has negligible effects on infauna

Alteration of physical habitat structure is a fundamental mechanism by which invaders produce ecosystem level effects. We assessed whether, along the east coast of Australia, the impacts of the non-native gastropod Maoricolpus roseus on soft-sediment habitats extend beyond the range of live populations as a result of shell export following death of animals. Sampling over an 18 month period revealed that M. roseus shells were temporally persistent in surface sediments of a coastal lagoon devoid of live populations of the gastropod. The well-preserved shells, of which 92% were entire, did not accumulate above a maximal density of 260 m^− 2 due to periodic burial. Manipulation of M. roseus shell densities indicated that at densities (140 m^− 2) presently experienced within surface sediments of the lagoon, the structure provided by the shells was weakly facilitative of some invertebrate species. Further increasing shell densities to the possible future scenario of 280 m^− 2, which may occur under continued expansion of nearby live populations did not, however, strengthen positive effects. To the contrary, plots with higher densities of M. roseus shells contained similar invertebrate assemblages to control plots, without shells. Consistent with the negligible effects of M. roseus shells on infauna, the foraging efficiency of the generalist predator, Carcinus maenas, and the naticid gastropod, Polinices sordidus, were not affected by addition of shells. Surprisingly, even an extreme scenario of 1600 m^− 2 did not affect their predation. Thus, in this dynamic coastal lagoon, that experiences considerable sediment movement and environmental variability, the structure introduced by import of non-native shells is unimportant in structuring soft-sediment communities. Whether this unrecognised aspect of molluscan invasions impacts the ecology of more stable coastal environments remains unclear and warrants further consideration.     

Countergradient variation in carotenoid use between sympatric morphs of sockeye salmon (Oncorhynchus nerka) exposes nonanadromous hybrids in the wild by their mismatched spawning colour

Understanding the mechanisms that decrease gene flow between diverging populations is critical to understanding speciation. Anadromous (sockeye) and nonanadromous (kokanee) morphs of the Pacific sockeye salmon Oncorhynchus nerka spawn sympatrically and interbreed, yet allele frequency differences at neutral loci indicate restricted gene flow. Disruptive natural selection associated with strong selective differences between anadromous and nonanadromous life histories is thought to maintain the genetic differentiation of the morphs. Recently, a putative third morph of O. nerka exhibiting green rather than red breeding colour has been found on the spawning grounds sympatric with sockeye and kokanee. We investigated the ancestry of these green fish in a 2‐year controlled breeding study by using previously documented heritable, countergradient variation in red breeding colour to distinguish pure and hybrid morphs. Stabilizing sexual selection for similar red breeding colour in sockeye and kokanee has led to adaptive differences in the efficiency of carotenoid uptake between the morphs given differences in carotenoid availability between marine and lacustrine habitats. On the same diet, offspring parented by the green fish were intermediate in colour and in the concentration of dietary carotenoid pigments in their flesh and skin to those parented by either sockeye or kokanee; they were most similar to those parented by known kokanee × sockeye hybrids. This countergradient variation in carotenoid use results in a genotype‐environment mismatch in nonanadromous hybrids that exposes them by their breeding colour on the spawning grounds. Given that red colour is important in mate choice, sexual selection will almost certainly reduce reproductive opportunities for these hybrids and promote sympatric divergence of these incipient species. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 84 , 287–305.     

The Invasive Woody Weed Ligustrum robustum subsp. walkeri Threatens Native Forests on La Réunion

One of the last primitive island ecosystems in the Indian Ocean has been invaded since 1969 by the Sri Lankan privet, Ligustrum robustum. L. robustum is still spreading in the forests of La Réunion Island, where only 30% of the original vegetation remains, but where 98% of the primary native vegetation of the Mascarene Islands still exists. On Mauritius, where L. robustum was introduced about 1895, it now forms dense, impenetrable thickets, and its presence has been correlated with the inability of native vegetation to re-establish. We assessed the potential impacts of L. robustum invasion on the native ecosystems of La Réunion and identified the factors of invasibility. We determined the degree of invasion in 12 plots of 156 m² and followed native flora and privet recruitment for 3 years in 12 seedling plots of 39 m². The data show that monocultural L. robustum stands now exist in human-disturbed primary forest patches (3.3 individuals/m² and 80% of total individuals) and high seedling densities (0.3–0.6/m²) occur in the least disturbed patches. L. robustum's rapid growth, high shade tolerance and seed production, bird-assisted seed dispersal and high seedling recruitment contribute to its invasiveness in intact forests. The conservation of the original ecosystems of La Réunion depends on the setting up of a long-term and immediate global control strategy. This revised version was published online in July 2006 with corrections to the Cover Date.     

Seasonal and interannual variations of sockeye salmon (<Emphasis Type="Italic">Oncorhynchus nerka</Emphasis>) microsatellite DNA in two Kamchatka lake-river systems

Seasonal and interannual variations in the sockeye salmon populations from two lake-river systems of the East and West Kamchatka were studied. Stability of allele and genotypic frequencies of six microsatellite DNA loci in the adjacent generations and spawning populations of the sockeye salmon of the Bol’shaya River was confirmed experimentally. The pairwise intersample differentiation (F _st) of the local sockeye salmon populations from the southwestern Kamchatka coast (Ozernaya and Bol’shaya Rivers) was almost 7 times higher than the corresponding values for the spawning populations of the Bol’shaya River sockeye salmon of the adjacent years; 15 times, for the adjacent Bol’shaya River sockeye salmon generations; and four times, for the seasonal races within the Kamchatka River.