Colonization of the Southern Patagonia Ocean by Exotic Chinook Salmon

Abstract:  Anadromous salmonids have been particularly successful at establishing wild populations in southern Patagonia, in contrast to their limited success elsewhere outside their native ranges. The most recent such discovery is a spawning population of Chinook salmon in the Santa Cruz River, which flows into the Atlantic Ocean from Argentina. We used mitochondrial DNA analysis to discriminate between alternative potential sources of this population and were able to discard in situ introductions of fish imported directly from California in the early twentieth century. Our results showed that the fish most likely came from Puget Sound, Washington, imported into southern Chile for salmon-ranching experiments in the 1980s. This finding provides concrete evidence of colonization of Atlantic rivers from Pacific locations. The southern Pacific and Atlantic oceans provide a favorable marine environment for the success of invading salmon. In particular, the waters associated with fjords, southern channels, and the inshore portion of the Patagonian shelf provide a rather bounded, continuous waterway for exotic anadromous salmonids, rich in diverse forage species.     

Reconstruction of Pacific salmon abundance from riparian tree-ring growth.

We use relationships between modern Pacific salmon (Oncorhynchus spp.) escapement (migrating adults counted at weirs or dams) and riparian tree-ring growth to reconstruct the abundance of stream-spawning salmon over 150-350 years. After examining nine sites, we produced reconstructions for five mid-order rivers and four salmon species over a large geographic range in the Pacific Northwest: chinook (O. tschwatcha) in the Umpqua River, Oregon, USA; sockeye (O. nerka) in Drinkwater Creek, British Columbia, Canada; pink (O. gorbuscha) in Sashin Creek, southeastern Alaska, USA; chum (O. keta) in Disappearance Creek, southeastern Alaska, USA; and pink and chum in the Kadashan River, southeastern Alaska, USA. We first derived stand-level, non-climatic growth chronologies from riparian trees using standard dendroecology methods and differencing. When the chronologies were compared to 18-55 years of adult salmon escapement we detected positive, significant correlations at five of the nine sites. Regression models relating escapement to tree-ring growth at the five sites were applied to the differenced chronologies to reconstruct salmon abundance. Each reconstruction contains unique patterns characteristic of the site and salmon species. Reconstructions were validated by comparison to local histories (e.g., construction of dams and salmon canneries) and regional fisheries data such as salmon landings and aerial surveys and the Pacific Decadal Oscillation climate index. The reconstructions capture lower-frequency cycles better than extremes and are most useful for determination and comparison of relative abundance, cycles, and the effects of interventions. Reconstructions show lower population cycle maxima in both Umpqua River chinook and Sashin Creek pink salmon in recent decades. The Drinkwater Creek reconstruction suggests that sockeye abundance since the mid-1990s has been 15-25% higher than at any time since 1850, while no long-term deviations from natural cycles are detected for salmon in the Kadashan River or in Disappearance Creek. Decadal-scale cycles in salmon abundance with periods of 25-68 years were detected in all of the reconstructions. This novel approach provides river-specific, long-term perspectives on salmon abundance and cycles. Additionally, it provides a new frame of reference for maintaining and rebuilding individual stocks and for striking a balance between societal demands and the limited, always-changing salmon resource.     

Salmon, wildlife, and wine: marine-derived nutrients in human-dominated ecosystems of central California.

Pacific salmon transfer large quantities of marine-derived nutrients to adjacent forest ecosystems with profound effects on plant and wildlife production. We investigated this process for two highly modified California wine country rivers, one with consistent salmon runs (Mokelumne River) and one without (Calaveras River). Mokelumne River Chinook salmon transported biomass and N comparable to Pacific Northwest salmon streams. Calaveras River levels were much less. Scavenger numbers correlated with salmon carcass counts over time on the Mokelumne River but not the Calaveras River. Likewise, salmon carcasses were consumed significantly faster on the Mokelumne River. Native riparian vegetation as well as cultivated wine grapes adjacent to Mokelumne River spawning sites received 18-25% of foliar N from marine sources, significantly higher than vegetation along the Calaveras River. These data suggest that robust salmon runs continue to provide important ecological services with high economic value, even in impaired watersheds. Loss of Pacific salmon can not only negatively affect stream and riparian ecosystem function, but can also affect local economies where agriculture and salmon streams coexist.     

Population interconnectivity and implications for recovery of a species of concern, the Pacific hake of Georgia Basin

To aid the recovery of a species, understanding the extent to which populations are connected is useful for targeting conservation efforts. Pacific hake within waters of Puget Sound, Washington State, USA, and Georgia Strait, British Columbia, Canada are listed as a species of concern under the U.S. Endangered Species Act due to dramatic declines in the Puget Sound population. To assess the role of dispersal in the recovery of Pacific hake, we sought to quantify patterns of connectivity between populations in Puget Sound and Georgia Strait. Using natural chemical markers from otoliths of fish sampled from these two populations, we linked natal signatures of fish to signatures of individuals from known spawning grounds. Results indicated that 82 % of individuals collected from Puget Sound (n = 78) were estimated to have originated there, while 40 and 92 % of the individuals collected from two cohorts within Georgia Strait (n = 9 and 24, respectively) had originated from Puget Sound. A trend of “population abandonment” of fish from Puget Sound suggests that recovery of this Pacific hake population will depend on local management practices.     

Mechanistic basis of individual mortality in Pacific salmon during spawning migrations

Reproductive-based migration is a challenging period for many animals, but particularly for Pacific salmonids, which must navigate from the high seas to freshwater natal streams. For the first time, we attempt to answer the question as to why some migratory adult Pacific salmon die en route to spawning grounds. Summer-run sockeye salmon (Oncorhynchus nerka) were used as a model, and the migration behavior of 301 fish was followed by intercepting them in the ocean about 215 km from the mouth of the Fraser River, British Columbia, Canada, and implanting a gastric radio transmitter. Before release, telemetered fish were also bio-sampled, which included drawing a blood sample, collecting a gill biopsy, and quantifying energetic status with a microwave energy meter. We tested the predictions that the fish that died prematurely would be characterized by low energy reserves, advanced reproductive development, elevated indicators of stress, and low osmoregulatory preparedness compared with fish that completed their river migration. Just over half (52.3%) of the sockeye tagged were subsequently detected in the Fraser River. Salmon that failed to enter the river had exhibited indicators of stress (e.g., elevated plasma lactate, glucose, and cortisol). Contrary to our prediction, fish that failed to enter the river tended to have higher gross somatic energy and be larger at the time of sampling in the ocean than fish that successfully entered the river. Of the fish that were detected in the river (i.e., 134 fish excluding fishery removals), 9.7% did not migrate beyond the lower reaches (approximately 250 km from ocean), and a further 14.2% reached the upper reaches but failed to reach natal sub-watersheds, whereas the remainder (76.1%) reached natal sub-watersheds. Of these, fish unsuccessful in the lower reaches tended to have a high plasma osmolality in the ocean, whereas fish failing in the upper reaches had lower levels of reproductive hormones in the ocean.     

Hierarchical Bayesian strategy for modeling correlated compositional data with observed zero counts

This article proposes a hierarchical multivariate conditional autoregressive model applied to a compositional response vector. We particularly focus on situations when the composition is discrete occurring when observations are based on small multinomial counts. We address drawbacks that exist in current modeling approaches for such data. Our hierarchical model will be demonstrated with data used to help manage a commercial sockeye salmon fishery in the Fraser River of British Columbia.     

Identifying interactions among salmon populations from observed dynamics

A simple direct correlation analysis of individual counts between different populations often fails to characterize the true nature of population interactions; however, the most common data type available for population studies is count data, and one of the most important objectives in population and community ecology is to identify interactions among populations. Here, I examine the dynamics of the spawning abundance of fall-run chinook salmon spawning within the California Central Valley and the Klamath Basin, California, and the Columbia River Basin, Oregon. I analyzed multiple time series from each watershed using a multivariate time-series technique called maximum autocorrelation factor analysis. This technique was used for finding common underlying trends in escapement abundance within each watershed. These trends were further investigated to identify potential resource-mediated interactions among the three groups of salmon. Each group is affected by multiple trends that are likely to be affected by environmental factors. In addition, some of the trends are coherent with each other, and the differences in population dynamics originate from variations in the relative importance of these trends among the three watershed groups.     

Does juvenile experience affect migration and spawning of adult Atlantic salmon?

Summary This paper documents differences in seasonal time of river ascent and descent, and instream behavior of adult wild and sea-ranched Atlantic salmon (Salmo salar) of the Norwegian River Imsa stock during the period 1981–1989. Wild fish use River Imsa as a nursery, and at an age of 2 years most of them migrate to the sea as smolts. The sea ranched fish are hatchery reared offspring of the River Imsa stock and are released as smolts at the mouth of the river. They are thus deprived of juvenile river life and a downstream smolt migration. Wild and sea ranched salmon feed for 1 or more years in the Norwegian Sea before homing as spawners. Both groups returned simultaneously to coastal Norway, but sea ranched fish ascended the river later and descended sooner after spawning than wild fish. All wild females and almost all wild males (96.2%) spawned in the river, whereas 13.5% and 36.7%, respectively, of the mature sea-ranched females and males left the river unspawned. The annual number, but not the proportion, of unspawned fish increased with increasing density of adult salmon in the river. Unspawned females were medium sized and small (45–70 cm); unspawned males were medium sized and large (50–90 cm). Independent of the density of spawners in the river, sea ranched fish moved up- and downstream the river more often than wild fish. More than 20% of the sea-ranched salmon and less than 1% of the wild salmon passed a trap 100 m above the river outlet more than once in each direction during the same spawning reason. Moreover, sea-ranched salmon were about twice as often seriously injured during spawning as wild fish. Lack of juvenile experience from the river may be the main reason for the behavioral differences between sea-ranched and wild fish. Offprint requests to: B. Jonsson     

Population genetic structure of escolar (<Emphasis Type="Italic">Lepidocybium flavobrunneum</Emphasis>)

Escolar (Lepidocybium flavobrunneum) is a large, mesopelagic fish that inhabits tropical and temperate seas throughout the world, and is a common bycatch in pelagic longline fisheries that target tuna and swordfish. Few studies have explored the biology and natural history of escolar, and little is known regarding its population structure. To evaluate the genetic basis of population structure of escolar throughout their range, we surveyed genetic variation over an 806 base pair fragment of the mitochondrial control region. In total, 225 individuals from six geographically distant locations throughout the Atlantic (Gulf of Mexico, Brazil, South Africa) and Pacific (Ecuador, Hawaii, Australia) were analyzed. A neighbor-joining tree of haplotypes based on maximum likelihood distances revealed two highly divergent clades (δ = 4.85%) that were predominantly restricted to the Atlantic and Indo-Pacific ocean basins. All Atlantic clade individuals occurred in the Atlantic Ocean and all but four Pacific clade individuals were found in the Pacific Ocean. The four Atlantic escolar with Pacific clade haplotypes were found in the South Africa collection. The nuclear ITS-1 gene region of these four individuals was subsequently analyzed and compared to the ITS-1 gene region of four individuals from the South Africa collection with Atlantic clade haplotypes as well as four representative individuals each from the Atlantic and Pacific collections. The four South Africa escolar with Pacific mitochondrial control region haplotypes all had ITS-1 gene region sequences that clustered with the Pacific escolar, suggesting that they were recent migrants from the Indo-Pacific. Due to the high divergence and geographic separation of the Atlantic and Pacific clades, as well as reported morphological differences between Atlantic and Indo-Pacific specimens, consideration of the Atlantic and Indo-Pacific populations as separate species or subspecies may be warranted, though further study is necessary.     

Public Willingness to Pay for Recovering and Downlisting Threatened and Endangered Marine Species

Abstract: Nonmarket valuation research has produced economic value estimates for a variety of threatened, endangered, and rare species around the world. Although over 40 value estimates exist, it is often difficult to compare values from different studies due to variations in study design, implementation, and modeling specifications. We conducted a stated‐preference choice experiment to estimate the value of recovering or downlisting 8 threatened and endangered marine species in the United States: loggerhead sea turtle (Caretta caretta), leatherback sea turtle (Dermochelys coriacea), North Atlantic right whale (Eubalaena glacialis), North Pacific right whale (Eubalaena japonica), upper Willamette River Chinook salmon (Oncorhynchus tshawytscha), Puget Sound Chinook salmon (Oncorhynchus tshawytscha), Hawaiian monk seals (Monachus schauinslandi), and smalltooth sawfish (Pristis pectinata). In May 2009, we surveyed a random sample of U.S. households. We collected data from 8476 households and estimated willingness to pay for recovering and downlisting the 8 species from these data. Respondents were willing to pay for recovering and downlisting threatened and endangered marine taxa. Willingness‐to‐pay values ranged from $40/household for recovering Puget Sound Chinook salmon to $73/household for recovering the North Pacific right whale. Statistical comparisons among willingness‐to‐pay values suggest that some taxa are more economically valuable than others, which suggests that the U.S. public's willingness to pay for recovery may vary by species.     

Bayesian Decision Analysis for Evaluating Management Options to Promote Recovery of a Depleted Salmon Population

Abstract:  The endangered population of sockeye salmon (Oncorhynchus nerka) in Cultus Lake, British Columbia, Canada, migrates through commercial fishing areas along with other, much more abundant sockeye salmon populations, but it is not feasible to selectively harvest only the latter, abundant populations. This situation creates controversial trade-offs between recovery actions and economic revenue. We conducted a Bayesian decision analysis to evaluate options for recovery of Cultus Lake sockeye salmon. We used a stochastic population model that included 2 sources of uncertainty that are often omitted from such analyses: structural uncertainty in the magnitude of a potential Allee effect and implementation uncertainty (the deviation between targets and actual outcomes of management actions). Numerous state-dependent, time-independent management actions meet recovery objectives. These actions prescribe limitations on commercial harvest rates as a function of abundance of Cultus Lake sockeye salmon. We also quantified how much reduction in economic value of commercial harvests of the more abundant sockeye salmon populations would be expected for a given increase in the probability of recovery of the Cultus population. Such results illustrate how Bayesian decision analysis can rank options for dealing with conservation risks and can help inform trade-off discussions among decision makers and among groups that have competing objectives.     

Conservation Status as a Biodiversity Trend Indicator: Recommendations from a Decade of Listing Species at Risk in British Columbia

Abstract:  Species conservation status is commonly used as a broad-scale indicator of the state of biological diversity. To learn about its value for tracking trends, we examined provincial lists of terrestrial vertebrate species and subspecies at risk in British Columbia, Canada, for 1992 and 2002 to see whether changes in these lists reflected changes in the status of the taxa they represent. Examination of the case histories of individual species and subspecies showed that 65% of additions and deletions to the British Columbia Red List were the result of improvement in knowledge of species status, changes in assessment procedures, and refinements in taxonomy rather than actual changes in a species' status. Comparison to an alternate set of rank scores provided by NatureServe for taxa that appeared on both 1992 and 2002 British Columbia Red Lists revealed changes in status that were not reflected by movement from the list. Estimates of historical conservation status for species on the 1992 British Columbia Red List demonstrated ambiguity around the natural baseline with regard to tracking changes in list composition over time. We discourage the continued use of indicators based solely on conservation status as a means of tracking biodiversity. Instead we recommend advancing strategic indicators around species at risk based on long-term monitoring data, deliberate and explicitly stated baselines, and consistent methods of conservation ranking.     

DNA markers indicate that distinct spawning cohorts and aggregations of Patagonian squid,<Emphasis Type="Italic"> Loligo gahi</Emphasis>, do not represent genetically discrete subpopulations

The Patagonian squid, Loligo gahi DOrbigny, has been described as having temporally and spatially identifiable spawning aggregations. Variation at six microsatellite loci was used to assess whether seasonal and geographical spawning groups around the Falkland Islands represent distinct sub-populations. Genetic variation at these loci is high in this species (mean expected heterozygosity=0.87; mean number of alleles=14.7). No evidence of significantly different allele frequencies was found, either between samples from putative spawning cohorts or geographical areas, indicating that L.gahi around the Falkland Islands comprise a single genetically homogeneous population. Age structure analysis of samples (from statolith growth increments) indicated substantial spread in hatching dates among individuals of similar size and maturity status, suggesting the potential for extensive interbreeding between seasonal cohorts. A sample of L.gahi from the SE Pacific (Peru) displayed distinctly different gene frequencies (and allele size distribution at one locus) from SW Atlantic samples, supporting the suggestion that SE Pacific and SW Atlantic populations may represent distinct subspecies.Contributed by J.P. Thorpe, Port Erin     

Correlations between forest fires in British Columbia, Canada, and sea surface temperature of the Pacific Ocean

Correlations and cross-correlations between forest fires in the province of British Columbia, Canada, and sea surface temperatures in the Pacific Ocean were evaluated. British Columbia has a long Pacific Ocean coastline; given that there may be teleconnections between the province's forest fires and climate variability over the ocean, significant correlations may exist between forest fires and the sea surface temperature of the Pacific Ocean. Fire occurrences and areas burned through lightning-caused and human-caused fires were analyzed against individual 1° × 1° grid cells of anomalies in the sea surface temperature to determine correlations for the period 1950-2006. Significant correlations (p < 0.05) for vast areas of the ocean were found between occurrences of lightning-caused fires and sea surface temperature anomalies for time lags of 1 and 2 years, whereas significant correlations between occurrences of human-caused fires and sea surface temperature anomalies occurred extensively for many time lags. To support the results of this approach, correlations between fire data and the Niño 3.4, Pacific Decadal Oscillation, and Arctic Oscillation indices were tested for the same period. Significant correlations were found between fire occurrences and these indices at certain time lags. Overall, fire occurrence appeared to be more extensively correlated with sea surface temperature anomalies than was area burned. These results support the hypothesis that teleconnections exist between fire activity in British Columbia and sea surface temperatures in the Pacific Ocean, and the correlations suggest that linear regression models or other regression techniques may be appropriate for predicting fire severity from the sea surface temperatures of one or more previous years.     

Does interspecific competition influence relationships between heterozygosity and fitness-related behaviors in juvenile Atlantic salmon (<Emphasis Type="Italic">Salmo salar</Emphasis>)?

Very few studies have investigated the effect of genetic diversity on the behavioral and phenotypic traits linked to the competitive ability of individuals. In this study, we reared juvenile Atlantic salmon (Salmo salar) alone or with the competitive rainbow trout (Oncorhynchus mykiss) in order to: (1) to assess correlations between heterozygosity and traits related to individual competitive ability [i.e., heterozygosity–fitness correlations (HFCs)] in Atlantic salmon, and (2) to evaluate the effect of the competitive rainbow trout on any such HFCs. We also tested whether a few loci had a disproportionately large effect (i.e., the local effect hypothesis) or, on the contrary, if all loci contributed equally (i.e., the global effect hypothesis) in explaining the observed HFCs. We found significant HFCs for phenotypic traits related to the competitive ability of juvenile Atlantic salmon, i.e., the growth rate and the distance to the feeding source. Some HFCs were nonlinear, suggesting that individuals with intermediate levels of heterozygosity were favored. In addition, we found that the competition exerted by rainbow trout only weakly modified these HFCs as the relationships were highly consistent across treatments. We demonstrated that the local-effect hypothesis best explained both linear and nonlinear HFCs. Overall, our results illustrated the importance of genetic diversity in explaining the behavioral variability observed within populations. Moreover, we provide evidence that, even if a competitive species can have strong ecological effects, the relationships between genetic diversity and fitness-related traits in juvenile Atlantic salmon were not influenced by such effects.     

Marine-derived nutrients, bioturbation, and ecosystem metabolism: reconsidering the role of salmon in streams

In coastal areas of the North Pacific Ocean, annual returns of spawning salmon provide a substantial influx of nutrients and organic matter to streams and are generally believed to enhance the productivity of recipient ecosystems. Loss of this subsidy from areas with diminished salmon runs has been hypothesized to limit ecosystem productivity in juvenile salmon rearing habitats (lakes and streams), thereby reinforcing population declines. Using five to seven years of data from an Alaskan stream supporting moderate salmon densities, we show that salmon predictably increased stream water nutrient concentrations, which were on average 190% (nitrogen) and 390% (phosphorus) pre-salmon values, and that primary producers incorporated some of these nutrients into tissues. However, benthic algal biomass declined by an order of magnitude despite increased nutrients. We also measured changes in stream ecosystem metabolic properties, including gross primary productivity (GPP) and ecosystem respiration (ER), from three salmon streams by analyzing diel measurements of oxygen concentrations and stable isotopic ratios (delta O-O2) within a Bayesian statistical model of oxygen dynamics. Our results do not support a shift toward higher primary productivity with the return of salmon, as is expected from a nutrient fertilization mechanism. Rather, net ecosystem metabolism switched from approximately net autotrophic (GPP > or = ER) to a strongly net heterotrophic state (GPP < ER) in response to bioturbation of benthic habitats by salmon. Following the seasonal arrival of salmon, GPP declined to <12% of pre-salmon rates, while ER increased by over threefold. Metabolism by live salmon could not account for the observed increase in ER early in the salmon run, suggesting salmon nutrients and disturbance enhanced in situ heterotrophic respiration. Salmon also changed the physical properties of the stream, increasing air-water gas exchange by nearly 10-fold during peak spawning. We suggest that management efforts to restore salmon ecosystems should consider effects on ecosystem metabolic properties and how salmon disturbance affects the incorporation of marine-derived nutrients into food webs.     

Bottom-up and climatic forcing on the worldwide population of leatherback turtles

Nesting populations of leatherback turtles (Dermochelys coriacea) in the Atlantic and western Indian Oceans are increasing or stable while those in the Pacific are declining. It has been suggested that leatherbacks in the eastern Pacific may be resource limited due to environmental variability derived from the El Ni?o Southern Oscillation (ENSO), but this has yet to be tested. Here we explored bottom-up forcing and the responding reproductive output of nesting leatherbacks worldwide. We achieved this through an extensive review of leatherback nesting and migration data and by analyzing the spatial, temporal, and quantitative nature of resources as indicated by net primary production at post-nesting female migration and foraging areas. Leatherbacks in the eastern Pacific were the smallest in body size and had the lowest reproductive output due to less productive and inconsistent resources within their migration and foraging areas. This derived from natural interannual and multidecadal climate variability together with an influence of anthropogenic climate warming that is possibly affecting these natural cycles. The reproductive output of leatherbacks in the Atlantic and western Indian Oceans was nearly twice that of turtles in the eastern Pacific. The inconsistent nature of the Pacific Ocean may also render western Pacific leatherbacks susceptible to a more variable reproductive output; however, it appears that egg harvesting on nesting beaches is their major threat. We suggest that the eastern Pacific leatherback population is more sensitive to anthropogenic mortality due to recruitment rates that are lower and more variable, thus accounting for much of the population differences compared to Atlantic and western Indian turtles.     

Ten years of varying lake level and selection on size-at-maturity in sockeye salmon

Despite the ubiquity of studies quantifying the strength and form of selection in nature, rarely is the ecological context for contemporary selection understood. Here we report a case where lake level is a selective factor acting on sockeye salmon body size-at-maturity because low lake levels cause large salmon to strand and die rather than reach the breeding grounds. As a result of a semelparous life history, death for salmon at this stage results in a lifetime fitness of zero. We combined information on the level of Lake Aleknagik (southwestern Alaska, USA) from 1952 through 2006 with a detailed comparison of the body size of mature salmon that died at the mouth of Hansen Creek vs. individuals that successfully ascended to the spawning grounds over 10 breeding seasons (1997-2006). The percentage of salmon stranding at the mouth varied among years: 2-42% in males and < 1-26% in females. Formal selection analyses indicated that the largest individuals were most susceptible to stranding mortality, especially in years when many salmon stranded, and these were years with low lake levels. Taken together, these results suggest that lake level was a strong and consistent selective force acting on this salmon population, acting synergistically with size-selective predation by bears. Salmon breeding in Hansen Creek tend to be smaller, younger, and more streamlined than conspecifics from neighboring populations, suggesting that selection against large individuals could be driving these patterns.     

Range-Wide Selection of Catchments for Pacific Salmon Conservation

Abstract:  Freshwater ecosystems are declining in quality globally, but a lack of data inhibits identification of areas valuable for conservation across national borders. We developed a biological measure of conservation value for six species of Pacific salmon ( Oncorhynchus spp.) in catchments of the northern Pacific across Canada, China, Japan, Russia, and the United States. We based the measure on abundance and life-history richness and a model-based method that filled data gaps. Catchments with high conservation value ranged from California to northern Russia and included catchments in regions that are strongly affected by human development (e.g., Puget Sound). Catchments with high conservation value were less affected by agriculture and dams than other catchments, although only 1% were within biodiversity reserves. Our set of high-value areas was largely insensitive to simulated error, although classification remained uncertain for 3% of catchments. Although salmon face many threats, we propose they will be most likely to exhibit resilience into the future if a complementary mosaic of conservation strategies can be proactively adopted in catchments with healthy salmon populations. Our analysis provides an initial map of where these catchments are likely to be located.     

Divergence between populations of a monogamous polychaete with male parental care: Premating isolation and chromosome variation

Low dispersal and sexual selection are characteristic of the coastal polychaeteNereis acuminata Ehlers 1868 [also known asNereis arenaceodentata Moore 1903 andNereis (Neanthes) caudata Delle Chiaje 1841]. We assessed levels of premating isolation between populations of this polychaete. Four North American populations were used, two from the Atlantic and two from the Pacific. Worms from all sites (1) were collected in 1987 and 1988 from the same habitat type, (2) were morphologically similar and keyed out asN. acuminata, and (3) reproduced monogamously and exhibited male parental care, an extremely rare reproductive mode in marine invertebrates. There was no evidence from 10-min or 36-h trials of premating isolation between the two Pacific populations. Incomplete premating isolation was found between the two Atlantic populations. High aggression and non-pairing occurred in some 10-min trials between males and females. However, in 36-h trials males and females from the two Atlantic populations always paired to mate. Complete premating isolation was found between Atlantic and Pacific populations. During 10-min trials, males and females from different oceans often attacked and then avoided each other, and they never paired to mate. Nor did they pair to mate in longer, 36-h trials. One Pacific and one Atlantic population were compared for tolerance to cold temperature. Pacific individuals were less able to tolerate cold water than Atlantic individuals. Two Atlantic populations studied had karyotypes with 11 pairs of small acrocentric chromosomes (2n=22), while the two Pacific populations had nine pairs of large metacentric or submetacentric chromosomes (2n=18). Such extreme dissimilarity in karyotype was not expected considering the similarity in morphology, habitat, and reproductive mode. Results suggest strongly that the Atlantic and Pacific populations have been allopatric for a long time, and are different species.