Early ocean distribution of juvenile Chinook salmon in an upwelling ecosystem

Extreme variability in abundance of California salmon populations is often ascribed to ocean conditions, yet relatively little is known about their marine life history. To investigate which ocean conditions influence their distribution and abundance, we surveyed juvenile Chinook salmon (Oncorhynchus tshawytscha) within the California Current (central California [37°30′N) to Newport, Oregon (44°00′N]) for a 2‐week period over three summers (2010–2012). At each station, we measured chlorophyll‐a as an indicator of primary productivity, acoustic‐based metrics of zooplankton density as an indicator of potential prey availability and physical characteristics such as bottom depth, temperature and salinity. We also measured fork lengths and collected genetic samples from each salmon that was caught. Genetic stock identification revealed that the majority of juvenile salmon were from the Central Valley and the Klamath Basin (91–98%). We constructed generalized logistic‐linear negative binomial hurdle models and chose the best model(s) using Akaike's Information Criterion (AIC) to determine which covariates influenced the salmon presence and, at locations where salmon were present, determined the variables that influenced their abundance. The probability of salmon presence was highest in shallower waters with a high chlorophyll‐a concentration and close to an individual's natal river. Catch abundance was primarily influenced by year, mean fork length and proximity to natal rivers. At the scale of sampling stations, presence and abundance were not related to acoustic indices of zooplankton density. In the weeks to months after ocean entry, California's juvenile Chinook salmon population appears to be primarily constrained to coastal waters near natal river outlets.     

Salish Sea Chinook salmon exhibit weaker coherence in early marine survival trends than coastal populations

Identifying factors that influence anadromous Pacific salmon (Oncorhynchus spp.) population dynamics is complicated by their diverse life histories and large geographic range. Over the last several decades, Chinook salmon (O. tshawytscha) populations from coastal areas and the Salish Sea have exhibited substantial variability in abundance. In some cases, populations within the Salish Sea have experienced persistent declines that have not rebounded. We analyzed a time series of early marine survival from 36 hatchery Chinook salmon populations spanning ocean entry years 1980–2008 to quantify spatial and temporal coherence in survival. Overall, we observed higher inter‐population variability in survival for Salish Sea populations than non‐Salish Sea populations. Annual survival patterns of Salish Sea populations covaried over smaller spatial scales and exhibited less synchrony among proximate populations relative to non‐Salish Sea populations. These results were supported by multivariate autoregressive state space (MARSS) models which predominantly identified region‐scale differences in survival trends between northern coastal, southern coastal, Strait of Georgia, and Puget Sound population groupings. Furthermore, Dynamic Factor Analysis (DFA) of regional survival trends showed that survival of southern coastal populations was associated with the North Pacific Gyre Oscillation, a large‐scale ocean circulation pattern, whereas survival of Salish Sea populations was not. In summary, this study demonstrates that survival patterns in Chinook salmon are likely determined by a complex hierarchy of processes operating across a broad range in spatial and temporal scales, presenting challenges to the management of mixed‐stock fisheries.     

Energy dynamics of subyearling Chinook salmon reveal the importance of piscivory to short‐term growth during early marine residence

Variation in prey quantity and quality can influence growth and survival of marine predators, including anadromous fish that migrate from freshwater systems. The objective of this study was to examine the energy dynamics of subyearling Chinook salmon (Oncorhynchus tshawytscha) following freshwater emigration. To address this objective, a population of Chinook salmon and their marine prey were repeatedly sampled from June to September over 2 years in coastal waters off Oregon and Washington. Subyearlings from the same population were also reared under laboratory conditions. Using a bioenergetics model evaluated in the laboratory, we found that growth rate variability in the field was associated more with differences in northern anchovy (Engraulis mordax) consumption and less with variation in diet energy density or ocean temperature. Highest growth rates (2.43–3.22% body weight/day) occurred in months when anchovy biomass peaked, and the timing of peak anchovy biomass varied by year. Our results support a general pattern among subyearling Chinook salmon occurring from Alaska to California that feeding rates contribute most to growth rate variability during early marine residence, although dominant prey types can differ seasonally, annually, or by ecosystem. In the northern California Current, faster growth appears to be associated with the availability of age‐0 anchovy. Identifying factors that influence the seasonal development of the prey field and regulate prey quantity and quality will improve understanding of salmon growth and survival during early marine residence.     

Effects of juvenile size at release and early marine growth on adult return rates for Hokkaido chum salmon (Oncorhynchus keta) in relation to sea surface temperature

Using path analyses, we investigated relationships between size at release from hatcheries, the early marine growth of juveniles, and adult return rates for chum salmon from five river stocks of Hokkaido, Japan, in relation to sea surface temperature during ocean residence. Marine growth was estimated using scales collected from 11 760 adults of age 0.3 (1980–2004). The growth and survival of each stock appeared to have a different suite of regulatory processes. Interannual variability in return rates was mainly regulated by size at release in two stocks from the Sea of Okhotsk. A similar relationship was found in one stock from the Sea of Japan, but growth during coastal residency also affected their return rates. In two stocks from the Pacific coast of Hokkaido, variability in return rates was not related to size at release or to the coastal growth of juveniles, but with offshore growth in the Sea of Okhotsk, the nursery area for juveniles after leaving Japanese coastal waters. Whereas coastal growth tended to be negatively correlated with size at release in some stocks, offshore growth was positively associated with the August–November sea surface temperature in all stocks. This study confirmed that mortality of juvenile salmon occurred in two phases, during the coastal residency and the late period of the growing season, but the relative importance of both phases varied by stock and region, which probably regulated year‐class strength of Hokkaido chum salmon.     

Infections by Renibacterium salmoninarum and Nanophyetus salmincola Chapin are associated with reduced growth of juvenile Chinook salmon,Oncorhynchus tshawytscha (Walbaum), in the Northeast Pacific Ocean

We examined 1454 juvenile Chinook salmon, Oncorhynchus tshawytscha (Walbaum), captured in nearshore waters off the coasts of Washington and Oregon (USA) from 1999 to 2004 for infection by Renibacterium salmoninarum, Nanophyetus salmincola Chapin and skin metacercariae. The prevalence and intensities for each of these infections were established for both yearling and subyearling Chinook salmon. Two metrics of salmon growth, weight residuals and plasma levels of insulin-like growth factor-1, were determined for salmon infected with these pathogens/parasites, both individually and in combination, with uninfected fish used for comparison. Yearling Chinook salmon infected with R. salmoninarum had significantly reduced weight residuals. Chinook salmon infected with skin metacercariae alone did not have significantly reduced growth metrics. Dual infections were not associated with significantly more severe effects on the growth metrics than single infections; the number of triple infections was very low and precluded statistical comparison. Overall, these data suggest that infections by these organisms can be associated with reduced juvenile Chinook salmon growth. Because growth in the first year at sea has been linked to survival for some stocks of Chinook salmon, the infections may therefore play a role in regulating these populations in the Northeast Pacific Ocean.     

Year-to-year variability in ocean recovery rate of Columbia River Upriver Bright fall Chinook salmon (Oncorhynchus tshawytscha)

Unusually large returns of several stocks of fall Chinook salmon (Oncorhynchus tshawytscha) from the U.S. Northwest commonly occurred during the late 1980s. These synchronous events seem to have been due to ocean rather than freshwater conditions because natal rivers of these stocks were geographically disconnected. We examined year‐to‐year variability in cohort strength of one of these stocks, Upriver Bright (URB) fall Chinook salmon from the Columbia River Hanford Reach for brood years 1976–99 (recovery years 1979–2002). We used the ocean recovery rate of coded‐wire‐tag (CWT) fish as an index of cohort strength. To analyse year‐to‐year variability in the ocean recovery rate, we applied a log‐linear model whose candidate explanatory variables were ocean condition variables, fishing effort, age of recovered fish, and fish rearing type (hatchery versus wild). Explanatory variables in the best model included fishing effort, and the quadratic term of winter sea surface temperature (SST) measured from coastal waters of British Columbia, Canada during the fish's first ocean year. The coefficient of the quadratic term of SST was significantly negative, so the model shape was convex. Our findings can be used to infer year‐to‐year variability in cohort strength of other fall Chinook salmon whose life history and ocean distributions are similar to the URB fish.     

Relationships between oceanic conditions and growth of Chinook salmon (Oncorhynchus tshawytscha) from California, Washington, and Alaska, USA

We model age‐specific growth rates of Chinook salmon (Oncorhynchus tshawytscha) with two life‐history behaviors from Alaska (i. Situk and ii.Taku Rivers), Puget Sound, Washington (iii., iv. Skagit River), and California (v. Smith River) relative to oceanic conditions in those regions. By analyzing over 20 yr of biological and physical data from the NE Pacific downwelling, upwelling, and transition zones, we are able to determine the factors affecting growth across much of the species’ range and between life‐history behaviors. With scale increment data from returning fish, we use path analysis and partial least squares regression to quantify the relationships between growth and regional‐ and large‐scale oceanic conditions (e.g., sea level height, sea surface temperature, upwelling). Alaskan fish with both ocean‐ and stream‐type behaviors were fit best by the environmental data from the winter in Alaska waters. Specifically, coastal and gyre factors such as sea surface temperature, river flow, and Ekman pumping positively correlated to growth, indicating a productive and strong Alaska Current promoted growth. Growth of fish from California was fit by local factors such as increased upwelling, lower coastal sea surface temperature, and wind stresses during summer and spring, indicating a productive and strong California Current promoted growth. For Puget Sound, Washington, growth of fish that migrate to sea in their first year was generally negatively correlated to a strong California Current. Puget Sound fish that spend a year in freshwater before migrating to sea were modeled well with environmental data from their source region for the first 2 yr at sea and by data from Alaska waters in their third year at sea. Results suggest that conditions in which the transition zone is dominated by neither the Alaska nor California Currents are best for increased growth of Puget Sound fish.     

Ocean distribution and habitat associations of yearling coho (Oncorhynchus kisutch) and Chinook (O. tshawytscha) salmon in the northern California Current

Yearling juvenile coho and Chinook salmon were sampled on 28 cruises in June and September 1981–85 and 1998–07 in continental shelf and oceanic waters off the Pacific Northwest. Oceanographic variables measured included temperature, salinity, water depth, and chlorophyll concentration (all cruises) and copepod biomass during the cruises from 1998–07. Juvenile salmonids were found almost exclusively in continental shelf waters, and showed a patchy distribution: half were collected in ～5% of the collections and none were collected in ～40% of the collections. Variance‐to‐mean ratios of the catches were high, also indicating patchy spatial distributions for both species. The salmon were most abundant in the vicinity of the Columbia River and the Washington coast in June; by September, both were less abundant, although still found mainly off Washington. In June, the geographic center‐of‐mass of the distribution for each species was located off Grays Harbor, WA, near the northern end of our sampling grid, but in September, it shifted southward and inshore. Coho salmon ranged further offshore than Chinook salmon: in June, the average median depth where they were caught was 85.6 and 55.0 m, respectively, and in September it was 65.5 and 43.7 m, respectively. Abundances of both species were significantly correlated with water depth (negatively), chlorophyll (positively) and copepod biomass (positively). Abundances of yearling Chinook salmon, but not of yearling coho salmon, were correlated with temperature (negatively). We discuss the potential role of coastal upwelling, submarine canyons and krill in determining the spatial distributions of the salmon.     

Spatial correlation patterns in coastal environmental variables and survival rates of salmon in the north-east Pacific Ocean

We examined spatial correlations for three coastal variables [upwelling index, sea surface temperature (SST), and sea surface salinity (SSS)] that might affect juvenile salmon ( Oncorhynchus spp.) during their early marine life. Observed correlation patterns in environmental variables were compared with those in survival rates of pink ( O. gorbuscha ), chum ( O. keta ), and sockeye ( O. nerka ) salmon stocks to help identify appropriate variables to include in models of salmon productivity. Both the upwelling index and coastal SST were characterized by strong positive correlations at short distances, which declined slowly with distance in the winter months, but much more rapidly in the summer. The SSS had much weaker and more variable correlations at all distances throughout the year. The distance at which stations were no longer correlated (spatial decorrelation scale) was largest for the upwelling index (> 1000 km), intermediate for SST (400–800 km in summer), and shortest for SSS (< 400 km). Survival rate indices of salmon showed moderate positive correlations among adjacent stocks that decreased to zero at larger distances. Spatial decorrelation scales ranged from approximately 500 km for sockeye salmon to approximately 1000 km for chum salmon. We conclude that variability in the coastal marine environment during summer, as well as variability in salmon survival rates, are dominated by regional scale variability of several hundred to 1000 km. The correlation scale for SST in the summer most closely matched the observed correlation scales for survival rates of salmon, suggesting that regional-scale variations in coastal SST can help explain the observed regional-scale covariation in survival rates among salmon stocks.     

Can research on the early marine life stage of juvenile chum salmon <Emphasis Type="Italic">Oncorhynchus keta</Emphasis> forecast returns of adult salmon? A case study from eastern Hokkaido,Japan

To examine the efficacy of juvenile salmon research as a tool for forecasting adult returns, the results from a study on the early marine life stage of juvenile chum salmon, conducted in the Nemuro Strait during 1999–2002 (i.e., 1998–2001 brood years), were compared with the return rates of adult salmon. Among the four brood years, the 2000 brood year (i.e., salmon migrating to the sea in 2001) was previously reported as showing higher abundance, higher growth rate and better somatic condition during the coastal residency period. Consequently, we expected it to have the highest return rate, under a hypothesis that juvenile survival in coastal residency regulates brood-year strength. Contrary to this expectation, the 2000 brood year had almost the lowest return rate. Alternatively, a statistical model in which sea surface temperature during the first year of marine life and size at release were utilized as explanatory variables reconstructed the actual variability in return rates more accurately than that based on the early marine life stage. Possible reasons for the discrepancy between the results of the juvenile salmon research and adult returns are discussed, and we suggest improvements for future research on juvenile salmon.     

Interannual variability in the feeding and condition of subyearling Chinook salmon off Oregon and Washington in relation to fluctuating ocean conditions

Chinook salmon (Oncorhynchus tshawytscha) is one of several economically‐important species of salmon found in the Northeast Pacific Ocean. The first months at sea are believed to be the most critical for salmon survival, with the highest rate of mortality occurring during this period. In the present study, we examined interannual diet composition and body condition trends for late‐summer subyearling Chinook salmon caught off Oregon and Washington from 1998 to 2012. Interannual variability was observed in juvenile salmon diet composition by weight of prey consumed. Juvenile subyearling Chinook salmon were mainly piscivorous, with northern anchovy (Engraulis mordax) being especially important, making up half the diet by weight in some years. Annual diets clustered into two groups, primarily defined by their proportion of invertebrate prey (14% versus 39% on average). Diet composition was found to influence adult returns, with salmon from high‐invertebrate years returning in significantly larger numbers 2–3 yrs later. However, years that had high adult returns had overall lower stomach fullness and poorer body condition as juveniles, a counterintuitive result potentially driven by the enhanced survival of less fit individuals in better ocean conditions (top‐down effect). Ocean conditions in years with a higher percentage of invertebrates in salmon diets were significantly cooler from May to August, and bottom‐up processes may have led to a fall plankton community with a larger proportion of invertebrates. Our results suggest that the plankton community assemblage during this first fall may be critical in predicting adult returns of Chinook salmon in the Pacific Northwest.     

Food habits and marine survival of juvenile Chinook and coho salmon from marine waters of Southeast Alaska

Little is known about the food habits of juvenile Chinook (Oncorhynchus tshawytscha) and coho (Oncorhynchus kisutch) salmon in marine environments of Alaska, or whether their diets may have contributed to extremely high marine survival rates for coho salmon from Southeast Alaska and much more modest survival rates for Southeast Alaskan Chinook salmon. To address these issues, we documented the spatial and temporal variability of diets of both species collected from marine waters of Southeast Alaska during summers of 1997–2000. Food habits were similar: major prey items of both species included fishes, crab larvae, hyperiid amphipods, insects, and euphausiids. Multivariate analyses of diet composition indicated that the most distinct groups were formed at the smallest spatial and temporal scales (the haul), although groups also formed at larger scales, such as by month or habitat type. Our expectations for how food habits would influence survival were only partially supported. As predicted, Southeast Alaskan coho salmon had more prey in their stomachs overall [1.8% of body weight (BW)] and proportionally far fewer empty stomachs (0.7%) than either Alaskan Chinook (1.4% BW, 5.1% empty) or coho salmon from other regions. However, contrary to our expectations, coho salmon diets contained surprisingly few fish (49% by weight). Apparently, Alaskan coho salmon achieved extremely high marine survival rates despite a diet consisting largely of small, less energetically‐efficient crustacean prey. Our results suggest that diet quantity (how much is eaten) rather than diet quality (what is eaten) is important to marine survival.     

Variability in the performance of juvenile Chinook salmon is explained primarily by when and where they resided in estuarine habitats

Estuarine habitats provide rearing opportunities for the juvenile life stage of anadromous fishes. Because survival is positively correlated with juvenile performance, these estuarine habitats play an important role in population abundance and productivity. To provide information for the recovery of several depressed stocks of Chinook salmon in the Columbia River Basin, we sought to identify the factors that explain variability in performance. Using otolith‐derived estimates of juvenile somatic growth rate as an index of recent performance, we observed a negative nonlinear relationship between growth rate and day of year, and a decreasing and increasing trend of growth rate over the 8 years of this study and distance from the river mouth respectively. Using a generalised linear modelling approach, we found that variability in juvenile somatic growth rate was best explained by where and when individuals were collected, their body size, contaminant loads, stock of origin, and whether a fish was hatchery produced or unmarked. Lastly, we argue that a considerable improvement to the growth rate of juveniles in estuarine habitats is physiologically possible. The results of this 8‐year study provide a baseline of the performance of juvenile Chinook salmon to evaluate habitat restoration programs and to compare against future anthropogenic conditions.     

Juvenile coho salmon movement,growth and survival in a coastal basin of southern Oregon

Juvenile salmonids display highly variable spatial and temporal patterns of early dispersal that are influenced by density‐dependent and density‐independent factors. Although juvenile coho salmon (Oncorhynchus kisutch) movement patterns in streams and their relationship with body mass and growth have been examined in previous studies, most observations were limited to one season or one stream section. In this study, we monitored the movement of juvenile coho salmon throughout their period of residence in a coastal basin to identify prevalent dispersal strategies and their relationships with body mass, growth rates and survival. Our results revealed seasonally and spatially variable movement patterns. Juvenile coho salmon that dispersed to tidally affected reaches soon after emergence remained more mobile and expressed lower site fidelity than those individuals that remained in upper riverine reaches. We did not detect significantly different growth rates between sedentary and mobile individuals. Although a greater proportion of sedentary than mobile fish survived winter to emigrate from the creek in the spring, reach of residence at the onset of winter influenced these survival estimates. Hence, apparent summer‐to‐smolt survival for mobile individuals was greater than for sedentary fish in tidally influenced reaches, whereas in riverine reaches the sedentary strategy seemed to be favoured. Our research identified complex movement patterns that reflect phenotypic and life history variation, and underscores the importance of maintaining diverse freshwater and estuarine habitats that support juvenile coho salmon before marine migration.     

Covariation between the average lengths of mature coho (Oncorhynchus kisutch) and Chinook salmon (O. tshawytscha) and the ocean environment

We used the average fork length of age‐3 returning coho (Oncorhynchus kisutch) and age‐3 ocean‐type and age‐4 stream‐type Chinook (Oncorhynchus tshawytscha) salmon along the northeast Pacific coast to assess the covariability between established oceanic environmental indices and growth. These indices included the Multivariate El Niño‐Southern Oscillation Index (MEI), Pacific Decadal Oscillation (PDO), Northern Oscillation Index, and Aleutian Low Pressure Index. Washington, Oregon, and California (WOC) salmon sizes were negatively correlated with the MEI values indicating that ultimate fish size was affected negatively by El Niño‐like events. Further, we show that the growth trajectory of WOC salmon was set following the first ocean winter. Returning ocean‐type British Columbia‐Puget Sound Chinook salmon average fork length was positively correlated with the MEI values during the summer and autumn of return year, which was possibly a result of a shallower mixed layer and improved food‐web productivity of subarctic Pacific waters. Size variation of coho salmon stocks south of Alaska was synchronous and negatively correlated with warm conditions (positive PDO) and weak North Pacific high pressure during ocean residence.     

Spatial distribution of ocean habitat of yearling Chinook (Oncorhynchus tshawytscha) and coho (Oncorhynchus kisutch) salmon off Washington and Oregon,USA

We determined the habitat usage and habitat connectivity of juvenile Chinook (Oncorhynchus tshawytscha) and coho (Oncorhynchus kisutch) salmon in continental shelf waters off Washington and Oregon, based on samples collected every June for 9 yr (1998–2006). Habitat usage and connectivity were evaluated using SeaWiFS satellite‐derived chlorophyll a data and water depth. Logistic regression models were developed for both species, and habitats were first classified using a threshold value estimated from a receiver operating characteristic curve. A Bernoulli random process using catch probabilities from observed data, i.e. the frequency of occurrence of a fish divided by the number of times a station was surveyed, was applied to reclassify stations. Zero‐catch probabilities of yearling Chinook and yearling coho salmon decreased with increases in chlorophyll a concentration, and with decreases in water depth. From 1998 to 2006, ～ 47% of stations surveyed were classified as unfavorable habitat for yearling Chinook salmon and ～ 53% for yearling coho salmon. Potentially favorable habitat varied among years and ranged from 9 856 to 15 120 km² (Chinook) and from 14 800 to 16 736 km² (coho). For both species, the smallest habitat area occurred in 1998, an El Niño year. Favorable habitats for yearling Chinook salmon were more isolated in 1998 and 2005 than in other years. Both species had larger and more continuous favorable habitat areas along the Washington coast than along the Oregon coast. The favorable habitats were also larger and more continuous nearshore than offshore for both species. Further investigations on large‐scale transport, mesoscale physical features, and prey and predator availability in the study area are necessary to explain the spatial arrangement of juvenile salmon habitats in continental shelf waters.     

Climate variability and marine survival of coho salmon (Oncorhynchus kisutch) in the Oregon production area

Time series of adult recruitment for natural runs of coho salmon from the Oregon coastal region (1970–94) and marine survival of hatchery-reared coho salmon from California to Washington (1960–94) are significantly correlated with a suite of meteorological and oceanographic variables related to the biological productivity of the local coastal region. These variables include strong upwelling, cool sea surface temperature (SST), strong wind mixing, a deep and weakly stratified mixed layer, and low coastal sea level, indicating strong transport of the California Current. Principal component analysis indicates that these variables work in concert to define the dominant modes of physical variability, which appear to regulate nutrient availability and biological productivity. Multiple regression analysis suggests that coho marine survival is significantly and independently related to the dominant modes acting over this region in the periods when the coho first enter the ocean and during the overwintering/spring period prior to their spawning migration. Linear relationships provided good fits to the data and were robust, capable of predicting randomly removed portions of the data set.     

Otolith microstructure of chum salmon <Emphasis Type="Italic">Oncorhynchus keta</Emphasis>: formation of sea entry check and daily deposition of otolith increments in seawater conditions

To apply otolith microstructure to examination of age and growth of juvenile chum salmon Oncorhynchus keta inhabiting coastal waters, formation of otolith increments was investigated for juveniles reared in a seawater aquarium and in net pens. In all otoliths examined, a distinctive check was formed at the time of sea entry of the fish. The deposition of otolith increments after the check was daily for rearing both in the aquarium (57 days) and in the net pens (26 days). Check formation associated with sea entry was also observed in otoliths of juvenile salmon collected 1 km off the coast of Shari, Hokkaido, Japan. Transmitted light observation of otoliths of those fish revealed a transition in otolith increment appearance from dark to light. Otolith Sr: Ca ratio remarkably changed from a low to a high level, coinciding with the transition in otolith appearance. It is suggested that the transition was associated with individual sea entry. This study demonstrated that the check and/or transition associated with sea entry are applicable to a benchmark for otolith increment counts of juvenile chum salmon inhabiting coastal waters.     

The density dilemma: limitations on juvenile production in threatened salmon populations

Density‐dependent processes have repeatedly been shown to have a central role in salmonid population dynamics, but are often assumed to be negligible for populations at low abundances relative to historical records. Density dependence has been observed in overall spring/summer Snake River Chinook salmon Oncorhynchus tshawytscha production, but it is not clear how patterns observed at the aggregate level relate to individual populations within the basin. We used a Bayesian hierarchical modelling approach to explore the degree of density dependence in juvenile production for nine Idaho populations. Our results indicate that density dependence is ubiquitous, although its strength varies between populations. We also investigated the processes driving the population‐level pattern and found density‐dependent growth and mortality present for both common life‐history strategies, but no evidence of density‐dependent movement. Overwinter mortality, spatial clustering of redds and limited resource availability were identified as potentially important limiting factors contributing to density dependence. The ubiquity of density dependence for these threatened populations is alarming as stability at present low abundance levels suggests recovery may be difficult without major changes. We conclude that density dependence at the population level is common and must be considered in demographic analysis and management.