A life‐stage population model of the European sturgeon (Acipenser sturio) in the Elbe River. Part I: general model outline and potential applications

The European sturgeon, Acipenser sturio, experienced a dramatic decline during the late 19^th century in Germany. Only after its confirmed extirpation in the wild were ex‐situ measures introduced as the basis of a recovery attempt. In order to achieve effective population restoration, it is important to evaluate and compare efficiency of both planned and ongoing remediation measures. For this purpose, a stochastic age‐structured model of this population was developed to conduct a population viability analysis and to assess different management and restoration measures. The model revealed a high susceptibility of the species to even very low fishery and by‐catch pressures of below 5%. Population recovery was shown to be a very slow process, with a minimum timeframe of 30–50 years. Continuous stocking represents an important initial management measure to accelerate population recovery, reducing by two to three times the time required in comparison to only initial stocking. Habitat remediation (e.g. spawning grounds) in the Elbe River seems necessary, as according to model estimations, the present carrying capacity could sustain no more than a few thousand adults. The model showed a clear correlation between spawning habitat availability and the sustained number of adults.     

Monitoring demographics of a commercially exploited population of shovelnose sturgeon in the Wabash River,Illinois/Indiana,USA

Shovelnose sturgeon (Scaphirhynchus platorynchus, Rafinesque, 1820) in the Wabash River, Illinois/Indiana, USA, provide an important recreational sport and commercial caviar fishery. In fact, it is one of the last commercially viable populations for sturgeon roe harvest. Due to increased demand in the caviar trade and endangered species legislation that protect shovelnose sturgeon in only a portion of their range, efforts of the roe harvest market may continue to divert toward unprotected populations like the shovelnose sturgeon in the Wabash River. Previous studies have shown that increased harvest pressure in this species can affect the age‐at‐maturation and result in recruitment overfishing. Therefore, it is important to closely and continuously monitor commercially exploited populations. Over the past decade (2007–2016), 13,170 shovelnose sturgeon were sampled with boat electroshocking, hoop nets, drift nets, trotlines, and benthic electrified trawls. Captured fish ranged from 61 to 910 mm fork length (FL; mean = 668 mm), with very few fish less than 550 mm FL. Although fish were found to be in a healthy condition (mean relative weight = 87), there was a decrease in the mean condition over time. In addition, we saw declines in mean FL, weight of roe‐per‐fish, and size‐at‐maturity for female fish directly impacted by harvest. The decline of these population parameters, coupled with an increase in total annual mortality and a truncated age frequency distribution, suggest that harvest is negatively impacting the demographics and recruitment of shovelnose sturgeon in the Wabash River. Considering the downward trajectory of population dynamics and high estimates of mortality, their resiliency to continued harvest and environmental changes will be limited.     

Population prediction and viability model for pallid sturgeon (Scaphirhynchus albus,Forbes and Richardson, ) in the lower Missouri River

Long‐term population trends of pallid sturgeon Scaphirhynchus albus in the lower Missouri River were evaluated via a discrete and stochastic age‐structure population viability model. The intent of this model was to (i) estimate the local pallid sturgeon population size, (ii) quantify the contribution of hatchery‐reared fish to the overall population, (iii) predict the level of natural production needed to create a self‐sustaining population, and (iv) determine the parameters that produce the largest model sensitivity. The model estimated that the wild, adult population size was approximately 6000 fish that remained in the lower Missouri River in 2012 compared to approximately 42 000 hatchery‐reared pallid sturgeon. Under the assumption of no natural recruitment, the population size will continue to decline at approximately 8% annually, with an annual egg to age‐1 survival rate of 0.00011 predicted to maintain a stable population. The model was most sensitive to survival rates of fish ≥ age‐1 and less sensitive to age‐0 survival rates and fecundity. Decreasing or increasing the female spawning interval by 1 year had minimal effect on the overall population trajectory. Recovery management planning for a species such as pallid sturgeon, which is slow‐growing, late‐maturing, and has intermittent spawning would require several years to access recovery potential and management decisions. Barring any unforeseen natural catastrophe, the pallid sturgeon population in the lower Missouri River is not in immediate danger of local extirpation; however, the population appears to be far from viable and self‐sustaining.     

Harvest of Mississippi River sturgeon drives abundance and reproductive success: a harbinger of collapse?

Within harvested populations, relationships between harvest intensity and reproductive responses are typically unclear, rendering regulatory decisions difficult. Harvest of the commercially important shovelnose sturgeon (Scaphirhynchus platorynchus) is increasing in the upper Mississippi River; standardized seasonal sampling revealed that adult abundance is declining. Relative density of annual cohorts varied negatively with historical harvest intensity (r² = 0.84), suggesting that removal of mature adults is reducing the contribution of cohorts to population density. The results of simulation modeling suggest that this currently unregulated fishery is experiencing both growth and recruitment overfishing. Further, the current proposed multi‐state minimum length regulation was insufficient to maintain a sustainable stock. Only a more conservative minimum length limit (685 mm) produced yields that were sustainable at the current level of mortality and provided room for the fishery to grow. The annual mortality rate of the sympatric, federally endangered pallid sturgeon (S. albus) was similar to that of the shovelnose sturgeon population, raising concerns that harvest‐induced mortality is affecting this congener's vital rates.     

Fecundity of Atlantic sturgeon (Acipenser oxyrinchus Mitchill, 1815) captured by the commercial fishery in the Saint John River,New Brunswick,Canada

Knowledge of fecundity in fishes is important for understanding life history, modeling population dynamics, developing fisheries management and determining conservation status. Sturgeon species are known for their high fecundity but to date only two populations of Atlantic sturgeon (Acipenser oxyrinchus Mitchill, 1815) have been examined for this aspect of their biology. The species is protected in most regions of eastern North America but the Saint John River, New Brunswick supports one of the two remaining commercial fisheries in Canada. Sixty females selected by this fishery were sampled for total length, weight, age, gonad weight and egg number. Absolute fecundity ranged from 153,630 to 1,306,626 eggs, with a mean fecundity of 582,832 ± 261,806 eggs. Absolute fecundity exhibited a significant positive relationship to increasing body weight but not to total length or age. Relative fecundity ranged from 129 to 216 eggs/g gonad weight and although it declined with both increasing body size and age the relationships were not significant. The Gonadosomatic Index (GSI) ranged from 1.3% to 27.0% with a mean of 7.8% ± 4.94 SD. Relative fecundity found in this study was higher and mean GSI lower than reported for other Atlantic sturgeon populations and other sturgeon species. Differences were probably due to location of the fishery downstream from spawning sites and selection of pre-spawning females by the commercial fishery.     

Genetic assessment of lake sturgeon (<Emphasis Type="Italic">Acipenser fulvescens</Emphasis>) population structure in the Ottawa River

Lake sturgeon (Acipenser fulvescens) are of conservation concern throughout their range. Many populations are dependent on fluvial habitats which have been increasingly impacted and fragmented by dams and human development. Although lake sturgeon were once abundant in the Ottawa River and its tributaries, historical commercial harvests and other anthropogenic factors caused severe declines and low contemporary numbers in lake sturgeon populations. Contemporary habitat fragmentation by dams may be increasing isolation among habitat patches and local rates of decline, raising concerns for persistence of local populations. We used microsatellite DNA markers to assess population structure and diversity of lake sturgeon in the Ottawa River, and analyzed samples from 10 sites that represent more than 500 km of riverine habitat. To test for evidence of anthropogenic fragmentation, patterns of genetic diversity and connectivity within and among river segments were tested for concordance with geographic location, separation by distance and obstacles to migration, considering both natural and artificial barriers as well as barrier age. Despite extensive habitat fragmentation throughout the Ottawa River, statistical analyses failed to refute panmixia of lake sturgeon in this system. Although the long generation time of lake sturgeon appears to have effectively guarded against the negative genetic impacts of habitat fragmentation and loss so far, evidence from demographic studies indicates that restoring connectivity among habitats is needed for the long-term conservation and management of this species throughout this river system.     

A synopsis of lake sturgeon management in Alberta, Canada

The lake sturgeon ( Acipenser fulvescens ) is resident in the North and South Saskatchewan rivers of Alberta. Because of their confined distribution, low abundance, and vulnerability to anthropogenic impacts, lake sturgeon in Alberta have been the focus of specific management actions for nearly 60 years.
Lake sturgeon harvest was prohibited in Alberta from 1940 to 1968, after which a limited harvest was again permitted, but only as a hook-and-line sport fishery. After being reopened the fishery was primarily managed as a "trophy" fishery. Sport fishery harvest statistics have been compiled annually since 1968, through mail-out questionnaires sent to all holders of sturgeon angling licenses.
Few research studies have been undertaken on the life history or habitat requirements of lake sturgeon in the province, and biologists have relied primarily on anglers to provide information for management decisions. Preliminary population information for the South Saskatchewan River, from mark-recapture data, suggests a population size of about 3700 fish. Recent studies in the North Saskatchewan River indicate a population of about 1300 fish. Studies indicate that a portion of the population in both rivers is trans-boundary, moving between the waters of the adjoining provinces of Alberta and Saskatchewan, and creating concerns because of differences in angling regulations. During recent decades, major consumptive uses of water have also contributed to depletion of lake sturgeon habitat in the province.
After reviewing past management strategies and actions, Alberta Environmental Protection implemented a number of regulatory changes in 1987 to further protect lake sturgeon populations in the province while continuing to provide angling opportunities and maintaining a controlled harvest.     

Potential responses of the Lower Missouri River Shovelnose Sturgeon (Scaphirhynchus platorynchus) population to a commercial fishing ban

We developed an age‐structured population matrix model to perform population viability analysis for Lower Missouri River (LMR) shovelnose sturgeon (Scaphirhynchus platorynchus). We investigated potential effects of the commercial fishing moratorium put in place to help protect the similar‐appearing pallid sturgeon (S. albus). The model applies different components of total variance in life history parameters at different levels: sampling variance (parameter uncertainty) between model iterations; temporal variance (temporal environmental fluctuations) between time steps within iterations; and individual variance (individual differences) within each time‐step. The model predicted annual rates of population increase of 1.96% under historic fishing and 2.67% with removal of historic fishing. We identified combinations of fishing and harvest size restrictions that would permit a sustainable harvest of shovelnose sturgeon. Overall, the ban on commercial fishing of shovelnose sturgeon in the LMR due to similarity of appearance to pallid sturgeon should help the LMR shovelnose sturgeon population begin to rebound while decreasing any negative effects it may have had on pallid sturgeon populations.     

Estimated population size of pallid sturgeon Scaphirhynchus albus in a novel reach of the lower Missouri River

The recovery criterion for pallid sturgeon Scaphirhynchus albus consists of attaining a self‐sustaining genetically diverse population for two generations. The suppressed pallid sturgeon population is theorized as a potential factor limiting recovery; therefore, the Pallid Sturgeon Conservation and Augmentation Program (PSCAP) was implemented to proliferate the overall population. The pallid sturgeon population has been quantified in the lower basin of the Missouri River (Gavins Point Dam [rkm 1,305.2] to the confluence of the Missouri and Mississippi rivers [rkm 0.0]), but previous population estimates have only covered less than ten percent of the lower basin. Therefore, the objectives of this study were to quantify the annual pallid sturgeon population (2015–2017) in a novel, unquantified 30.1 rkm reach of the lower Missouri River basin and compare these results to previously published population estimates in the lower Missouri River basin. The study area included a six‐bend reach in the upper channelized Missouri River, approximately 226.3 rkm below the last main‐stem dam. Population estimates (N_sp) for the “super‐population” of pallid sturgeon within the 30.1 rkm study area varied from 593 (95% CI 471–716) in 2015 to 471 (373–569) in 2016 and 608 (482–734) in 2017. The population estimates (fish/rkm ± SE) ranged from 15.6 ± 1.0 to 20.2 ± 1.3 fish/rkm. This study aligned with a previously published estimate in the same proximal reach but was two or three times higher compared to an estimate reported from approximately 500 rkm downstream. Understanding the temporal and spatial variations of the pallid sturgeon population is critical as recovery efforts continue, especially to seed input parameters into population prediction models that provide management guidance.     

Mitochondrial DNA variation, effective female population size and population history of the endangered Chinese sturgeon, Acipenser sinensis

The anadromous Chinese sturgeon (Acipenser sinensis), mainly endemic to the Yangtze River in China, is an endangered fish species. The natural population has declined since the Gezhouba Dam blocked its migratory route to the spawning grounds in 1981. In the near future, the completion of the Three Gorges Dam, the world's largest hydroelectric project, may further impact this species by altering the water flow of the Yangtze River. Little is currently known about the population genetic structure of the Chinese sturgeon. In this study, DNA sequence data were determined from the control region (D-loop) of the mitochondrial genome of adult sturgeons (n = 106) that were collected between 1995–2000. The molecular data were used to investigate genetic variation, effective female population size and population history of the Chinese sturgeon in the Yangtze River. Our results indicate that the reduction in abundance did not change genetic variation of the Chinese sturgeon, and that the population underwent an expansion in the past. AMOVA analysis indicated that 98.7% of the genetic variability occurred within each year's spawning populations, the year of collection had little influence on the diversity of annual temporary samples. The relative large effective female population size (N _ef) indicates that good potential exists for the recovery of this species in the future. Strikingly, the ratio of N _ef to the census female population size (N _f) is unusually high (0.77–0.93). This may be the result of a current bottleneck in the population of the Chinese sturgeon that is likely caused by human intervention. This revised version was published online in July 2006 with corrections to the Cover Date.     

Population viability analysis of the Danube sturgeon populations in a Vortex simulation model

Populations of six sturgeon species in the Danube River (beluga, Russian sturgeon, stellate sturgeon, sterlet, ship sturgeon and Atlantic sturgeon) have experienced severe decline during the last several decades, mostly due to the unsustainable fishery, river fragmentation and water pollution. Present lack of knowledge on basic sturgeon demography, life history and relative effects of different negative factors is further hindering implementation of efficient policy and management measures. In the present study, population viability analysis in a Vortex simulation model has been conducted in order to assess the state of the six Danube sturgeon species, their future risk of extinction and to determine the most suitable conservation and management measures. Population viability analysis has revealed a large sensitivity of the Danube sturgeon populations to changes in the natural mortality, fecundity, age at maturity and spawning frequency. It was also confirmed that the sturgeons are highly susceptible to even moderate levels of commercial fishery, and that their recovery is a multi-decadal affair. Stocking with adult individuals was shown to produce considerably greater effect on population persistence than stocking with juveniles, but the latter approach is probably still preferable since it avoids many inherent problems of aquaculture cultivation. This study represents the first population viability analysis of the Danube sturgeons.     

Effects of harvest and length limits on shovelnose sturgeon in the upper Wabash River, Indiana

Shovelnose sturgeon Scaphirhynchus platorynchus are one of the few sturgeon species that currently support sustainable commercial harvest. However, harvest closures for many Eurasian sturgeons have resulted in increased exploitation of this fishery, thereby raising concerns about the sustainability of shovelnose sturgeon resources. As a result, the maintenance of self‐sustaining shovelnose sturgeon populations will require the estimation of appropriate harvest levels. This study used an age‐structured population model to examine the effects of harvest (u = 0.15–0.75) and length restrictions on population abundance, mean length‐at‐harvest, biomass, yield, and reproductive potential of female shovelnose sturgeon in the upper Wabash River, Indiana. Model simulations for four hypothetical length‐restriction scenarios (610‐ to 813‐mm reverse slot limit, and a 610‐, 635‐, and 660‐mm minimum length limit) were compared to outputs with no restriction. All population parameters within each length‐restriction scenario declined with increases in harvest level. For each harvest level, all population parameters increased as length limits became more restrictive. The reverse slot limit and 610‐mm minimum length limit provided adequate protection to allow population parameters to increase through an annual harvest level of 0.55. However, these length restrictions were not sufficiently conservative to warrant implementation due to their similarity to length‐at‐maturity of female shovelnose sturgeon. The implementation of a 635‐mm minimum length limit would protect female shovelnose sturgeon from harvest rates >0.75, allow 92% of the females to remain available for harvest, and minimize short‐term (<30 years) declines in yield. Further, sensitivity and robustness analyses suggested that the 635‐mm minimum length limit would allow population parameters to increase even at the worst‐case scenario. As a result, the 635‐mm minimum length limit was recommended as the most appropriate regulation to promote conservation and sustainable harvest of shovelnose sturgeon in the upper Wabash River.     

Documentation of lake sturgeon (Acipenser fulvescens Rafinesque, 1817) recovery and spawning success from a restored population in the Mississippi River,Missouri, USA

Lake sturgeon Acipenser fulvescens are considered rare and were nearly extirpated in the Mississippi River in Missouri by 1931 as a result of overfishing and habitat fragmentation. Propagation efforts have been implemented by the Missouri Department of Conservation since 1984 as means to restore the lake sturgeon population. Although recent population increases have been observed, a formalized evaluation to determine if lake sturgeon are self‐sustaining in the Missouri portion of the Mississippi River has not been completed. Therefore, the objectives of this study were to: (i) determine the proportion of reproductive individuals, (ii) evaluate seasonal movement patterns of adults, and (iii) validate purported spawning locations within the Mississippi River in Missouri. Lake sturgeon catch data indicated that approximately 11 percent of the population are reproductively mature. Additionally, telemetry data confirms that the greatest movement by adult lake sturgeon occurs during spring, which suggests spawning behavior. Finally, it was possible to document lake sturgeon embryos and emergent fry larvae below Melvin Price Locks and Dam 26 in the Upper Mississippi River near St. Louis, Missouri. Water velocity, depth, and substrate size were measured at this location and embryos were collected and hatched in the laboratory. River gage data suggest that spawning behavior may have been elicited by a large influx of water during a drawdown period of water above the dam. This study represents the first documented spawning of A. fulvescens in the Mississippi River and highlights the success of recovery efforts in Missouri.     

Reproductive biology of female shovelnose sturgeon in the upper Wabash River, Indiana

Shovelnose sturgeon Scaphirhynchus platorynchus are commercially important, supporting a viable roe fishery throughout much of the Mississippi River drainage. We examined the reproductive attributes of stage‐5 female shovelnose sturgeon captured from the upper Wabash River, Indiana, from March to June 2004. Shovelnose sturgeon were collected using boat electrofishing and experimental gill nets, measured for fork length (FL) and wet weight, and sexed externally if possible. Size‐ and age‐at‐maturity, absolute and relative fecundity, relative egg size, and gonadosomatic index (GSI) were determined for 49 female shovelnose sturgeon (range, 601‐ to 858‐mm FL). Female shovelnose sturgeon reached sexual maturity at approximately 600 mm and age‐at‐maturity ranged from ages 6 to 12 (median age = 9). Relative fecundity ranged from 11 220 to 23 956 eggs kg^?1 (mean = 18 156 eggs kg^?1). Absolute fecundity ranged from 14 294 to 65 490 eggs female^?1 (mean = 30 397 eggs female^?1) and was positively related to FL (r² = 0.76) and wet weight (r² = 0.82). The number of eggs g^?1 of ovary weight ranged from 72 to 170 (mean = 98 eggs g^?1) and was negatively correlated with GSI. GSI values ranged from 9.4 to 27.2 (mean = 19.3) and were positively correlated to FL (r² = 0.18). Our results increase our understanding of shovelnose sturgeon reproductive biology and recruitment dynamics and provide input for models to evaluate the effects of harvest on this species.     

Population viability analysis of Lower Missouri River shovelnose sturgeon with initial application to the pallid sturgeon

Demographic models for the shovelnose (Scaphirhynchus platorynchus) and pallid (S. albus) sturgeons in the Lower Missouri River were developed to conduct sensitivity analyses for both populations. Potential effects of increased fishing mortality on the shovelnose sturgeon were also evaluated. Populations of shovelnose and pallid sturgeon were most sensitive to age‐0 mortality rates as well as mortality rates of juveniles and young adults. Overall, fecundity was a less sensitive parameter. However, increased fecundity effectively balanced higher mortality among sensitive age classes in both populations. Management that increases population‐level fecundity and improves survival of age‐0, juveniles, and young adults should most effectively benefit both populations. Evaluation of reproductive values indicated that populations of pallid sturgeon dominated by ages ≥35 could rapidly lose their potential for growth, particularly if recruitment remains low. Under the initial parameter values portraying current conditions the population of shovelnose sturgeon was predicted to decline by 1.65% annually, causing the commercial yield to also decline. Modeling indicated that the commercial yield could increase substantially if exploitation of females in ages ≤12 was highly restricted.     

Entrainment of shovelnose sturgeon by towboat navigation in the Upper Mississippi River

Estimated number of shovelnose sturgeon Scaphirhynchus platorynchus impacted annually by towboat entrainment in navigation pools of the Upper Mississippi River were compared against estimates of fishery harvest and ambient population densities to evaluate the relevance of entrainment at the population level. Mean number of sturgeon entrained per kilometer of navigation was estimated at 0.02, and mean number entrained annually considering towboat traffic was estimated at 0.38 sturgeon/ha. Losses associated with entrainment were mostly lower than fishery harvest, although differences were not large. The two sources of mortality combined could potentially reduce the mature adult population to a level where it no longer has the reproductive capacity to replenish itself. Thus, through a combination of entrainment and fishing mortality shovelnose sturgeon may be looming near unsustainable population levels. These estimates are preliminary considering the many uncertainties associated with quantifying entrainment and its effects. Additional research is needed not only to derive better estimates, but also to develop options for managing entrainment.     

Age,growth, mortality,and abundance of lake sturgeon in the Grasse River,New York,USA

An increased understanding of lake sturgeon (Acipenser fulvescens) population dynamics is a key requirement for successful management efforts. Little is known regarding the Grasse River population of lake sturgeon except that it is one of a few populations in New York State where spawning has been documented. Thus our purpose was to assess the current status of lake sturgeon in the Grasse River system, including age, growth, mortality, and abundance. Age was determined for 196 of 211 lake sturgeon by examination of sectioned pectoral fin rays. Ages ranged from 0 to 32 years and the annual mortality rate for fish between ages 7 and 14 was 16.8%. The weight (W, g) to total length (TL, mm) relationship was W = 1.281 × 10^?6TL^3.202. The von Bertalanffy growth equation was TL = 1913(1?e^{?0.0294(t+9.5691)}). While the range of observed ages was similar to that of nearby St. Lawrence River populations, mean weight at age for an individual at 1000 mm TL was lower than that observed for lake sturgeon within Lake St. Francis of the St. Lawrence River. Predicted growth based on von Bertalanffy parameters was similar to that observed for the nearby Lake St. Francis. An open population estimator using the POPAN sub‐module in the Program MARK produced an abundance estimate of 793 lake sturgeon (95% CI = 337–1249).     

Long‐term breeding demography and density dependence in an increasing population of Golden Eagles Aquila chrysaetos

Few studies have quantified the dynamics of recovering populations of large raptors using long‐term, spatially explicit studies. Using data collected over 37 years in the western Italian Alps, we assessed the trends in distribution, abundance, fecundity and breeding population structure of Golden Eagles Aquila chrysaetos. Using the spatial distribution of territory centroids in 2007, we found that the spatial distribution of eagle territories was over‐dispersed up to 3 km. Although population size and total productivity increased from 1972 to 2008, the proportion of pairs that laid eggs showed a strong decline, falling to no more than 50% after 2003. On average, 15% of successful nests produced two fledglings, and productivity also declined over time. No significant relationship between population growth rate and total population size was detected, but the percentage of pairs that bred and breeding success showed evidence of density dependence, as they declined significantly with increasing density. Our results suggest that density dependence, operating across heterogeneous habitats, is currently regulating this population, while the carrying capacity may still be increasing. This may explain the apparent paradox of reduced breeding effort despite increasing total productivity.     

Scaling the consequences of interactions between invaders from the individual to the population level

The impact of human‐induced stressors, such as invasive species, is often measured at the organismal level, but is much less commonly scaled up to the population level. Interactions with invasive species represent an increasingly common source of stressor in many habitats. However, due to the increasing abundance of invasive species around the globe, invasive species now commonly cause stresses not only for native species in invaded areas, but also for other invasive species. I examine the European green crab Carcinus maenas, an invasive species along the northeast coast of North America, which is known to be negatively impacted in this invaded region by interactions with the invasive Asian shore crab Hemigrapsus sanguineus. Asian shore crabs are known to negatively impact green crabs via two mechanisms: by directly preying on green crab juveniles and by indirectly reducing green crab fecundity via interference (and potentially exploitative) competition that alters green crab diets. I used life‐table analyses to scale these two mechanistic stressors up to the population level in order to examine their relative impacts on green crab populations. I demonstrate that lost fecundity has larger impacts on per capita population growth rates, but that both predation and lost fecundity are capable of reducing population growth sufficiently to produce the declines in green crab populations that have been observed in areas where these two species overlap. By scaling up the impacts of one invader on a second invader, I have demonstrated that multiple documented interactions between these species are capable of having population‐level impacts and that both may be contributing to the decline of European green crabs in their invaded range on the east coast of North America.