Density dependence and density independence during the early life stages of four marine fish stocks

Recruitment variability caused by density-dependent and density-independent processes is an important area within the study of fish dynamics. These processes can exhibit nonlinearities and nonadditive properties that may have profound dynamic effects. We investigate the importance of population density (i.e., density dependence) and environmental forcing (i.e., density independence) on the age-0 and age-1 abundance of capelin (Mallotus villosus), northeast Arctic cod (Gadus morhua), northeast Arctic haddock (Melanogrammus aeglefinus), and Norwegian spring spawning herring (Clupea harengus) in the Barents Sea. We use statistical methods that explicitly account for nonlinearities and nonadditive interactions between internal and external variables in the abundance of these two pre-recruitment stages. Our results indicate that, during their first five months of life, cod, haddock, and herring experience higher density-dependent survival than capelin. The abundance of age-0 cod depends on the mean age and biomass of the spawning stock, a result which has implications for the management of the entire cod stock. Temperature is another important factor influencing the abundance at age-0 and age-1 of all four species, except herring at age-1. Between age-0 and age-1, there is an attenuation of density-dependent survival for cod and herring, while haddock and capelin experience density dependence at high and low temperatures, respectively. Predation by subadult cod is important for both capelin and cod at age-1. We found strong indications for interactions among the studied species, pointing to the importance of viewing the problem of species recruitment variability as a community, rather than as a population phenomenon.     

Feeding opportunities of larval and juvenile cod (Gadus morhua) in a Greenlandic fjord: temporal and spatial linkages between cod and their preferred prey

Feeding of fish depends on a spatial and temporal match with prey, and since larval and juvenile feeding can be highly selective, their preferences for given prey sizes and taxa should be considered when quantifying the actual availability of potential prey. We investigated the diet and prey preferences of the early-life stages of Atlantic cod (Gadus morhua) to quantify the availability of prey during a spring-summer season in a West Greenlandic fjord. We hypothesized that abundances of larval and juvenile cod at size were synchronized to optimal availability of preferred prey in space and time. The present analysis is based on nine cruises each covering 5 stations visited between 24 May and 5 August 2010 comparing zooplankton abundance, cod gut content and distribution patterns. Cod 4–25 mm in length preferred prey of about 5 % of their own length. During ontogeny, their preferences changed from calanoid nauplii towards Pseudocalanus spp. and Calanus spp. copepodites. The larvae/juvenile had an exceptionally high dietary contribution from cladocerans, which were highly preferred by cod larger than 9 mm, while the abundant Metridia longa and the non-calanoid copepods contributed less. These findings stress the importance of focusing on abundance of preferred prey when assessing the actual prey availability to young fish. We found a spatio-temporal overlap between cod and their preferred prey, and observations suggest that advection of both zooplankton and cod contributed to this overlap. Hence, the larval feeding opportunities might be sensitive to climate-related changes affecting the circulation patterns in this fjord.     

Predicting abundance of desert riparian birds: validation and calibration of the Effective Area Model.

Reliable prediction of the effects of landscape change on species abundance is critical to land managers who must make frequent, rapid decisions with long-term consequences. However, due to inherent temporal and spatial variability in ecological systems, previous attempts to predict species abundance in novel locations and/or time frames have been largely unsuccessful. The Effective Area Model (EAM) uses change in habitat composition and geometry coupled with response of animals to habitat edges to predict change in species abundance at a landscape scale. Our research goals were to validate EAM abundance predictions in new locations and to develop a calibration framework that enables absolute abundance predictions in novel regions or time frames. For model validation, we compared the EAM to a null model excluding edge effects in terms of accurate prediction of species abundance. The EAM outperformed the null model for 83.3% of species (N=12) for which it was possible to discern a difference when considering 50 validation sites. Likewise, the EAM outperformed the null model when considering subsets of validation sites categorized on the basis of four variables (isolation, presence of water, region, and focal habitat). Additionally, we explored a framework for producing calibrated models to decrease prediction error given inherent temporal and spatial variability in abundance. We calibrated the EAM to new locations using linear regression between observed and predicted abundance with and without additional habitat covariates. We found that model adjustments for unexplained variability in time and space, as well as variability that can be explained by incorporating additional covariates, improved EAM predictions. Calibrated EAM abundance estimates with additional site-level variables explained a significant amount of variability (P < 0.05) in observed abundance for 17 of 20 species, with R2 values >25% for 12 species, >48% for six species, and >60% for four species when considering all predictive models. The calibration framework described in this paper can be used to predict absolute abundance in sites different from those in which data were collected if the target population of sites to which one would like to statistically infer is sampled in a probabilistic way.     

Regional-scale variation in the distribution and abundance of farming damselfishes on Australia’s Great Barrier Reef

Territorial damselfishes that manipulate (“farm”) the algae in their territories can have a marked effect on benthic community structure and may influence coral recovery following disturbances. Despite the numerical dominance of farming species on many reefs, the importance of their grazing activities is often overlooked, with most studies only examining their roles over restricted spatial and temporal scales. We used the results of field surveys covering 9.5° of latitude of the Great Barrier Reef to describe the distribution, abundance and temporal dynamics of farmer communities. Redundancy analysis revealed unique subregional assemblages of farming species that were shaped by the combined effects of shelf position and, to a lesser extent, by latitude. These spatial patterns were largely stable through time, except when major disturbances altered the benthic community. Such disturbances affected the functional guilds of farmers in different ways. Since different guilds of farmers modify benthic community structure and affect survival of juvenile corals in different ways, these results have important implications for coral recovery following disturbances.     

Spatial scaling of avian population dynamics: population abundance, growth rate, and variability

Synchrony in population fluctuations has been identified as an important component of population dynamics. In a previous study, we determined that local-scale (<15-km) spatial synchrony of bird populations in New England was correlated with synchronous fluctuations in lepidopteran larvae abundance and with the North Atlantic Oscillation. Here we address five questions that extend the scope of our earlier study using North American Breeding Bird Survey data. First, do bird populations in eastern North America exhibit spatial synchrony in abundances at scales beyond those we have documented previously? Second, does spatial synchrony depend on what population metric is analyzed (e.g., abundance, growth rate, or variability)? Third, is there geographic concordance in where species exhibit synchrony? Fourth, for those species that exhibit significant geographic concordance, are there landscape and habitat variables that contribute to the observed patterns? Fifth, is spatial synchrony affected by a species' life history traits? Significant spatial synchrony was common and its magnitude was dependent on the population metric analyzed. Twenty-four of 29 species examined exhibited significant synchrony in population abundance: mean local autocorrelation (rho)= 0.15; mean spatial extent (mean distance where rho=0) = 420.7 km. Five of the 29 species exhibited significant synchrony in annual population growth rate (mean local autocorrelation = 0.06, mean distance = 457.8 km). Ten of the 29 species exhibited significant synchrony in population abundance variability (mean local autocorrelation = 0.49, mean distance = 413.8 km). Analyses of landscape structure indicated that habitat variables were infrequent contributors to spatial synchrony. Likewise, we detected no effects of life history traits on synchrony in population abundance or growth rate. However, short-distance migrants exhibited more spatially extensive synchrony in population variability than either year-round residents or long-distance migrants. The dissimilarity of the spatial extent of synchrony across species suggests that most populations are not regulated at similar spatial scales. The spatial scale of the population synchrony patterns we describe is likely larger than the actual scale of population regulation, and in turn, the scale of population regulation is undoubtedly larger than the scale of individual ecological requirements.     

Range-wide effects of breeding- and nonbreeding-season climate on the abundance of a Neotropical migrant songbird

Geographic variation in the population dynamics of a species can result from regional variability in climate and how it affects reproduction and survival. Identifying such effects for migratory birds requires the integration of population models with knowledge of migratory connectivity between breeding and nonbreeding areas. We used Bayesian hierarchical models with 26 years of Breeding Bird Survey data (1982-2007) to investigate the impacts of breeding- and nonbreeding-season climate on abundance of American Redstarts (Setophaga ruticilla) across the species range. We focused on 15 populations defined by Bird Conservation Regions, and we included variation across routes and observers as well as temporal trends and climate effects. American Redstart populations that breed in eastern North America showed increased abundance following winters with higher plant productivity in the Caribbean where they are expected to overwinter. In contrast, western breeding populations showed little response to conditions in their expected wintering areas in west Mexico, perhaps reflecting lower migratory connectivity or differential effects of winter rainfall on individuals across the species range. Unlike the case with winter climate, we found few effects of temperature prior to arrival in spring (March-April) or during the nesting period (May-June) on abundance the following year. Eight populations showed significant changes in abundance, with the steepest declines in the Atlantic Northern Forest (-3.4%/yr) and the greatest increases in the Prairie Hardwood Transition (4%/yr). This study emphasizes how the effects of climate on populations of migratory birds are context dependent and can vary depending on geographic location and the period of the annual cycle. Such knowledge is essential for predicting regional variation in how populations of a species might vary in their response to climate change.     

Drivers of euphausiid species abundance and numerical abundance in the Atlantic Ocean

Mid-ocean ridges are common features of the world’s oceans but there is a lack of understanding as to how their presence affects overlying pelagic biota. The Mid-Atlantic Ridge (MAR) is a dominant feature of the Atlantic Ocean. Here, we examined data on euphausiid distribution and abundance arising from several international research programmes and from the continuous plankton recorder. We used a generalized additive model (GAM) framework to explore spatial patterns of variability in euphausiid distribution on, and at either side of, the MAR from 60°N to 55°S in conjunction with variability in a suite of biological, physical and environmental parameters. Euphausiid species abundance peaked in mid-latitudes and was significantly higher on the ridge than in adjacent waters, but the ridge did not influence numerical abundance significantly. Sea surface temperature (SST) was the most important single factor influencing both euphausiid numerical abundance and species abundance. Increases in sea surface height variance, a proxy for mixing, increased the numerical abundance of euphausiids. GAM predictions of variability in species abundance as a function of SST and depth of the mixed layer were consistent with present theories, which suggest that pelagic niche availability is related to the thermal structure of the near surface water: more deeply-mixed water contained higher euphausiid biodiversity. In addition to exposing present distributional patterns, the GAM framework enables responses to potential future and past environmental variability including temperature change to be explored.     

Distribution,abundance and survival of juvenile hermatypic corals (Scleractinia) and the importance of life history strategies in the parent coral community

The distribution and abundance of juvenile corals were examined at depths from 3 to 37 m on the reefs of Curaçao and Bonaire (Netherlands Antilles). Juveniles of Agaricia agaricites were most abundant (60.6%), followed by Helioseris cucullata (8.3%). The large massive corals such as Montastrea annularis, M. cavernosa and branched species such as Madracis mirabilis and Acropora palmata had few juveniles. This, combined with species characteristics, shows that these species employ very different life history strategies. In some species the abundance of juveniles over the reef paralleled that of larger colonies, but not for example in Agaricia agaricites. The composition of the coral community was apparently no direct function of juvenile abundance. A change in angle of settlement of A. agaricites juveniles with increasing depth, from vertical to horizontal surfaces, seems to reflect the preferred light intensity. We studied the survival of juvenile corals during a half-year period. One-third remained unharmed, one-third died or disappeared, and one-third was limited in growth by factors such as spatial competition. This was the same for all depths, but factors influencing survival varied with depth.     

Thermal reaction norms for growth vary among cohorts of Pacific cod (Gadus macrocephalus)

While much effort has been directed at determining the spatial scales of adaptation in thermal reaction norms for growth, it is widely assumed that these reaction norms have high temporal stability. Water temperatures in the Gulf of Alaska in 2007 were the coldest on record since the mid-1970s and we present evidence that the thermal reaction norm for growth of age-0 Pacific cod (Gadus macrocephalus) in this cohort differed significantly from two adjacent cohorts. In addition to exhibiting higher growth potential at low temperatures, the 2007 cohort had a higher mean vertebral count, consistent with the widespread thermal effect known as “Jordan’s Rule.” Variation among cohorts in these physiological and morphological traits suggests a persistent response to environmental history (epigenetic effect). Temperature-induced phenotypic plasticity in the reaction norm for growth has significant implications for using growth rates to evaluate habitat quality and illustrates the complex responses of fishes to climate variability.     

Reef size and isolation determine the temporal stability of coral reef fish populations

Temporal variance in species abundance, a potential driver of extinction, is linked to mean abundance through Taylor's power law, the empirical observation of a linear log-log relationship with a slope between 1 and 2 for most species. Here we test the idea that the slope of Taylor's power law can vary both among species and spatially as a function of habitat area and isolation. We used the world's most extensive database of coral reef fish communities comprising a 15-year series of fish abundances on 43 reefs of Australia's Great Barrier Reef. Greater temporal variances were observed at small and isolated reefs, and lower variances at large and connected ones. The combination of reef area and isolation was associated with an even greater effect on temporal variances, indicating strong empirical support for the idea that populations on small and isolated reefs will succumb more frequently to local extinction via higher temporal variability, resulting in lower resilience at the community level. Based on these relationships, we constructed a regional predictive map of the dynamic fragility of coral reef fish assemblages on the Great Barrier Reef.     

The Paradox of the Long‐Term Positive Effects of a North American Crayfish on a European Community of Predators

Abstract: Invasions of non‐native species are one of the major causes of losses of native species. In some cases, however, non‐natives may also have positive effects on native species. We investigated the potential facilitative effects of the North American red swamp crayfish (Procambarus clarkii) on the community of predators in southwestern Spain. To do so, we examined the diets of predators in the area and their population trends since introduction of the crayfish. Most predator species consumed red swamp crayfish, which sometimes occurred in over 50% of their diet samples. Moreover, the abundance of species preying on crayfish increased significantly in the area as opposed to the abundance of herbivores and to predator populations in other areas of Europe, where those predators are even considered threatened. Thus, we report the first case in which one non‐native species is both beneficial because it provides prey for threatened species and detrimental because it can drive species at lower trophic levels to extinction. Increases in predator numbers that are associated with non‐native species of prey, especially when some of these predators are also invasive non‐natives, may increase levels of predation on other species and produce cascading effects that threaten native biota at longer temporal and larger spatial scales. Future management plans should include the complexity of interactions between invasive non‐natives and the entire native community, the feasibility of successful removal of non‐native species, and the potential social and economic interests in the area.     

Gizzard shad put a freeze on winter mortality of age-0 yellow perch but not white perch.

Four decades of observations on the limnology and fishes of Oneida Lake, New York, USA, provided an opportunity to investigate causes of mortality during winter, a period of resource scarcity for most juvenile fishes, in age-0 yellow perch (Perca flavescens) and age-0 white perch (Morone americana). This time series contains several environmental (e.g., winter severity) and biological (e.g., predator abundance) signals that can be used to disentangle multiple effects on overwinter mortality of these fishes. A multiple regression analysis indicated that age-0 yellow perch winter mortality was inversely related to fish length in autumn and to the abundance of alternative prey (gizzard shad [Dorosoma cepedianum] and white perch). However, no length-selective predation of yellow perch by one of the main predators, adult walleye (Sander vitreus), was detected. In contrast, white perch mortality was directly associated with total predator biomass and abundance of white perch in autumn, and inversely related to yellow perch abundance as a potential buffer species, but not to the abundance of gizzard shad. Winter severity was not a significant predictor of mortality for either perch species. Predicted winter starvation mortality, from a model described in the literature, was much lower than observed mortality for yellow perch. Similar models for white perch were correlated with observed mortality. These results collectively suggest that predation is the main mechanism shaping winter mortality of yellow perch, while both predation and starvation may be important for white perch. This analysis also revealed that gizzard shad buffer winter mortality of yellow perch. Although winter duration determines the northern limit of fish distributions, in mid-latitude Oneida Lake and for these species, predator-prey interactions seem to exert a greater influence on winter mortality than starvation.     

Spatiotemporal variation in survival rates: implications for population dynamics of yellow-bellied marmots

Spatiotemporal variation in age-specific survival rates can profoundly influence population dynamics, but few studies of vertebrates have thoroughly investigated both spatial and temporal variability in age-specific survival rates. We used 28 years (1976-2003) of capture-mark-recapture (CMR) data from 17 locations to parameterize an age-structured Cormack-Jolly-Seber model, and investigated spatial and temporal variation in age-specific annual survival rates of yellow-bellied marmots (Marmota flaviventris). Survival rates varied both spatially and temporally, with survival of younger animals exhibiting the highest degree of variation. Juvenile survival rates varied from 0.52 +/- 0.05 to 0.78 +/- 0.10 among sites and from 0.15 +/- 0.14 to 0.89 +/- 0.06 over time. Adult survival rates varied from 0.62 +/- 0.09 to 0.80 +/- 0.03 among sites, but did not vary significantly over time. We used reverse-time CMR models to estimate the realized population growth rate (lamda), and to investigate the influence of the observed variation in age-specific survival rates on lamda. The realized growth rate of the population closely covaried with, and was significantly influenced by, spatiotemporal variation in juvenile survival rate. High variability in juvenile survival rates over space and time clearly influenced the dynamics of our study population and is also likely to be an important determinant of the spatiotemporal variation in the population dynamics of other mammals with similar life history characteristics.     

Environmental stability and seasonality of a harpacticoid copepod community

The southwest monsoon on the west coast of India brings about dynamic changes in estuaries and coastal waters. The response of the meiofauna to monsoonal rain is obvious, but the impact of such environmental changes on the community structure of harpacticoid copepod species and their seasonality in the estuaries influenced by the tropical monsoon is poorly understood. In this study the spatial and temporal variability in abundance and community structure of meiobenthic copepods was investigated over an annual cycle (June 1983 to June 1984), in an estuary influenced by the tropical monsoon. Total meiofaunal abundance showed wide variations in space and time. Minimum and maximum densities were observed in the monsoon and pre-monsoon seasons, respectively. Quick recovery of harpacticoid populations in the early post-monsoon season indicated their recuperative power under adverse conditions. Of the 25 species recorded from lower, middle and upper reaches, eight comprised over 70% of the total copepod population. The peak and low occurrence of dominant species displayed striking correlations with the summer and rainy seasons, respectively. While other species were restricted in distribution, Stenhelia longifurca was recorded from all salinity regimes. The harpacticoid community was greatly influenced by the onset of the monsoon period, and their spatial and temporal variabilities were related with physico-chemical parameters and the variability of these parameters in the estuary.     

Effects of small-scale habitat fragmentation on predator–prey interactions in a temperate sea grass system

During the last decades, fragmentation has become an important issue in ecological research. Habitat fragmentation operates on spatial scales ranging over several magnitudes from patches to landscapes. We focus on small-scale fragmentation effects relevant to animal foraging decision making that could ultimately generate distribution patterns. In a controlled experimental environment, we tested small-scale fragmentation effects in artificial sea grass on the feeding behaviour of juvenile cod (Gadus morhua). Moreover, we examined the influence of fragmentation on the distribution of one of the juvenile cod’s main prey resources, the grass shrimp (Palaemon elegans), in association with three levels of risk provided by cod (no cod, cod chemical cues and actively foraging cod). Time spent by cod within sea grass was lower in fragmented landscapes, but total shrimp consumption was not affected. Shrimp utilised vegetation to a greater extent in fragmented treatments in combination with active predation. We suggest that shrimp choose between sand and vegetation habitats to minimize risk of predation according to cod habitat-specific foraging capacities, while cod aim to maximize prey-dependent foraging rates, generating a habitat-choice game between predator and prey. Moreover, aggregating behaviour in grass shrimp was only found in treatments with active predation. Hence, we argue that both aggregation and vegetation use are anti-predator defence strategies applied by shrimp. We therefore stress the importance of considering small-scale behavioural mechanisms when evaluating consequences from habitat fragmentation on trophic processes in coastal environments.     

Detecting conservation benefits in spatially protected fish populations with meta-analysis of long-term monitoring data

Marine protected areas (MPA) produce a positive effect on fish populations, but this may be difficult to identify due to the high temporal variability of populations. Meta-analysis is an option for analysing data from different sources and sampling designs and it can address problems related to temporal and spatial variability in fish populations. We analysed fish abundance data from visual counts conducted in summer, from 1996 to 2002, in the MPA of Tabarca (Alicante, Spain). The results showed an overall positive effect of protection at the species and family levels. Overall abundance of fishes inside the reserve was, on average, 1.22 times higher than outside the reserve boundaries. Positive effect of protection was found for Boops boops, Diplodus annularis, Diplodus cervinus, Epinephelus marginatus, Epinephelus costae and Epinephelus aenus. Species of Labrids were not affected by protection, except for Thalassoma pavo and Symphodus ocellatus. Meta-analysis of temporal data allows evaluation of the protection MPA provide and is particularly useful when data sources have different experimental designs or sampling programs. The Tabarca MPA has benefited fish populations by increasing their abundance and we suggest that meta-analysis is a complementary tool for the management of MPAs.     

Using individual-based movement information to identify spatial conservation priorities for mobile species

The optimal design of reserve networks and fisheries closures depends on species occurrence information and knowledge of how anthropogenic impacts interact with the species concerned. However, challenges in surveying mobile and cryptic species over adequate spatial and temporal scales can mask the importance of particular habitats, leading to uncertainty about which areas to protect to optimize conservation efforts. We investigated how telemetry-derived locations can help guide the scale and timing of fisheries closures with the aim of reducing threatened species bycatch. Forty juvenile speartooth sharks (Glyphis glyphis) were monitored over 22 months with implanted acoustic transmitters and an array of hydrophone receivers. Using the decision-support tool Marxan, we formulated a permanent fisheries closure that prioritized areas used more frequently by tagged sharks and considered areas perceived as having high value to fisheries. To explore how the size of the permanent closure compared with an alternative set of time-area closures (i.e., where different areas were closed to fishing at different times of year), we used a cluster analysis to group months that had similar arrangements of selected planning units (informed by shark movements during that month) into 2 time-area closures. Sharks were consistent in their timing and direction of migratory movements, but the number of tagged sharks made a big difference in the placement of the permanent closure; 30 individuals were needed to capture behavioral heterogeneity. The dry-season (May–January) and wet-season (February–April) time-area closures opened 20% and 25% more planning units to fishing, respectively, compared with the permanent closure with boundaries fixed in space and time. Our results show that telemetry has the potential to inform and improve spatial management of mobile species and that the temporal component of tracking data can be incorporated into prioritizations to reduce possible impacts of spatial closures on established fisheries.     

Urbanized landscapes favored by fig-eating birds increase invasive but not native juvenile strangler fig abundance

Propagule pressure can determine the success or failure of invasive plant range expansion. Range expansion takes place at large spatial scales, often encompassing many types of land cover, yet the effect of landscape context on propagule pressure remains largely unknown. Many studies have reported a positive correlation between invasive plant abundance and human land use; increased propagule pressure in these landscapes may be responsible for this correlation. We tested the hypothesis that increased rates of seed dispersal by fig-eating birds, which are more common in urban habitats, result in an increase in invasive strangler fig abundance in landscapes dominated by human land use. We quantified abundance of an invasive species (Ficus microcarpa) and a native species (F. aurea) of strangler fig in plots spanning the entire range of human land use in South Florida, USA, from urban parking lots to native forest. We then compared models that predicted juvenile fig abundance based on distance to adult fig seed sources and fig-eating bird habitat quality with models that lacked one or both of these terms. The best model for juvenile invasive fig abundance included both distance to adult and fig-eating bird habitat terms, suggesting that landscape effects on invasive fig abundance are mediated by seed-dispersing birds. In contrast, the best model for juvenile native fig abundance included only presence/absence of adults, suggesting that distance from individual adult trees may have less effect on seed limitation for a native species compared to an invasive species undergoing range expansion. However, models for both species included significant effects of adult seed sources, implying that juvenile abundance is limited by seed arrival. This result was corroborated by a seed addition experiment that indicated that both native and invasive strangler figs were strongly seed limited. Understanding how landscape context affects the mechanisms of plant invasion may lead to better management techniques. Our results suggest that prioritizing removal of adult trees in sites with high fig-eating bird habitat may be the most effective method to control F. microcarpa abundance.