Abstract. The study of population dynamics requires unbiased, precise estimates of abundance and vital rates that account for the demographic structure inherent in all wildlife and plant populations. Traditionally, these estimates have only been available through approaches that rely on intensive mark-recapture data. We extended recently developed Nmixture models to demonstrate how demographic parameters and abundance can be estimated for structured populations using only stage-structured count data. Our modeling framework can be used to make reliable inferences on abundance as well as recruitment, immigration, stage-specific survival, and detection rates during sampling. We present a range of simulations to illustrate the data requirements, including the number of years and locations necessary for accurate and precise parameter estimates. We apply our modeling framework to a population of northern dusky salamanders (Desmognathus fuscus) in the mid-Atlantic region (USA) and find that the population is unexpectedly declining. Our approach represents a valuable advance in the estimation of population dynamics using multistate data from unmarked individuals and should additionally be useful in the development of integrated models that combine data from intensive (e.g., mark-recapture) and extensive (e.g., counts) data sources.
Linear and heterogeneous habitat makes headwater stream networks an ideal ecosystem in which to test the influence of environmental factors on spatial genetic patterns of obligatory aquatic species. We investigated fine-scale population structure and influence of stream habitat on individual-level genetic differentiation in brook trout (Salvelinus fontinalis) by genotyping eight microsatellite loci in 740 individuals in two headwater channel networks (7.7 and 4.4 km) in Connecticut, USA. A weak but statistically significant isolation-by-distance pattern was common in both sites. In the field, many tagged individuals were recaptured in the same 50-m reaches within a single field season (summer to fall). One study site was characterized with a hierarchical population structure, where seasonal barriers (natural falls of 1.5-2.5 m in height during summer base-flow condition) greatly reduced gene flow and perceptible spatial patterns emerged because of the presence of tributaries, each with a group of genetically distinguishable individuals. Genetic differentiation increased when pairs of individuals were separated by high stream gradient (steep channel slope) or warm stream temperature in this site, although the evidence of their influence was equivocal. In a second site, evidence for genetic clusters was weak at best, but genetic differentiation between individuals was positively correlated with number of tributary confluences. We concluded that the population-level movement of brook trout was limited in the study headwater stream networks, resulting in the fine-scale population structure (genetic clusters and clines) even at distances of a few kilometres, and gene flow was mitigated by 'riverscape' variables, particularly by physical barriers, waterway distance (i.e. isolation-by-distance) and the presence of tributaries.
Climate change affects seasonal weather patterns, but little is known about the relative importance of seasonal weather patterns on animal population vital rates. Even when such information exists, data are typically only available from intensive fieldwork (e.g., mark-recapture studies) at a limited spatial extent. Here, we investigated effects of seasonal air temperature and precipitation (fall, winter, and spring) on survival and recruitment of brook trout (Salvelinus fontinalis) at a broad spatial scale using a novel stage-structured population model. The data were a 15-year record of brook trout abundance from 72 sites distributed across a 170-km-long mountain range in Shenandoah National Park, Virginia, USA. Population vital rates responded differently to weather and site-specific conditions. Specifically, young-of-year survival was most strongly affected by spring temperature, adult survival by elevation and per-capita recruitment by winter precipitation. Low fall precipitation and high winter precipitation, the latter of which is predicted to increase under climate change for the study region, had the strongest negative effects on trout populations. Simulations show that trout abundance could be greatly reduced under constant high winter precipitation, consistent with the expected effects of gravel-scouring flows on eggs and newly hatched individuals. However, high-elevation sites would be less vulnerable to local extinction because they supported higher adult survival. Furthermore, the majority of brook trout populations are projected to persist if high winter precipitation occurs only intermittently (≤3 of 5 years) due to density-dependent recruitment. Variable drivers of vital rates should be commonly found in animal populations characterized by ontogenetic changes in habitat, and such stage-structured effects may increase population persistence to changing climate by not affecting all life stages simultaneously. Yet, our results also demonstrate that weather patterns during seemingly less consequential seasons (e.g., winter precipitation) can have major impacts on animal population dynamics.
1. Defining functional connectivity between habitats in spatially heterogeneous landscapes is a particular challenge for small-bodied aquatic species. Traditional approaches (e.g. mark-recapture studies) preclude an assessment of animal movement over the life cycle (birth to reproduction), and movement of individuals may not represent the degree of gene movement for fecund species. 2. We investigated the degree of habitat connectivity (defined as the exchange of individuals and genes between mainstem and tributary habitats) in a stream brook trout (Salvelinus fontinalis) population using mark-recapture [passive integrated transponder (PIT) tags], stationary PIT-tag antennae and genetic pedigree data collected over 4 years (3425 marked individuals). We hypothesised that: (i) a combination of these data would reveal higher estimates of animal movement over the life cycle (within a generation), relative to more temporally confined approaches, and (ii) movement estimates of individuals within a generation would differ from between-generation movement of genes because of spatial variation in reproductive success associated with high fecundity of this species. 3. Over half of PIT-tagged fish (juveniles and adults) were recaptured within 20 m during periodic sampling, indicating restricted movement. However, continuous monitoring with stationary PIT-tag antennae revealed distinct peaks in trout movements in June and October-November, and sibship data inferred post-emergence movements of young-of-year trout that were too small to be tagged physically. A combination of these methods showed that a moderate portion of individuals (28-33%) moved between mainstem and tributary habitats over their life cycle. 4. Patterns of reproductive success varied spatially and temporally. The importance of tributaries as spawning habitat was discovered by accounting for reproductive history. When individuals born in the mainstem reproduced successfully, over 50% of their surviving offspring were inferred to have been born in tributaries. This high rate of gene movement to tributaries was cryptic, and it would have been missed by estimates based only on movement of individuals. 5. This study highlighted the importance of characterising animal movement over the life cycle for inferring habitat connectivity accurately. Such movements of individuals can contribute to substantial gene movements in a fecund species characterised by high variation in reproductive success.
Summary Abundance of the young‐of‐the‐year (YOY) fish can vary greatly among years and it may be driven by several key biological processes (i.e. adult spawning, egg survival and fry survival) that span several months. However, the relative influence of seasonal weather patterns on YOY abundance is poorly understood. We assessed the importance of seasonal air temperature (a surrogate for stream temperature) and precipitation (a surrogate for stream flow) on brook trout (Salvelinus fontinalis) YOY summer abundance using a 29‐year data set from 115 sites in Shenandoah National Park, Virginia, U.S.A. We used a Bayesian hierarchical model that allowed the effect of seasonal weather covariates to vary among sites and accounted for imperfect detection of individuals. Summer YOY abundance was affected by preceding seasonal air temperature and precipitation, and these regional‐scale drivers led to spatial synchrony in YOY abundance dynamics across the 170‐km‐long study area. Mean winter precipitation had the greatest effect on YOY abundance and the relationship was negative. Mean autumn precipitation, and winter and spring temperature had significantly positive effects on YOY abundance, and mean autumn temperature had a significant negative effect. In addition, the effect of summer precipitation differed along a latitudinal gradient, with YOY abundance at more northern sites being more responsive to inter‐annual variation in summer precipitation. Strong YOY years resulted in high abundance of adults (>age 1 + fish) in the subsequent year at more than half of sites. However, higher adult abundance did not result in higher YOY abundance in the subsequent year at any of the study sites (i.e. no positive stock–recruitment relationship). Our results indicate that YOY abundance is a key driver of brook trout population dynamics that is mediated by seasonal weather patterns. A reliable assessment of climate change impacts on brook trout needs to account for how alternations in seasonal weather patterns impact YOY abundance and how such relationships may differ across the range of brook trout distribution.
Previous studies of climate change impacts on stream fish distributions commonly project the potential patterns of habitat loss and fragmentation due to elevated stream temperatures at a broad spatial scale (e.g. across regions or an entire species range). However, these studies may overlook potential heterogeneity in climate change vulnerability within local stream networks. We examined fine-scale stream temperature patterns in two headwater brook trout Salvelinus fontinalis stream networks (7.7 and 4.4 km) in Connecticut, USA, by placing a combined total of 36 pairs of stream and air temperature loggers that were approximately 300 m apart from each other. Data were collected hourly from March to October 2010. The summer of 2010 was hot (the second hottest on record) and had well below average precipitation, but stream temperature was comparable with those of previous 2 years because streamflow was dominated by groundwater during base-flow conditions. Nonlinear regression models revealed stream temperature variation within local stream networks, particularly during warmest hours of the day (i.e. late afternoon to evening) during summer. Thermal variability was primarily observed between stream segments, versus within a stream segment (i.e. from confluence to confluence). Several cold tributaries were identified in which stream temperature was much less responsive to air temperature. Our findings suggested that regional models of stream temperature would not fully capture thermal variation at the local scale and may misrepresent thermal resilience of stream networks. Groundwater appeared to play a major role in creating the fine-scale spatial thermal variation, and characterizing this thermal variation is needed for assessing climate change impacts on headwater species accurately.
The electrofishing distance needed to estimate fish species richness at the stream or river reach scale is an important question in fisheries science. This distance is governed by the shape of the species accumulation curve, which, in turn, is influenced by a combination of factors, including the number of species, their overall abundances, habitat associations, the efficiency of the sampling method, and the occurrence of rare species. In this study we document the influence of rare species on the species accumulation curves from stream and river sites in data sets from five dispersed regions of the USA. Spatial discontinuity (i.e., a noncontinuous distribution within reaches) was observed in four of the five data sets, and the four data sets contained numerically rare species represented by one or two individuals (termed singletons and doubletons, respectively). Numerically rare species were typically proportionately rare (i.e., ,1% of the total number of individuals captured), but proportionately rare species were not always numerically rare and were dependent on the total number of fish captured. Species richness asymptotes were reached at shorter electrofishing distances when singletons and doubletons were removed. The number of singletons and doubletons in the samples remained relatively constant with increasing sampling effort (i.e., sampling distance and total abundance). Simulation modeling indicated that individual aggregation within species was not a plausible reason for spatially discontinuous species distributions. When accurately detecting the presence of species is a sampling goal, the presence and prevalence of numerically rare species may need to be considered in determining sampling protocols.
Brook trout Salvelinus fontinalis populations have declined in much of the native range in eastern North America and populations are typically relegated to small headwater streams in Connecticut, USA. We used sibship reconstruction to infer mating systems, dispersal and effective population size of resident (non-anadromous) brook trout in two headwater stream channel networks in Connecticut. Brook trout were captured via backpack electrofishing using spatially continuous sampling in the two headwaters (channel network lengths of 4.4 and 7.7 km). Eight microsatellite loci were genotyped in a total of 740 individuals (80-140 mm) subsampled in a stratified random design from all 50 m-reaches in which trout were captured. Sibship reconstruction indicated that males and females were both mostly polygamous although single pair matings were also inferred. Breeder sex ratio was inferred to be nearly 1:1. Few large-sized fullsib families ([3 individuals) were inferred and the majority of individuals were inferred to have no fullsibs among those fish genotyped (family size = 1). The median stream channel distance between pairs of individuals belonging to the same large-sized fullsib families ([3 individuals) was 100 m (range: 0-1,850 m) and 250 m (range: 0-2,350 m) in the two study sites, indicating limited dispersal at least for the size class of individuals analyzed. Using a sibship assignment method, the effective population size for the two streams was estimated at 91 (95%CI: 67-123) and 210 (95%CI: 172-259), corresponding to the ratio of effectiveto-census population size of 0.06 and 0.12, respectively. Both-sex polygamy, low variation in reproductive success, and a balanced sex ratio may help maintain genetic diversity of brook trout populations with small breeder sizes persisting in headwater channel networks.
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