Patterns of natural mortality in stream‐living brown trout (Salmo trutta)

1. We tested the hypothesis that lifetime mortality patterns and their corresponding rates and causal factors differ among populations of stream‐living salmonids. To this end, we examined the lifetime mortality patterns of several successive cohorts of two stream‐living brown trout (Salmo trutta) populations in Spain and Denmark. 2. In the southern population, we observed a consistent two‐phase pattern, in which mortality was negligible during the first half of the lifetime and severe during the rest of the lifetime. In contrast, the northern population demonstrated a three‐phase pattern with an earlier phase varying from negligible to severe, followed by a second stage of weak mortality, and lastly by a third life stage of severe mortality. 3. Despite substantial differences in the mortality patterns between the two populations, the combined effect of recruitment (as a proxy of the density‐dependent processes occurring during the lifetime) and mean body mass (as a proxy of growth experienced by individuals in a given cohort) explained c. 89% of the total lifetime mortality rates across cohorts and populations. 4. A comparison with other published data on populations of stream‐living brown trout within its native range highlighted lifetime mortality patterns of one, two, three and four phases, but also suggested that common patterns may occur in populations that experience similar individual growth and population density.     

Density‐dependent mortality in adults,but not juveniles,of stream‐resident brown trout (Salmo trutta)

1. This study investigates when and where density dependence operates on the mortality rates of stream‐resident brown trout Salmo trutta. To this aim, I explored populations in habitats of different quality containing high, low or intermediate densities over broad scales of space and time. The study is based on census data of 170 cohorts quantified from recruitment to the total disappearance at 12 sites in four contrasting tributaries of the Rio Esva drainage (north‐western Spain), over the years 1986–2007. 2. Log₁₀‐transformed survivor density over time highlighted a consistent pattern for the 170 cohorts characterised by the occurrence of only two life stages. An early stage starts at recruitment, lasts about half the lifetime and shows no or negligible mortality. A threshold time at 425–620 days after emergence preceded a second stage of continuous and constant mortality until the final disappearance of the cohorts. Consequently, in all scenarios, mortality only occurred in the adult component and no effect of season, year, age‐class and/or reproductive stage was detected. 3. Substantial spatial and temporal variations typified both recruitment (range R = 0.01–1.62 ind m^?2) and adults’ mortality rates (range Z = 0.03–0.38 day^?1). Nested anova s revealed strong effects of site and year on both recruitment and mortality with sites interspersed along the stream gradients where recruitment and mortality were typically high relative to other sites located either nearby in the same stream or distant in another stream, where both recruitment and mortality rates were typically low or intermediate. 4. Adult mortality rates plotted against recruitment for the 170 cohorts pooled revealed a continuous, positive power relationship that explained 45.3% of variation in mortality rates over the whole range of recruitment values. Similarly, highly significant power relationships were elucidated for site‐specific mortality rates averaged across years and for annual‐specific mortality rates averaged across sites against the corresponding mean recruitment averaged across years and sites, respectively. These relationships support the hypothesis that the operation of density dependence is scale independent and context independent but operates in a continuous manner across all scenarios examined. 5. A chronic effect of density dependence on adult losses induces temporally persistent populations maintained by a low number of spawners. Apparently, the operation of density dependence adjusts the number of spawners to the availability of rearing and spawning habitat. This dynamic process may also help to explain the small effective population size (Ne) recently documented by genetic studies of stream‐living brown trout and other salmonids.     

Dynamics of a harvested moose population in a variable environment

1. Population size, calves per female, female mean age and adult sex ratio of a moose ( Alces alces ) population in Vefsn, northern Norway were reconstructed from 1967 to 1993 using cohort analysis and catch-at-age data from 96% (6752) of all individuals harvested.
2. The dynamics of the population were influenced mainly by density-dependent harvesting, stochastic variation in climate and intrinsic variation in the age-structure of the female segment of the population.
3. A time delay in the assignment of hunting permits in relation to population size increased fluctuations in population size.
4. Selective harvesting of calves and yearlings increased the mean age of adult females in the population, and, because fecundity in moose is strongly age-specific, the number of calves per female concordantly increased. However, after years with high recruitment, the adult mean age decreased as large cohorts entered the adult age-groups. This age-structure effect generated cycles in the rate of recruitment to the population and fluctuations introduced time-lags in the population dynamics.
5. An inverse relationship between recruitment rate and population density, mediated by a density-dependent decrease in female body condition, could potentially have constituted a regulatory mechanism in the dynamics of the population, but this effect was counteracted by a density-dependent increase in the mean age of adult females.
6. Stochastic variation in winter snow depth and summer temperature had delayed effects on recruitment rate and in turn population growth rate, apparently through effects on female body condition before conception.     

Higher treefall rates on slopes and waterlogged soils result in lower stand biomass and productivity in a tropical rain forest

1.  Relationships between tropical rain forest biomass and environmental factors have been determined at regional scales, e.g. the Amazon Basin, but the reasons for the high variability in forest biomass at local scales are poorly understood. Interactions between topography, soil properties, tree growth and mortality rates, and treefalls are a likely reason for this variability.
2.  We used repeated measurements of permanent plots in lowland rain forest in French Guiana to evaluate these relationships. The plots sampled topographic gradients from hilltops to slopes to bottomlands, with accompanying variation in soil waterlogging along these gradients. Biomass was calculated for >175 tree species in the plots, along with biomass productivity and recruitment rates. Mortality was determined as standing dead and treefalls.
3.  Treefall rates were twice as high in bottomlands as on hilltops, and tree recruitment rates, radial growth rates and the abundance of light-demanding tree species were also higher.
4.  In the bottomlands, the mean wood density was 10% lower than on hilltops, the basal area 29% lower and the height:diameter ratio of trees was lower, collectively resulting in a total woody biomass that was 43% lower in bottomlands than on hilltops.
5.  Biomass productivity was 9% lower in bottomlands than on hilltops, even though soil Olsen P concentrations were higher in bottomlands.
6.   Synthesis . Along a topographic gradient from hilltops to bottomlands there were higher rates of treefall, which decreased the stand basal area and favoured lower allocation to height growth and recruitment of light-demanding species with low wood density. The resultant large variation in tree biomass along the gradient shows the importance of determining site characteristics and including these characteristics when scaling up biomass estimates from stand to local or regional scales.     

Recruitment rates in forest plots: Gf estimates using growth rates and size distributions

1 In censuses of tree populations in permanent plots, short census intervals and small population size lead to uncertainty in the observed recruitment rate of a minimum size. Increasing the census interval, however, underestimates the rate because of unrecorded 'recruit and die' events.
2 We propose a new Gf procedure for estimation of recruitment rates. Recruitment rate per area is obtained by multiplication of the density in the smallest size class (f) and the average size growth rate in that class (G) divided by the width of the class. This procedure is valid when the size distribution of the population examined is continuous with size.
3 When tree size structure is negative‐exponentially distributed, as is often the case in natural rain forest populations, the Gf estimate of the recruitment rate for a given size class was least biased close to the midpoint size of this class.
4 Gf estimates agreed well with census estimates of recruitment rate from permanent plots in rain forests. A tendency for Gf estimates to be larger than census estimates disappeared when census estimates were corrected for mortality after recruitment.
5 The effects of plot size, census interval and variation in growth rate on estimates of recruitment rate were simulated using model populations. Small plot size caused substantially more among‐plot deviation for the census count of recruitment events than for the Gf estimate. The census recruitment rate also showed larger variation among plots for shorter intervals than the Gf estimate, which was independent of census interval. The Gf estimates were therefore more accurate than census counts in many situations. More than several tens of trees were needed in a size class to allow a reliable Gf estimates.     

Culling Versus Density Effects in Management of a Deer Population

Abstract: Wildlife managers often manipulate hunting regulations to control deer populations. However, few empirical studies have examined the level of hunting effort (hunter-days) required to limit population growth and demographic effects through harvesting of females. Moreover, the relative importance of density effects on population growth has not been quantified. We reconstructed a sika deer [Cervus nippon] population over a period of 12 years (1990–2001) using age- and sex-specific harvest data. Using cohort analysis, we analyzed population dynamics, focusing on 1) the relationship between hunting effort and hunting-induced mortality rate, 2) relative contributions of hunting mortality and recruitment of yearlings to annual changes in population growth rate, and 3) annual variation in recruitment rate. Population size increased until 1998 and declined thereafter. The population growth rate changed more in response to annual changes in recruitment rate than hunting mortality rate. Temporal variation in recruitment rate was not controlled by birth rate alone; direct density dependence, intensities of hunting mortality for fawns, and for females (≥2 yr of age), which accounted for the fawn survival rate, were required as factors to explain temporal variation. Density effects on the recruitment rate were not strong enough to regulate the population within the study period; high hunting mortality, with intensive female harvesting, was necessary to prevent population growth. Hunting effort was a good predictor of the hunting mortality rate, and female harvest had a negative effect on the recruitment rate through fawn survival. We suggest that >3,500 hunter-days and prioritization of female harvesting are required to prevent increases in this deer population.     

A study of the dynamics of a natural population of the freshwater crayfish, Austropotamobius pallipes

SUMMARY. 1. Population size, survival and recruitment were estimated from mark—recapture data at frequent intervals.
2. Two types of mortality were identified: (1) moult-associated—an intense but short duration mortality, and (2) overwinter mortality, which operated at a lower estimated weekly rate but over some 30 weeks.
3. Population size reached a mid-summer peak as recruitment into the catchable size classes occurred. Thereafter numbers declined slowly to the end of the growth season, followed by an overwinter decline of some 50% of total.
4. Annual production was estimated at 196.5 kg wet wt ha⁻¹ in 1977, 116.8 kg wet wt ha⁻¹ in 1978 and 87.79 kg wet wt ha⁻¹ in 1979.     

Inter-biome comparison of factors controlling stream metabolism

1. We studied whole-ecosystem metabolism in eight streams from several biomes in North America to identify controls on the rate of stream metabolism over a large geographic range. The streams studied had climates ranging from tropical to cool-temperate and from humid to arid and were all relatively uninfluenced by human disturbances.
2. Rates of gross primary production (GPP), ecosystem respiration (R) and net ecosystem production (NEP) were determined using the open-system, two-station diurnal oxygen change method.
3. Three general patterns in metabolism were evident among streams: (1) relatively high GPP with positive NEP (i.e. net oxygen production) in early afternoon, (2) moderate primary production with a distinct peak in GPP during daylight but negative NEP at all times and (3) little or no evidence of GPP during daylight and a relatively constant and negative NEP over the entire day.
4. Gross primary production was most strongly correlated with photosynthetically active radiation (PAR). A multiple regression model that included log PAR and stream water soluble reactive phosphorus (SRP) concentration explained 90% of the variation in log GPP.
5. Ecosystem respiration was significantly correlated with SRP concentration and size of the transient storage zone and, together, these factors explained 73% of the variation in R. The rate of R was poorly correlated with the rate of GPP.
6. Net ecosystem production was significantly correlated only with PAR, with 53% of the variation in log NEP explained by log PAR. Only Sycamore Creek, a desert stream in Arizona, had positive NEP (GPP: R  > 1), supporting the idea that streams are generally net sinks rather than net sources of organic matter.
7. Our results suggest that light, phosphorus concentration and channel hydraulics are important controls on the rate of ecosystem metabolism in streams over very extensive geographic areas.     

Assessment of the effects of environmental change on the performance and density of Bistorta vivipara: the use of multivariate analysis and experimental manipulation

1 Studies of plant performance in relation to local variability in environmental conditions can be used to predict the responses of species to environmental change.
2 We used redundancy analysis to study how interpopulation variation in average plant weight, flower and bulbil number, bulbil weight and population density among 28 populations of Bistorta vivipara was related to variation in 15 environmental factors. We also examined the responses of plants to a 4-year experimental increase in temperature.
3 Significant differences in average plant performance were found between populations. Variance partitioning of the response data showed that environmental factors explained 45% of the variation in plant performance and density between populations, whereas variation due to transect position was small (10.8%). Soil pH, altitude and season length were the most influential of the environmental variables, and explained 23%, 21% and 14%, respectively, of the variation. Plant performance was in general negatively correlated with these variables, whereas plant density increased along the pH and altitude gradients, suggesting that environmental factors associated with elevation (temperature and vegetation cover) and pH (soil fertility) had opposing effects on individual performance and density. Season length was highly important for average bulbil weight.
4 Plants growing in open top chambers had significantly enhanced growth and produced significantly heavier bulbils than those in control plots, whereas flower and bulbil number were unaffected by experimentally increased temperatures. Plant density was equal for warmed and control plots.
5 Although warming may increase bulbil weight and plant size in B. vivipara , the response to climate change is complicated by the fact that population densities may decrease due to intensified competition for light caused by a denser vegetation cover.     

Population dynamics of the gastropod Theodoxus jordani (Sowerby) in the Shatt Al-Arab river

SUMMARY. 1. Measurements of the population density of Theodoxus jordani were made for the period July 1980 until June 1981. Numbers showed a well-defined pattern of fluctuation and varied between 390 in February and 1804 snails m⁻² in May after recruitment.
2. The population age structure revealed that the snail was semelparous, and that one generation dominated the population for most of the year.
3. The breeding season was long. Snails laid eggs from early December to the end of April. Eggs were laid in protective capsules and one snail developed per capsule. Total population natality estimated from field samples was 9813 snails m⁻².
4. Age specific mortality was high immediately after recruitment, low during the juvenile stages, and increased sharply at the adult stage with the onset of reproduction.     

Population studies of the beech leaf mining weevil (Rhynchaenus fagi) in Ireland and Scotland

Abstract. 1. Population budgets for Rhynchaenus fagi, based for the most part on samples of beech leaves taken in June, were constructed for a 10-year period (Northern Ireland) and a 3-year period (Scottish borders).
2. There were years of sustained population growth and sustained decline, but not enough evidence to suggest a cycle. Similar population levels were found at both sites in all but one year.
3. Key factors were associated with the adult stage, although egg mortality may also make a significant contribution to population change.
4. Unknown sources of mortality embodied in the residual mortality were also suspected of acting in a density dependent manner, compensating for variation in parasitism.
5. Death rate of larvae within mines was related to May temperature in Northern Ireland, but not so clearly in Scotland.     

Environmental constraints on oviposition limit egg supply of a stream insect at multiple scales

Species with complex life cycles pose challenges for understanding what processes regulate population densities, especially if some life stages disperse. Most studies of such animals that are thought to be recruitment limited focus on the idea that juvenile mortality limits the density of recruits (and hence population density), fewer consider the possibility that egg supply may be important. For species that oviposit on specific substrata, environmental constraints on oviposition sites may limit egg supply. Female mayflies in the genus Baetis lay egg masses on the underside of stream rocks that emerge above the water’s surface. We tested the hypothesis that egg mass densities are constrained by emergent rock densities within and between streams, by counting egg masses on emergent rocks. All emergent rocks were counted along 1-km lengths of four streams, revealing significant variation in emergent rock density within streams and a more than three-fold difference between streams. In each stream, egg mass density increased with the density of emergent rocks in 30-m stretches. We used regression equations describing these small-scale relationships, coupled with the large-scale spatial variation of emergent rocks, to estimate egg mass densities for each 1-km stream length, a scale relevant to population processes. Scaled estimates were positively associated with emergent rock density and provided better estimates than methods that ignored environmental variation. Egg mass crowding was inversely related to emergent rock density at the stream scale, a pattern consistent with the idea that oviposition substrata were in short supply in streams with few emergent rocks, but crowding did not compensate entirely for differences in emergent rock densities. The notion that egg supply, not larval mortality, may limit population density is an unusual perspective for stream insects. Environmental constraints on egg supply may be widespread among other species with specialised oviposition behaviours.     

Regulation of damseifly populations: the effects of larval density on larval urvival, development rate and size in the field

SUMMARY. 1. The popuhttion density of Coenagrion pttella larvae was monitored in five populations, and of Ischntira elegans in two populations, between October 1982 and May 1983.
2. There was no measurable mortality of larvae over winter and no larval growth until April. Larvae in high density populations were smaller than those in low density populations and were more likely to have a semi- voltine life history.
3. The population density of C. ptiella was also monitored (more frequently) in two populations with differenl initial densities between July and November 1983. In the high density population there was a constant rate of larval mortality, while in the low density population there was no detectable larval mortality, indicating that larval mortality may be density dependent. Larvae in the high density population were again smaller, and more likely to be semi-voitine, than those in the low density population.
4. The role of density dependent larval growth, development and mortality in the regulation of damseifly populations is discussed.     

The effect of growth pattern, sampling interval and number of size classes on benthic invertebrate production estimated by the size-frequency method

SUMMARY. 1. A hypothetical leech population with known initial density, initial weight, final weight and cohort production interval (CPI) was established. Production estimated by the size-frequency method for various growth patterns, mortalities, number of samples per CPI and number of size classes was compared with actual production estimated from daily growth and mortality by the increment-summation method. The population had either perfectly continuous reproduction or a perfectly synchronous cohort.
2. When size-classes were delimited in order to equalize the time spent in each size class, the deviations from actual production increased with decreasing number of size-classes and increasing mortality. For a population with perfectly continuous reproduction, production was only overestimated by 32% with an extreme mortality of 2.0% day⁻¹ and three size-classes. For a perfectly synchronous cohort, production was either underestimated or overestimated, depending on the first day of sampling. The deviations from actual production increased considerably with decreasing number of size-classes, increasing mortality and decreasing number of samples per CPI.
3. Differences between actual and assumed growth patterns may give underestimates or overestimates of more than one order of magnitude at high mortalities and few size-classes. It is concluded that knowing the actual growth pattern, the size frequency method will give realistic estimates of production in cases when normal cohort methods cannot be used. The estimate can be improved significantly by increasing the number of size classes and the number of samples per CPI.     

The effects of population density and lake characteristics on growth and size structure of brook trout Salvelinus fontinalis (Mitchill, 1815) in boreal forest lakes in Canada

This paper examines the effect of lake characteristics on population density and how this variation affects growth, mortality and population size structure of brook trout, Salvelinus fontinalis. The study was conducted on 17 recreationally fished, reproductively isolated boreal forest lakes in Newfoundland, Canada from 1993 to 2000. A standardized sampling program, the Fyke Littoral Index Netting program (FLIN) was used to collected data that describes brook trout population parameters and life history attributes. Regression analyses showed significant relationships between fish density and biomass and characteristics of the lakes. Variation in fish density and biomass was explained by lake surface area and littoral habitat area. Significant relationships were found when growth, mortality and size structure were regressed against density. The proportional stock distribution and theoretical maximum size of brook trout were negatively related to density, and natural mortality was positively related to density. The largest maximum length and highest proportional stock densities occurred at brook trout densities of less than 30 fish/ha. In general, the higher the proportion of littoral habitat area the higher the densities of brook trout, which correspondingly had important effects on growth, natural mortality and size structure of the brook trout populations. This information is critical to the development of management strategies aimed at altering size distribution to produce specific fisheries management outcomes.     

Recruitment density can determine adult morphology and fecundity in the barnacle, Semibalanus balanoides

Although consequences of the settlement preference of larvae have been well documented, the consequence of these settlement choices on subsequent mortality, morphology and fecundity has been little studied. The aim of this study was to determine the relationship between recruit and adult density and to determine the effect of recruitment on adult morphology and egg tissue mass. This study follows 48,718 barnacles (Semibalanus balanoides) from recruitment at the end of the settling season to reproductively mature adults at a field site in the Clyde Sea (UK). Overall survivorship of the recruits to adulthood was 8.5%, although survivorship was up to 42% on low density settlement panels. In low density colonies (< 10 recruits cm-2), recruitment density was related to adult density (P < 0.001), whereas no relationship was found for higher density colonies. A shell morphology index measured at adulthood was related to recruitment density for low density recruited colonies (P < 0.001) but not high density colonies. Using ANCOVA, variations between the colonies in shell and egg tissue mass were not explained by mass of somatic tissue. However, egg mass was explained by recruitment density (P < 0.01). These results show that adult density is not a reliable indicator of the previous population density of the colony. Moreover, there are marked differences in population development between colonies with high and low recruit densities in terms of impact upon shell morphology and egg production. The dynamics that operate between recruits at the end of the settlement season and sexually mature adults to create the patterns elucidated in this paper, and other literature, remain unclear.     

Contributions of forage competition, harvest, and climate fluctuation to changes in population growth of northern white-tailed deer

Recently there has been considerable interest in determining the relative roles of endogenous (density-dependent) and exogenous (density-independent) factors in driving the population dynamics of free-ranging ungulates. We used time-series analysis to estimate the relative contributions of density-dependent forage competition, climatic fluctuation, and harvesting on the population dynamics of white-tailed deer (Odocoileus virginianus) in Nova Scotia, Canada, from 1983 to 2000. A model incorporating the population density 2 years previous, an interaction term for the harvest of females and population density 2 years previous, and the total snowfall during the previous 2 winters explained 80% of the variation in inter-annual population growth rate. Natality of adult females was negatively related to deer density during the present winter, whereas that of yearlings may have been correlated with the snowfall of three winters previous. Natality of fawns was related to deer density and total snowfall during the previous winter. Coyotes (Canis latrans) prey extensively on deer fawns in northeastern North America and the annual harvest of snowshoe hares (Lepus americanus), the major alternate prey of coyotes, explained 48% of the inter-annual variation in fawn recruitment. The proportions of fawn, yearling, and adult deer suffering from severe malnutrition during late winter were all correlated with deer density during the present winter. We conclude that the limiting effects of winter weather on over-winter survival of deer may be cumulative over two consecutive winters. During the late 1980s, density dependence and winter severity acted in concert to effect substantial declines in deer population growth both by effecting winter losses directly and by exacerbating predation by coyotes. During this period liberal harvesting did not relieve density-dependent forage competition and probably accelerated the decline.     

Temporal changes in the strength of density-dependent mortality and growth in intertidal barnacles

1. In demographically open marine systems, the extent to which density-dependent processes in the benthic adult phase are required for population persistence is unclear. At one extreme, represented by the recruitment limitation hypothesis, larval supply may be insufficient for the total population size to reach a carrying capacity and density-independent mortality predominates. At the opposite extreme, populations are saturated and density-dependent mortality is sufficiently strong to reshape patterns established at settlement. 2. We examined temporal variation in the way density-independent and density-dependent mortality interact in a typical sessile marine benthic invertebrate, the acorn barnacle Semibalanus balanoides (L.), over a 2-year period. 3. Recruitment was manipulated at two high recruitment sites in north Wales, UK to produce recruit densities covering the range naturally found in this species. Following manipulation, fixed quadrats were monitored using digital photography and temporal changes in mortality and growth rate were examined. 4. Over a 2-year period there was a clear, spatially consistent, over-compensatory relationship between the density of recruits and adult abundance indicating strong density-dependent mortality. The strength of density dependence intensified with increasing recruitment. 5. Density-dependent mortality did not operate consistently over the study period. It only operated in the early part of the benthic phase, but the pattern of adult abundance generated was maintained throughout the whole 2-year period. Thus, early life-history processes dictated adult population abundance and dynamics. 6. Examination of the natural recruitment regime in the area of study indicated that both positive and negative effects of recruitment will occur over scales varying from kilometres to metres.