Bifurcation theory, adaptive dynamics and dynamic energy budget-structured populations of iteroparous species

In this paper, we describe a technique to evaluate the evolutionary dynamics of the timing of spawning for iteroparous species. The life cycle of the species consists of three life stages, embryonic, juvenile and adult whereby the transitions of life stages (gametogenesis, birth and maturation) occur at species-specific sizes. The dynamics of the population is studied in a semi-chemostat environment where the inflowing food concentration is periodic (annual). A dynamic energy budget-based continuous-time model is used to describe the uptake of the food, storage in reserves and allocation of the energy to growth, maintenance, development (embryos, juveniles) and reproduction (adults). A discrete-event process is used for modelling reproduction. At a fixed spawning date of the year, the reproduction buffer is emptied and a new cohort is formed by eggs with a fixed size and energy content. The population consists of cohorts: for each year one consisting of individuals with the same age which die after their last reproduction event. The resulting mathematical model is a finite-dimensional set of ordinary differential equations with fixed 1-year periodic boundary conditions yielding a stroboscopic map. We will study the evolutionary development of the population using the adaptive dynamics approach. The trait is the timing of spawning. Pairwise and mutual invasibility plots are calculated using bifurcation analysis of the stroboscopic map. The evolutionary singular strategy value belonging to the evolutionary endpoint for the trait allows for an interpretation of the reproduction strategy of the population. In a case study, parameter values from the literature for the bivalve Macoma balthica are used.     

花鼠种群年龄和繁殖的初步研究

1991年3-10月在山西省中条山区猎获花鼠202只。以眼晶体干重为指标划分出4个年龄组,分析了种群年龄组成的季节变化。该鼠每年繁殖1次, 3月下旬-1月中旬怀孕,产仔期在4月中旬-5月中旬,胎仔数5.08±0.16只, 6月上旬幼鼠出现。幼鼠于翌年春季出蛰后参加繁殖。此外,还将晶体干重、齿高指数和臼齿磨损程度3种年龄指标所划分的结果作了比较,也证实体重、体长和尾长不能用于该鼠的年龄鉴定。     

The cause of extreme year class variation in a population of roach, Rutilus rutilus L., from a eutrophic lake in southern England

Investigations into the biology of the roach population at Slapton Ley, Devon, U.K. were carried out between October 1982 and December 1984 and data collected from the lake since 1977 have been re-analysed to assist interpretation. The roach population has been characterized in recent years by extreme year class variation and a tendency for year classes to alternate in strength. The survival of the roach was very poor, and the spawning population therefore consisted largely of 2-and some 3-year-old fish. Consequently, the size of the spawning stock was small and dependent on the strengths of the year classes that it contained. Recruitment to many year classes was probably limited by the size of the spawning stock. The occurrence of a poor year class in the population would therefore set up an autonomous cycle of poor recruitment to alternate year classes in a manner similar to many tropical marine fisheries. Observation of other British roach populations showed that a short life-span may be associated with conditions that promote very rapid growth rates. The growth rate of individual roach at Slapton was found to be one of the fastest in Britain, and this was apparently due to the very high productivity of Slapton Ley together with the persistence of the roach population at very low densities.     

The migration of spawning stocks of graylingThymallus thymallus, in Lake Mjøsa, Norway

Synopsis The migration of grayling was studied in Lake Mjøsa and 13 of its tributaries in which grayling spawn. The study demonstrate that grayling spawn in creeks in May/June and leave for Lake Mjøsa shortly after spawning to return the next spring. Once mature, the grayling spawn every year. Grayling mix in the lake and are found at a mean distance of 11.9 km from the estuary of their spawning creek in September. Grayling embryos hatch in June and the juveniles stay in the tributaries until September/October. Grayling spend the winter months in the lake. Out of 1599 grayling tagged in tributaries during the spawning season, 240 of 284 recaptures (84.5%) were made in subsequent spawning seasons in the same tributary in which they were first tagged. Forty-four grayling were recaptured in adjacent tributaries. These results demonstrate that, despite the fact that all grayling leave the tributaries and mix in the lake, there is a high precision of homing. It is suggested that the grayling population in lake Mjøsa is composed of stocks using specific tributaries for spawning.     

Biological characteristics of European Atlantic sturgeon, Acipenser sturio, as the basis for a restoration program in France

The European Atlantic Sturgeon, Acipenser sturio, has received increased attention in France because of population declines due to overfishing and deterioration of spawning grounds. Conservation of this species requires many actions, including publicizing the necessity to protect this fish and its habitats, investigations on catches and probable spawning grounds, and on artificial reproduction, which is still in an experimental stage. During its sea life, European Atlantic sturgeon occur from the Bay of Biscay to the Bristol Channel and North Sea. Presently, the number of young fish in the Gironde Estuary during summertime is low and the population has a unimodal age structure. At the time this paper was written, the last recorded reproduction of sturgeon in the Gironde system occurred in 1988 (new evidence of reproduction was discovered in 1995). Growth of young fish from the 1988 cohort was faster than that previously determined for others cohorts. Over our study period (1980–1994), the availability of wild broodfish declined. Successful artificial reproduction of wild-caught females requires an optimal physiological state; any delay decreases their reproductive potential. Acclimatization of wild-caught juveniles to fresh water was most successful when fish were transported and held upon arrival in low salinity water. For such wild-caught juveniles, the first food intake usually occurs several months after capture, but remains irregular. These findings will be used to improve ongoing efforts to restore A. sturio.     

Testing for the Effects of Seasonal and Lunar Periodicity on the Reproduction of the Edible Sea Urchin <Emphasis Type="Italic">Tripneustes gratilla</Emphasis> (L) in Kenyan Coral Reef Lagoons

The annual and lunar reproductive cycle of the widely distributed edible sea urchin Tripneustes gratilla (L) was examined through measurements of gonad index, histological examination of gametogenesis, and induction of spawning with KCl injections. The population density and morphological characteristics of urchins at Diani, Kanamai, and Vipingo reef lagoons were also studied as well as the effects of seawater temperature and light on reproduction. Gonad growth started early during the northeast monsoon and reached a peak in June at the beginning of the southeast monsoon followed by a sharp decrease in gonad size of 50% in July and August towards the end of the southeast monsoons. Histological examination of gonads, revealed many different stages of gametogenesis with gametes present throughout the year, indicating continuous reproduction. There was a significant relationship between gonad index and lunar day with spawning occurring between lunar day 7 and 21, but spawning was not in perfect synchrony in the population. The population density of urchins at each reef is variable from year to year and was highest on average at Vipingo. Urchins at Kanamai had the lowest gonad indices, the largest jaws and smallest individuals an indication of food limitation. The gonads (roe) of T. gratilla at all three sites, were perpetually ‘runny’ an attribute that is not suitable for urchin fisheries. Studies to develop techniques to improve roe quality are recommended.     

Climate change and condition of herring (Clupea harengus) explain long-term trends in extent of skipped reproduction

It is commonly assumed that iteroparous fish, once mature, normally reproduce in all consecutive seasons. Recent work has suggested, however, that in Norwegian spring-spawning herring—a population that undertakes extensive spawning migrations—almost one in two adults may skip their second spawning migration. Why should herring not return to spawn the year after first spawning, but instead wait an extra year? For herring, participation in distant, energetically costly, and risky spawning migrations will only pay off in terms of fitness if individuals are sufficiently large, and in sufficient condition, to both successfully migrate and spawn. Changes in the environment and individual condition should therefore affect the likelihood of skipped spawning. This paper describes long-term changes in the extent to which the second reproductive season is skipped in this herring population. These are shown to be linked to the size and condition of herring as first-time spawners, and to climatic factors possibly related to food availability. The findings corroborate the hypothesis that skipped reproduction results from trade-offs between current and future reproduction, growth and survival.     

长江鲥鱼种群生长和繁殖特性的研究

本文研究了长江鲥鱼的生长参数和繁殖特性:(1)采用Von Bertalanffy生长方程计算长江鲥鱼的生长,求得渐近体长值L_∞()=58.12厘米,L_∞()=52.22厘米,渐近体重值W_∞()=2874.64克,W_∞()=1989.99克,生长拐点t=1.7年,t=1.3年。(2)长江鲥鱼主要集中在赣江产卵,产卵时的水温为24.2—33.5℃,流速为0.31—1.52米/秒。(3)分析和比较了长江鲥鱼不同年际间种群数量和繁殖群体结构的特点,指出性成熟年龄提前是鲥鱼资源衰退的主要标征。     

Life-history traits of Amur sleeper, <Emphasis Type="Italic">Perccottus glenii</Emphasis>, in the invaded Vistula River: early investment in reproduction but reduced growth rate

In recent decades, Amur sleeper is one of the most invasive fish species in Eastern and Central Europe. Generally, it is assumed that success of an invasive species can largely depend on the plasticity of its life-history traits, e.g., indicated by higher investment in reproduction in the initial stage of its invasion. On the other hand, such energy allocation to production of gonads should negatively impact somatic growth rate. The aim of this article was to explore this phenomenon in a non-native population of Amur sleeper inhabiting artificial reservoir on a large lowland river in Central Europe. Through analysis of the population age structure, sex-dependent growth rate and life-history traits we assessed investment in reproduction and its possible relationship with growth pattern as well as compared the results with literature data from native and introduced range. Samples collected monthly from April 2005 to March 2006 were used to estimate sex ratio, GSI, fecundity, eggs diameter frequency over the year, duration of spawning season, and mode of spawning. Age structure was calculated from scales and the same data were also used to back-calculate standard length (SL) at age. The population in the Włocławski Reservoir was represented by eight age groups, and its life span was one of the longest recorded both in its natural range and introduced areas. The weight–length relationship showed that the Amur sleeper grew isometrically and there was no difference between females and males. Although the Amur sleeper growth rate was relatively slow in the Włocławski Reservoir, its increment in the first year of life was comparable to that in other introduced areas and higher than in its natural range. The female age of maturation (1+) found in the reservoir was earlier than observed in its natural range as well as in most introduced areas (2+, 3+). Female length at maturation was similar to that reported elsewhere. The back-calculated standard length (SL) of females and males showed that in the first two age classes (1 and 2) males achieved larger SL than females. Meanwhile, at age 3, 4 and 5, females were larger than males but these differences were insignificant, except for class 5. For both sexes the average annual increment of SL decreased with age but in females the increment was always higher than in males (P < 0.05). The growth rate differences between females and males resulted possibly from their unequal investment in reproduction. Multiple spawning in the Włocławski Reservoir lasted from April to the end of August and was almost 3 months longer than in other regions. Thus, this higher investment in reproduction displayed by, e.g., earlier maturation of females and longer spawning season in the Włocławski Reservoir, may contribute to invasive success of Amur sleeper in newly colonized areas; however, the costs of this strategy result in slower growth in older age classes.     

Postmetamorphic growth,survival, and age at first reproduction of the salamander,hynobius nebulosus tokyoensis Tago in relation to a consideration on the optimal timing of first reproduction

The postmetamorphic growth and survival of the salamander Hynobius nebulosus tokyoentisTago were surveyed in the study site located in Habu village of Hinodemachi, a suburb of Tokyo City, during 1975–1981. A laboratory experiment on the growth rate of juveniles was conducted in parallel with the field survey. The result indicated that this salamander grew at the rate of 8,mm in s.v.l. per year during the juvenile stage, but its growth rate decreased markedly as low as 1.8 mm for males and 1.1 mm for females, once it had attained sexual maturity. According to the “capture-recapture” procedure the annual survival rate after metamorphosis was found to be quite high; that is, approximately 0.7. By using the growth rate of juveniles and the difference between the sizes at metamorphosis and sexual maturity, the age at first reproduction was estimated to be 4 year for males and 5 year for females. From the data obtained in this study, the intrinsic rates of increase (r) were calculated for various values of age at first reproduction under different survival schedules, and the relationship between the age at first reproduction and fitness as measured by r was examined. The result indicated that an optimal age maximizing fitness always existed under respective survival schedules, and the observed age at first reproduction of this salamandei was found to coincide well with the predicted optimal age.     

Year class splitting in the woodlouse Philoscia muscorum explained through studies of growth and survivorship

Year class splitting has been observed in a population of the terrestrial isopod Philoscia muscorum (Scopoli) in a fixed dune grassland. In this population, individuals born in one season differentiate into two forms, one of which grows to maturity in one year, the other in two.
Laboratory investigations of growth, fecundity and survivorship of individuals showed fundamental positive relationships between rate of growth and the ambient temperature/day light regime and between parental size and the number of offspring produced. Size also affected survivorship, with adults surviving better than juveniles. A link between survivorship and the temperature/day light regime was only obvious in small juveniles.
No direct physiological cause for the year class split could be derived from the laboratory studies, but a trade-off between the need for delayed or early breeding could result in the patterns of growth and reproduction observed in the field population. Delayed breeding allows continued growth of adults and hence larger numbers of offspring. Early breeding allows more growth of juveniles which greatly improves their chances of winter survival.     

Demography of persistent juveniles in Fraxinus excelsior

Seedling emergence, survival, growth, and juvenile development of Fraxinus excelsior were followed over 10 yr in permanent plots m a dense Alnus-Fraxinus woodland in the outer archipelago of Stockholm, Sweden Number of germinated seedlings varied between years, due to variation in total seed crop but was also affected by weather conditions Seedlings and juveniles showed high survivorship 39 2% of the emerged seedlings of the two oldest cohorts were alive in the 9th growth season Mortality was somewhat raised during the first two years, but was age-independent from the third year The density of the field-layer and climatic factors were the most important determinants of mortality The juveniles produced 1 2 pairs of leaves per year and had an annual height increment of 7 1 mm, on average, from their second growth season The ontogenetic development, e g the increase m leaf complexity, was correspondingly slow and included regressions to more simple leaves
Fraxinus excelsior shows a regenerative strategy involving persistent juveniles The divergence between this and previous studies may be due to the lack of rabbit grazing     

Effects of egg size, parental origin and feeding conditions on growth of larval and juvenile cod Gadus morhua

An experimental study was performed to disentangle parental and environmental effects on the growth of Atlantic cod Gadus morhua larvae and juveniles. Eggs were collected during the spawning season from spawning pairs (families) kept separately in specially designed spawning compartments. Newly hatched larvae were released simultaneously into two mesocosms of 2500 and 4400 m³. Larval growth was monitored by sampling over a 10 week period, after which juveniles were transferred to on-growing tanks, where they were tagged and kept for up to 2 years. Maternal origin was determined by individual microsatellite genotyping of the larvae ( n = 3949, 24 families) and juveniles ( n = 600). The results showed significant positive correlations between egg size and larval size during the whole mesocosm period. Correlations, however, weakened with time and were no longer significant at the first tank-rearing sampling at an age of 9 months. Significant family-specific differences in growth were observed. The coefficient of variation ( c.v. ) was calculated in order to examine variation in standard length of larvae during the mesocosm period. Inter-family c.v. was on average 69% of intra-family c.v. Differences in zooplankton densities between the two mesocosms were reflected in larval growth, condition factor and c.v. Low food abundance appeared to reduce c.v. and favour growth of larvae that showed relatively slow growth at high food abundance. It is suggested that genetically determined variation in growth potential is maintained by environmental variability.     

Aspects of the ecology of bream, Abramis brama (L.), in a gravel pit lake and the effects of reducing the population density

The growth of bream, Abrumis brumu (L.), in a gravel pit lake was considered to be good in 1982/1983 (L_x=663cm, K=0.105). Theadult breampopulationwasdominated by the 1973and 1976 year-classes. These year classes still formed most of the spawning population in 1987, but had grown very little (<2 cm) in the previous 5 years. In November 1987, 158 kg ha⁻¹ of bream were removed from this lake and in I988 the remaining bream population showed an increase in growth (mean length by 2.7 cm and mean weight by 0.5 kg) and condition (1.87 in 1987 to 2.09 in 1988) in one growing season. This additional growth of an ageing stunted bream population is considered to be the result of an increased food supply after reducing the fish density.     

Growth under uncertainty: The influence of marine variability on early development of Galapagos sea lions

Development implies a change in allocation of resources from somatic growth to reproduction. In a highly variable environment, growth can vary from year to year thereby influencing the long‐term life history perspective. The Galapagos sea lion (Zalophus wollebaeki) lives in a highly unpredictable marine environment in which food abundance varies not only seasonally, but also annually due to El Niño. Galapagos sea lions are restricted to a patch of cold upwelling waters surrounding the archipelago and are closely tied to land as nursing females alternate between foraging at sea and nursing ashore. Therefore, their offspring are especially vulnerable to ocean warming causing reduced food abundance. We found a significant correlation between sea surface temperature (SST) and early growth: Both mass at birth and linear growth within the first 2 mo of life correlated negatively with SST. Absolute mass gain was higher for males, but both sexes gained equally 1.9% of birth mass per day. Until the age of 3 yr male and female juveniles showed similar growth to an asymptotic mass of 40 and 35 kg, respectively. As a consequence of the highly variable environment, the plasticity in growth strategy of Galapagos sea lion juveniles appears wider than that of all other sea lions allowing them to cope with poor conditions.     

Fish reproduction and the impact of acidification in the Kyrönjoki River estuary in the Baltic Sea

Synopsis Several freshwater species use the Kyrönjoki River estuary as a spawning and nursery area. The main reasons for this seem to be the morphology of the estuary, the abundance of shelter provided by aquatic macrophytes, high food production and favourable temperature conditions. Acidification of the estuary due to drainage from acidic soils has made part of the estuary unsuitable for fish reproduction. In addition, year to year fluctuations in the acidity of the estuarine water have affected the reproductive success of several species. The severity of the effects of the acidification at the population level is determined by the spatial and temporal distribution of the larvae and juveniles.     

Growth of young-of-the-year mackerel in the Bay of Biscay

The first growth season of young-of-the-year (0+ year) mackerel Scomber scombrus , sampled in the Bay of Biscay, was parameterized to determine growth patterns. Daily increments were identified on sagittae otoliths, for calculation of age and growth of 92 larvae and 54 juveniles over the range 3·6–215·0 mm standard length ( L _S). A Gompertz curve was fitted to the length-at-age data. At the end of the first year of growth L _S was 194·2 mm, with a maximum growth increment of c . 2 mm day⁻¹, observed 62 days after hatching. Backcalculated growth increments for mackerel juveniles, during their larval stage, were higher than those observed for sampled larvae; only 10·9% of sampled larvae were estimated to survive. Growth for north-eastern Atlantic mackerel was slower than that published for north-western Atlantic mackerel. Backcalculated hatching dates for mackerel were consistent with the typical temporal distribution of mackerel spawning in the Bay of Biscay.     

Population dynamics of the perennial Plantago media in semi‐natural grasslands

Abstract. In this study we investigated population dynamics of a perennial forb, Plantago media, in semi‐natural grasslands in southeastern Sweden. Plantago media is a rather common inhabitant of grasslands in this region, but it has been shown to experience dispersal limitations both among managed grassland sites and at potential dispersal routes along road verges. The demography of P. media was studied in 22 permanent plots at two sites over five years. A seed sowing experiment was also performed at each site. The life‐cycle of P. media includes seedlings, juveniles, small vegetative rosettes, large vegetative rosettes and flowering rosettes. The population growth rates (λ‐values) were negative for both populations, with one exception: the first year transition matrix at one site. The projected life span for individuals surviving from seedlings to flowering rosettes varied between 19.6 yr and 227.8 yr. Elasticity analysis showed that remaining in the large rosette stages, both vegetative and flowering, was the most important factor influencing population growth rate. LTRE analysis gave similar results, indicating that spatial and temporal variation have similar effects on the population growth rate. The expected time to extinction in populations with an initial size of between 100 and 1000 individuals varied between 60 and 200 yr. The seed sowing experiment showed that seedling emergence was enhanced by both seed addition and disturbance, suggesting that recruitment is limited by seed and microsite availability. Recruitment was not found to have much impact on population growth rate. However, long‐term population maintenance may depend on occasional small‐scale disturbances that enhance pulses of recruitment. Fragmented landscape is likely to effect plant populations including rare and endangered species as well as more common species which have limited dispersal mechanisms.     

Mortality and production of 0+ perch, Perca fluviatilis L., in two Scottish lakes

Mortality of 0+ Eurasian perch was measured directly from the decline in catch in a high-speed plankton sampler and described by a negative exponential model. A mean mortality of 0.063 day ¹ (range=0.020–0.089) was calculated; for 0+ juveniles it was 0.021 day ¹. Mean larval production was 0.85–1.30 mg dry weight m⁻² day⁻¹ in Loch Kinord and 0.08–0.12 in Loch Davan. Annual production of 0+ juveniles was 1.97–7.55 kg wet weight ha ⁻¹ at L. Kinord and 1.01 kg at L. Davan. The contribution of 0+ perch, including the egg stages, to total perch production was 88–96%. Availability of food items on transition to exogenous feeding and through the larval period did not limit survival. Cannibalism by adults accounted for most of larval mortality, and predation by pike and adult perch were responsible for the majority of juvenile losses. The population structure of adult perch, for 12 year classes, was stable at L. Kinord, while year classes at L. Davan exhibited fluctuations in abundance and in two sampling years were eliminated in the first summer. Survival in the eleutheroembryonic phase was variable at L. Davan and observed to be low compared with L. Kinord. It is suggested that this was a result of water turbulence on the exposed and shallow spawning areas at L. Davan.     

Growth allocation between height and stem diameter in nonsuppressed reproducing Abies mariesii trees

To understand the growth patterns with respect to competition and leaf‐mass increase in reproducing trees, growth allocation between height and stem diameter was examined for nonsuppressed reproducing Abies mariesii trees in a subalpine forest in northern Honshu, Japan. The growth allocation was analyzed by dividing the relative growth rate of the stem volume into the relative contributions of height and stem‐diameter growth. During a 9‐yr period, height growth and seed‐cone production showed obvious annual variation, while stem‐diameter growth recorded moderate variation. For two of three years of seed‐cone production during the 9‐yr period, trees with larger seed‐cone production were associated with less height growth in the following year of seed‐cone production; however, there was no trend of height growth in the year of seed‐cone production. In the following year of mast seeding, trees with larger stem‐volume growth were associated with less height growth. This trend was also shown for the relationship between the cumulative stem‐volume growth during the 9‐yr period and growth allocation to height, suggesting that trees with a larger biomass increase depress the allocation of photosynthate to competition with a large expenditure for reproduction. In contrast to this, trees with a smaller biomass increase might allocate photosynthate to competition with surrounding trees. The results of this study suggest that an increase in reproductive organs during life history and annual variation in reproduction are closely associated with the growth patterns of the stem in A. mariesii trees.