Life-history traits and potential invasiveness of introduced pumpkinseed Lepomis gibbosus populations in northwestern Europe

To evaluate the potential invasiveness of pumpkinseed Lepomis gibbosus introduced to northwestern European inland waters, growth and reproduction traits were examined in ten populations along a trajectory spanning northwestern Europe (Norway, England, Holland, Belgium and France) and evaluated in light of published dataset from Europe. In the 848 pumpkinseed captured, maximum age was 3–4 years, with a sex ratio near unity in all but one population. Significant variations with increasing latitude were observed in adult growth (age 2–3 increment in total length, TL) and mean age at maturity (A _M), with non-significant variations observed in juvenile growth (TL at age 2), sex ratio and gonado-somatic index. As observed elsewhere in Europe, mean A _M decreased significantly with increasing TL at age 2. Using this relationship, which has been proposed elsewhere as a potential predictive model of pumpkinseed invasiveness, eight of the ten populations could be provisionally categorized as ‘non-invasive’ (five populations), ‘transitional’ (one population) and ‘potentially invasive’ (two populations), with two populations not categorized due to insufficient data. Based on the available knowledge on each population, the relationship between juvenile growth and age at maturity appeared to predict reasonably the status of pumpkinseed in northwestern Europe and its applicability to other species should be tested.     

A review of growth and life-history traits of native and non-native European populations of black bullhead <Emphasis Type="Italic">Ameiurus melas</Emphasis>

North American black bullhead, Ameiurus melas, which were introduced to Europe in the nineteenth and twentieth centuries, have received relatively little study. With focus on growth and reproduction, this extensive review, which includes new European data, aims to inform the risk analysis process concerning this non-native species in Europe. Surprisingly, the new data for Europe were more comprehensive than for native populations, with data available mainly from Oklahoma, and North and South Dakota (USA). In terms of relative growth, juvenile A. melas were found to have a relatively uniform body shape regardless of the population’s origin, whereas adults developed different phenotypes depending upon location. Overall growth trajectory was significantly faster for native than for non-native populations. Growth index values decreased significantly with increasing latitude in non-native but not native populations—the latter decreasing weakly with increasing altitude in the populations located at latitudes <40°. Mean general condition (slope ‘b’), mean sex ratio and mean egg diameter did not differ significantly between native and non-native populations. Absolute fecundity was slightly (but not significantly) higher in non-native than native populations. GSI data, which were very scarce for native populations, suggest gonad production may be slightly higher in native than in non-native populations. Precise data on age at maturity (AaM) are lacking for the native range, where 2–5 years is reported. Whereas, in the introduced range the greatest AaM was 3.5 years, and AaM decreases with increasing juvenile growth (TL at age 3). The populations with fastest juvenile growth tended to be from warmer water bodies where they are considered to be invasive. The great growth and life-history plasticity of black bullhead affords the species great potential to invade and establish viable populations in new areas.     

Old world versus new world: life-history alterations in a successful invader introduced across Europe

We examined differences in pumpkinseed (Lepomis gibbosus) life-history traits between native North American and introduced European populations, and tested three life-history predictions related to the effect of temperature, growth, waterbody size, and the presence/absence of predators on native and non-native populations. Pumpkinseed populations exhibit more ‘opportunistic’ traits (earlier maturity, smaller size at maturity, and higher reproductive allocation) in their introduced European range than those in their native range. Predictions of life-history traits were improved when indicators of juvenile growth rate (mean length at age 2), waterbody size (surface area), and thermal regime (air temperature degree-days above 10 °C) were incorporated into models along with continental location, but European pumpkinseed populations exhibit more opportunistic life-history traits than North American populations even when these factors are accounted for. Native pumpkinseed in waterbodies containing piscivores mature later and at a larger size, and have lower gonadosomatic indices than those in waterbodies lacking piscivores, whereas there is no significant difference in the same three life-history traits between European waterbodies containing or lacking piscivores. Because congeneric competitors of the pumpkinseed are absent from Europe, the apparent absence of a predator life-history effect there could also be due to the absence of the major sunfish competitors. In either case, the evolution and maintenance of more opportunistic traits in European pumpkinseed can likely be attributed to enemy release, and this may explain the successful establishment and spread of pumpkinseed in many parts of Europe.     

Relationships of reproductive traits and body size with attainment of sexual maturity and age in Blanding's turtles (Emydoidea blandingi)

To place associations among body size, age at maturity, age, and reproductive traits of a long-lived organism in the context of current life history models based on the concept of norms of reaction, we examined data from a mark-recapture study of Blanding's turtles (Emydoidea blandingi) in southeastern Michigan during 24 of the years between 1953 and 1988. Females matured between 14 and 20 years of age. Both the smallest and largest adult females in the population were reproducing for the first time in their lives. This result suggests that a combination of differences in juvenile growth rates and ages at maturity, and not indeterminate growth, are the primary cause of variation in body size among adults. Body size variation among individuals was not related to age at sexual maturity. Females that had slower growth rates as juveniles matured later at similar mean body size compared to those with more rapid growth that matured at an earlier age. As a result, a linear model of age at sexual maturity with growth rates of primiparous females between hatching and maturity was significant and negative (R² = 0.76). Frequency of reproduction of the largest and smallest females was not significantly different. Clutch size did not vary significantly with age among either primiparous or multiparous females. Clutch sizes of primiparous females and multiparous females were not significantly different. However, older females (>55 years minimum age) reproduced more frequently than did younger females (minimum age <36 y).     

Demographic and environmental influences on life-history traits of isolated populations of the Andean catfish Astroblepus ubidiai

In this study, we examined the influence of demographic and environmental variables on the life histories of six remnant populations of the Andean catfishAstroblepus ubidiai (Actinopterygii; Siluriform) located in isolated refuges, and tested six predictions on the relationships between age at maturity, generation time, population density and juvenile and adult growth and survival rates. The three populations inhabiting the watershed with a direct connection to a major lake (Imbakucha) exhibited later maturity, a longer generation time, higher adult survivorship, a higher adult-to-juvenile survival ratio and lower biomass density than those of the adjacent, non-lacustrine watersheds. Across all six populations, there was a strong correlation between mean age at maturity and both the adult-to-juvenile survival ratio and biomass density. Study populations also showed a pattern of inverse density-dependence associated with Allee effects. We conclude that without limitations in the availability of environmental resources, higher population density can be negatively related to age at maturity. This relationship would be expected to become positive when carrying capacity of the systems is reached. Delayed maturity in the Imbakucha populations may be an adaptive response for increasing juvenile survival under stressful conditions.     

Latitudinal clines in life-history variables of anadromous brown trout in Europe

Based on published data, we reviewed clinal variations in life-history characteristics of anadromous brown trout Salmo trutta from 102 European rivers at latitudes between 54 and 70° N. Growth rate in fresh water, mean smolt age, mean sea age at first sexual maturity, proportion of repeal spawners among adults, longevity, and length of adult life span exhibited latitudinal clines. Brown trout grew faster in fresh water, smolted and matured younger, lived fewer years but spawned more times in the south than in the north. The life-history traits studied were often correlated. Longevity, smolt age and sea age at maturity were negatively and smolt length and proportion of repeat spawners among adults were positively correlated with growth rate in fresh water. Longevity was positively correlated with smolt age and sea age at maturity. The latter also increased with increasing smolt age. None of these significant correlations among life history variables, except for those between smolt age and parr growth and proportion of repeat spawners and parr growth, are latitudinal effects. We do not know to what extent the latitudinal variation in life–history traits reflects phenotypic plasticity and to what extent it is caused by inherited differences among populations.     

Life-history traits of introduced Iberian pumpkinseed Lepomis gibbosus relative to native populations. Can differences explain colonization success?

To assess the colonization success of pumpkinseed Lepomis gibbosus on the Iberian Peninsula, life-history attributes of pumpkinseed populations from Spanish water bodies were compared to populations in the northern and southern parts of its native range, as well as to those of English water bodies where introduced populations have shown minimal natural range expansion. Discriminate function analysis using five population characteristics [mean age at maturity, mean total length ( L _T) at maturity, gonado-somatic index ( I _G), mean L _T at age 2 years as an indication of juvenile growth rate and relative body condition] strongly differentiated populations from the four regional study areas. Spanish populations were early maturing, showed moderate juvenile growth rate and L _T at maturity and high I _G relative to the other population groups. Spanish populations matured significantly earlier than Canadian and English populations, but not southern U.S.A. populations. Spanish populations, however, had a significantly higher I _G than southern U.S.A. populations, suggesting greater reproductive output. Considering these differences in the context of the Winemiller and Rose triangular life-history strategy model, the pumpkinseed exhibits 'equilibrium' life-history traits that have made other non-native fishes successful invaders in areas where habitat alteration has resulted in more predictable discharge regimes and water levels. Populations in Iberia, however, appear to have achieved their success by adopting a more 'opportunistic' life-history strategy than their native counterparts. High levels of life-history plasticity appear to contribute to the success of this species on the Iberian Peninsula.     

Demography of squirrel monkeys (Saimiri sciureus) in captive environments and its effect on population growth

Understanding which life-history variables have the greatest influence on population growth rate has great ecological and conservation importance. Applying models of population regulation and demographic mechanisms can aid management and conservation of both wild and captive populations. By comparisons of sensitivity, elasticity, and life-table response analyses, we identified demographic processes that were most likely to produce changes in population size (via prospective analyses) and the traits that actually influenced population changes (via retrospective analyses) among sexes, zoological facilities, and generations of captive squirrel monkey populations (Saimiri sciureus). Variation in life-history traits occurs within each group analyzed. Those traits that vary the most include age at maturity, age at last reproduction, and fertility. Zoos with increasing population growth rates maintain earlier ages of maturity, later ages of last reproduction, high rates of juvenile and adult survival, and most importantly greater fertility, reflecting shorter inter-birth intervals. Using prospective analyses, juvenile and adult survivals were predicted to be demographic traits with the greatest effect on population growth. Surprisingly, and despite predictions, retrospective analyses revealed that fertility was the life-history characteristic trait that contributed the most to changes in population size.     

The relative importance of life-history variables to population growth rate in mammals: Cole's prediction revisited

The relative importance of life-history variables to population growth rate (lambda) has substantial consequences for the study of life-history evolution and for the dynamics of biological populations. Using life-history data for 142 natural populations of mammals, we estimated the elasticity of lambda to changes in age at maturity (alpha), age at last reproduction (omega), juvenile survival (Pj), adult survival (Pa), and fertility (F). Elasticities were then used to quantify the relative importance of alpha, omega, Pj, Pa, and F to lambda and to test theoretical predictions regarding the relative influence on lambda of changes in life-history variables. Neither alpha nor any other single life-history variable had the largest relative influence on lambda in the majority of the populations, and this pattern did not change substantially when effects of phylogeny and body size were statistically removed. Empirical support for theoretical predictions was poor at best. However, analyses of elasticities on the basis of the magnitude (F) and onset (alpha) of reproduction revealed that alpha, followed by F, had the largest relative influence on lambda in populations characterized by early maturity and high reproductive rates, or when F/alpha > 0.60. When maturity was delayed and reproductive rates were low, or when F/alpha < 0.15, survival rates were overwhelmingly most influential, and reproductive parameters (alpha and F) had little relative influence on lambda. Population dynamic consequences of likely responses of biological populations to perturbations in life-history variables are examined, and predictions are made regarding the numerical dynamics of age-structured populations on the basis of values of the F/alpha ratio.     

Indeterminate growth in long-lived freshwater turtles as a component of individual fitness

Although evidence that reptiles exhibit indeterminate growth remains equivocal and based on inadequate data, the assumption that they do is still widely accepted as a general trait of reptiles. We examined patterns of variation in adult growth using long-term mark-recapture data on 13 populations of 9 species representing 3 families of freshwater turtles located in South Carolina, Michigan, and Arizona in the USA and in Ontario, Canada. Across 13 study populations, growth rates of all adults and only those that grew averaged 1.5 and 1.9 mm/yr respectively. Sources of variation in growth rates included species, population, sex, age, and latitude. Most adults of both sexes with recapture intervals greater than 10 years grew, but across all populations an average of 19 % of individuals did not grow (some with recapture intervals up to 30 years). For known-age adults of three species, the highest growth rates occurred during the 10 years following sexual maturity, and the proportions of non-growing individuals increased with age. Growth rates of adults were on average 92 % lower than those of juveniles. Based on linear relationships of clutch size and body size of females at average juvenile and adult growth rates it would take 0.7 (0.2–1.2) years and 8.6 (min–max = 2.3–18.5) years, respectively, to grow enough to increase clutch size by one egg. The majority of within population variation in adult body size in 3 species appeared to be a combination of differences in ages at maturity and juvenile and early adult growth, rather than indeterminate growth. The results from our study populations indicate that increases in body size (and associated reproductive output) that results from indeterminate growth are not substantial enough to represent a major factor in the evolution of life histories in general or the evolution of longevity and aging specifically.     

Growth and life history traits of Aegean chub,Squalius fellowesii (Günther, 1868) in streams in Muğla Province,Aegean coast,Turkey

To aid in species' conservation, the aim of this study was to provide initial findings on age, growth and reproduction of an endemic species, Aegean chub Squalius fellowesii (Günther, 1868) populations from streams in the Aegean region of Mu?la Province, Turkey. The species is relatively short‐lived (maximum 6 years), attaining a size of about 200 mm total length with a rapid growth to first maturity (≈60 mm TL), and relatively little growth thereafter. The male:female ratio was 1.0 : 0.6, males significantly outnumbering females in the majority of the streams. General condition values of individual fish varied between 2.9 and 3.4. Sexual maturity was usually achieved later and at larger sizes in females than in males. Sexual maturation in most populations was at the age of 2 years in females and 1 year in males. The species spawns between early April and late May. Mean absolute and relative fecundity were about 4440 eggs and 57 eggs·g^?1, respectively. Mean egg diameter was 1.00 ± 0.03 mm, ranging from 0.70 to 1.20 mm. Suggestions for the conservation of Aegean chub are discussed.     

Pathways of spread of the introduced ascidian Styela clava (Tunicata) in Northern Europe, as revealed by microsatellite markers

Styela clava, a solitary ascidian native to the NW Pacific, has become a conspicuous member of fouling communities in NW European waters. As its natural dispersal appears to be limited, the wide distribution of S. clava along coasts within its introduced range may be attributed to secondary spread assisted by human activities. Here, we used six microsatellite loci to examine the genetic diversity and extent of gene flow among S. clava populations in its European introduced range. Samples were collected from 21 populations within Europe (N = 808), 4 populations within the USA and two populations within the native range (Japan). Large variation in genetic diversity was observed among the European populations but were not explained either by the geographic distance from the first introduction area (i.e. Plymouth, UK) nor by the time elapsed since the introduction. No founder effect was observed in the introduced populations, except possibly in Puget Sound (USA). At least two different introductions occurred in Europe, identified as distinct genetic clusters: northern Danish populations (resembling one Japanese population), and the rest of Europe; a sample from Shoreham (England) possibly represents a third introduction. In North America, the population from the Atlantic was genetically similar to the majority of European populations, suggesting a European origin for populations on this seaboard, while populations from the Pacific coast were genetically similar to the same Japanese population as the Danish populations.     

THE EFFECTS OF PREDATION ON THE AGE AND SIZE OF MATURITY OF PREY

The effects of nonselective predation on the optimal age and size of maturity of their prey are investigated using mathematical models of a simple life history with juvenile and adult stages. Fitness is measured by the product of survival to the adult stage and expected adult reproduction, which is usually an increasing function of size at maturity. Size is determined by both age at maturity and the value of costly traits that increase mean growth rate (growth effort). The analysis includes cases with fixed size but flexible time to maturity, fixed time but flexible size, and adaptively flexible values of both variables. In these analyses, growth effort is flexible. For comparison with previous theory, models with a fixed growth effort are analyzed. In each case, there may be indirect effects of predation on the prey's food supply. The effect of increased predation depends on (1) which variables are flexible; (2) whether increased growth effort requires increased exposure to predators; and (3) how increased predator density affects the abundance of food for juvenile prey. If there is no indirect effect of predators on prey food supply, size at maturity will generally decrease in response to increased predation. However, the indirect effect from increased food has the opposite effect, and the net result of predation is often increased size. Age at maturity may either increase or decrease, depending on functional forms and parameter values; this is true regardless of the presence of indirect effects. The results are compared with those of previous theoretical analyses. Observed shifts in life history in response to predation are reviewed, and the role of size-selective predation is reassessed.     

Life history and body size evolution in Holopedium gibberum Zaddach (Crustacea, Cladocera)

SUMMARY. 1. Body size, development, sheath size and relative egg size were examined in two populations of Holopedium gibberum Zaddach. The populations were chosen because of differences in juvenile versus adult mortality.
2. The population subject to food limitation and predation from icthyoplankton (high juvenile mortality) exhibited large size at birth and maturity, large sheath size, and eggs which were relatively large compared to the adult. In contrast, individuals in the population exposed to predation by golden shiner (high adult mortality) were smaller at birth and maturity, had a smaller sheath, and produced relatively small eggs.
3. I contrast body size and relative egg sizes of these H. gibberum populations with literature values for all Cladocera, and propose a general hypothesis for the evolution of relative egg size in Cladocera.     

Early Life Conditions and Physiological Stress following the Transition to Farming in Central/Southeast Europe: Skeletal Growth Impairment and 6000 Years of Gradual Recovery

Early life conditions play an important role in determining adult body size. In particular, childhood malnutrition and disease can elicit growth delays and affect adult body size if severe or prolonged enough. In the earliest stages of farming, skeletal growth impairment and small adult body size are often documented relative to hunter-gatherer groups, though this pattern is regionally variable. In Central/Southeast Europe, it is unclear how early life stress, growth history, and adult body size were impacted by the introduction of agriculture and ensuing long-term demographic, social, and behavioral change. The current study assesses this impact through the reconstruction and analysis of mean stature, body mass, limb proportion indices, and sexual dimorphism among 407 skeletally mature men and women from foraging and farming populations spanning the Late Mesolithic through Early Medieval periods in Central/Southeast Europe (~7100 calBC to 850 AD). Results document significantly reduced mean stature, body mass, and crural index in Neolithic agriculturalists relative both to Late Mesolithic hunter-gatherer-fishers and to later farming populations. This indication of relative growth impairment in the Neolithic, particularly among women, is supported by existing evidence of high developmental stress, intensive physical activity, and variable access to animal protein in these early agricultural populations. Among subsequent agriculturalists, temporal increases in mean stature, body mass, and crural index were more pronounced among Central European women, driving declines in the magnitude of sexual dimorphism through time. Overall, results suggest that the transition to agriculture in Central/Southeast Europe was challenging for early farming populations, but was followed by gradual amelioration across thousands of years, particularly among Central European women. This sex difference may be indicative, in part, of greater temporal variation in the social status afforded to young girls, in their access to resources during growth, and/or in their health status than was experienced by men.     

GENETIC AND MORPHOMETRIC DIVERGENCE IN ANCESTRAL EUROPEAN AND DESCENDENT NEW ZEALAND POPULATIONS OF CHAFFINCHES (FRINGILLA COELEBS)

Descendent populations of chaffinches (Fringilla coelebs) introduced to New Zealand about 120 years ago were compared with “ancestral” populations in northern Europe and with those in a broader region of Europe (including Iberia) using protein electrophoresis at 42 loci and 12 skeletal measurements. The New Zealand populations exhibit very small scale differentiation in genetics (F_st = 0.040) and morphometrics, and the haphazard pattern of among-population variation does not align with environmental variation nor is it predicted by the geographic proximity of populations. Thus random drift is implicated in the differentiation among the descendent populations. The New Zealand chaffinches have diverged only slightly in morphometrics from an extant population in southern England, and constant heritability rate tests suggest that random drift alone could account for this small shift. In sharp contrast, the European populations are subdivided genetically (F_st = 0.222) and morphometrically, and this subdivision coincides with the Pyrenees mountains between Iberia and northern Europe which act as a barrier to gene flow between these regions. Iberian populations have smaller skulls and longer wings on average than northern European populations and are characterized by high frequencies of alternative common alleles at Ada and Np. Within both the Iberian and northern European regions, however, populations are effectively panmictic in protein-encoding genes, indicating that homogenizing gene flow is apparently extensive enough to prevent among-population differentiation in allozymes by drift. Variation in body size as represented by PC I is related to environmental productivity across Europe, unlike in New Zealand. These observations jointly suggest that longer term adaptive differentiation via selection for optimal body size has evolved in Europe. Because multilocus evolution is expected to proceed slowly in populations subject to the opposing forces of selection and homogenizing gene flow, I argue that local adaptation within “ancestral” populations in northern Europe may still be evolving.     

Correlated contemporary evolution of life history traits in New Zealand Chinook salmon, Oncorhynchus tshawytscha

Size at age and age at maturity are important life history traits, affecting individual fitness and population demography. In salmon and other organisms, size and growth rate are commonly considered cues for maturation and thus age at maturity may or may not evolve independently of these features. Recent concerns surrounding the potential phenotypic and demographic responses of populations facing anthropogenic disturbances, such as climate change and harvest, place a premium on understanding the evolutionary genetic basis for evolution in size at age and age at maturity. In this study, we present the findings from a set of common-garden rearing experiments that empirically assess the heritable basis of phenotypic divergence in size at age and age at maturity in Chinook salmon (Oncorhynchus tshawytscha) populations introduced to New Zealand. We found consistent evidence of heritable differences among populations in both size at age and age at maturity, often corresponding to patterns observed in the wild. Populations diverged in size and growth profiles, even when accounting for eventual age at maturation. By contrast, most, but not all, cases of divergence in age at maturity were driven by the differences in size or growth rate rather than differences in the threshold relationship linking growth rate and probability of maturation. These findings help us understand how life histories may evolve through trait interactions in populations exposed to natural and anthropogenic disturbances, and how we might best detect such evolution.     

Seasonal trends in body size of adult male mosquitofish,Gambusia affinis,with evidence for their social control

Synopsis Collections from a natural population of mosquitofish,Gambusia affinis, in southern Indiana showed that males born early in the breeding season mature rapidly at a small body size, while males born later in the breeding season delay maturation and achieve larger body sizes. A field experiment, involving removal of males from pond populations, was conducted to test the hypothesis that delayed maturation by late summer males is, at least in part, under social control. Mean total length (TL) of adult and maturing males in late summer in ponds from which large numbers of males had previously been removed was significantly lower than mean TL of males in control ponds. These data support the hypothesis of social control and support Sohn's competition-predation hypothesis for control of adult male body size inGambusia.     

Biology of the marbled lungfish, Protopterus aethiopicus Heckel, in Lake Baringo, Kenya

In this paper we provide life history characteristics for an introduced marbled lungfish (Protopterus aethiopicus) population in Lake Baringo, Kenya, including their length–weight relationship, body condition, fecundity, sex ratio, reproduction and food. These data are compared with those of other populations. Samples were obtained from the commercial fishery between January and October 2001. The lungfish exhibited positive allometric growth, with the length–weight relationship described by the equation: logW = ?6.41 + 3.52 logTL. The mean relative condition factor was close to unity, did not vary much between months and was not significantly different between sexes. The overall sex ratio was skewed in favour of females. The size of fish at first maturity was 70–76 cm total length (TL) for females and 82–88 cm TL for males. Fish in maturity stages III–IV occurred in all monthly samples suggesting year round spawning. The marbled lungfish in Lake Baringo is largely piscivorous and its impact on other fishes certainly warrants further research.     

Variation in growth and age at maturity in bluegill sunfish: genetic or environmental effects?

Populations of bluegill sunfish Lepomis macrochirus , experiencing heavy juvenile predation, showed increased growth rates and increased age and size at maturity relative to populations experiencing decreased predation on juveniles but increased predation on adults. This study examined bluegills experimentally from both types of populations and a cross between them in a common environment to determine if variation in growth and age at maturity is genetically or environmentally induced. Two factorial experiments, varying strain of bluegills and resource availability, were used to evaluate differences in growth rate. One experiment, varying strain of bluegills, was used to assess differences in age at maturity. Growth was strongly influenced by resource level, but growth rate did not vary among populations. Nearly all bluegills in each population matured at 1 year of age in a common environment. Thus, variation observed in source populations must be mostly attributable to differences in the environment between populations. At least three factors could potentially cause differences in growth and age at maturity: (1) variation in resource availability; (2) variation in demographic structure; and (3) variation in size-specific mortality rates caused by differences in predator abundance between populations. Observed patterns of variation between populations are best explained by effects of differences in predator populations.