Differential mortality in Turkana agriculturalists and pastoralists

Nomadic pastoral populations appear to have much lower rates of growth than the otherwise very high growth rates now characteristic of populations in developing nations. Because dramatic declines in infant mortality have been a primary contributor to increased population growth rates in these countries, it has been assumed that nomadic pastoral populations are still characterized by high levels of mortality in the first few years of life. Few studies, however, have been undertaken to estimate demographic parameters for nomadic pastoral populations, and even fewer of a comparative nature have been undertaken to document the impact of subsistence strategy on demographic processes. This study compares indirect childhood mortality estimates for Turkana nomadic pastoralists with childhood mortality in a settled agricultural group within the same population and finds that pastoralists have substantially higher levels of mortality. Based on the childhood mortality estimates, model life tables are selected for pastoral and agricultural groups from which values for mean life expectancy and infant mortality are estimated and compared. Recent improvements in primary health care for the settled agricultural group are ruled out as being an important cause of their lower mortality levels, and some aspects of life-style associated with subsistence strategy are discussed as likely determinants of the mortality differences.     

Using matrix models to explore the influence of temperature on population growth of arthropod pests

1 Population growth of four arthropod pests on agricultural crops, the pea aphid, the bird‐cherry oat aphid, the Mediterranean fruit fly, and the two‐spotted spider mite, was modelled using stage matrix models. 2 Temperature was included as a variable in the model, affecting fecundity, mortality and growth rate. Linear temperature dependence was used to describe fecundity and individual growth rate in the matrix model. Life table data collected at various constant experimental temperatures were used to parameterize the model. 3 Sensitivity analysis identified the most influential demographic parameters determining the growth rate of the population at different temperature regimes. Mortality of young adults and immature stages were found to be the most important parameters in determining population growth for all species presented. However, high temperatures increased the relative impact of individual growth rates and fecundity on the growth of the population.     

农业区游牧民族饮食文化的滞后性——基于大同东信广场北魏墓群人骨的稳定同位素研究

不同地域的人群具有不同的饮食传统和饮食文化。然而,当人们持续不断的直接或间接接触时,一方或双方原有的饮食文化将会产生变化,如拓跋鲜卑的饮食文化就经历了从游牧向农业的转变。本文对山西大同东信广场北魏墓群26例人骨进行了C、N稳定同位素分析,结果 (δ~(13)C值为-11.5±2.7‰,n=26;δ~(15)N值为9.8±1.2‰,n=26)显示,先民的食物类型差异较大,并包含大量的肉食资源。研究表明,以C_3类食物为主的先民主要放牧或狩猎,以C_4类食物为主的先民主要以种植粟、黍等C_4型农作物喂养家畜。大同东信广场北魏墓群人群中两种不同饮食传统的存在,显示游牧民族新的饮食习惯(以粟作农业产品为食)的形成滞后性于北魏政权的建立,这可能与气候变冷及他们的排斥有关。     

Altitudinal patterns for longevity, fecundity and senescence in Drosophila buzzatii

We tested for variation in longevity, senescence rate and early fecundity of Drosophila buzzatii along an elevational transect in Argentina, using laboratory-reared flies in laboratory tests performed to avoid extrinsic mortality. At 25 °C, females from lowland populations lived longer and had a lower demographic rate of senescence than females from highland populations. Minimal instead of maximal temperature at the sites of origin of population best predicted this cline. A very different pattern was found at higher test temperature. At 29.5 °C, longevity of males increased with altitude of origin of population. No clinal trend was apparent for longevity of females at 29.5 °C. There was evidence for a trade-off between early fecundity and longevity at non-stressful temperature (25 °C) along the altitudinal gradient. This trait association is consistent with evolutionary theories of aging. Population-by-temperature and sex-by-temperature interactions indicate that senescence patterns are expressed in environment specific ways.     

Adaptation of <Emphasis Type="Italic">Drosophila melanogaster</Emphasis> to unfavorable growth medium affects lifespan and age-related fecundity

Experimental adaptation of Drosophila melanogaster to nutrient-deficient starch-based (S) medium resulted in lifespan shortening, increased early-life fecundity, accelerated reproductive aging, and sexually dimorphic survival curves. The direction of all these evolutionary changes coincides with the direction of phenotypic plasticity observed in non-adapted flies cultured on S medium. High adult mortality rate caused by unfavorable growth medium apparently was the main factor of selection during the evolutionary experiment. The results are partially compatible with Williams’ hypothesis, which states that increased mortality rate should result in relaxed selection against mutations that decrease fitness late in life, and thus promote the evolution of shorter lifespan and earlier reproduction. However, our results do not confirm Williams’ prediction that the sex with higher mortality rate should undergo more rapid aging: lifespan shortening by S medium is more pronounced in naive males than females, but it was female lifespan that decreased more in the course of adaptation. These data, as well as the results of testing of F₁ hybrids between adapted and control lineages, are compatible with the idea that the genetic basis of longevity is different in the two sexes, and that evolutionary response to increased mortality rate depends on the degree to which the mortality is selective. Selective mortality can result in the development of longer (rather than shorter) lifespan in the course of evolution. The results also imply that antagonistic pleiotropy of alleles, which increase early-life fecundity at the cost of accelerated aging, played an important role in the evolutionary changes of females in the experimental lineage, while accumulation of deleterious mutations with late-life effects due to drift was more important in the evolution of male traits.     

Selection for long lifespan in men: benefits of grandfathering?

Life-history theory suggests that individuals should live until their reproductive potential declines, and the lifespan of human men is consistent with this idea. However, because women can live long after menopause and this prolonged post-reproductive life can be explained, in part, by the fitness enhancing effects of grandmothering, an alternative hypothesis is that male lifespan is influenced by the potential to gain fitness through grandfathering. Here we investigate whether men, who could not gain fitness through reproduction after their wife's menopause (i.e. married only once), enhanced their fitness through grandfathering in historical Finns. Father presence was associated with reductions in offspring age at first reproduction and birth intervals, but generally not increases in reproductive tenure lengths. Father presence had little influence on offspring lifetime fecundity and no influence on offspring lifetime reproductive success. Overall, in contrast to our results for women in the same population, men do not gain extra fitness (i.e. more grandchildren) through grandfathering. Our results suggest that if evidence for a 'grandfather' hypothesis is lacking in a monogamous society, then its general importance in shaping male lifespan during our more promiscuous evolutionary past is likely to be negligible.     

Allopatric combination of Fasciola hepatica and Lymnaea truncatula is more efficient than sympatric ones

Parasites are capable of rapid evolutionary changes relative to their hosts, due to short life cycle, short generation time, and high fecundity. The direction of the evolution of parasite virulence can be studied in cross-transfer experiments, combining hosts and parasites from different localities, and comparing the outcome of established (sympatric and potentially locally adapted) and novel (allopatric) combinations of hosts and parasites. We aimed to compare the compatibility with snails hosts, the infectivity of metacercariae in rabbits and rats, and the fitness among different combinations (French-FF and Spanish-SS sympatries and allopatry-FS). The first isolate of Fasciola hepatica and its corresponding intermediate host, Lymnaea truncatula originated from Lugo's northwestern Spain. The second isolate of parasite and snail was collected in the Limoges area in central France. The Spanish snails were more susceptible to their sympatric trematode than the French snails. The Spanish flukes were more infective to intermediate hosts (snails) than the French flukes, but subsequent definitive hosts (rats or rabbits) infections remained similar. The estimated fitness was low in sympatric infections and highly similar (from 4.7 to 5.3). The fitness similarity corresponds, however, to different variations in life-history traits that could represent different strategies among the host-parasite local combinations. The infection rate in snails, metacercarial productivity, metacercarial infectivity, and the estimated fitness were better for allopatric combination (FS). The susceptibility data showed a higher efficiency of flukes in the allopatric snail population than in their local snail population. However, our results were obtained after one generation and from a single isolate and it remains to be determined if all allopatric fluke-snail isolates may present a better fitness. Nevertheless our results indicate that introduction of liver fluke-infected cattle should be monitored carefully, as it could result in the introduction of more efficient parasites.     

Sexual cannibalism and population viability

Some behaviours that typically increase fitness at the individual level may reduce population persistence, particularly in the face of environmental changes. Sexual cannibalism is an extreme mating behaviour which typically involves a male being devoured by the female immediately before, during or after copulation, and is widespread amongst predatory invertebrates. Although the individual‐level effects of sexual cannibalism are reasonably well understood, very little is known about the population‐level effects. We constructed both a mathematical model and an individual‐based model to predict how sexual cannibalism might affect population growth rate and extinction risk. We found that in the absence of any cannibalism‐derived fecundity benefit, sexual cannibalism is always detrimental to population growth rate and leads to a higher population extinction risk. Increasing the fecundity benefits of sexual cannibalism leads to a consistently higher population growth rate and likely a lower extinction risk. However, even if cannibalism‐derived fecundity benefits are large, very high rates of sexual cannibalism (>70%) can still drive the population to negative growth and potential extinction. Pre‐copulatory cannibalism was particularly damaging for population growth rates and was the main predictor of growth declining below the replacement rate. Surprisingly, post‐copulatory cannibalism had a largely positive effect on population growth rate when fecundity benefits were present. This study is the first to formally estimate the population‐level effects of sexual cannibalism. We highlight the detrimental effect sexual cannibalism may have on population viability if (1) cannibalism rates become high, and/or (2) cannibalism‐derived fecundity benefits become low. Decreased food availability could plausibly both increase the frequency of cannibalism, and reduce the fecundity benefit of cannibalism, suggesting that sexual cannibalism may increase the risk of population collapse in the face of environmental change.     

NATURAL GENETIC VARIATION OF LIFE SPAN,REPRODUCTION, AND JUVENILE GROWTH IN DAPHNIA

The evolutionary theory of senescence predicts that high extrinsic mortality in natural populations should select for accelerated reproductive investment and shortened life span. Here, we test the theory with natural populations of the Daphnia pulex-pulicaria species complex, a group of freshwater zooplankton that spans an environmental gradient of habitat permanence. We document substantial genetic variation in demographic life-history traits among parent and hybrid populations of this complex. Populations from temporary ponds have shorter life spans, earlier and faster increases of intrinsic mortality risk, and earlier and steeper declines in fecundity than populations from permanent lakes. We also examine the age-specific contribution to fitness, measured by reproductive value, and to expected lifetime reproduction; these traits decline faster in populations from temporary ponds. Despite having more rapid senescence, pond Daphnia exhibit faster juvenile growth and higher early fitness, measured as population growth rate (r). Among populations within this species complex we observed negative genetic correlations between r and indices of life-history timing, suggesting trade-offs between early- and late-life performance. Our results cannot be explained by a trade-off between survival and fecundity or by nonevolutionary theories of senescence. Instead, our data are consistent with the evolutionary theory of senescence because the genetic variation in life histories we observed is roughly congruent with the temporal scale of environmental change in the field.     

Life-history correlates of evolution under high and low adult mortality

Abstract. Life-history theory predicts evolutionary changes in reproductive traits and intrinsic mortality rates in response to differences in extrinsic mortality rates. Trade-offs between life- history traits play a pivotal role in these predictions, and such trade-offs are mediated, at least in part, by physiological allocations. To gain insight into these trade-offs, we have been performing a long-term experiment in which we allow fruitflies, Drosophila melanogaster , to evolve in response to high (HAM) and low (LAM) adult mortality rates. Here we analyze the physiological correlates of the life-history trade-offs. In addition to changing development time and early fecundity in the direction predicted, high adult mortality affected three traits expressed early in life–body size, growth rate, and ovariole number–but had little or no effect on body composition (relative fat content), viability, metabolic rate, activity, starvation resistance, or desiccation resistance. Correlations among lines revealed trade-offs between early fecundity, late fecundity, and starvation resistance, which appear to be mediated by differential allocation of lipids.     

Food limitation of a Delaware salt marsh population of the mummichog,Fundulus heteroclitus (L.)

Summary The relationship of the mummichog, Fundulus heteroclitus, to its food supply was examined using 8 m x 20 m enclosures under field conditions. Density effects were examined by stocking these enclosures with fish at densities ranging from 1/8 to four times the natural population density. Mean growth rate in each pen was estimated from measurements on individually marked fish. Fish enclosed at normal density grew at the same rate as fish from the natural population, indicating that caging had minimal effects on growth rate. When enclosed below natural density, mummichogs had a growth rate that was 2–3 times higher than that of the natural population. Fish enclosed at four times normal density had a negative mean growth rate, a higher mortality rate than fish enclosed at natural density, and no egg production.In further experiments, food was added to the same type of enclosure at fish densities ranging from one-half to four times natural density. Food additions had a positive effect on growth rate at all densities. Food additions also caused egg production of fish enclosed at the highest density to increase to a rate equivalent with the natural population. We conclude that the total food supply may regulate the maximum size of the Canary Creek mummichog population by affecting the individual growth rate, mortality rate, and fecundity.     

Can a spider web be too sticky? Tensile mechanics constrains the evolution of capture spiral stickiness in orb-weaving spiders

Owing to habitat loss and fragmentation, large mammal populations all over the world are becoming increasingly small and isolated. It is therefore a conservation priority to understand mechanisms influencing the demography of such populations, which can easily be driven to extinction. The Przewalski's horse Equus ferus przewalskii remains one of the world's most endangered species and reintroduced animals are still vulnerable. Over 9 years, we analysed factors affecting mortality and female fecundity at the individual level in a predator-free, closed population of Przewalski's horses, which grew from 11 to 55 individuals. Similar to other wild equids, the annual growth rate of the population was r =0.169. Typically, adult mortality was much lower than juvenile mortality, the latter being correlated with neither inbreeding coefficient of foals nor population density. We found no link between female fecundity and operational sex ratio of the herd, or inbreeding coefficient, lactation status and body condition of the mares. Although food therefore seemed not to be limiting in this population, density (number of horses ha⁻¹) clearly reduced fecundity, especially in subadult mares. Thus, our results show that space can slow the growth rate of a population before resources become limited, a potential source of concern for increasingly shrinking habitats of endangered large mammals. Possible mechanisms causing this may be found in incest avoidance or other social parameters. Finally, in large herbivores, population density is said to exert influence in a sequential order: juvenile survival first, followed by fecundity of young females, then adult females, and adult survival last. Although we observed no link between density and juvenile survival in the studied population, our results otherwise support this hypothesis.     

Selection for increased brood size in historical human populations

Human twinning rates are considered to either reflect the direct fitness effects of twinning in variable environments, or to be a maladaptive by-product of selection for other maternal reproductive traits (e.g., polyovulation). We used historical data (1710-1890) of Sami populations from Northern Scandinavia to contrast these alternative hypotheses. We found that women who produced twins started their reproduction younger, ceased it later, had higher lifetime fecundity, raised more offspring to adulthood, and had higher fitness (individual lambda) than mothers of singletons in all populations studied. For example, an average of 1.2 offspring survived to adulthood from a twin delivery, irrespective of its sex ratio, whereas only 0.8 offspring survived to adulthood from a singleton delivery. Only if mothers started reproduction at very late age (> 37 yr), or had a very long reproductive life span (> 20 yr), was it more beneficial to produce only singletons. These findings suggest that twin deliveries among Sami could not be explained as a maladaptive by-product of selection for other maternal reproductive traits. In contrast, our results suggest that twinning was under natural selection, although the strength of selection was likely to have been context dependent.     

Compensation in individual growth rates and its influence on lake trout population dynamics in the Michigan waters of Lake Superior

The role of compensatory mechanisms in the population dynamics of lake trout in the Michigan waters of Lake Superior was explored during three time periods: the pre-sea lamprey period, prior to 1950 when lake trout were at a relatively high abundance and the fishery was the primary source of lake trout mortality; the sea lamprey dominant period, from 1951 to 1961 when lake trout were at a very low abundance due to sea lamprey predation and overexploitation; and currently, from 1985 to 1993 when wild lake trout abundance was at a moderate level. The role of compensatory changes in growth and fecundity rates of lake trout in the Michigan waters of Lake Superior was evaluated using a life table approach. Individual growth and fecundity rates were calculated and compared between time periods. These rates were used to determine age-specific fecundity which, along with age-specific survival, were incorporated into a Leslie projection matrix to calculate the finite rate of population increase (λ). Individual growth rates and age-specific fecundity rates changed in response to the different levels of lake trout abundance during each of the study periods. Lake trout during the sea lamprey dominant period, which experienced the lowest abundance and highest mortality levels, exhibited the fastest individual growth rates and the highest age-specific fecundity. These high rates contributed to the relatively large compensatory scope exhibited by lake trout during the sea lamprey dominant period as compared to lake trout during the pre-sea lamprey or the current periods which are associated with higher levels of abundance.     

Life history divergence of sympatric diploid and polyploid populations of brine shrimp Artemia parthenogenetica

In order to study how polyploidy affects life history patterns in animals, we have examined sympatric diploid and polyploid brine shrimp (Artemia parthenogenetica) from China, Italy and Spain under laboratory conditions. At optimal temperature and salinity (25°C and 90 ppt), diploids from the three populations had much higher intrinsic rates of increase, higher fecundity, faster developmental rates, and larger brood sizes than their sympatric polyploids. The Chinese and Italian populations were selected for further analysis to determine the life history responses of diploids and polyploids to temperature and salinity changes. Under intermediate and high salinities, Chinese and Italian polyploids produced most of their offspring as dormant cysts while their sympatric diploids produced most of their offspring as nauplii. This relationship is reversed in the Spanish diploid-polyploid complex. For the Chinese population at 25° C, pentaploid clones had higher developmental rates than diploid clones at 35 ppt; at 90 ppt, diploid clones had higher developmental rates than the pentaploids. Italian diploids and tetraploids had different responses to variation in both temperature (25° C and 31° C) and salinity (30 ppt and 180 ppt). Our results demonstrate that relative fitness of the two cytotypes is a function of environmental conditions and that sympatric diploids and polyploids respond differently to environmental changes. Chinese and Italian polyploids are expected to have lower fitness than their sympatric diploids when the physical environment is not stressful and when intraspecific competition is important. However, polyploids may have advantages over sympatric diploids in stressful habitats or when they encounter short-term lethal temperatures. These results suggest that polyploid Artemia have evolved a suite of life-history characteristics adapting them to environments that contrast to those of their sympatric diploids.     

Causes of early human population growth

The archaeological record indicates large increases in human population coincident with the emergence of food production about 10,000 years ago. The cause of the growth is unclear. Extreme views attribute the change to increases in the birth rate or to decreases in the death rate. Many argue that sedentism led to improved ovarian function and higher fertility through higher caloric intakes or reduced activity levels. Similarly, shortened lactation periods may have reduced birth spacing and increased fertility. Others attribute the rise in population to decreases in mortality, arguing that the evidence from skeletal populations indicates improvements in health and the expectation of life at birth, though others use the same evidence to argue that mortality increased. An analysis presented here draws on findings that indicate substantial increases in the survival of young children as populations switch from nomadic to sedentary lives. Projections indicate that this improvement in child survival is so critical that it may be followed by substantially larger decreases in survival at later ages, yet result in higher population growth rates and reduced expectation of life at birth. Increases in the birth rate are not necessary for population growth, even when overall mortality increases. Large increases in overall mortality can be associated with large increases in population. Because positive population growth can occur while the expectation of life at birth declines, this analysis shows that this summary statistic is not an appropriate indicator of population fitness. © 1996 Wiley-Liss, Inc.     

Contrasting growth strategies of pond versus marine populations of nine-spined stickleback (Pungitius pungitius): a combined effect of predation and competition?

Gigantism in isolated ponds in the absence of sympatric fish species has previously been observed in nine-spined sticklebacks (Pungitius pungitius). Patterns in sexual size dimorphism suggested that fecundity selection acting on females might be responsible for the phenomenon. However, the growth strategy behind gigantism in pond sticklebacks has not been studied yet. Here, we compared von Bertalanffy growth parameters of four independent nine-spined stickleback populations reared in a common laboratory environment: two coastal marine (typical size) and two pond (giant size) populations. We found that both pond populations had larger estimated final size than marine populations, which in turn exhibited higher intrinsic growth rates than the pond populations. Female growth strategies were more divergent among marine and pond populations than those of males. Asymptotic body size and intrinsic growth rate were strongly negatively correlated. Hence, pond versus marine populations exhibited different growth strategies along a continuum. Our data suggest that quick maturation—even with the cost of being small (low fecundity)—is favoured in marine environments. On the contrary, growth to a giant final size (high fecundity)—even if it entails extended growth period—is favoured in ponds. We suggest that the absence (ponds) versus presence (marine environment) of sympatric predatory fish species, and the consequent change in the importance of intraspecific competition are responsible for the divergence in growth strategies. The sex-dependence of the patterns further emphasizes the role of females in the body size divergence in the species. Possible alternative hypotheses are also discussed.     

The responses of a perch, Perca fluviatilis L., population to eutrophication and associated changes in fish fauna in a small lake

A number of parameters were measured in a population of perch whose numbers had declined steeply in a habitat subjected to increasing eutrophication, and these measurements were compared with those taken before the population decline. Contrary to expectation, no adaptive responses were identified. The growth rate of perch up to the age of III had only declined slightly, and there was a marked preponderance of young fish. Relative growth rate was consistent despite the population decline. No dominant year class was apparent. Mean instantaneous mortality rate and annual survival rate were calculated, and the length-weight relationship was analysed. Feeding strategy had not altered since an earlier study. A decline was seen in the percentage of perch maturing at early age. Ripeness coefficients and fecundity estimates were calculated. A previously documented spawning migration was confirmed. The absence of the expected adaptive changes is discussed.     

Similarity of growth among Great tits (Parus major) and Blue tits (P. caeruleus)

Differences in morphology among species are proximately caused by changes in the ontogeny of individuals. It is therefore of importance to analyse possible differences in growth parameters among closely related species in order to understand what parameters are most and least likely, respectively, to change in evolution. In this paper I analyse growth in two closely related sympatric species, namely Great tit, Parus major, and Blue tit, P. caeruleus. The former is considerably larger than the latter in all external traits. The growth rates of the two species were found to be very similar for all traits, thus excluding differences in growth rate as a potential cause of evolutionary size changes. Offset of growth occurred at relatively similar times in the two species, excluding this factor as a major cause of the final size differences. However, size differences at hatching were pronounced and remained so throughout ontogeny, pointing to initial size (egg size or hatching size) as the target of factors promoting change. Bivariate allometric relations of traits vs. body size (mass) were similar between the two species at all ontogenetic stages. There was a high correlation among traits especially at intermediate age stages (5 and 8 days), but these correlations became weaker at older age and approached the low pattern of integration found in adults. All this suggests the operation of a general growth factor affecting all parts of the phenotype simultaneously, which has its major influence at the time of maximal growth. If closely related species in general have highly similar growth patterns, strong evolutionary allometry as found in many avian taxa is to be expected.     

Egg Size Versus Number of Offspring Trade-Off: Female Age Rather Than Size Matters in a Domesticated Arctic Charr Population

Understanding how organisms adjust reproductive allocation trade-offs between offspring size versus number (OSN trade-off) is a central question in evolutionary biology. In organisms with indeterminate growth, changes in OSN according to maternal size or age have been reported in numerous taxa. The relative contribution of age and size remains largely unclear, as they are often highly correlated. In this study, we investigated how females adjust the offspring size versus number trade-off and analyzed the relative contribution of female age and size in a domesticated population of Arctic charr Salvelinus alpinus (Linnaeus, 1758) that exhibit large variation in size within five age classes. Our results show that the reproductive output (i.e. as measured by the clutch mass), was strongly correlated to female mass and age suggesting that the proportion of resources allocated to reproduction do not vary along lifetime. Egg mass and fecundity (egg number) increased with female mass overall. However, within an age class, larger females had higher fecundity but egg mass was poorly related to female mass. At the population level, a positive relationship was observed between fecundity and egg mass but within each class age the relation was negative revealing a OSN trade-off. Overall, our results show that, in our model Arctic charr population, allocation trade-off to reproduction and the way females allocate to egg mass and fecundity is largely determined by their age rather than mass.