Bias in population growth rate estimation: sparse data,partial life cycle analysis and Jensen's inequality

Demographic matrix models have become an integral part of population viability analysis for threatened and endangered species, but their use is often limited by data availability. A common solution to this problem is to assume constant annual rates within a multi‐year stage. Partial life cycle analysis (PLC), which incorporates only juvenile and adult stages, is a noteworthy example of this approach because it has been described in the literature as a reliable approximation of age‐structured populations. However, we predict from Jensen's Inequality that the required lumping of age classes leads to over‐ or underestimation of population fitness when survival rates are truly age‐dependent. We illuminate this problem by comparing fitness estimates from Leslie matrix and PLC models for theoretical populations having different levels of age‐dependence in their survival rates. We also propose a modification of the PLC approach to address this problem and demonstrate its applicability using data from a published long‐term study of red deer Cervus elephas.     

Brothers delay menarche and the onset of sexual activity in their sisters

The higher costs of sons compared with daughters extends to a negative effect of brothers on the lifetime reproductive success of their siblings in subsistence and preindustrial societies. In societies with fewer resource constraints, one might expect that these effects would be limited or non-existent. This study investigates the costs of brothers and sisters in a contemporary western society of adult Australians. Girls with elder brothers had a delayed age at menarche. Younger brothers were associated with delayed onset of sexual activity in sisters, but not in brothers. Neither younger nor elder brothers influenced fitness parameters (number of pregnancies, number of children, age at first pregnancy or age at first birth) in siblings of either sex. This study provides evidence that brothers negatively affect their sisters'' onset of reproductive maturity and sexual activity; however, this delay is not associated with a fitness cost in contemporary Australia. We suggest this is due to the long period of independence prior to child bearing.     

Ecological succession,patch age and the evolution of social behaviour and terminal investment

Recent years have witnessed a growing interest in understanding the evolution of social behaviour in heterogeneous spatially structured populations. These studies, however, have neglected the impact of extinction–colonisation dynamics and ecological succession on the dynamical expression of social behaviour over time. Here, I present a kin‐selection model in which patches are structured into age‐classes. I show that ecological succession and patch age lead to highly plastic social phenotypes that vary dramatically as societies age since their initial establishment until their ultimate collapse. I find that the mode of colonisation following dispersal strongly influences the patch age‐dependent trajectories of social phenotypes. When patches are colonised by a random collection of immigrants, aggression is favoured during the build‐up of a society, but it slowly subsides until it eventually gives place to cooperation throughout the later stages of a society's lifespan. When newly established societies are formed by collectives of close relatives, cooperation is favoured during the build‐up of the society as well as when the society nears its eventual collapse. At intermediate societal ages, the genetic structure of the society is sufficiently resilient to the influx of immigrants such that cooperation remains relatively high. Moreover, I report a novel form of social terminal investment, whereby cooperative effort rises when patches approach their collapse. When dispersal is allowed to co‐evolve with cooperation, we observe a sudden rise in dispersal phenotypes before a patch's collapse, and the surprising result that clonal colonisation does not yield significantly higher levels of cooperation than the individual mode of colonisation. More generally, my results show that ecological succession strongly determines the dynamics of kin selection after colonisation, and therefore I expect that these findings will be valuable for understanding behavioural syndromes during range expansion or biological invasions.     

On the concept of limited polygyny: A reply to Frost

The main thrust of Frost's comment on my article (MacDonald 1990) is that polygyny is “limited” or nonexistent among human societies with the exception of those in Subsaharan Africa and New Guinea. The issue raised is an important one for an evolutionary account of human societies. Resource polygyny follows naturally from the evolutionary theory of sex. Roughly, males are expected to benefit much more than females by having multiple mates, so that under certain ecological conditions, males are expected to compete for females as a limiting resource. I argue here that an evolutionary/ ecological approach is not incompatible with supposing that there are ecological circumtances in which polygyny is absent or highly limited. The point is that these conditions did not occur in the populations of Western Europe, so that there is no ecological reason to suppose that the polygyny which did occur was limited in any interesting sense. There are, however, non-ecological reasons which might have limited polygyny, and these will be considered as well.     

Climate change impacts on the conservation outlook of populations on the poleward periphery of species ranges: A case study of Canadian black‐tailed prairie dogs (Cynomys ludovicianus)

Given climate change, species' climatically suitable habitats are increasingly expected to shift poleward. Some imperilled populations towards the poleward edge of their species' range might therefore conceivably benefit from climate change. Interactions between climate and population dynamics may be complex, however, with climate exerting effects both indirectly via influence over food availability and more directly, via effects on physiology and its implications for survival and reproduction. A thorough understanding of these interactions is critical for effective conservation management. We therefore examine the relationship between climate, survival and reproduction in Canadian black‐tailed prairie dogs, a threatened keystone species in an imperilled ecosystem at the northern edge of the species' range. Our analyses considered 8 years of annual mark–recapture data (2007–2014) in relation to growing degree days, precipitation, drought status and winter severity, as well as year, sex, age and body mass. Survival was strongly influenced by the interaction of drought and body mass class, and winter temperature severity. Female reproductive status was associated with the interaction of growing degree days and growing season precipitation, with spring precipitation and with winter temperature severity. Results related to body mass suggested that climatic variables exerted their effects via regulation of food availability with potential linked effects of food quality, immunological and behavioural implications, and predation risk. Predictions of future increases in drought conditions in North America's grassland ecosystems have raised concerns for the outlook of Canadian black‐tailed prairie dogs. Insights gained from the analyses, however, point to mitigating species management options targeted at decoupling the mechanisms by which climate exerts its negative influence. Our approach highlights the importance of understanding the interaction between climate and population dynamics in peripheral populations whose viability might ultimately determine their species' ability to track climatically suitable space.     

Effect of penicillin and the habitat medium in the body of bacterial carriers on the intercellular bonds in populations of the meningococcus and pertussis microbe

The relationship of the bacterial cells in populations and their adhesion activity is at present one of the research priorities in microbiological studies. The stimulating effect of penicillin on the development of morphologically different intercellular bonds (IB) in populations of the pertussis causative agent and first of all derivatives or evaginates of the cell wall membranes was observed. Morphologically similar systems and polytubular IB were detected in populations of meningococcal strains isolated from carriers having no signs of the disease. Correlation between the after-effect of penicillin and the presence of the causative agent in bacterial carriers was shown. Unknown systems of interlacing tubular structures not directly bound with the cells, the walls of which were single contour membranes were determined in the meningococcal populations treated with penicillin. IB were observed in the population in the form of transpopulation cords. Morphologically different IB playing the role of specialized organelles might be considered as factors of the functional unity of the bacterial population as a multicellular system.     

Impact of Empire Expansion on Household Diet: The Inka in Northern Chile's Atacama Desert

The impact of expanding civilization on the health of American indigenous societies has long been studied. Most studies have focused on infections and malnutrition that occurred when less complex societies were incorporated into more complex civilizations. The details of dietary change, however, have rarely been explored. Using the analysis of starch residues recovered from coprolites, here we evaluate the dietary adaptations of indigenous farmers in northern Chile''s Atacama Desert during the time that the Inka Empire incorporated these communities into their economic system. This system has been described as “complementarity” because it involves interaction and trade in goods produced at different Andean elevations. We find that as local farming societies adapted to this new asymmetric system, a portion of their labor had to be given up to the Inka elite through a corvée tax system for maize production. In return, the Inka system of complementarity introduced previously rare foods from the Andean highlands into local economies. These changes caused a disruption of traditional communities as they instituted a state-level economic system on local farmers. Combined with previously published infection information for the same populations under Inka rule, the data suggest that there may have been a dual health impact from disruption of nutrition and introduction of crowd disease.     

Hatching of the winter eggs of the fruit-tree red spider mite (Panonychus ulmi (Koch))

Observations on the hatching of winter eggs of Panonychus ulmi (Koch) from apple orchards in England over the 5 years 1962-6 are described. Differences of 2–3 weeks between orchards in the date at which 50% hatch was reached were quite common and this was out of step with petal fall in the majority of orchards. There was a continuous range between early and late hatching populations and a tendency for the populations to hatch in the same order every year. In 1966 populations on Cox's Orange Pippin hatched earlier than those on Worcester Pearmain but the difference was unlikely to be of practical importance. Neither recording accumulated day degrees for populations hatched out-of-doors nor hatching in an incubator after chilling was considered to give a sufficiently reliable forecast of 50% hatch but observations in 1966 suggested that this might be predicted from the total eggs hatched in an incubator expressed as a percentage of total healthy eggs.     

Spread of cattle led to the loss of matrilineal descent in Africa: a coevolutionary analysis

Matrilineal descent is rare in human societies that keep large livestock. However, this negative correlation does not provide reliable evidence that livestock and descent rules are functionally related, because human cultures are not statistically independent owing to their historical relationships (Galton''s problem). We tested the hypothesis that when matrilineal cultures acquire cattle they become patrilineal using a sample of 68 Bantu- and Bantoid-speaking populations from sub-Saharan Africa. We used a phylogenetic comparative method to control for Galton''s problem, and a maximum-parsimony Bantu language tree as a model of population history. We tested for coevolution between cattle and descent. We also tested the direction of cultural evolution--were cattle acquired before matriliny was lost? The results support the hypothesis that acquiring cattle led formerly matrilineal Bantu-speaking cultures to change to patrilineal or mixed descent. We discuss possible reasons for matriliny''s association with horticulture and its rarity in pastoralist societies. We outline the daughter-biased parental investment hypothesis for matriliny, which is supported by data on sex, wealth and reproductive success from two African societies, the matrilineal Chewa in Malawi and the patrilineal Gabbra in Kenya.     

Individual experience alone can generate lasting division of labor in ants

Division of labor, the specialization of workers on different tasks, largely contributes to the ecological success of social insects [1, 2]. Morphological, genotypic, and age variations among workers, as well as their social interactions, all shape division of labor [1-12]. In addition, individual experience has been suggested to influence workers in their decision to execute a task [13-18], but its potential impact on the organization of insect societies has yet to be demonstrated [19, 20]. Here we show that, all else being equal, ant workers engaged in distinct functions in accordance with their previous experience. When individuals were experimentally led to discover prey at each of their foraging attempts, they showed a high propensity for food exploration. Conversely, foraging activity progressively decreased for individuals who always failed in the same situation. One month later, workers that previously found prey kept on exploring for food, whereas those who always failed specialized in brood care. It thus appears that individual experience can strongly channel the behavioral ontogeny of ants to generate a lasting division of labor. This self-organized task-attribution system, based on an individual learning process, is particularly robust and might play an important role in colony efficiency.     

Mortality-rate crossovers and maximum lifespan in advantaged and disadvantaged populations: accelerated-mortality and sudden-death models

One population is advantaged relative to another by our definition if its survival function is greater at all ages. A population has a lifespan maximum if there is an age at which its survival function becomes exactly zero. Earlier work concerned conditions under which the mortality-rate functions of advantaged and disadvantaged populations displaying lifespan maxima always crossed. Here two survival models of populations having lifespan maxima are presented in which mortality-rate crossings between advantaged and disadvantaged subpopulations may fail to appear. One, the accelerated-mortality model, has a continuous survival function; in the other, the sudden-death model, the survival function is discontinuous. Both differ from examples examined previously in that their mortality-rate functions become infinite at their lifespan maxima.     

Fitness Consequences of Advanced Ancestral Age over Three Generations in Humans

A rapid rise in age at parenthood in contemporary societies has increased interest in reports of higher prevalence of de novo mutations and health problems in individuals with older fathers, but the fitness consequences of such age effects over several generations remain untested. Here, we use extensive pedigree data on seven pre-industrial Finnish populations to show how the ages of ancestors for up to three generations are associated with fitness traits. Individuals whose fathers, grandfathers and great-grandfathers fathered their lineage on average under age 30 were ~13% more likely to survive to adulthood than those whose ancestors fathered their lineage at over 40 years. In addition, females had a lower probability of marriage if their male ancestors were older. These findings are consistent with an increase of the number of accumulated de novo mutations with male age, suggesting that deleterious mutations acquired from recent ancestors may be a substantial burden to fitness in humans. However, possible non-mutational explanations for the observed associations are also discussed.     

What drives selection on flowering time? An experimental manipulation of the inherent correlation between genotype and environment

The optimal timing of the seasonal switch from somatic growth to reproduction can depend on an individual's condition at reproduction, the quality of the environment in which it will reproduce, or both. In annual plants, vegetative size (a function of age at flowering) affects resources available for seed production, whereas exposure to mutualists, antagonists, and abiotic stresses in the environment (functions of Julian date of flowering) influences success in converting resources into offspring. The inherent tight correlation between age, size, and environment obscures their independent fitness contributions. We isolated the fitness effects of these factors by experimentally manipulating the correlation between age at flowering and date of flowering in Brassica rapa. We staggered the planting dates of families with differing ages at flowering to produce experimental populations in which age at flowering and date of flowering were positively, negatively, or uncorrelated. In all populations, plants with an early date of flowering produced more seed than those flowering late, regardless of age or size at flowering onset. The temporal environment was thus the principal driver of selection on flowering time, but its importance relative to that of age and size varied with the presence/absence of herbivores and seed predators.     

Local Adaptation and the Effects of Isolation and Population Size – the Semelparous Perennial <Emphasis Type="Italic">Carlina vulgaris</Emphasis> as a Study Case

We made a reciprocal transplantation experiment with Carlina vulgaris, among 12 semi-natural grassland sites situated in southwest Sweden. The fate of seeds and seedlings were followed during 2 years. Local adaptation was investigated both by native superiority over non-natives, and by comparing the observed performance of a population to the fitted value of a reduced statistical model that showed the populations’ performance at all sites and the performance of all other populations at its home site. The latter method indicates presence of local adaptation even when natives are inferior to introduced populations as long as the negative difference in fitness between the populations is smaller at the native population’s home site. The strength of local adaptation was measured as the ratio of the observed to the expected performance in reduced statistical models and regressed on the degree of isolation and population size. We found no evidence of local adaptation in terms of native superiority compared to non-natives, but with the relative method we found one of six fitness components, juvenile survival, to be 6% higher for natives at their home sites compared to how they performed at other sites and how others performed at their site. Further, our results indicate that both isolation and population size have a positive effect on the process of local adaptation. Co-ordinating editor: J. Tuomi     

Longitudinal study of dominance relations among captive patas monkeys

Eight and a half years of dominance relations within a captive group of patas monkeys were analyzed. It was found that matrilineal kinship significantly influenced individuals' ranks. In contrast, with the exception of certain intramatriline changes, increasing age had no predictable effect on overall rank, at least for females (this was untestable for males). Offspring typically challenged maternal dominance and in eight of twelve dyads, offspring either rose fully over their mothers (three cases, all daughters) or at least achieved dominance ambiguity with them. Additionally, two of the four younger sisters with an opportunity to rise in rank over an older sister did so. The group dominance hierarchy was unstable for 75% of the study due to a combination of agonistically induced and demographically induced rank changes. Concentration of the highest ranks in a single matriline showed a stronger association with group hierarchy stability than did the presence of an adult, nonnatal male. Group hierarchy stability was associated with increased affiliation (sitting close and sitting touching), but otherwise there were no behavioral correlations. Individuals' ranks within the group hierarchy were unrelated to their chances of being wounded or having diarrhea. Adult females' ranks were over twice as stable as the group hierarchy (57.1% stability), but stability/instability was not correlated with any behavioral changes. Available evidence suggests that dominance relations play only a minor role in the organization of patas monkeys' intragroup behavior. Am. J. Primatol. 42:41–51, 1997. © 1997 Wiley-Liss, Inc.     

The effect of epistasis between linked genes on quantitative trait locus analysis

The effect of epistasis between linked genes on quantitative trait locus (QTL) analysis was studied as a function of their contribution to the phenotypic variance and their genetic distance by simulation of F2 (at least 200 individuals) and recombinant inbred line (RIL) populations. Data sets were replicated 100 times. For F2 populations, the presence of epistasis improves the detection of QTLs having effects in opposite directions. Epistasis between linked QTLs (26.5 cM) was poorly detected even when its contribution was relatively high compared to the main effects, and was null for heritabilities lower than 0.10. The detection of false-positive main effects is strongly affected by the distance between epistatic QTLs. The closer they are (≤11.5 cM), the higher the probability of detecting false-positive main-effect QTLs and the lower the probability of detecting epistatic effects. In this case, the presence of main-effect QTLs is due to the deviation of the heterozygote from the homozygotes at each linked interacting QTL and is algebraically explained by the joint effect of the linkage and the additive-by-additive interaction, resulting in a heterosis at a single genomic region in the absence of simulated dominant genetic effects. The number of false-positive main effects only reached nominal levels at about 100 cM. For RIL populations, the number of false positives or the detection of existing epistasis does not depend on the distance, and the power to detect epistatic QTLs is much higher even with small sample sizes and low contributions to the trait. RIL populations are highly recommended to detect epistatic QTLs and to better infer the genetic architecture of a quantitative trait.     

The evolution of British Biology: Presidential Address to the Linnean Society delivered at the Anniversary Meeting, 24 May 1983

British biology is multiply divided between 115 national societies and 900 local ones, as well as divisions between amateurs and professionals, pure and applied, universities and research institutes. It is in fact composed of four strands, three of them (natural history, scientific biology and preservation/conservation) having arisen independently, and all having given rise to the fourth (international) strand. Natural history declined at the end of the nineteenth century with the growth of professional biology, government funding, and new techniques imported from abroad; it has recovered spectacularly since the 1939–45 war. Scientific biology grew mainly from continental science, and came of age as a professional discipline around 1900. Concern for conservation has risen as human pressures on the environment have increased.
The question asked is whether this serendipitous structure is the most efficient one, in a time when the conditions which gave rise to much of it have changed: personal motivation, the role of scientific meetings, ease of travel, accessibility of libraries, sources of funding, etc. The trends have always been to split, and virtually all attempts at co-ordination have failed. This process leads to long-term inefficiency, and for scientific, political and personal advantage, the future will almost inevitably require some foci for biologists beyond narrow subject societies. The Linnean Society is peculiarly fitted to become an active focus, perhaps catalysing an amalgamation or federal sybiosis along the lines of the Royal Society of Chemistry.     

The impact of paternity on male–infant association in a primate with low paternity certainty

In multimale groups where females mate promiscuously, male–infant associations have rarely been studied. However, recent studies have shown that males selectively support their offspring during agonistic conflicts with other juveniles and that father's presence accelerates offspring maturation. Furthermore, it was shown that males invest in unrelated infants to enhance future mating success with the infant's mother. Hence, infant care might provide fitness gain for males. Here, we investigate male–infant associations in rhesus macaques (Macaca mulatta), a primate with low paternity certainty as females mate with multiple partners and males ensure paternity less efficiently through mate‐guarding. We combined behavioural data with genetic paternity analyses of one cohort of the semi‐free‐ranging population of Cayo Santiago (Puerto Rico) and recorded affiliative and aggressive interactions between focal subjects and adult males from birth to sexual maturation (0–4 years) of focal subjects. Our results revealed that 9.6% of all interactions of focal subjects involved an adult male and 94% of all male–infant interactions were affiliative, indicating the rareness of male–infant aggression. Second and most interestingly, sires were more likely to affiliate with their offspring than nonsires with unrelated infants. This preference was independent of mother's proximity and emphasized during early infancy. Male–infant affiliation rose with infant age and was pronounced between adult males and male rather than female focal subjects. Overall, our results suggest that male–infant affiliation is also an important component in structuring primate societies and affiliation directed towards own offspring presumably represent low‐cost paternal care.     

Prevalence of symptoms of Raynaud's phenomenon in general practice.

OBJECTIVE--To investigate the prevalence of Raynaud''s phenomenon in the populations of five general practices. DESIGN--Two populations studied. A questionnaire was given to all new patients attending five general practices over four weeks, and the same questionnaire was sent by post to a random sample of adults from two of the practices. SETTING--General practices in inner London, Merseyside, and Cheshire. SUBJECTS--1532 Patients who completed questionnaires (1119 who attended the surgeries (response rate unknown) and 413 respondents to the postal survey (response rate 69%)). MAIN OUTCOME MEASURES--Response to questionnaire on symptoms of Raynaud''s phenomenon: patients were regarded as having the disease if they had episodes of blanching of the fingers that were precipitated by cold and accompanied by sensory symptoms (pins and needles or numbness). Subsequent interview and clinical appraisal of patients with the disease according to their responses to the questionnaire. RESULTS--The prevalence of Raynaud''s phenomenon was 11% (26/231) and 19% (34/182) respectively in men and women who completed the postal questionnaire and 16% (56/357) and 21% (157/762) respectively in those who completed the questionnaire when attending their general practice. Thus the overall rates were slightly higher in women, but there was no effect of age even after adjustment of the rates for practice and method of survey. CONCLUSION--The prevalence of Raynaud''s phenomenon is high compared with the low number of patients who seek treatment for the disease.     

Human population growth and the demographic transition

The world and most regions and countries are experiencing unprecedentedly rapid demographic change. The most obvious example of this change is the huge expansion of human numbers: four billion have been added since 1950. Projections for the next half century expect a highly divergent world, with stagnation or potential decline in parts of the developed world and continued rapid growth in the least developed regions. Other demographic processes are also undergoing extraordinary change: women''s fertility has dropped rapidly and life expectancy has risen to new highs. Past trends in fertility and mortality have led to very young populations in high fertility countries in the developing world and to increasingly older populations in the developed world. Contemporary societies are now at very different stages of their demographic transitions. This paper summarizes key trends in population size, fertility and mortality, and age structures during these transitions. The focus is on the century from 1950 to 2050, which covers the period of most rapid global demographic transformation.