Optimizing α for better statistical decisions: A case study involving the pace‐of‐life syndrome hypothesis

Setting optimal significance levels that minimize Type I and Type II errors allows for more transparent and well‐considered statistical decision making compared to the traditional α = 0.05 significance level. We use the optimal α approach to re‐assess conclusions reached by three recently published tests of the pace‐of‐life syndrome hypothesis, which attempts to unify occurrences of different physiological, behavioral, and life history characteristics under one theory, over different scales of biological organization. While some of the conclusions reached using optimal α were consistent to those previously reported using the traditional α = 0.05 threshold, opposing conclusions were also frequently reached. The optimal α approach reduced probabilities of Type I and Type II errors, and ensured statistical significance was associated with biological relevance. Biologists should seriously consider their choice of α when conducting null hypothesis significance tests, as there are serious disadvantages with consistent reliance on the traditional but arbitrary α = 0.05 significance level.     

Macroevolutionary evidence suggests trait‐dependent coevolution between behavior and life‐history

Species with fast life‐histories typically prioritize current over future reproductive events, compared to species with slow life‐histories. These species therefore require greater energetic input into reproduction, and also likely have less time to realize their reproductive potential. Hence, behaviors that increase access to both resources and mating opportunities, at a cost of increased mortality risk, could coevolve with the pace of life‐history. However, whether this prediction holds across species, remains untested under standardized conditions. Here, we test how risky behaviors, which facilitate access to resources and mating opportunities (i.e., activity, boldness, and aggression), along with metabolic rate, coevolve with the pace of life‐history across 20 species of killifish that present remarkable divergences in the pace of life‐history. We found a positive association between the pace of life‐history and aggression, but interestingly not with other behavioral traits or metabolic rate. Aggression is linked to interference competition, and in killifishes is often employed to secure mates, while activity and boldness are more relevant for exploiting energetic resources. Our results suggest that the trade‐off between current and future reproduction plays a more prominent role in shaping mating behavior, while behaviors related to energy acquisition may be influenced by ecological factors.     

Environmental effects on the covariation among pace‐of‐life traits

Pace‐of‐life syndromes (POLSs) are suites of life‐history, physiological and behavioural traits that arise due to trade‐offs between allocation to current and future reproduction. Traits generally show covariation that can arise from genetic and environmental influences on phenotypes and constrain the independent evolution of traits, resulting in fitness consequences and impacts on population dynamics. The notion that correlations among traits may vary among populations along environmental gradients suggests an important role for the environment in shaping and maintaining POLSs. However, no synthesis has been attempted of the myriad ways in which environmental factors should influence POLSs. Here, we formulate a series of hypotheses targeting the critical interfaces of the environment and life‐history ‐ behaviour associations across different organisms. We discuss the hypotheses in light of findings from a systematic review of studies that measured changes in the association between behaviour and life‐history traits as a function of environmental conditions. The review revealed that POLSs are often shaped by environmental variation, where harshness of the environment in early life has the most consistent effects on POLS. However, only partial or no effects of environmental variation were found in a number of studies, which may result from the highly variable study systems, traits and environments studied. We highlight promising directions arising from the available studies and identify knowledge gaps that, if unaddressed, will impede progress in the field.     

Potential ‘costs of reproduction’ in a skink: Inter‐ and intrapopulational variation

Reproductive costs are important determinants of reproductive effort in squamate reptiles. Consequently, differences in costs of reproduction between populations of geographically or climatically widespread species are likely to result in different patterns of reproductive effort. In the present study, the effect of pregnancy on sprint speed was examined in a small viviparous skink, Niveoscincus ocellatus (Gray 1845), from two populations at the climatic extremes of its distribution. Decreased sprint speed has the potential to be an important cost of reproduction in this species, through a reduced ability to avoid predation and/or decreased foraging efficiency. Lizards inhabiting the colder site were larger than those from the warmer site and, contrary to predictions from life history theory, had a higher reproductive effort. In both populations, sprint speed was lower in pregnant lizards than in either the same individuals after birth or non‐pregnant control lizards. Within each population, sprint speed was unrelated to the level of reproductive effort of the female in terms of either absolute mass of the reproductive burden or the burden relative to her post‐partum body mass. However, within each population, the mass of the clutch that an individual female was carrying relative to snout–vent length was an important determinant of her sprint speed while pregnant. Thus, within each population, a relatively high reproductive burden may potentially increase costs of reproduction in this species. Despite this relationship and predictions from life history theory suggesting that annual reproductive effort will be lower in populations with a large body size and delayed maturity, it is suggested that a higher reproductive effort at the cold site is possible because they have a higher absolute sprint speed because of their larger size and a relatively higher abundance of cover at the cold site, and differences in predation pressure may alter selective pressures on reproductive investment.     

Revisiting the ‘bank of microscopic forms’ in macroalgal‐dominated ecosystems

Theoretical ecological models, such as succession and facilitation, were defined in terrestrial habitats, and subsequently applied to marine and freshwater habitats in intertidal and then subtidal realms. One such model is the soil seed bank, defined as all viable seeds (or fruits) found near the soil surface that facilitate community restoration/recovery. “Banks of microscopic forms” have been hypothesized in aquatic habitats and recent work from aquaculture has highlighted dormancy in algal life cycle stages. To reinvigorate the discussions about these algal banks, we discuss differences in life cycles, dispersal, and summarize research on banks of macroalgal stages in aquatic ecosystems that may be easier to explore with modern advances in molecular technology. With focus on seminal work in global kelp forest ecosystems, we present a pilot study in northern California as proof of concept that Nereocystis luetkeana and Alaria marginata stages can be detected within kelp forests in the biofilm of rocks and bedrock using targeted primers long after zoospore release. Considering the increased interest in algae as an economic resource, [blue] carbon sink, and as ecosystem engineers, the potential for “banking” macroalgal forms could be a mechanism of resilience and recovery in aquatic populations that have complex life cycles and environmental cues for reproduction. Molecular barcoding is becoming an important tool for identifying banks of macroalgal forms in marine communities. Understanding banks of macroalgal stages, especially in deforested habitats with intense disturbance and grazer pressure, will allow researchers and marine resource managers to facilitate this natural process in recovery of the aquatic system.     

Life‐history strategy,with ctenidial and pallial larval brooding,of the troglodytic ‘living fossil’ Congeria kusceri (Bivalvia: Dreissenidae) from the subterranean Dinaric Alpine karst of Croatia

Among freshwater bivalves, the brooding of embryos and larvae within the maternal ctenidia is well known. Exceptions to this generalization are the non‐brooding freshwater and estuarine species of Dreissena and Mytilopsis, respectively. It was reported that the freshwater troglodytic cousin, Congeria kusceri Bole, 1962, of these dreissenids does not brood either. It is herein demonstrated that C. kusceri undergoes one reproductive cycle each year. Sexes are separate, with an early male and later female bias. A small percentage (2.14%) of individuals is hermaphroditic. The gonads mature over summer from May to November. Spawning commences in September, when females release mature oocytes into their ctenidia and inhale sperm from mature males. Here the oocytes are fertilized, and develop within interfilamentary marsupia. Ctenidial tissues glandularize, and may provide a source of maternal nutrition for the embryos. At the late prodissoconch‐1 or prodissoconch‐2 stage (PR2, ~220 μm), larvae are released into the infrabranchial chamber via a birth channel along the outer edge of the ventral marginal food groove of both inner demibranchs. Here, they are brooded further in mantle pouches located beneath the inhalant siphon. Subsequently, after the PR2 stage (nepioconch/dissoconch), they are released from the inhalant siphon and assume an independent life as crawling juveniles. Such juveniles may be found amongst clusters of adults. Not only is C. kusceri unique amongst the Dreissenidae in possessing the capacity to brood internally fertilized ova, but it is also exceptional amongst the Bivalvia in possessing the described methods of brooding and birth. Explanations for both lie in its troglodytic lifestyle, decadal length longevity and habitat: that of byssal attachment to the hard surfaces of underground freshwater rivers, caves, pits, and sinkholes in the Tethyan arc of the Dinaric karst. Internal fertilization of a few large yolky eggs, lecithotrophic larvae, ctenidial brooding, and secondary pallial parental care represent relatively recent, Late Miocene, evolutionary adaptations from a Tethyan lentic ancestor.     

Re‐examining extreme longevity of the cave crayfish Orconectes australis using new mark–recapture data: a lesson on the limitations of iterative size‐at‐age models

1. Centenarian species, defined as those taxa with life spans that frequently exceed 100 years, have long been of interest to ecologists because they represent an extreme end point in a continuum of life history strategies. One frequently reported example of a freshwater centenarian is the obligate cave crayfish Orconectes australis, with a maximum longevity reported to exceed 176 years. As a consequence of its reported longevity, O. australis has been used as a textbook example of life history adaptation to the organic‐carbon limitation that characterises many cave‐stream food webs. 2. Despite being widely reported, uncertainties surround the original estimates of longevity for O. australis, which were based on a single study dating from the mid‐1970s. In the present study, we re‐evaluated the growth rate, time‐to‐maturity, female age‐at‐first‐reproduction and longevity of O. australis using a mark–recapture study of more than 5 years based upon more than 3800 free‐ranging individuals from three isolated cave streams in the south‐eastern United States. 3. The results of our study indicate that accurate estimates of the longevity of O. australis are ≤22 years, with only a small proportion of individuals (<5%) exceeding this age. Our estimates for female time‐to‐maturity (4–5 years) and age‐at‐first‐reproduction (5–6 years) are also substantially lower than earlier estimates. 4. These new data indicate that the age thresholds for life history events of O. australis are comparable to other estimates for a modest assemblage of cave and surface species of crayfish for which credible age estimate exists, suggesting that a cave environment per se is not required for the evolution of extreme longevity in crayfish.     

A mother’s legacy: the strength of maternal effects in animal populations

Although mothers influence the traits of their offspring in many ways beyond the transmission of genes, it remains unclear how important such ‘maternal effects’ are to phenotypic differences among individuals. Synthesizing estimates derived from detailed pedigrees, we evaluated the amount of phenotypic variation determined by maternal effects in animal populations. Maternal effects account for half as much phenotypic variation within populations as do additive genetic effects. Maternal effects most greatly affect morphology and phenology but, surprisingly, are not stronger in species with prolonged maternal care than in species without. While maternal effects influence juvenile traits more than adult traits on average, they do not decline across ontogeny for behaviour or physiology, and they do not weaken across the life cycle in species without maternal care. These findings underscore maternal effects as an important source of phenotypic variation and emphasise their potential to affect many ecological and evolutionary processes.     

A review of predation as a limiting factor for bird populations in mesopredator‐rich landscapes: a case study of the UK

The impact of increasing vertebrate predator numbers on bird populations is widely debated among the general public, game managers and conservationists across Europe. However, there are few systematic reviews of whether predation limits the population sizes of European bird species. Views on the impacts of predation are particularly polarised in the UK, probably because the UK has a globally exceptional culture of intensive, high‐yield gamebird management where predator removal is the norm. In addition, most apex predators have been exterminated or much depleted in numbers, contributing to a widely held perception that the UK has high numbers of mesopredators. This has resulted in many high‐quality studies of mesopredator impacts over several decades. Here we present results from a systematic review of predator trends and abundance, and assess whether predation limits the population sizes of 90 bird species in the UK. Our results confirm that the generalist predators Red Fox (Vulpes vulpes) and Crows (Corvus corone and C. cornix) occur at high densities in the UK compared with other European countries. In addition, some avian and mammalian predators have increased numerically in the UK during recent decades. Despite these high and increasing densities of predators, we found little evidence that predation limits populations of pigeons, woodpeckers and passerines, whereas evidence suggests that ground‐nesting seabirds, waders and gamebirds can be limited by predation. Using life‐history characteristics of prey species, we found that mainly long‐lived species with high adult survival and late onset of breeding were limited by predation. Single‐brooded species were also more likely to be limited by predation than multi‐brooded species. Predators that depredate prey species during all life stages (i.e. from nest to adult stages) limited prey numbers more than predators that depredated only specific life stages (e.g. solely during the nest phase). The Red Fox and non‐native mammals (e.g. the American Mink Neovison vison) were frequently identified as numerically limiting their prey species. Our review has identified predator–prey interactions that are particularly likely to result in population declines of prey species. In the short term, traditional predator‐management techniques (e.g. lethal control or fencing to reduce predation by a small number of predator species) could be used to protect these vulnerable species. However, as these techniques are costly and time‐consuming, we advocate that future research should identify land‐use practices and landscape configurations that would reduce predator numbers and predation rates.     

Toll‐like receptor variation in the bottlenecked population of the Seychelles warbler: computer simulations see the ‘ghost of selection past’ and quantify the ‘drift debt’

Balancing selection can maintain immunogenetic variation within host populations, but detecting its signal in a postbottlenecked population is challenging due to the potentially overriding effects of drift. Toll‐like receptor genes (TLRs) play a fundamental role in vertebrate immune defence and are predicted to be under balancing selection. We previously characterized variation at TLR loci in the Seychelles warbler (Acrocephalus sechellensis), an endemic passerine that has undergone a historical bottleneck. Five of seven TLR loci were polymorphic, which is in sharp contrast to the low genomewide variation observed. However, standard population genetic statistical methods failed to detect a contemporary signature of selection at any TLR locus. We examined whether the observed TLR polymorphism could be explained by neutral evolution, simulating the population's demography in the software DIYABC. This showed that the posterior distributions of mutation rates had to be unrealistically high to explain the observed genetic variation. We then conducted simulations with an agent‐based model using typical values for the mutation rate, which indicated that weak balancing selection has acted on the three TLR genes. The model was able to detect evidence of past selection elevating TLR polymorphism in the prebottleneck populations, but was unable to discern any effects of balancing selection in the contemporary population. Our results show drift is the overriding evolutionary force that has shaped TLR variation in the contemporary Seychelles warbler population, and the observed TLR polymorphisms might be merely the ‘ghost of selection past’. Forecast models predict immunogenetic variation in this species will continue to be eroded in the absence of contemporary balancing selection. Such ‘drift debt’ occurs when a gene pool has not yet reached its new equilibrium level of polymorphism, and this loss could be an important threat to many recently bottlenecked populations.     

Cycles of coral reef ‘turn‐on’, rapid growth and ‘turn‐off’ over the past 8500 years: a context for understanding modern ecological states and trajectories

Human activities threaten reef ecosystems globally, forcing ecological change at rates and scales regarded as unprecedented in the Holocene. These changes are so profound that a cessation of reef accretion (reef ‘turn‐off’) and net erosion of reef structures is argued by many as the ultimate and imminent trajectory. Here, we use a regional scale reef growth dataset, based on 76 core records (constrained by 211 radiometric dates) from 22 reefs along and across the inner‐shelf of the Great Barrier Reef, Australia, to examine the timing of different phases of reef initiation (‘turn‐on’), growth and ‘turn‐off’ during the Holocene. This dataset delineates two temporally discrete episodes of reef‐building over the last 8500 years: the first associated with the Holocene transgression‐early highstand period [～8.5–5.5 k calibrated years bp (cal ybp )]; the second since ～2.3 k cal ybp . During both periods, reefs accreted rapidly to sea level before entering late evolutionary states – states naturally characterized by reduced coral cover and low accretion potential – and a clear hiatus occurs between these reef‐building episodes for which no records of reef initiation exist. These transitions mimic those projected under current environmental disturbance regimes, but have been driven entirely by natural forcing factors. Our results demonstrate that, even through the late Holocene, reef health and growth has fluctuated through cycles independent of anthropogenic forcing. Consequently, degraded reef states cannot de facto be considered to automatically reflect increased anthropogenic stress. Indeed, in many cases degraded or nonaccreting reef communities may reflect past reef growth histories (as dictated by reef growth–sea level interactions) as much as contemporary environmental change. Recognizing when changes in reef condition reflect these natural ‘turn‐on’– growth –‘turn‐off’ cycles and how they interact with on‐going human disturbance is critical for effective coral reef management and for understanding future reef ecological trajectories.     

Non‐specificity of ethylene inhibitors: ‘double‐edged’ tools to find out new targets involved in the root morphogenetic programme

In the last decade, genetic and pharmacological approaches have been used to explore ethylene biosynthesis and perception in order to study the role of ethylene and ethylene/auxin interaction in root architecture development. However, recent findings with pharmacological approaches highlight the non‐specificity of commonly used inhibitors. This suggests that caution is required for interpreting these studies and that the use of pharmacological agents is a ‘double‐edged’ tool. On one hand, non‐specific effects make interpretation difficult unless other experiments, such as with different mutants or with multiple diversely acting chemicals, are conducted. On the other hand, the non‐specificity of inhibitors opens up the possibility of uncovering some ligands or modulators of new receptors such as plant glutamate‐like receptors and importance of some metabolic hubs in carbon and nitrogen metabolism such as the pyridoxal phosphate biosynthesis involved in the regulation of the root morphogenetic programme. Identification of such targets is a critical issue to improve the efficiency of absorption of macronutrients in relation to root the morphogenetic programme.     

The evolutionary dynamics of timing of maternal immunity: evaluating the role of age‐specific mortality

If a female survives an infection, she can transfer antibodies against that particular pathogen to any future offspring she produces. The resulting protection of offspring for a period after their birth is termed maternal immunity. Because infection in newborns is associated with high mortality, the duration of this protection is expected to be under strong selection. Evolutionary modelling structured around a trade‐off between fertility and duration of maternal immunity has indicated selection for longer duration of maternal immunity for hosts with longer lifespans. Here, we use a new modelling framework to extend this analysis to consider characteristics of pathogens (and hosts) in further detail. Importantly, given the challenges in characterizing trade‐offs linked to immune function empirically, our model makes no assumptions about costs of longer lasting maternal immunity. Rather, a key component of this analysis is variation in mortality over age. We found that the optimal duration of maternal immunity is shaped by the shifting balance of the burden of infection between young and old individuals. As age of infection depends on characteristics of both the host and the pathogen, both affect the evolution of duration of maternal immunity. Our analysis provides additional support for selection for longer duration of maternal immunity in long‐lived hosts, even in the absence of explicit costs linked to duration of maternal immunity. Further, the scope of our results provides explanations for exceptions to the general correlation between duration of maternal immunity and lifespan, as we found that both pathogen characteristics and trans‐generational effects can lead to important shifts in fitness linked to maternal immunity. Finally, our analysis points to new directions for quantifying the trade‐offs that drive the development of the immune system.     

A dubious defense of ‘after‐birth abortion’: A reply to Räsänen

Scholars have offered various critiques of Alberto Giubilini and Francesca Minerva's controversial article, ‘After‐birth abortion: Why should the baby live?’ My book The Ethics of Abortion: Women's Rights, Human Life, and the Question of Justice presents four such critiques. First, Giubilini and Minerva argue from the deeply controversial to the even more controversial. Second, they presuppose a false view of personal identity called body‐self dualism. Third, their view cannot secure human equality. And fourth, their account of harm cannot account for harm found in some cases of murder. In the article, ‘Pro‐life arguments against infanticide and why they are not convincing’, J. Räsänen examines and finds wanting these four critiques. This essay responds to Räsänen's defense of infanticide and argues that his responses to the four objections fail.     

The temporal ‘structure’ and function of the insect photoperiodic clock: a tribute to Colin S. Pittendrigh

In 1936, Erwin Bünning suggested that photoperiodic time measurement was a function of the circadian system. Colin Pittendrigh became an ardent supporter of Bünning's hypothesis, drawing parallels between photoperiodism and his own group's investigations of adult eclosion rhythmicity in the fruit fly Drosophila pseudoobscura. They developed several more modern versions of Bünning's general hypothesis based on the entrainment of circadian oscillations to the light cycle, including ‘external coincidence’, which is a derivation of Bünning's original model, and ‘internal coincidence’, which relied upon seasonal changes in the mutual phase relationship of oscillators within a multi‐oscillator circadian system. This review considers the experimental evidence for the central role of the circadian system in photoperiodic timing and, in some species, for both external and internal coincidence. Pittendrigh, however, pursued the idea of internal coincidence further with his analysis of the pacemaker–slave organization of eclosion rhythmicity in D. pseudoobscura and proposed a similar theoretical model for photoperiodism comprising a group of slave oscillators driven by a light‐sensitive pacemaker. In this model, the phase relationships of the slaves to the pacemaker were affected by (i) the relative periods of the pacemaker and slave(s); (ii) the strength(s) of the coupling between the two; and (iii) the dampening coefficients of the various slaves. Manipulation of these variables showed that the slaves adopted different internal phase relationships (both to each other and to the pacemaker) under the influence of changes in daily photophase, the period of the Zeitgeber and phase shifts of the entraining light cycle.     

The North Atlantic subpolar gyre and the marine migration of Atlantic salmon Salmo salar: the ‘Merry‐Go‐Round’ hypothesis

One model for marine migration of Atlantic salmon Salmo salar proposes that North American and southern European stocks (<62° N) move directly to feeding grounds off west Greenland, then overwinter in the Labrador Sea, whereas northern European stocks (>62° N) utilize the Norwegian Sea. An alternate model proposes that both North American and European stocks migrate in the North Atlantic Subpolar Gyre (NASpG) where S. salar enter the NASpG on their respective sides of the Atlantic, and travel counterclockwise within the NASpG until returning to natal rivers. A review of data accumulated during the last 50 years suggests a gyre model is most probable. Freshwater parr metamorphose into smolts which have morphological, physiological and behavioural adaptations of epipelagic, marine fishes. Former high‐seas fisheries were seasonally sequential and moved in the direction of NASpG currents, and catches were highest along the main axis of the NASpG. Marking and discrimination studies indicate mixed continental origin feeding aggregations on both sides of the Atlantic. Marked North American smolts were captured off Norway, the Faroe Islands, east and west Greenland, and adults tagged at the Faroes were recovered in Canadian rivers. Marked European smolts were recovered off Newfoundland and Labrador, west and east Greenland, and adults tagged in the Labrador Sea were captured in European rivers. High Caesium‐137 (¹³⁷Cs) levels in S. salar returning to a Quebec river suggested 62·3% had fed at or east of Iceland, whereas levels in 1 sea‐winter (SW) Atlantic Canada returnees indicated 24·7% had fed east of the Faroes. Lower levels of ¹³⁷ Cs in returning 1SW Irish fish suggest much of their growth occurred in the western Atlantic. These data suggest marine migration of S. salar follows a gyre model and is similar to other open‐ocean migrations of epipelagic fishes.     

Sensitive and selective detection of Cu(II) ion: A new effective 1,8‐naphthalimide‐based fluorescence ‘turn off’ sensor

The present study reports the development of a new 1,8‐naphthalimide‐based fluorescent sensor V for monitoring Cu(II) ions. The sensor exhibited pH independence over a wide pH range 2.52–9.58, and indicated its possible use for monitoring Cu(II) ions in a competitive pH medium. The sensor also showed high selectivity and sensitivity towards the Cu(II) ions over other competitive metal ions in DMSO–HEPES buffer (v/v, 1:1; pH 7.4) with a fluorescence ‘turn off’ mode of 79.79% observed. A Job plot indicated the formation of a 1:1 binding mode of the sensor with Cu(II) ions. The association constant and detection limit were 1.14 × 10⁶ M^–1 and 4.67 × 10^–8 M, respectively. The fluorescence spectrum of the sensor was quenched due to the powerful paramagnetic nature of the Cu(II) ions. Potential application of this sensor was also demonstrated when determining Cu(II) ion levels in two different water samples.     

Hypofolins A – L,ent‐Labdane Diterpenoids from the Roots of Hypoestes phyllostachya ‘Pink Splash’

Twelve new ent‐labdane diterpenoids, hypofolins A – F ( 1 – 6 ) and hypofolins G – L ( 7a / 7b , 8a / 8b , and 9a / 9b ), were isolated from the roots of Hypoestes phyllostachya ‘Pink Splash’. Their structures were elucidated by extensive 1D‐ and 2D‐NMR spectroscopic and HR‐MS data. The absolute configurations of 1 , 2 , 5 , and 7a / 7b were determined by single crystal X‐ray diffraction and ECD analysis, as well as chemical transformations. Compounds 7a / 7b , 8a / 8b , and 9a / 9b were isolated as three pairs of interconverting mixture of two isomers between ketone and hemiketal types. Compound 1 showed weak cytotoxicity against SMMC‐7721 cell line with IC₅₀ value of 31.40 μm .