Selection on defensive traits in a sterile caste – caste evolution: a mechanism to overcome life‐history trade‐offs?

SUMMARY During development and evolution individuals generally face a trade-off between the development of weapons and gonads. In termites, characterized by reproductive division of labor, a caste evolved—the soldiers—which is completely sterile and which might be released from developmental trade-offs between weapons and testes. These soldiers are exclusively dedicated to defense. First, we investigated whether defensive traits are under selection in sterile termite soldiers using allometric analyses. In soldiers of the genus Cryptotermes phragmotic traits such as a sculptured and foreshortened head evolve rapidly but were also lost twice. Second, we compared the scaling relationships of these weapons with those in solitary insects facing a trade-off between weapons and gonads. Defensive traits consistently had lower slopes than nondefensive traits which supports the existence of stabilizing selection on soldier phragmotic traits in order to plug galleries. Moreover, soldier head widths were colony specific and correlated with the minimum gallery diameter of a colony. This can proximately be explained by soldiers developing from different instars. The scaling relationships of these termite soldiers contrast strikingly with those of weapons of solitary insects, which are generally exaggerated (i.e., overscaling) male traits. These differences may provide important insights into trait evolution. Trade-offs constraining the development of individuals may have been uncoupled in termites by evolving different castes, each specialized for one function. When individuals in social insect are "released" from developmental constraints through the evolution of castes, this certainly contributed to the ecological and evolutionary success of social insects.     

A review of urban impacts on avian life‐history evolution: Does city living lead to slower pace of life?

The concept of a pace‐of‐life syndrome describes inter‐ and intraspecific variation in several life‐history traits along a slow‐to‐fast pace‐of‐life continuum, with long lifespans, low reproductive and metabolic rates, and elevated somatic defences at the slow end of the continuum and the opposite traits at the fast end. Pace‐of‐life can vary in relation to local environmental conditions (e.g. latitude, altitude), and here we propose that this variation may also occur along an anthropogenically modified environmental gradient. Based on a body of literature supporting the idea that city birds have longer lifespans, we predict that urban birds have a slower pace‐of‐life compared to rural birds and thus invest more in self maintenance and less in annual reproduction. Our statistical meta‐analysis of two key traits related to pace‐of‐life, survival and breeding investment (clutch size), indicated that urban birds generally have higher survival, but smaller clutch sizes. The latter finding (smaller clutches in urban habitats) seemed to be mainly a characteristic of smaller passerines. We also reviewed urbanization studies on other traits that can be associated with pace‐of‐life and are related to either reproductive investment or self‐maintenance. Though sample sizes were generally too small to conduct formal meta‐analyses, published literature suggests that urban birds tend to produce lower‐quality sexual signals and invest more in offspring care. The latter finding is in agreement with the adult survival hypothesis, proposing that higher adult survival prospects favour investment in fewer offspring per year. According to our hypothesis, differences in age structure should arise between urban and rural populations, providing a novel alternative explanation for physiological differences and earlier breeding. We encourage more research investigating how telomere dynamics, immune defences, antioxidants and oxidative damage in different tissues vary along the urbanization gradient, and suggest that applying pace‐of‐life framework to studies of variation in physiological traits along the urbanization gradient might be the next direction to improve our understanding of urbanization as an evolutionary process.     

Has a river dam affected the life‐history traits of a freshwater prawn?

In recent years, species richness and diversity in aquatic ecosystems has declined as environments are increasingly impacted by anthropic actions. Freshwater prawns are well adapted to survive in a disturbed and heterogeneous environment. For instance, Amazon river prawn (Macrobrachium amazonicum) populations vary in migratory behavior between rivers and estuaries, depending on factors such as dams. However, there is limited information on the influence of environmental conditions on life‐history traits of this species, which we investigate here using two distinct and unconnected aquatic systems, a dammed river and an estuary, in the eastern Brazilian Amazon. The biological characteristics of M. amazonicum populations in the two environments were compared and related to environmental parameters, which differed significant differences between the two environments and between seasons. Dissolved oxygen, precipitation, and temperature varied most significantly with the seasons in both the estuary and river. M. amazonicum prawns in the estuary were larger and heavier than those in the river during rainy periods. The mass–length ratios and condition factor varied significantly between the M. amazonicum populations in the estuary and river, with negative allometric growth (grows faster in length than in weight) predominating in both populations, and condition factor was better in the estuary for males and in the river for females. The relative frequencies of occurrence of the different female maturation stages and the male morphotypes were related to precipitation and turbidity in both environments and also to salinity in the estuary. In these two distinct aquatic systems, the abiotic parameters determined by the seasonal precipitation cycle profoundly influenced the development of this crustacean, despite its ecological plasticity. Overall, the study showed that river damming triggered environmental changes in the freshwater river ecosystem and played a key role in determining the life‐history characteristics of M. amazonicum in these contrasting aquatic systems.     

Germline stem cell maintenance as a proximate mechanism of life‐history trade‐offs?

We suggest that the commonly observed trade-offs between early- and late-life reproduction may be mediated by genetic variation in germline stem cell maintenance. Stem cell biology provides a natural framework and experimental methods for understanding the mechanistic basis of life-history evolution. At the same time, natural variation in life-history strategies can serve as a powerful tool for identifying the genes and molecular pathways involved in the maintenance of stem cells in aging adults. We illustrate the connections between life-history and stem cells with examples drawn primarily from Drosophila melanogaster and Caenorhabditis elegans, and suggest a number of testable hypotheses and avenues for future investigation that can be addressed with existing models and tools.     

The evolution of ecological specialization in southern African ungulates: competition‐ or physical environmental turnover?

Using long‐term diet reconstructions spanning the past one million years, we contrast hypotheses that biotic interactions versus physical environmental changes are primary drivers of evolutionary turnover in mammals. We use stable carbon (δ¹³C) and oxygen (δ¹⁸O) isotope ratios in tooth enamel carbonate to trace herbivore niche shifts through the Late Quaternary Land Mammal Ages (LMAs) of grassland savannas in the South African interior (Cornelian‐1.0 to 0.6 Ma; Florisian‐500 to 10 ka; and Holocene/modern). Data reveal niche separation amongst closely related coeval taxa, and dispersals through time into empty niche spaces following extinctions. This suggests a primary role of competitive exclusion and niche displacement for speciation and extinctions in these early grassland environments. However, niche changes through time show a similar trend in many taxa, entailing increased δ¹³C (elevated C₄ grass consumption) from the Cornelian to the Florisian, and from the Florisian to the Holocene/modern, and elevated δ¹⁸O in Holocene/modern taxa that reflect global aridification around the terminal Pleistocene. Commonality in isotopic trends implies universal environmental forcing of ecological, and ultimately macroevolutionary, turnover. Yet some taxa shift from a mixed C₃/C₄ diet in the Florisian to a near‐pure C₃ diet today. Indeed, we find that while δ¹³C data are normally distributed for Cornelian fossils, non‐normal distributions characterize more recent time intervals. Such distributions are in line with the bimodal distribution of δ¹³C and diet in contemporary African ungulates. Thus, while environmental forcing did not, by necessity, lead to increases in C₄ intake, the results show changes from mixed to more specialized diets. We propose that this niche specialization was a function of long‐term exposure to C₄ grasslands, consistent with predictions that relatively high metabolic demands of C₄ grazing in subtropical environments forced the differentiation of herbivores into one of two highly specialized feeding niches, i.e. C₃ browsing or C₄ grazing.     

Multispecies comparisons of adaptability to climate change: A role for life‐history characteristics?

Phenological advancement allows individuals to adapt to climate change by timing life‐history events to the availability of key resources so that individual fitness is maximized. However, different trophic levels may respond to changes in their environment at different rates, potentially leading to a phenological mismatch. This may be especially apparent in the highly seasonal arctic environment that is experiencing the effects of climate change more so than any other region. During a 14‐year study near Utqia?vik (formerly Barrow), Alaska, we estimated phenological advancement in egg laying in relation to snowmelt for eight arctic‐breeding shorebirds and investigated potential linkages to species‐specific life‐history characteristics. We found that snowmelt advanced 0.8 days/year—six times faster than the prior 60‐year period. During this same time, six of the eight species exhibited phenological advancement in laying dates (varying among species from 0.1 to 0.9 days earlier per year), although no species appeared capable of keeping pace with advancing snowmelt. Phenological changes were likely the result of high phenotypic plasticity, as all species investigated in this study showed high interannual variability in lay dates. Commonality among species with similar response rates to timing of snowmelt suggests that nesting later and having an opportunistic settlement strategy may increase the adaptability of some species to changing climate conditions. Other life‐history characteristics, such as migration strategy, previous site experience, and mate fidelity did not influence the ability of individuals to advance laying dates. As a failure to advance egg laying is likely to result in greater phenological mismatch, our study provides an initial assessment of the relative risk of species to long‐term climatic changes.     

Do differences in life‐history traits and the timing of peak mating activity between host‐associated populations of Chilo suppressalis have a genetic basis?

The development of host races, genetically distinct populations of the same species with different hosts, is considered to be the initial stage of ecological speciation. Ecological and biological differences consistent with host race formation have been reported between water‐oat and rice‐associated populations of Chilo suppressalis. In order to confirm whether these differences have a genetic basis, we conducted experiments to determine the extent to which various life‐history traits and the time of peak mating activity of these populations were influenced by the species of host plant larvae were raised on. Individuals from each population were reared for three consecutive generations on either water‐oat fruit pulp or rice seedlings. Descendants of both populations had higher larval survival rates, shorter larval developmental periods, higher pupal weight, and longer adult forewings, when reared on water‐oats than when reared on rice. The time of peak of mating activity differed between the descendants of each population, irrespective of whether they were raised on water‐oats or rice. These results indicate that although some life‐history traits of host‐associated populations of C. suppressalis are influenced by the host plant larvae are raised on, time of peak mating activity is not. Because it is a stable, objective, phenotypic trait, further research on difference in the time of peak mating activity between host‐associated populations of C. suppressalis should be conducted to clarify the mechanism responsible for host race formation in this species.     

Do different disparity proxies converge on a common signal? Insights from the cranial morphometrics and evolutionary history of Pterosauria (Diapsida: Archosauria)

Disparity, or morphological diversity, is often quantified by evolutionary biologists investigating the macroevolutionary history of clades over geological timescales. Disparity is typically quantified using proxies for morphology, such as measurements, discrete anatomical characters, or geometric morphometrics. If different proxies produce differing results, then the accurate quantification of disparity in deep time may be problematic. However, despite this, few studies have attempted to examine disparity of a single clade using multiple morphological proxies. Here, as a case study for this question, we examine the disparity of the volant Mesozoic fossil reptile clade Pterosauria, an intensively studied group that achieved substantial morphological, ecological and taxonomic diversity during their 145+ million-year evolutionary history. We characterize broadscale patterns of cranial morphological disparity for pterosaurs for the first time using landmark-based geometric morphometrics and make comparisons to calculations of pterosaur disparity based on alternative metrics. Landmark-based disparity calculations suggest that monofenestratan pterosaurs were more diverse cranially than basal non-monofenestratan pterosaurs (at least when the aberrant anurognathids are excluded), and that peak cranial disparity may have occurred in the Early Cretaceous, relatively late in pterosaur evolution. Significantly, our cranial disparity results are broadly congruent with those based on whole skeleton discrete character and limb proportion data sets, indicating that these divergent approaches document a consistent pattern of pterosaur morphological evolution. Therefore, pterosaurs provide an exemplar case demonstrating that different proxies for morphological form can converge on the same disparity signal, which is encouraging because often only one such proxy is available for extinct clades represented by fossils. Furthermore, mapping phylogeny into cranial morphospace demonstrates that pterosaur cranial morphology is significantly correlated with, and potentially constrained by, phylogenetic relationships.     

Co‐evolution of marine worms and their chemoautotrophic bacterial symbionts: unexpected host switches explained by ecological fitting?

Mutualistic associations of bacteria and invertebrates are widespread and encompass an enormous diversity on the side of both partners. The advantages gained from the symbiosis favour reciprocal adaptations that increase the stability of the association and can lead to codiversification of symbiont and host. While numerous examples of a strictly vertical transfer of the symbionts from parent to offspring among intracellular associations abound, little is known about the fidelity of the partners in extracellular associations, where symbionts colonize the surface or body cavity of their host. In this issue of Molecular Ecology, Zimmermann et al. ( 2016 ) investigated the evolutionary history of the symbiotic association between a monophyletic clade of sulphur‐oxidizing Gammaproteobacteria and two distantly related lineages of marine worms (nematodes and annelids). The study supports the surprising conclusion that partner fidelity does not necessarily increase with the intimacy of the association. Ectosymbionts on the cuticle of the nematodes seem to be cospeciating with their hosts, whereas endosymbionts housed in the body cavity of the annelids must have originated multiple times, probably by host switching, from ectosymbionts of sympatric nematodes. This excellent case study on the evolutionary history of invertebrate–microbe interactions supports the emerging concept that the co‐evolutionary processes shaping such mutualistic symbioses include both codiversification and ecological fitting.     

A new perspective on phylogeny and evolution of tetraodontiform fishes (Pisces: Acanthopterygii) based on whole mitochondrial genome sequences: Basal ecological diversification?

Background  

The order Tetraodontiformes consists of approximately 429 species of fishes in nine families. Members of the order exhibit striking morphological diversity and radiated into various habitats such as freshwater, brackish and coastal waters, open seas, and deep waters along continental shelves and slopes. Despite extensive studies based on both morphology and molecules, there has been no clear resolution except for monophyly of each family and sister-group relationships of Diodontidae + Tetraodontidae and Balistidae + Monacanthidae. To address phylogenetic questions of tetraodontiform fishes, we used whole mitochondrial genome (mitogenome) sequences from 27 selected species (data for 11 species were newly determined during this study) that fully represent all families and subfamilies of Tetraodontiformes (except for Hollardinae of the Triacanthodidae). Partitioned maximum likelihood (ML) and Bayesian analyses were performed on two data sets comprising concatenated nucleotide sequences from 13 protein-coding genes (all positions included; third codon positions converted into purine [R] and pyrimidine [Y]), 22 transfer RNA and two ribosomal RNA genes (total positions = 15,084).     

Parasite-host relations and new reports of helminths for Rhinella dorbignyi (Duméril & Bibron, 1841) (Anura: Bufonidae) from Neotropical region

This study aimed to analyze helminth assemblage in Rhinella dorbignyi regarding host gender, size and mass in two sampling sites in southern Brazil, and to report new parasite associations. Anurans (n = 100) were collected in two localities from Rio Grande do Sul (RS) state, Brazil, from 2017 to 2020. Nineteen taxa (adults and larval forms) belonging to Nematoda, Acanthocephala, Digenea, and Cestoda were found in different infection sites. Cosmocercidae gen. spp., Physaloptera liophis, Catadiscus sp., and Cylindrotaenia americana were the dominant taxa in helminth assemblage. Female anurans showed higher helminth species richness than males considering the total sample (two localities) or one of the two localities. However, prevalence and mean intensity of infection did not show any significant difference between genders. Mean intensity of infection was significantly higher (19.52) in Laranjal locality. Host body size does not influence helminth abundance, as infections did not show significant correlation snout-vent length (SVL) nor body mass (BM) of anurans. The findings reveal anurans of R. dorbignyi may be intermediate, paratenic and definitive host for these parasites. Plagiorchioidea helminths (Digenea), Physaloptera liophis, larvae of Acuariidae and Spiroxys sp. (Nematoda), and cystacanth of Lueheia sp. (Acanthocephala) constitute new records for R. dorbignyi. Additionally, this is the first record of Cylindrotaenia americana larvae in this host species. Resulting information increases the knowledge on biodiversity and parasite-host relations and may help future conservation programs developed in ecosystems in the extreme south of Brazil.     

Temporal change in inbreeding depression in life‐history traits in captive populations of guppy (Poecilia reticulata): evidence for purging?

Inbreeding depression, which generally affects the fitness of small populations, may be diminished by purging recessive deleterious alleles when inbreeding persists over several generations. Evidence of purging remains rare, especially because of the difficulties of separating the effects of various factors affecting fitness in small populations. We compared the expression of life-history traits in inbred populations of guppy (Poecilia reticulata) with contemporary control populations over 10 generations in captivity. We estimated inbreeding depression as the difference between the two types of populations at each generation. After 10 generations, the inbreeding coefficient reached a maximum value of 0.56 and 0.16 in the inbred and control populations, respectively. Analysing changes in the life-history traits across generations showed that inbreeding depression in clutch size and offspring survival increased during the first four to six generations in the populations from the inbred treatment and subsequently decreased as expected if purging occurred. Inbreeding depression in two other traits was weaker but showed similar changes across generations. The loss of six populations in the inbred treatment indicates that removal of deleterious alleles also occurred by extinction of populations that presumably harboured high genetic load.     

Recovery after African Olive invasion: can a ‘bottom‐up’ approach to ecological restoration work?

African Olive (Olea europaea ssp. cuspidata) is a densely crowned evergreen small tree, native to eastern Africa that is highly invasive in areas where it has been introduced, including Hawaii and Australia. Invasion by African Olive threatens Cumberland Plain Woodland, a critically endangered grassy eucalypt woodland from western Sydney, Australia, through the formation of a dense mid‐canopy excluding the regeneration of native species. We established a 3‐year field experiment to determine the effectiveness of direct seeding and fire, as techniques for early stage restoration of a 2 ha historically cleared and degraded Cumberland Plain Woodland site after the removal of African Olive. Direct seeding was able to re‐establish a native perennial grass cover which was resistant to subsequent weed invasion and could be managed as an important first stage in woodland restoration with fire and selective herbicide. Fire was able to stimulate some germination of colonising native species from the soil seed bank after 15 years of African Olive invasion; however, germination and establishment of native shrubs from the applied seed mix was poor. We propose a ‘bottom‐up’ model of ecological restoration in such highly degraded sites that uses a combination of direct seeding and stimulation of the soil seed bank by fire, which could be applicable to other degraded grassy woodland sites and plant communities.     

Too hot to handle? Phenological and life‐history responses to simulated climate change of the southern green stink bug Nezara viridula (Heteroptera: Pentatomidae)

The effect of simulated climate change on Nezara viridula was studied close to the species' northern range limit in Japan. Insects from the same egg masses were reared for 15 months in 10 consecutive series under quasi‐natural (i.e. outdoor) conditions and in a transparent incubator, in which climate warming was simulated by adding 2.5 °C to the outdoor temperature. The warming strongly affected all life‐history and phenological parameters. In the spring, the simulated warming advanced the timing of postdiapause body colour changes and reproduction. In the early summer, it increased egg production and accelerated nymphal development. In the late summer (the hottest season), the effect of the simulated warming was strongly deleterious: nymphs developed slowly, suffered higher mortality and had difficulties during final moulting; the emerged females were smaller, some exhibited abnormal cuticle, produced fewer eggs and had a decreased life span. In the autumn, the warming accelerated nymphal development, resulted in larger female size, affected the timing of the diapause‐associated adult body colour change from green to russet and enhanced preparation for overwintering. Larger females had higher winter survival rate than smaller females. The warming strongly increased survival rate in both size classes and allowed smaller females to reach the same winter survival rate as larger females had under the quasi‐natural conditions. The winter survival also differed between the green and dark‐coloured females under the quasi‐natural, but not under the warming conditions. However, under the warming conditions, green females survived the winter even better than dark‐coloured females did under the quasi‐natural conditions. The warming also shortened the life span of females from the summer generations and prolonged it in those from the autumn generation. It is concluded that even a moderate temperature increase (+2.5 °C) in the future is likely to have a complex influence upon insects, strongly affecting many of their life‐history and phenological parameters.     

What explains patterns of species richness? The relative importance of climatic‐niche evolution,morphological evolution,and ecological limits in salamanders

A major goal of evolutionary biology and ecology is to understand why species richness varies among clades. Previous studies have suggested that variation in richness among clades might be related to variation in rates of morphological evolution among clades (e.g., body size and shape). Other studies have suggested that richness patterns might be related to variation in rates of climatic‐niche evolution. However, few studies, if any, have tested the relative importance of these variables in explaining patterns of richness among clades. Here, we test their relative importance among major clades of Plethodontidae, the most species‐rich family of salamanders. Earlier studies have suggested that climatic‐niche evolution explains patterns of diversification among plethodontid clades, whereas rates of morphological evolution do not. A subsequent study stated that rates of morphological evolution instead explained patterns of species richness among plethodontid clades (along with “ecological limits” on richness of clades, leading to saturation of clades with species, given limited resources). However, they did not consider climatic‐niche evolution. Using phylogenetic multiple regression, we show that rates of climatic‐niche evolution explain most variation in richness among plethodontid clades, whereas rates of morphological evolution do not. We find little evidence that ecological limits explain patterns of richness among plethodontid clades. We also test whether rates of morphological and climatic‐niche evolution are correlated, and find that they are not. Overall, our results help explain richness patterns in a major amphibian group and provide possibly the first test of the relative importance of climatic niches and morphological evolution in explaining diversity patterns.     

Genomic‐scale comparison of sequence‐ and structure‐based methods of function prediction: Does structure provide additional insight?

A function annotation method using the sequence-to-structure-to-function paradigm is applied to the identification of all disulfide oxidoreductases in the Saccharomyces cerevisiae genome. The method identifies 27 sequences as potential disulfide oxidoreductases. All previously known thioredoxins, glutaredoxins, and disulfide isomerases are correctly identified. Three of the 27 predictions are probable false-positives. Three novel predictions, which subsequently have been experimentally validated, are presented. Two additional novel predictions suggest a disulfide oxidoreductase regulatory mechanism for two subunits (OST3 and OST6) of the yeast oligosaccharyltransferase complex. Based on homology, this prediction can be extended to a potential tumor suppressor gene, N33, in humans, whose biochemical function was not previously known. Attempts to obtain a folded, active N33 construct to test the prediction were unsuccessful. The results show that structure prediction coupled with biochemically relevant structural motifs is a powerful method for the function annotation of genome sequences and can provide more detailed, robust predictions than function prediction methods that rely on sequence comparison alone.     

Expression of the 22 nucleotide let‐7 heterochronic RNA throughout the Metazoa: a role in life history evolution?

The 22 nucleotide let-7 small temporal RNA has been found consistently in samples from diverse bilateria but not from sponge or cnidarians. Here we further examine the phylogenetic distribution of this regulatory RNA by sampling representatives of diverse metazoan lineages. The 22 nucleotide let-7 RNA is detectable in triclad and polyclad platyhelminths, nemertean, and chaetognath but not ctenophore or acoel metazoans. These results support recent arguments that acoels are distinct from other acoelomate platyhelminths. We argue that let-7 is not a bilaterian or triploblast synapomorphy but instead evolved later in metazoan evolution, perhaps in association with complex life history traits.     

Can life‐history traits predict the fate of introduced species? A case study on two cyprinid fish in southern France

1. The ecological and economic costs of introduced species can be high. Ecologists try to predict the probability of success and potential risk of the establishment of recently introduced species, given their biological characteristics.
2. In 1990 gudgeon, Gobio gobio , were released in a drainage canal of the Rhône delta of southern France. The Asian topmouth gudgeon, Pseudorasbora parva, was found for the first time in the same canal in 1993. Those introductions offered a unique opportunity to compare in situ the fate of two closely related fish in the same habitat.
3. Our major aims were to assess whether G. gobio was able to establish in what seemed an unlikely environment, to compare populations trends and life-history traits of both species and to assess whether we could explain or could have predicted our results, by considering their life-history strategies.
4. Data show that both species have established in the canal and have spread. Catches of P. parva have increased strongly and are now higher than those of G. gobio .
5. The two cyprinids have the same breeding season and comparable traits (such as short generation time, small body, high reproductive effort), so both could be classified as opportunists. The observed difference in their success (in terms of population growth and colonization rate) could be explained by the wider ecological and physiological tolerance of P. parva .
6. In conclusion, our field study seems to suggest that invasive vigour also results from the ability to tolerate environmental changes through phenotypic plasticity, rather than from particular life-history features pre-adapted to invasion. It thus remains difficult to define a good invader simply on the basis of its life-history features.