Life History Tactics Research Articles

Population genomics, life-history tactics, and mixed-stock subsistence fisheries in the northernmost American Atlantic salmon populations.

While Atlantic salmon (Salmo salar) of the northernmost American populations is alimentary, economically, and culturally important for Ungava Inuit communities (Nunavik, Canada) and might play a key role in the persistence of the species in a global warming context, many mysteries remain about those remote and atypical populations. Thus, our first aim was to document the genomic structure of the Nunavik populations. The second objective was to determine whether salmon only migrating to the estuary without reaching the sea, apparently unique to those populations, represent distinct populations from the typical anadromous salmons and subsequently explore the genetic basis of migratory life-history tactics in the species. Finally, the third goal was to quantify the contribution of each genetically distinct population and life-history tactic in the mixed-stock subsistence fishery of the Koksoak R. estuary. We used Genotyping-by-Sequencing to genotype 14,061 single nucleotide polymorphisms in the genome of 248 individuals from 8 source populations and 280 individuals from the Koksoak estuary mixed-stock fishery. Life-history tactics were identified by a visual assessment of scales. Results show a hierarchical structure mainly influenced by isolation-by-distance with 7 populations out of the 8 studied rivers. While no obvious structure was detected between marine and estuarine salmon within the population, we have identified genomic regions putatively associated with those migration tactics. Finally, all salmon captured in the Koksoak estuary originated from the Koksoak drainage and mostly from 2 tributaries, but no inter-annual variation in the contribution of these tributaries was found. Our results indicate, however, that both marine and estuarine salmon contribute substantially to estuarine fisheries and that there is inter-annual variation in this contribution. These findings provide crucial information for the conservation of salmon populations in a rapidly changing ecosystem, as well as for fishery management to improve the food security of Inuit communities.

Alternative life-history strategy contributions to effective population size in a naturally spawning salmon population.

Alternative life-history tactics are predicted to affect within-population genetic processes but have received little attention. For example, the impact of precocious males on effective population size (N e) has not been quantified directly in Pacific salmon Oncorhynchus spp., even though they can make up a large percentage of the total male spawners. We investigated the contribution of precocial males ("jacks") to N e in a naturally spawning population of Coho Salmon O. kisutch from the Auke Creek watershed in Juneau, Alaska. Mature adults that returned from 2009 to 2019 (~8000 individuals) were genotyped at 259 single-nucleotide polymorphism (SNP) loci for parentage analysis. We used demographic and genetic methods to estimate the effective number of breeders per year (N b). Jack contribution to N b was assessed by comparing values of N b calculated with and without jacks and their offspring. Over a range of N b values (108-406), the average jack contribution to N b from 2009 to 2015 was 12.9% (SE = 3.8%). Jacks consistently made up over 20% of the total male spawners. The presence of jacks did not seem to influence N b/N. The linkage disequilibrium N e estimate was lower than the demographic estimate, possibly due to immigration effects on population genetic processes: based on external marks and parentage data, we estimated that immigrant spawners produced 4.5% of all returning offspring. Our results demonstrate that jacks can influence N b and N e and can make a substantial contribution to population dynamics and conservation of threatened stocks.

Blood-based gene expression as non-lethal tool for inferring salinity-habitat history of European eel (Anguilla anguilla)

The European eel is a facultative catadromous species, meaning that it can skip the freshwater phase or move between marine and freshwater habitats during its continental life stage. Otolith microchemistry, used to determine the habitat use of eel or its salinity history, requires the sacrifice of animals. In this context, blood-based gene expression may represent a non-lethal alternative. In this work, we tested the ability of blood transcriptional profiling to identify the different salinity-habitat histories of European eel. Eels collected from different locations in Norway were classified through otolith microchemistry as freshwater residents (FWR), seawater residents (SWR) or inter-habitat shifters (IHS). We detected 3451 differentially expressed genes from blood by comparing FWR and SWR groups, and then used that subset of genes in a machine learning approach (i.e., random forest) to the extended FWR, SWR, and IHS group. Random forest correctly classified 100% of FWR and SWR and 83% of the IHS using a minimum of 30 genes. The implementation of this non-lethal approach may replace otolith-based microchemistry analysis for the general assessment of life-history tactics in European eels. Overall, this approach is promising for the replacement or reduction of other lethal analyses in determining certain fish traits.

Genome-wide DNA methylation predicts environmentally driven life history variation in a marine fish.

Epigenetic modifications are thought to be one of the molecular mechanisms involved in plastic adaptive responses to environmental variation. However, studies reporting associations between genome-wide epigenetic changes and habitat-specific variations in life history traits (e.g., lifespan, reproduction) are still scarce, likely due to the recent application of methylome resequencing methods to non-model species. In this study, we examined associations between whole genome DNA methylation and environmentally driven life history variation in 2 lineages of a marine fish, the capelin (Mallotus villosus), from North America and Europe. In both lineages, capelin harbor 2 contrasting life history tactics (demersal vs. beach-spawning). Performing whole genome and methylome sequencing, we showed that life history tactics are associated with epigenetic changes in both lineages, though the effect was stronger in European capelin. Genetic differentiation between the capelin harboring different life history tactics was negligible, but we found genome-wide methylation changes in both lineages. We identified 9,125 European and 199 North American differentially methylated regions (DMRs) due to life history. Gene ontology (GO) enrichment analysis for both lineages revealed an excess of terms related to neural function. Our results suggest that environmental variation causes important epigenetic changes that are associated with contrasting life history tactics in lineages with divergent genetic backgrounds, with variable importance of genetic variation in driving epigenetic variation. Our study emphasizes the potential role of genome-wide epigenetic variation in adaptation to environmental variation.

Life-history variation as a source of diversity for endemic white charr (Salmonidae) of the lower Kamchatka River.

Sympatric fish morphs diversifying in their feeding pattern in fresh waters typically implement alternative migratory tactics so that omnivores become migrants and specialists-residents. Charrs of the genus Salvelinus populating the lower Kamchatka River (Northeast Asia) are a rare example wherein two related sympatric morphs both implement a variety of life-history tactics in parallel. Here the authors analyse the ecological diversity in the endemic piscivorous "white" morph that exploits resources of the Kamchatka River in sympatry with the partially anadromous invertivorous "Dolly Varden" (DV) morph. Eco-morphological criteria allowed the authors to validate the morph identification. The white charr (WC) was found to subdivide into the small-sized (up to 1.6 kg) fish inhabiting the lacustrine part of the ecosystem and the large-sized (up to 3.4 kg) fish inhabiting the main river channel. The persistent spatial segregation of the sub-groups was confirmed by significant differences in the life span, muscle δ13 С signature and parasite load. According to contrasting patterns of strontium accumulation in otoliths, the riverine WC is represented by resident and semi-anadromous individuals. At the same time, the lack of microsatellite DNA differentiation and allometric nature of the morphometric discrepancy point to the intra-population source of the WC polymorphism. The authors suggest that WC diverged from DV as a result of feeding specialization on the threespine stickleback numerous in the ecosystem since the temporary flooding by marine waters in the middle Holocene. The modern stickleback division into local stocks following the ecosystem differentiation into a river, side lake and estuary resulted in the WC life-history split and ecological radiation.

Fertility Dynamics and Life History Tactics Vary by Socioeconomic Position in a Transitioning Cohort of Postreproductive Chilean Women

Globally, mortality and fertility rates generally fall as resource abundance increases. This pattern represents an evolutionary paradox insofar as resource-rich ecological contexts can support higher numbers of offspring, a component of biological fitness. This paradox has not been resolved, in part because the relationships between fertility, life history strategies, reproductive behavior, and socioeconomic conditions are complex and cultural-historically contingent. We aim to understand how we might make sense of this paradox in the specific context of late-twentieth-century, mid–demographic transition Chile. We use distribution-specific generalized linear models to analyze associations between fertility-related life-history traits—number of offspring, ages at first and last reproduction, average interbirth interval, and average number of live births per reproductive span year—and socioeconomic position (SEP) using data from a cohort of 6,802 Chilean women born between 1961 and 1970. We show that Chilean women of higher SEP have shorter average interbirth intervals, more births per reproductive span year, later age at first reproduction, earlier ages at last reproduction, and, ultimately, fewer children than women of lower SEP. Chilean women of higher SEP consolidate childbearing over a relatively short time span in the middle of their reproductive careers, whereas women of lower SEP tend to reproduce over the entirety of their reproductive lifespans. These patterns may indicate that different SEP groups follow different pathways toward declining fertility during the demographic transition, reflecting different life-history trade-offs in the process.

Metabarcoding prey DNA from fecal samples of adult dragonflies shows no predicted sex differences, and substantial inter-individual variation, in diets.

Sexes often differ in foraging and diet, which is associated with sex differences in size, trophic morphology, use of habitats, and/or life history tactics. Herein, strikingly similar diets were found for adult sexes of a dragonfly (Leucorrhinia intacta), based on comparing 141 dietary taxa identified from the metabarcoding of mitochondrial DNA archived in feces. Arthropods in > 5% of samples included five species of dipterans, two hemipterans, two spider species and one parasitic mite. The mite was not traditional prey as its presence was likely due to DNA contamination of samples arising through parasitism or possibly via accidental consumption during grooming, and therefore the mite was excluded from diet characterizations. Common prey species were found with statistically indistinguishable frequencies in male and female diets, with one exception of an aphid more often found in male diets, although this pattern was not robust to corrections for multiple statistical tests. While rare prey species were often found in diets of only one sex, instances of this were more frequent in the more oft-sampled females, suggesting sampling artefact. Sexes did not differ in the mean prey species richness in their diets. Overall, sexes showed statistically indistinguishable diets both on a prey species-by-species basis and in terms of multivariate characterizations of diet composition, derived from presence-absence data of prey species analyzed via PERMANOVA and accumulation curves. Males and females may have similar diets by being both opportunistic and generalist predators of arthropods, using the same foraging habitats and having similar sizes and flight agilities. Notably, similarities in diet between sexes occur alongside large interindividual differences in diet, within sexes. Researchers intending on explaining adaptive sex differences in diet should consider characteristics of species whose sexes show similar diets.

Personality, spatiotemporal ecological variation and resident/explorer movement syndromes in the sleepy lizard.

Individual variation in movement is profoundly important for fitness and offers key insights into the spatial and temporal dynamics of populations and communities. Nonetheless, individual variation in fine-scale movement behaviours is rarely examined even though animal tracking devices offer the long-term, high-resolution, repeatable data in natural conditions that are ideal for studying this variation. Furthermore, of the few studies that consider individual variation in movement, even fewer also consider the internal traits and environmental factors that drive movement behaviour which are necessary for contextualising individual differences in movement patterns. In this study, we GPS tracked a free-ranging population of sleepy lizards Tiliqua rugosa, each Austral spring over 5years to examine consistent among-individual variation in movement patterns, as well as how these differences were mediated by key internal and ecological factors. We found that individuals consistently differed in a suite of weekly movement traits, and that these traits strongly covaried among-individuals, forming movement syndromes. Lizards fell on a primary movement continuum, from 'residents' that spent extended periods of time residing within smaller core areas of their home range, to 'explorers' that moved greater distances and explored vaster areas of the environment. Importantly, we also found that these consistent differences in lizard movement were related to two ecologically important animal personality traits (boldness and aggression), their sex, key features of the environment (including food availability, and a key water resource), habitat type and seasonal variation (cool/moist vs. hot/drier) in environmental conditions. Broadly, these movement specialisations likely reflect variation in life-history tactics including foraging and mating tactics that ultimately underlie key differences in space use. Such information can be used to connect phenotypic population structure to key ecological and evolutionary processes, for example social networks and disease-transmission pathways, further highlighting the value of examining individual variation in movement behaviour.

Many lifetime growth trajectories for a single mammal.

Despite their importance in shaping life history tactics and population dynamics, individual growth trajectories have only been rarely explored in the wild because their analysis requires multiple measurements of individuals throughout their lifetime and some knowledge of age, a key timer of body growth. The availability of long‐term longitudinal studies of two wild boar populations subjected to contrasting environments (rich vs. poor) provided an opportunity to analyze individual growth trajectories. We quantified wild boar growth trajectories at both the population and the individual levels using standard growth models (i.e., Gompertz, logistic, and monomolecular models) that encompass the expected range of growth shapes in determinate growers. Wild boar is a rather altricial species, with a polygynous mating system and is strongly sexually dimorphic in size. According to current theories of life history evolution, we thus expect wild boar to display a sex‐specific Gompertz type growth trajectory and lower sexual size dimorphism in the poorer environment. While wild boar displayed the expected Gompertz type trajectory in the rich site at the population level, we found some evidence for potential differences in growth shapes between populations and individuals. Asymptotic body mass, growth rate and timing of maximum growth rate differed as well, which indicates a high flexibility of growth in wild boar. We also found a cohort effect on asymptotic body mass, which suggests that environmental conditions early in life shape body mass at adulthood in this species. Our findings demonstrate that body growth trajectories in wild boar are highly diverse in relation to differences of environmental context, sex and year of birth. Whether the intermediate ranking of wild boar along the precocial–altricial continuum of development at birth may explain the ability of this species to exhibit this high diversity of growth patterns remains to be investigated.

Differential gene expression associated with behavioral variation in ecotypes of Lake Superior brook trout (Salvelinus fontinalis)

Associations between behaviors and the development of different life history tactics have been documented in several species of salmon, trout, and charr. While it is well known that such behaviors are heritable the genes and molecular pathways connected to these behaviors remain unknown. We used an RNA-seq approach to identify genes and molecular pathways differentially regulated in brain tissue between “shy” and “bold” brook trout (Salvelinus fontinalis). A small number of genes were differentially expressed between the behavioral types at several months after hatching and two years of age. Pathway analysis revealed that EIF2 signaling differed consistently between shy and bold individuals suggesting large-scale differences in protein synthesis between behavioral types in the brain. Additionally, the RNA-seq data were used to find polymorphisms within the brook trout genome and a GWAS approach was used to test for statistical associations between genetic variants and behavior type. One allele located in a transcription factor (TSHZ3) contained a protein-coding non-synonymous SNP suggesting that functional variation within TSHZ3 is connected to the development of different behaviors. These results suggest that the molecular basis of behavioral development is complex and due to the differential expression of many genes involved in a wide-range of different molecular pathways.

Every-other-day clutch-initiation synchrony as an adaptive response to egg cannibalism in Glaucous-winged Gulls (Larus glaucescens)

Sea surface temperatures (SST) in the Pacific Northwest of North America rose 1 °C during the last half of the twentieth century. Cannibalism, a behavior observed in diverse taxa, is often associated with low food supplies, which for marine animals can be precipitated by high SSTs. In an 8 year study, we found that in years of higher sea surface temperatures, Glaucous-winged Gulls (Larus glaucescens) breeding in a colony in Washington State, Salish Sea, USA, tended to exhibit higher rates of egg cannibalism by males, higher levels of every-other-day clutch-initiation synchrony by females, and longer egg-laying seasons than in years of lower SST. Clutch-initiation synchrony increased the odds that an egg survived cannibalism and may serve as an adaptive response to egg cannibalism. Short-term climate and resource fluctuations associated with El Niño Southern Oscillation (ENSO) events may select for behavioral plasticity in marine organisms, allowing long-lived individuals such as marine birds to switch between alternative life history tactics. The implications for long-term SST warming, however, remain unknown.

Positive Feedback between Behavioral and Hormonal Dynamics Leads to Differentiation of Life-History Tactics.

AbstractCompetitive interaction among individuals of a single population may result in the differentiation of two or more distinct life-history tactics. For example, although they exhibit unimodal size distribution, male juveniles of salmonids differentiate into those going down to the ocean to grow and returning to the natal stream after several years to reproduce (migratory tactic) and those staying in the stream and reproducing for multiple years (resident tactic). In this study, we developed a simple mathematical model for the positive feedback between hormonal and behavioral dynamics, with the expectation of establishing multiple discrete clusters of hormone levels leading to differentiation of life-history tactics. The assumptions were that probability of winning in fighting depends both on the body size and hormone level of the two contestants. An individual with a higher hormone level would be more likely to win the competition, which further enhanced hormone production, forming a positive feedback loop between hormone level and fighting ability. If the positive feedback was strong but not excessive, discrete clusters of hormone levels emerged from a continuous distribution. In contrast, no clear clustering structure appeared in the distribution of hormone levels if the probability of winning in fighting was controlled by the body size.

One in the Hand Worth Two in the Bush? Reproductive Effort of Young Males Is Not Affected by the Presence of Adult Males

Reproduction is a costly endeavor, and most large, long-lived, and iteroparous mammals exhibit conservative life-history tactics wherein an individual may forego or abandon a reproduction event for the sake of survival. Nevertheless, risks and benefits associated with reproduction are not equal across males and females, nor across their life. Whereas expenditure for females is associated with rearing young (e.g., lactation), expenditure for males occurs with securing mating opportunities. Young males may be more successful when dominant males are lacking, but it is less clear whether—and at what cost—young males will expend effort when those opportunities arise. We designed an experiment to quantify reproductive effort (e.g., food intake, somatic loss [body mass and fat]) of male white-tailed deer (Odocoileus virginianus) to better understand the reproductive ecology of male ungulates, with an emphasis on determining how adult males (≥4.5-years old) affect timing and extent of reproductive effort expended by yearling males (1.5-years old). Food intake, hormone levels, body mass, and somatic loss during rut were similar between yearling males that interacted with adult males and those that did not. Somatic loss by all males was greatest during peak estrus of females, but forage intake relative to metabolic body mass for yearling males was nearly twice that of adult males. Testosterone levels were lower for yearling than adult males early in rut and were related negatively to forage intake. Whereas adult males lost 20% (23.5kg) of body mass and 31% (5.3 percentage points [ppt]) of body fat during the rut, yearling males lost 12% (9.3kg) of body mass and 22% (4.7ppt) of body fat. Reproductive effort by young males was not influenced by the presence of adult males, though young males expended less reproductive effort than adults. Instead, reproductive allocation occurred in a state-dependent manner, where pre-season levels of somatic reserves dictated reproductive effort, regardless of age. Like female ungulates, male deer displayed risk-sensitive reproductive allocation wherein current reproductive allocation occurred as a function of resources garnered during the prior season and were expended in a way that should have avoided over-investing and creating a tradeoff between reproduction and survival.

Fisheries-induced evolution of alternative male life history tactics in Coho salmon.

Fisheries‐induced evolution (FIE) can result when harvest imposes artificial selection on variation in heritable phenotypic traits. While there is evidence for FIE, it remains difficult to disentangle the contributions of within‐generation demographic adjustment, phenotypic plasticity, and genetic adaption to observed changes in life history traits. We present evidence for FIE using dozens of Coho salmon (Oncorhynchus kisutch) populations in which males adopt one of two age‐invariant, heritable life history tactics: most mature as large three‐year‐old “hooknose” and typically fight for spawning opportunities, while some mature as small two‐year‐old “jacks” and fertilize eggs through sneaking. The closure of a fishery targeting three‐year‐old fish provided an experimental test of the prediction that fishery‐imposed selection against hooknose males drives an evolutionary increase in the proportion of males adopting the jack tactic. The data support the prediction: 43 of 46 populations had higher jack proportions during than after the fishery. The data further suggest that changes in jack proportion were not solely the result of demographic adjustments to harvest. We suggest that systems where fisheries differentially exploit phenotypically discrete, age‐invariant life histories provide excellent opportunities for detecting FIE.

Contrasting size-dependent life history strategies of an insular lizard.

In many species with continuous growth, body size is an important driver of life-history tactics and its relative importance is thought to reflect the spatio-temporal variability of selective pressures. We developed a deterministic size-dependent integral projection model for 3 insular neighboring lizard populations with contrasting adult body sizes to investigate how size-related selective pressures can influence lizard life-history tactics. For each population, we broke down differences in population growth rates into contributions from size-dependent body growth, survival, and fecundity. A life table response experiment (LTRE) was used to compare the population dynamics of the 3 populations and quantify the contributions of intrinsic demographic coefficients of each population to the population growth rate (λ). Perturbation analyses revealed that the largest adults contributed the most to the population growth rate, but this was not true in the population with the smallest adults and size-independent fertility. Although we were not able to identify a single factor responsible for this difference, the combination of the demographic model on a continuous trait coupled with an LTRE analysis revealed how individuals from sister populations of the same species follow different life strategies and showed different compensatory mechanisms among survival, individual body growth, and fertility. Our results indicate that body size can play a contrasting role even in closely-related and closely-spaced populations.

Food and temperature stressors have opposing effects in determining flexible migration decisions in brown trout (Salmo trutta).

With rapid global change, organisms in natural systems are exposed to a multitude of stressors that likely co-occur, with uncertain impacts. We explored individual and cumulative effects of co-occurring environmental stressors on the striking, yet poorly understood, phenomenon of facultative migration. We reared offspring of a brown trout population that naturally demonstrates facultative anadromy (sea migration), under different environmental stressor treatments and measured life history responses in terms of migratory tactics and freshwater maturation rates. Juvenile fish were exposed to reduced food availability, temperatures elevated to 1.8°C above natural conditions or both treatments in combination over 18months of experimental tank rearing. When considered in isolation, reduced food had negative effects on the size, mass and condition of fish across the experiment. We detected variable effects of warm temperatures (negative effects on size and mass, but positive effect on lipids). When combined with food restriction, temperature effects on these traits were less pronounced, implying antagonistic stressor effects on morphological traits. Stressors combined additively, but had opposing effects on life history tactics: migration increased and maturation rates decreased under low food conditions, whereas the opposite occurred in the warm temperature treatment. Not all fish had expressed maturation or migration tactics by the end of the study, and the frequency of these 'unassigned' fish was higher in food deprivation treatments, but lower in warm treatments. Fish showing migration tactics were smaller and in poorer condition than fish showing maturation tactics, but were similar in size to unassigned fish. We further detected effects of food restriction on hypo-osmoregulatory function of migrants that may influence the fitness benefits of the migratory tactic at sea. We also highlight that responses to multiple stressors may vary depending on the response considered. Collectively, our results indicate contrasting effects of environmental stressors on life history trajectories in a facultatively migratory species.

Life History Variation Within and Among Populations of the Ninespine Stickleback (Pungitius pungitius) in Alaska: Lake-Stream Contrasts

Intra- and inter-population comparisons of size at reproduction, clutch size, and egg mass revealed differences between one stream and one lake population of ninespine stickleback (Pungitius pungitius) in Alaska. These results are compared with published data for a second lake population. Ninespine stickleback in the stream appeared to reproduce at a smaller overall average size and to produce greater numbers of smaller eggs per clutch than lake-dwelling ninespine stickleback. This pattern of life history traits is consistent with that reported for populations of the threespine stickleback (Gasterosteus aculeatus) in published studies from Alaska and Central Europe, suggesting stream environments favor a different set of life history tactics than lakes in both species. Further research is needed to test this hypothesis and to elucidate the causal mechanisms.

Development of sexual dimorphism in two sympatric skinks with different growth rates.

Sexual size dimorphism (SSD) is widespread in animals, especially in lizards (Reptilia: Squamata), and is driven by fecundity selection, male–male competition, or other adaptive hypotheses. However, these selective pressures may vary through different life history periods; thus, it is essential to assess the relationship between growth and SSD. In this study, we tracked SSD dynamics between a “fading‐tail color skink” (blue tail skink whose tail is only blue during its juvenile stage: Plestiodon elegans) and a “nonfade color” tail skink (retains a blue tail throughout life: Plestiodon quadrilineatus) under a controlled experimental environment. We fitted growth curves of morphological traits (body mass, SVL, and TL) using three growth models (Logistic, Gompertz, and von Bertalanffy). We found that both skinks have male‐biased SSD as adults. Body mass has a higher goodness of fit (as represented by very high R 2 values) using the von Bertalanffy model than the other two models. In contrast, SVL and TL for both skinks had higher goodness of fit when using the Gompertz model. Two lizards displayed divergent life history tactics: P. elegans grows faster, matures earlier (at 65 weeks), and presents an allometric growth rate, whereas P. quadrilineatus grows slower, matures later (at 106 weeks), and presents an isometric growth rate. Our findings imply that species‐ and sex‐specific trade‐offs in the allocation of energy to growth and reproduction may cause the growth patterns to diverge, ultimately resulting in the dissimilar patterns of SSD.

Novel life history tactic observed in fall-run Chinook Salmon.

Novel life history tactic observed in fall-run Chinook Salmon.

Reproduction and mortality rates in ecologically distinct species of murid rodents.

The trade-off between reproduction and somatic maintenance is one of the most studied concepts of modern evolutionary ecology. This theory predicts a negative relationship between maximum species longevity and total reproductive output. However, studies performed on natural animal populations have found contradictory results, probably due to the unlikelihood of wild animals gaining both maximum longevity and maximum potential fecundity. A comparison of the mortality rates and reproductive output of four ecologically distinct rodent species of Cricetidae family that were maintained in the laboratory in controlled conditions revealed the different life-history tactics of subterranean social mole voles and three related aboveground species: hydrophilic water voles, arid dwarf hamsters and steppe lemmings. Regardless of the relatively higher mortality rates at early ages in mole voles, this species has considerably higher maximum species longevity and smaller litter sizes that do not depend on calendar age, whereas in dwarf hamsters and water voles clear negative correlations between female age and litter size were detected. Steppe lemmings, as a semi-social arid species, shared some life-history tactics with both mole voles and aboveground non-social rodents.