Divergent Life History Strategies Research Articles

Genetic trade-offs underlie divergent life history strategies for local adaptation in white clover.

Local adaptation is common in plants, yet characterization of its underlying genetic basis is rare in herbaceous perennials. Moreover, while many plant species exhibit intraspecific chemical defence polymorphisms, their importance for local adaptation remains poorly understood. We examined the genetic architecture of local adaptation in a perennial, obligately-outcrossing herbaceous legume, white clover (Trifolium repens). This widespread species displays a well-studied chemical defence polymorphism for cyanogenesis (HCN release following tissue damage) and has evolved climate-associated cyanogenesis clines throughout its range. Two biparental F2 mapping populations, derived from three parents collected in environments spanning the U.S. latitudinal species range (Duluth, MN, St. Louis, MO and Gainesville, FL), were grown in triplicate for two years in reciprocal common garden experiments in the parental environments (6,012 total plants). Vegetative growth and reproductive fitness traits displayed trade-offs across reciprocal environments, indicating local adaptation. Genetic mapping of fitness traits revealed a genetic architecture characterized by allelic trade-offs between environments, with 100% and 80% of fitness QTL in the two mapping populations showing significant QTL×E interactions, consistent with antagonistic pleiotropy. Across the genome there were three hotspots of QTL colocalization. Unexpectedly, we found little evidence that the cyanogenesis polymorphism contributes to local adaptation. Instead, divergent life history strategies in reciprocal environments were major fitness determinants: selection favoured early investment in flowering at the cost of multiyear survival in the southernmost site versus delayed flowering and multiyear persistence in the northern environments. Our findings demonstrate that multilocus genetic trade-offs contribute to contrasting life history characteristics that allow for local adaptation in this outcrossing herbaceous perennial.

Distinct nutritional and endocrine regulation of prothoracic gland activities underlies divergent life history strategies in Manduca sexta and Drosophila melanogaster

Life history trade-offs lead to various strategies that maximize fitness, but the developmental mechanisms underlying these alternative strategies continue to be poorly understood. In insects, trade-offs exist between size and developmental time. Recent studies in the fruit fly Drosophila melanogaster have suggested that the steroidogenic prothoracic glands play a key role in determining the timing of metamorphosis. In this study, the nutrient-dependent growth and transcriptional activation of prothoracic glands were studied in D. melanogaster and the tobacco hornworm Manduca sexta. In both species, minimum viable weight (MVW) was associated with activation of ecdysteroid biosynthesis genes and growth of prothoracic gland cells. However, the timing of MVW attainment in M. sexta is delayed by the presence of the sesquiterpenoid hormone, juvenile hormone (JH), whereas in D. melanogaster it is not. Moreover, in D. melanogaster, the transcriptional regulation of ecdysteroidogenesis becomes nutrient-independent at the MVW/critical weight (CW) checkpoint. In contrast, in M. sexta, starvation consistently reduced transcriptional activation of ecdysteroid biosynthesis genes even after CW attainment, indicating that the nature of CW differs fundamentally between the two species. In D. melanogaster, the prothoracic glands dictate the timing of metamorphosis even in the absence of nutritional inputs, whereas in M. sexta, prothoracic gland activity is tightly coupled to the nutritional status of the body, thereby delaying the onset of metamorphosis before CW attainment. We propose that selection for survival under unpredictable nutritional availability leads to the evolution of increased modularity in both morphological and endocrine traits.

Landscape edges shape dispersal and population structure of a migratory fish.

Many freshwater organisms have a life-history stage that can disperse through seawater. This has obvious benefits for colonization and connectivity of fragmented sub-populations, but requires a physiologically challenging migration across a salinity boundary. We consider the role of landscape boundaries between freshwater and seawater habitats, and evaluate their potential effects on traits and developmental histories of larvae and juveniles (i.e., dispersing life-history stages) of an amphidromous fish, Galaxias maculatus. We sampled juvenile fish on their return to 20 rivers in New Zealand: 10 rivers had abrupt transitions to the sea (i.e., emptying to an open coastline); these were paired with 10 nearby rivers that had gradual transitions to the sea (i.e., emptying into estuarine embayments). We reconstructed individual dispersal histories using otolith microstructure, otolith microchemistry, and stable isotope analysis. We found that fish recruiting to embayment rivers had distinct dispersal and foraging histories, were slower growing, smaller in size, and older than fish recruiting to nearby non-embayment rivers. Our results indicate that landscape edges can affect dispersal capabilities of aquatic organisms, potentially leading to divergent life-history strategies (i.e., limited- versus widespread-dispersal). Patterns also suggest that dispersal potential among landscape boundaries can create heterogeneity in the traits of individuals, with implications for metapopulation dynamics.

A genetics-based approach confirms immune associations with life history across multiple populations of an aquatic vertebrate (Gasterosteus aculeatus).

Understanding how wild immune variation covaries with other traits can reveal how costs and trade‐offs shape immune evolution in the wild. Divergent life history strategies may increase or alleviate immune costs, helping shape immune variation in a consistent, testable way. Contrasting hypotheses suggest that shorter life histories may alleviate costs by offsetting them against increased mortality, or increase the effect of costs if immune responses are traded off against development or reproduction. We investigated the evolutionary relationship between life history and immune responses within an island radiation of three‐spined stickleback, with discrete populations of varying life histories and parasitism. We sampled two short‐lived, two long‐lived and an anadromous population using qPCR to quantify current immune profile and RAD‐seq data to study the distribution of immune variants within our assay genes and across the genome. Short‐lived populations exhibited significantly increased expression of all assay genes, which was accompanied by a strong association with population‐level variation in local alleles and divergence in a gene that may be involved in complement pathways. In addition, divergence around the eda gene in anadromous fish is likely associated with increased inflammation. A wider analysis of 15 populations across the island revealed that immune genes across the genome show evidence of having diverged alongside life history strategies. Parasitism and reproductive investment were also important sources of variation for expression, highlighting the caution required when assaying immune responses in the wild. These results provide strong, gene‐based support for current hypotheses linking life history and immune variation across multiple populations of a vertebrate model.

Can reproductive traits help to explain the coexistence of mud crabs Panopeus (Decapoda: Panopeidae)? A case of two sympatric species inhabiting an impacted mangrove area of Southern Brazil

Coexistence among species is commonly related to niche divergence. However, congenerics usually are very similar in their microhabitat selection and food consumption. Thus, divergent life history strategies may represent the mechanism that allows sympatry in related species. Here, we describe and compare reproductive features in two sympatric mud crabs Panopeus americanus and P. occidentalis in an impacted mangrove area in Southern Brazil. As these species are ecologically similar, we hypothesize that these species diverge in their reproductive traits, which could explain their coexistence. Crabs were collected every two months from September 2004 to July 2006. Reproductive features such as number and size of ovigerous females, breeding season, fecundity, reproductive output, and embryo volume were assessed. Panopeus americanus produced embryos during the entire sampled period, while P. occidentalis produced only between September and March. Panopeus americanus produced more embryos considering the size of the species, had significantly lower embryo volume, and higher reproductive output than P. occidentalis. These data permit to classify P. americanus as an r-strategist and P. occidentalis as a K-strategist regarding their reproductive traits. In conclusion, our results support the hypothesis that divergent reproductive features may allow coexistence of these mud crabs.

Multi‐scale environmental filters and niche partitioning govern the distributions of riparian vegetation guilds

Across landscapes, riparian plant communities assemble under varying levels of disturbance, environmental stress, and resource availability, leading to the development of distinct riparian life‐history guilds over evolutionary timescales. Identifying the environmental filters that exert selective pressures on specific riparian vegetation guilds is a critical step in setting baseline expectations for how riparian vegetation may respond to environmental conditions anticipated under future global change scenarios. In this study, we ask: (1) What riparian plant guilds exist across the interior Columbia and upper Missouri River basins? (2) What environmental filters shape riparian guild distributions? (3) How does resource partitioning among guilds influence guild distributions and co‐occurrence? Woody species composition was measured at 703 stream reaches and each species' morphological and functional attributes were extracted from a database in four categories: (1) life form, (2) persistence and growth, (3) reproduction, and (4) resource use. We clustered species into guilds by morphological characteristics and attributes related to environmental tolerances, modeling these guilds' distributions as a function of environmental filters—regional climate, watershed hydrogeomorphic characteristics, and stream channel form—and guild co‐existence. We identified five guilds: (1) a tall, deeply rooted, long‐lived, evergreen tree guild, (2) a xeric, disturbance tolerant shrub guild, (3) a hydrophytic, thicket‐forming shrub guild, (4) a low‐statured, shade‐tolerant, understory shrub guild, and (5) a flood tolerant, mesoriparian shrub guild. Guilds were most strongly discriminated by species' rooting depth, canopy height and potential to resprout and grow following biomass‐removing disturbance (e.g., flooding, fire). Hydro‐climatic variables, including precipitation, watershed area, water table depth, and channel form attributes reflective of hydrologic regime, were predictors of guilds whose life history strategies had affinity or aversion to flooding, drought, and fluvial disturbance. Biotic interactions excluded guilds with divergent life history strategies and/or allowed for the co‐occurrence of guilds that partition resources differently in the same environment. We conclude that the riparian guild framework provides insight into how disturbance and bioclimatic gradients shape riparian functional plant diversity across heterogeneous landscapes. Multiple environmental filters should be considered when the riparian response guild framework is to be used as a decision‐support tool framework across large spatial extents.

Symbiotic state influences life-history strategy of a clonal cnidarian.

Along the North American Pacific coast, the common intertidal sea anemone Anthopleura elegantissima engages in facultative, flexible symbioses with Symbiodinium muscatinei (a dinoflagellate) and Elliptochloris marina (a chlorophyte). Determining how symbiotic state affects host fitness is essential to understanding the ecological significance of engaging in such flexible relationships with diverse symbionts. Fitness consequences of hosting S. muscatinei, E. marina or negligible numbers of either symbiont (aposymbiosis) were investigated by measuring growth, cloning by fission and gonad development after 8.5-11 months of sustained exposure to high, moderate or low irradiance under seasonal environmental conditions. Both symbiotic state and irradiance affected host fitness, leading to divergent life-history strategies. Moderate and high irradiances led to a greater level of gonad development in individuals hosting E. marina, while high irradiance and high summer temperature promoted cloning in individuals hosting S. muscatinei and reduced fitness of aposymbiotic anemones. Associating with S. muscatinei may contribute to the success of A. elegantissima as a spatial competitor on the high shore: (i) by offsetting the costs of living under high temperature and irradiance conditions, and (ii) by promoting a high fission rate and clonal expansion. Our results suggest that basic life-history characteristics of a clonal cnidarian can be affected by the identity of the endosymbionts it hosts.

Age-related changes in urinary testosterone levels suggest differences in puberty onset and divergent life history strategies in bonobos and chimpanzees

Research on age-related changes in morphology, social behavior, and cognition suggests that the development of bonobos (Pan paniscus) is delayed in comparison to chimpanzees (Pan troglodytes). However, there is also evidence for earlier reproductive maturation in bonobos. Since developmental changes such as reproductive maturation are induced by a number of endocrine processes, changes in hormone levels are indicators of different developmental stages. Age-related changes in testosterone excretion are an indirect marker for the onset of puberty in human and non-human primates. In this study we investigated patterns of urinary testosterone levels in male and female bonobos and chimpanzees to determine the onset of puberty. In contrast to other studies, we found that both species experience age-related changes in urinary testosterone levels. Older individuals of both sexes had significantly higher urinary testosterone levels than younger individuals, indicating that bonobos and chimpanzees experience juvenile pause. The males of both species showed a similar pattern of age-related changes in urinary testosterone levels, with a sharp increase in levels around the age of eight years. This suggests that species-differences in aggression and male mate competition evolved independently of developmental changes in testosterone levels. Females showed a similar pattern of age-related urinary testosterone increase. However, in female bonobos the onset was about three years earlier than in female chimpanzees. The earlier rise of urinary testosterone levels in female bonobos is in line with reports of their younger age of dispersal, and suggests that female bonobos experience puberty at a younger age than female chimpanzees.

Divergent immunity and energetic programs in the gills of migratory and resident Oncorhynchus mykiss

Divergent life history strategies occur in steelhead or rainbow trout Oncorhynchus mykiss, and many populations produce both migrant (anadromous fish that move to the ocean after rearing) and resident (do not migrate and remain in fresh water) individuals. Mechanisms leading to each type are only partially understood; while the general tendency of a population is heritable, individual tendency may be plastic, influenced by local environment. Steelhead hatchery programmes aim to mitigate losses in wild stocks by producing trout that will migrate to the ocean and not compete with wild trout for limited freshwater resources. To increase our understanding of gill function in these migratory or resident phenotypes, here we compare gill transcriptome profiles of hatchery-released fish either at the release site (residents) or five river kilometres downstream while still in full fresh water (migrants). To test whether any of these genes can be used as predictive markers for smoltification, we compared these genes between migrant-like and undifferentiated trout while still in the hatchery in a common environment (prerelease). Results confirmed the gradual process of smoltification, and the importance of energetics, gill remodelling and ion transport capacity for migrants. Additionally, residents overexpressed transcripts involved in antiviral defences, potentially for immune surveillance via dendritic cells in the gills. The best smoltification marker candidate was protein s100a4, expression of which was highly correlated with Na(+) , K(+) ATPase (NKA) activity and smolt-like morphology in pre- and postrelease trout gills.

Does plant performance under stress explain divergent life history strategies? The effects of flooding and nutrient stress on two wetland sedges

Novel hydrology and nutrient loads are common environmental filters in disturbed wetlands. These filters impact plant growth and influence the composition and productivity of wetland plant communities by selecting for flooding-tolerant species that can efficiently acquire and use nutrients. We performed a factorial greenhouse trial to elucidate how nutrients and flooding affect the growth and morphology of Carex obnupta and Carex stipata, two wetland sedges with divergent life history strategies. Individual seedlings of each species were exposed to a combination of flooding (deep or shallow) and fertilization (low or high) treatments. These treatment combinations create a stress gradient from high fertilization and shallow flooding (lowest stress) to low fertilization and deep flooding (highest stress). Plant biomass accumulation and allocation were measured after 77-days. Total biomass, leaf area, and shoot biomass were greatest in high fertilizer treatments, regardless of flooding treatment. There was an initial plant size effect in C. obnupta: small plants accumulated less biomass than large plants under high flooding. C. stipata had higher biomass and specific leaf areas than C. obnupta in all treatments. C. obnupta had higher net assimilation rates per unit biomass and leaf area than C. stipata. These stress responses illustrate that each species has a distinct life history strategy. C. obnupta is a stress tolerator that can persist across a range of flooding conditions. C. stipata is a ruderal species that outgrows flooding stress by allocating carbon to leaf growth. These species’ niches likely diverge based on how each species grows and allocates biomass under environmental stress

Evolution of Divergent Life History Strategies in Marine Alphaproteobacteria

ABSTRACTMarine bacteria in the Roseobacter and SAR11 lineages successfully exploit the ocean habitat, together accounting for ~40% of bacteria in surface waters, yet have divergent life histories that exemplify patch-adapted versus free-living ecological roles. Here, we use a phylogenetic birth-and-death model to understand how genome content supporting different life history strategies evolved in these related alphaproteobacterial taxa, showing that the streamlined genomes of free-living SAR11 were gradually downsized from a common ancestral genome only slightly larger than the extant members (~2,000 genes), while the larger and variably sized genomes of roseobacters evolved along dynamic pathways from a sizeable common ancestor (~8,000 genes). Genome changes in the SAR11 lineage occurred gradually over ~800 million years, whereas Roseobacter genomes underwent more substantial modifications, including major periods of expansion, over ~260 million years. The timing of the first Roseobacter genome expansion was coincident with the predicted radiation of modern marine eukaryotic phytoplankton of sufficient size to create nutrient-enriched microzones and is consistent with present-day ecological associations between these microbial groups. We suggest that diversification of red-lineage phytoplankton is an important driver of divergent life history strategies among the heterotrophic bacterioplankton taxa that dominate the present-day ocean.

Does low daily energy expenditure drive low metabolic capacity in the tropical robin, Turdus grayi?

Temperate and tropical birds possess divergent life history strategies. Physiological parameters including energy metabolism correlate with the life history such that tropical species with a slower 'pace of life' have lower resting and maximal metabolic rates than temperate congeners. To better understand the physiological mechanisms underlying these differences, we investigated the relationship of metabolic capacity, muscle oxidative capacity and activity patterns to variation in life history patterns in American robins (Turdus migratorius), while resident in central North America and Clay-colored robins (Turdus grayi) resident in Panama. We measured summit metabolism [Formula: see text] in birds from both tropical and temperate habitats and found that the temperate robins have a 60% higher metabolic capacity. We also measured the field metabolic rate (FMR) of free-living birds using heart rate (HR) telemetry and found that temperate robins' daily energy expenditure was also 60% higher. Thus, [Formula: see text] and FMR both reflect life history differences between the species. Further, both species operate at a nearly identical ~50% of their thermogenic capacity throughout a given day. As a potential mechanism to explain differences in activity and metabolic capacity, we ask whether oxidative properties of flight muscle are altered in accordance with life history variation and found minimal differences in oxidative capacity of skeletal muscle. These data demonstrate a close relationship between thermogenic capacity and daily activity in free-living birds. Further, they suggest that the slow pace of life in tropical birds may be related to the maintenance of low activity rather than functional capacity of the muscle tissue.

The physiological implications of primary xylem organization in two ferns

Xylem structure and function are well described in woody plants, but the implications of xylem organization in less-derived plants such as ferns are poorly understood. Here, two ferns with contrasting phenology and xylem organization were selected to investigate how xylem dysfunction affects hydraulic conductivity and stomatal conductance (g(s)). The drought-deciduous pioneer species, Pteridium aquilinum, exhibits fronds composed of 25 to 37 highly integrated vascular bundles with many connections, high g(s) and moderate cavitation resistance (P50 = -2.23 MPa). By contrast, the evergreen Woodwardia fimbriata exhibits sectored fronds with 3 to 5 vascular bundles and infrequent connections, low g(s) and high resistance to cavitation (P50 = -5.21 MPa). Xylem-specific conductivity was significantly higher in P. aqulinium in part due to its wide, efficient conduits that supply its rapidly transpiring pinnae. These trade-offs imply that the contrasting xylem organization of these ferns mirrors their divergent life history strategies. Greater hydraulic connectivity and g(s) promote rapid seasonal growth, but come with the risk of increased vulnerability to cavitation in P. aquilinum, while the conservative xylem organization of W. fimbriata leads to slower growth but greater drought tolerance and frond longevity.

SEXUAL SELECTION AFFECTS THE EVOLUTION OF LIFESPAN AND AGEING IN THE DECORATED CRICKETGRYLLODES SIGILLATUS

Recent work suggests that sexual selection can influence the evolution of ageing and lifespan by shaping the optimal timing and relative costliness of reproductive effort in the sexes. We used inbred lines of the decorated cricket, Gryllodes sigillatus, to estimate the genetic (co)variance between age-dependent reproductive effort, lifespan, and ageing within and between the sexes. Sexual selection theory predicts that males should die sooner and age more rapidly than females. However, a reversal of this pattern may be favored if reproductive effort increases with age in males but not in females. We found that male calling effort increased with age, whereas female fecundity decreased, and that males lived longer and aged more slowly than females. These divergent life-history strategies were underpinned by a positive genetic correlation between early-life reproductive effort and ageing rate in both sexes, although this relationship was stronger in females. Despite these sex differences in life-history schedules, age-dependent reproductive effort, lifespan, and ageing exhibited strong positive intersexual genetic correlations. This should, in theory, constrain the independent evolution of these traits in the sexes and may promote intralocus sexual conflict. Our study highlights the importance of sexual selection to the evolution of sex differences in ageing and lifespan in G. sigillatus.

EVIDENCE FOR STRONG INTRALOCUS SEXUAL CONFLICT IN THE INDIAN MEAL MOTH, PLODIA INTERPUNCTELLA

Males and females share a genome and express many shared phenotypic traits, which are often selected in opposite directions. This generates intralocus sexual conflict that may constrain trait evolution by preventing the sexes from reaching their optimal phenotype. Furthermore, if present across multiple loci, intralocus sexual conflict can result in a gender load that may diminish the benefits of sexual selection and help maintain genetic variation for fitness. Despite the importance of intralocus sexual conflict, surprisingly few empirical studies conclusively demonstrate its operation. We show that the pattern of multivariate selection acting on three sexually dimorphic life-history traits (development time, body size, and longevity) in the Indian meal moth, Plodia interpunctella, is opposing for the sexes. Moreover, we combined our estimates of selection with the additive genetic variance-covariance matrix (G) to predict the evolutionary response of the life-history traits in the sexes and showed that the angle between the vector of responses and the vector of sexually antagonistic selection was almost orthogonal at 84.70°. Thus, G biases the predicted response of life-history traits in the sexes away from the direction of sexually antagonistic selection, confirming the presence of strong intralocus sexual conflict in this species. Despite this, sexual dimorphism has evolved in all of the life-history traits examined suggesting that mechanism(s) have evolved to resolve this conflict and allow the sexes to reach their life-history optima. We argue that intralocus sexual conflict is likely to play an important role in the evolution of divergent life-history strategies between the sexes in this species.

Growth and longevity in freshwater mussels: evolutionary and conservation implications

The amount of energy allocated to growth versus other functions is a fundamental feature of an organism's life history. Constraints on energy availability result in characteristic trade-offs among life-history traits and reflect strategies by which organisms adapt to their environments. Freshwater mussels are a diverse and imperiled component of aquatic ecosystems but little is known about their growth and longevity. Generalized depictions of freshwater mussels as 'long-lived and slow-growing' may give an unrealistically narrow view of life-history diversity which is incongruent with the taxonomic diversity of the group and can result in development of inappropriate conservation strategies. We investigated relationships among growth, longevity, and size in 57 species and 146 populations of freshwater mussels using original data and literature sources. In contrast to generalized depictions, longevity spanned nearly two orders of magnitude, ranging from 4 to 190 years, and the von Bertalanffy growth constant, K, spanned a similar range (0.02-1.01). Median longevity and K differed among phylogenetic groups but groups overlapped widely in these traits. Longevity, K, and size also varied among populations; in some cases, longevity and K differed between populations by a factor of two or more. Growth differed between sexes in some species and males typically reached larger sizes than females. In addition, a population of Quadrula asperata exhibited two distinctly different growth trajectories. Most individuals in this population had a low-to-moderate value of K (0.15) and intermediate longevity (27 years) but other individuals showed extremely slow growth (K = 0.05) and reached advanced ages (72 years). Overall, longevity was related negatively to the growth rate, K, and K explained a high percentage of variation in longevity. By contrast, size and relative shell mass (g mm⁻¹ shell length) explained little variation in longevity. These patterns remained when data were corrected for phylogenetic relationships among species. Path analysis supported the conclusion that K was the most important factor influencing longevity both directly and indirectly through its effect on shell mass. The great variability in age and growth among and within species shows that allocation to growth is highly plastic in freshwater mussels. The strong negative relationship between growth and longevity suggests this is an important trade-off describing widely divergent life-history strategies. Although life-history strategies may be constrained somewhat by phylogeny, plasticity in growth among populations indicates that growth characteristics cannot be generalized within a species and management and conservation efforts should be based on data specific to a population of interest.

The role of insulin-like growth factor-1 (IGF-1) in growth and reproduction in female brown house snakes ( Lamprophis fuliginosus)

Insulin-like growth factor-1 (IGF-1) is a peptide hormone critically involved in the regulation of key life-history traits such as growth and reproduction. Its structure and function are well-characterized among diverse mammal, fish, and bird species; however, little is known regarding the activities of IGF-1 in non-avian reptiles, particularly snakes and lizards. Nevertheless, several unique characteristics of reptiles, such as high metabolic flexibility and remarkable diversity in life-history strategy, suggest that they are of great interest in the study of endocrinological mechanisms underlying the regulation and evolution of life-history traits. Here we test for a relationship between IGF-1 and individual feeding rate, growth rate and reproductive stage in lab-reared female offspring of wild-caught oviparous house snakes, Lamprophis fuliginosus. We confirm a positive correlation between IGF-1 and both feeding and growth rates in sexually immature snakes, similar to that reported in other taxa. We also show a family effect on IGF-1, suggesting that IGF-1 levels may be heritable in these snakes, and serve as an important target of selection to produce divergent life-history strategies. Furthermore, we provide evidence that suggests that IGF-1 may peak rapidly after first mating, and subsequently decline prior to egg-laying, a phenomenon not previously reported in other taxa. These findings suggest that further comparative study of IGF-1 in snakes may reveal both the extent to which IGF-1 function is conserved across major taxonomic groups, as well as novel and intriguing roles for IGF-1 in the regulation of reproductive activities.

Dispersal provided resilience to range collapse in a marine mammal: insights from the past to inform conservation biology

Population loss is often a harbinger of species extinction, but few opportunities exist to follow a species' demography and genetics through both time and space while this occurs. Previous research has shown that the northern fur seal (Callorhinus ursinus) was extirpated from most of its range over the past 200-800 years and that some of the extirpated populations had unique life history strategies. In this study, widespread availability of subfossils in the eastern Pacific allowed us to examine temporal changes in spatial genetic structure during massive population range contraction and partial recovery. We sequenced the mitochondrial control region from 40 ancient and 365 modern samples and analyzed them through extensive simulations within a serial Approximate Bayesian Computation framework. These analyses suggest that the species maintained a high abundance, probably in subarctic refugia, that dispersal rates are likely 85% per generation into new breeding colonies, and that population structure was not higher in the past. Despite substantial loss of breeding range, this species' high dispersal rates and refugia appear to have prevented a loss of genetic diversity. High dispersal rates also suggest that previous evidence for divergent life history strategies in ancient populations likely resulted from behavioral plasticity. Our results support the proposal that panmictic, or nearly panmictic, species with large ranges will be more resilient to future disturbance and environmental change. When appropriately verified, evidence of low population structure can be powerful information for conservation decision-making.

Above- and below-ground vertebrate herbivory may each favour a different subordinate species in an aquatic plant community

At least two distinct trade-offs are thought to facilitate higher diversity in productive plant communities under herbivory. Higher investment in defence and enhanced colonization potential may both correlate with decreased competitive ability in plants. Herbivory may thus promote coexistence of plant species exhibiting divergent life history strategies. How different seasonally tied herbivore assemblages simultaneously affect plant community composition and diversity is, however, largely unknown. Two contrasting types of herbivory can be distinguished in the aquatic vegetation of the shallow lake Lauwersmeer. In summer, predominantly above-ground tissues are eaten, whereas in winter, waterfowl forage on below-ground plant propagules. In a 4-year exclosure study we experimentally separated above-ground herbivory by waterfowl and large fish in summer from below-ground herbivory by Bewick’s swans in winter. We measured the individual and combined effects of both herbivory periods on the composition of the three-species aquatic plant community. Herbivory effect sizes varied considerably from year to year. In 2 years herbivore exclusion in summer reinforced dominance of Potamogeton pectinatus with a concomitant decrease in Potamogeton pusillus, whereas no strong, unequivocal effect was observed in the other 2 years. Winter exclusion, on the other hand, had a negative effect on Zannichellia palustris, but the effect size differed considerably between years. We suggest that the colonization ability of Z. palustris may have enabled this species to be more abundant after reduction of P. pectinatus tuber densities by swans. Evenness decreased due to herbivore exclusion in summer. We conclude that seasonally tied above- and below-ground herbivory may each stimulate different components of a macrophyte community as they each favoured a different subordinate plant species.

Monozygotic Twin Wolves with Divergent Life Histories

Genetic evidence for monozygotic (identical) twinning in mammalian species is rare in the literature. Here we report what may be the first pair of monozygotic twins identified in a wild caniform carnivore, the grey wolf ( Canis lupus ). One of these individuals remained in its natal population of Banks Island, Northwest Territories, Canada, while its twin migrated across the polar sea ice to the mainland. This suggests divergent life history strategies in genetically identical individuals, making this incidence of twinning particularly interesting.