Cyst bank life-history model for a fairy shrimp from ephemeral ponds

1. Ephemeral wetland habitats provide a useful model system for studying how life‐history patterns enable populations to persist despite high environmental variation. One important life‐history trait of both plants and crustaceans in such habitats involves hatching/germination of only some of the eggs/seeds at any time. This bet‐hedging leads to the development of a bank composed of dormant propagules of many ages. 2. The San Diego fairy shrimp, Branchinecta sandiegonensis (Crustacea: Anostraca), a dominant faunal element of ephemeral ponds in San Diego, California, is a suitable organism for studying the consequences of highly fluctuating environmental conditions. As a result of large‐scale habitat loss, the species is also endangered, and this motivated our specific study towards understanding the hatching dynamics of its egg bank for planning conservation efforts such as pool restoration and re‐creation. 3. We formulated a matrix population model using egg age within the bank to study the relationship between adult survival and reproduction, and survival in and hatching from the egg bank. As vital rates for fairy shrimp are only poorly known, we generated 48 matrices with parameters encompassing ranges of likely values for the vital rates of B. sandiegonensis. We calculated population growth rates and eigenvalue elasticities both for a static model and a model with periodic reproductive failure. 4. The model shows that in good filling events, population growth rate is very high and the egg bank is increased dramatically. While population growth rate is insensitive to long‐term survival in the egg bank in our static deterministic model, it becomes sensitive to survival in the egg bank when a regime of periodically failed reproductive events is imposed. 5. Under favourable conditions, it was best for shrimp to hatch from eggs as soon as possible. However, under a regime where failed reproductive events were common, it was best to hatch after several pool fillings. Because conditions change from favourable to unfavourable unpredictably, variation in age within the egg bank appears to be critical for the persistence of the population. This attribute needs to be carefully considered when restoring or creating new pools for conservation purposes.     

Drought and aquatic community resilience: the role of eggs and seeds in sediments of temporary wetlands

1. A long‐lived bank of propagules consisting of eggs, seeds and spores is one mechanism that allows aquatic communities to survive drought. A drying (drought) event is, for aquatic organisms in a temporary wetland, a phase from which communities must recover. Such a dry phase is often considered a disturbance but should not be considered adverse or catastrophic for the organisms that have evolved to live in temporarily wet habitats. 2. This paper explores the parallels between the egg bank of zooplankton and the seed bank of aquatic plants as means of survival in temporary wetlands. The resilience of communities in temporary wetland ecosystems is assessed by examining dormancy, hatching, germination, establishment and reproduction of animals and plants from the egg and seed banks of wetlands with a range of wetting and drying regimes. 3. Both the zooplankton and aquatic plants of the temporary wetlands studied rely on their egg and seed banks as a means for surviving drying. These communities recover after the disturbance of drying by means of specific patterns of dormancy, dormancy breakage, hatching, germination, establishment and reproduction. Spatial and temporal patterns of species richness allow resilience through dormancy, as not all species are present at all sites and not all species hatch and germinate at the same time. Multiple generations in the egg and seed bank and complexity of environmental cues for dormancy breakage also contribute to the ecosystem's ability to recover after a drying event. A persistent egg and seed bank allows species‐rich communities to hatch, germinate and develop rapidly once dormancy is broken. Rapid establishment of species‐rich communities that reproduce rapidly and leave many propagules in the egg and seed bank also facilitates community recovery on flooding of a temporary wetland after a drying event. 4. To maintain the diversity of temporary wetland communities through droughts and floods we need to manage the dry and wet phases of wetlands. To conserve a wide range of wetland types, we need to maintain a variety of hydrological patterns across the landscape.     

Predictions of climate change infer increased environmental harshness and altered connectivity in a cluster of temporary pools

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Effect of habitat permanence on life-history: extending the Daphnia model into new climate spaces

We use an Australian freshwater invertebrate species, Daphnia carinata, to assess whether variation in habitat permanence influences life-history traits in subpopulations. Using a life table experiment, we measure the life-history traits of populations from both permanent and temporary pools. We show that these habitat classes are associated with clear differences in important life-history traits and evidence of trade-offs in important traits influencing reproduction, diapause, and growth rate and suggest this is evidence for local adaptation. Here we use Daphnia from Australian populations spanning semi-arid and temperate climates generating results that are in broad agreement with similar studies in the northern hemisphere, and so extend these results to a new continent and its particular climate. Variation in habitat permanence, it appears, is a very general driver of life-history divergence.

     

Foraging tactics in alternative heterochronic salamander morphs: trophic quality of ponds matters more than water permanency

1. In lentic freshwater habitats, the composition of animal assemblages shifts along a gradient from temporary to permanent basins. When habitats with different degrees of permanence are at the scale of the home range of species, they constitute alternatives in terms of energy acquisition through feeding. 2. In this context, previous studies showed an advantage of metamorphic over paedomorphic tiger salamanders (Ambystoma tigrinum) in temporary ponds which are only available to metamorphs. The aim of this study was to establish whether salamanders obtain similar benefits in ponds that do not differ in water permanence and whether salamanders shifted from detrimental to advantageous ponds. To this end, we determined the feeding habits, body condition and movement patterns of the two morphs in a complex of four permanent and four temporary ponds. 3. Consistent with previous studies, metamorphs consumed higher‐quality diets than paedomorphs in term of energy intake. However, these differences occurred because metamorphs consumed fairy shrimp in a single temporary pond. Individual movement patterns confirmed that most of the metamorphs used different aquatic habitats both within and between years and that most of them moved from permanent ponds for breeding towards the most profitable temporary pond in terms of foraging. 4. These results indicate that habitat selection by salamanders is optimal in term of energy intake in metamorphs that use high quality ponds independently of hydroperiod. It seems that both spatial and temporal variation can influence the relative foraging success of each morph.     

Community structure in temporary freshwater pools: disentangling the effects of habitat size and hydroregime

1. Hydroregime (duration, frequency and predictability of the aquatic phase) is a key feature of temporary aquatic habitats that not only moulds community structure and diversity (species sorting) but also life history characteristics of the inhabitants (natural selection). However, since hydroregime is a complex multidimensional entity that cannot be estimated from short term observations, morphometric variables are commonly used as proxies for hydroregime, making it impossible to separate effects of habitat size and hydroregime on biota.
2. We have used a simple hydrological model, validated with recent (average r ² = 96%) and historic water level observations (average r ² = 81%), to accurately reconstruct hydroregime based on historical rainfall and evaporation data in a cluster of 36 temporary rock pools in central South Africa.
3. Using the model output, we demonstrated that both hydroregime and habitat size had unique and shared effects on temporary pool biota and that these effects depended on the dispersal modes of the taxa. Model-generated hydrological data explained more variation in community patterns than short-term observations of hydroperiod. Hydroregime was more important for passive dispersers than for active dispersers that can migrate when pools dry up. Basin morphometry was a relatively poor predictor of pool hydroregime. We concluded that simple modelling may greatly improve the resolution of studies linking hydroregime to biological variables.
4. An accurate account of hydrological variation provides a firm foundation to understand community and population structure and dynamics in temporary aquatic habitats. Since many of these habitats have been destroyed or degraded in recent decades, our findings and tools may contribute to the development of reliable conservation guidelines.     

Dormancy and dispersal as mediators of zooplankton population and community dynamics along a hydrological disturbance gradient in inland temporary pools

At some stage in their life cycle, most zooplankton in temporary waters produce dormant eggs that assemble in a persistent egg bank to cope with unfavourable conditions. As part of a risk-spreading strategy, only a fraction of the egg bank hatches during a single inundation. Besides this dispersal in time, resistant dormant eggs also disperse in space via vectors including wind, water and animals. The structure and functioning of the dormant egg bank has important consequences for (meta) population and (meta) community structure and dynamics. Here, we merge empirical and theoretical data into a conceptual framework for the study of population and community responses in temporary ponds along a gradient in hydrological disturbance. Overall, we conclude that changes in hydrological disturbance may compromise both the abiotic (i.e. water quality) and biotic (i.e. population and community processes) integrity of temporary pools which is especially relevant in light of ongoing anthropogenic alterations in the hydrology of inland waters.     

Diapause, quiescence, hatching requirements: what we can learn from large freshwater branchiopods (Crustacea: Branchiopoda: Anostraca, Notostraca, Conchostraca)

The extent to which dormancy in large freshwater branchiopods is controlled endogenously (diapause) or exogenously (quiescence) is not always clear. It is assumed that both processes occur even within the same brood. Based on the effectiveness of common diapause-deactivating processes such as desiccation, hibernation, and resting, it can be stated that diapause is not a general process controlling responsiveness of large freshwater branchiopod eggs. Only in limited cases unequivocal evidence for the positive influence of these treatments is found.With few exceptions, hatching of activated cysts is effected by specific conditions that may even differ among conspecific populations. Generally, each species (or even population) has a specific temperature range or regime for optimal hatching performance. In a suitable thermal environment with sufficient light and oxygen, hatching is invariably invoked by a low osmotic medium.The erratic hatching pattern in most species is thought to be an adaptation to the variable temporary habitat. Hatching is generally spread over several days or even weeks, but the highest peak usually occurs on the first or second day of hatching. Low hatching percentages were found only in subtropical/desert species and may be a reflection of the low chances for successful reproduction. Generation carry-over of propagules in the egg bank by dormancy, and hatching at low conductivity, are, together with obligate oviparity and absence of an asexual life cycle phase, thought to be highly adaptive to the temporary environment. This enabled large branchiopods to survive since the Upper Cambrian.     

Life history adaptations of phyllopods in response to predators,vegetation, and habitat duration in north-eastern Natal

Phyllopod populations were monitored in three temporary pools differing in the amount of submerged, peripheral vegetation present, surface area and duration. The effects of these factors on the life history strategies employed by phyllopods were investigated. Triops granarius, various conchostracan species and the anostracan Branchipodopsis sp. inhabited the periphery of two pools where rooted, submerged vegetation was abundant while three Streptocephalus species dominated the central, unvegetated regions of the pools and the unvegetated pool. This distribution pattern appeared to be related to the animals' morphology and feeding habits. The peripheral regions of the pools were stressful habitats since they were colonized by large numbers of predators and competitors 30–40 days after inundation and they dried out sooner than the centre. The peripheral species exhibited typical r-selected life history strategies; they grew rapidly, reproduced early and had short lifespans and in this way they overcame the threats presented by their habitat. The central species took advantage of their predator-free, more stable habitat and exhibited life history patterns which tended towards the K-end of the r-K continuum. A degree of intraspecific variation in growth and reproduction was obvious and appeared to be related to differences in habitat duration of the three pools.     

Female fecundity traits in wild populations of African annual fish: the role of the aridity gradient

The evolution of life history is shaped by life expectancy. Life‐history traits coevolve, and optimal states for particular traits are constrained by trade‐offs with other life‐history traits. Life histories contrast among species, but may also diverge intraspecifically, at the level of populations. We studied the evolution of female reproductive allocation strategy, using natural populations of two sympatric species of African annual fishes, Nothobranchius furzeri and Nothobranchius orthonotus. These species inhabit pools in the Mozambican savanna that are formed in the rainy season and persist for only 2–10 months. Using 207 female N. furzeri from 11 populations and 243 female N. orthonotus from 14 populations, we tested the effects of genetic background (intraspecific lineage) and life expectancy (position on the aridity gradient determining maximum duration of their temporary habitat) on female fecundity traits. First, we found that variation in female body mass was small within populations, but varied considerably among populations. Second, we found that fecundity was largely defined by female body mass and that females spawned most of their eggs in the morning. Third, we found that the trade‐off between egg size and egg number varied among lineages of N. furzeri and this outcome has been confirmed by data from two separate years. Overall, we demonstrate that local conditions were important determinants for Nothobranchius growth and fecundity and that eggs size in arid region was less limited by female fecundity than in humid region.     

Better desiccated than eaten by fish: Distribution of anurans among habitats with different risks to offspring

1. Freshwater species with complex life cycles face a trade-off between the risks of offspring mortality due to desiccation in temporary habitats and due to predators common in long-duration habitats, especially fish. In real-world conditions, intermediate-gradient areas that are highly suitable for some ecological specialists are often limited. I examined the relative significance of drivers of the permanence–predation gradient in habitat selection by pond-breeding anurans.
2. Anuran oviposition was investigated at the community level in fish pond landscapes with only three types of habitat, permanent fish-containing ponds and highly desiccation-prone pools, either fishless or recently emptied of fish and with fish odour (fish cue). Altogether, 65 ponds/pools, interspersed in four clusters, were examined for the presence of egg masses.
3. Bufo bufo was the only species preferring permanent ponds. Egg masses of Rana sp. (Rana arvalis/Rana temporaria) occurred in all fishless pools, and less frequently in the ponds. Bufotes viridis and long larval period species Pelobates fuscus and Hyla orientalis oviposited almost exclusively in fishless pools. Fish-cue pools were avoided by all species. Anuran richness was higher in fishless pools than in permanent ponds. Species distribution between the two habitats was nested, with both common and rare species occurring in fishless pools.
4. The results indicate the primacy of offspring predation risk over hydroperiod constraints in oviposition decision-making by fish-intolerant anurans. The absence of some species from ponds and pools with fish or fish cues shows that non-consumptive interactions may better explain the scarcity of anuran larvae in waters dominated by fish than actual consumption. The strict avoidance of fish habitats despite their proximity to fishless patches indicates fine-scale assessment of predation risk. However, rigid habitat selection against fish predation on offspring may prove maladaptive if habitats with high desiccation risk are the main alternative and are indiscriminately preferred. Maintenance of fishless wetlands with a hydroperiod sufficient to allow completion of metamorphosis should be a conservation priority for anuran diversity in areas where fishless habitats are limited.

     

NATURAL GENETIC VARIATION OF LIFE SPAN,REPRODUCTION, AND JUVENILE GROWTH IN DAPHNIA

The evolutionary theory of senescence predicts that high extrinsic mortality in natural populations should select for accelerated reproductive investment and shortened life span. Here, we test the theory with natural populations of the Daphnia pulex-pulicaria species complex, a group of freshwater zooplankton that spans an environmental gradient of habitat permanence. We document substantial genetic variation in demographic life-history traits among parent and hybrid populations of this complex. Populations from temporary ponds have shorter life spans, earlier and faster increases of intrinsic mortality risk, and earlier and steeper declines in fecundity than populations from permanent lakes. We also examine the age-specific contribution to fitness, measured by reproductive value, and to expected lifetime reproduction; these traits decline faster in populations from temporary ponds. Despite having more rapid senescence, pond Daphnia exhibit faster juvenile growth and higher early fitness, measured as population growth rate (r). Among populations within this species complex we observed negative genetic correlations between r and indices of life-history timing, suggesting trade-offs between early- and late-life performance. Our results cannot be explained by a trade-off between survival and fecundity or by nonevolutionary theories of senescence. Instead, our data are consistent with the evolutionary theory of senescence because the genetic variation in life histories we observed is roughly congruent with the temporal scale of environmental change in the field.     

Wind-borne short-range egg dispersal in anostracans (Crustacea: Branchiopoda)

Evidence is provided for the first time that at least part of the egg bank of the southern African fairy shrimp Branchipodopsis wolfi is dispersed over short distances by wind. A total of 423 sticky surfaces were mounted around and between individual basins at three rock pool sites, to trap any dispersing eggs during a 3 day period. Eight viable eggs were found in different egg traps (1.9% of the total), seven of which were located at one site with shallow basins. Given the proximity of vast egg banks to the egg traps, the low observed dispersal rate cautions against overestimating the importance of wind dispersal for population genetic processes. By generating egg banks to hedge against drought catastrophes, and producing egg types with different potential dispersibility, B. wolfi is a strong bet-hedger which has established a means of escaping temporal and spatial stress. These life history functions enable populations to persist in small desert rock-pools, which are the most extreme of temporary habitats.     

水蚤滞育的数量遗传学研究:遗传与环境控制及遗传作用

在单性生殖循环水蚤群体中,滞育卵由有性生殖产生．在一系列实验中用到了不同种群和种类的水蚤,通过这些实验来观察：1)有性生殖和滞育卵复苏的遗传和环境控制;2)生活在相同区域中,但有不同生存微环境的近缘种类的有性生殖的光周期反应;3)在群体遗传结构上有性生殖的遗传效应(基因型均值和生活史性状遗传方差)．结果发现：1)遗传作用和环境作用,以及两者的相互作用都对有性生殖和滞育卵的孵化有显著的影响．GE显著的相互作用对环境中观察到的有性生殖来说,有助于维持其较高的遗传方差;2)在相同区域中,不同生存微环境的近缘种类的有性生殖的光周期反应有所不同．这有助于进一步区别近缘的水蚤种类,这也可能是一个水环境中同素异形的物种形成的例子;3)在有性生殖上,生活史性状平均值和遗传方差变化与前代选择造成的均值和遗传方差相反(遗传滑阻),这会造成暂时的适应不良(遗传滑阻和隐藏的遗传变异的表达),应对它补偿滞育的进化优势．     

Landscape‐scale life‐history gradients in New Zealand freshwater fish

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Microgeographic life history variation in an alpine caddisfly: plasticity in response to seasonal time constraints

1. Temporally constrained environments, such as habitats with short growth seasons or short hydroperiods, cause potentially strong selection on life histories. Depending on the predictability of these events and the extent of spatial and temporal heterogeneity, local populations could become adapted either via a fixed phenotype or via life history plasticity in response to these environmental cues. 2. We used a common garden experiment to investigate microgeographic variation in life history responses to combined changes in photoperiod (ambient/late) and hydroperiod (constant/drying) time constraint cues in an alpine caddisfly (Trichoptera). We compared six populations (three permanent/three temporary streams) originating from a small, alpine floodplain and which spanned an expected gradient in growth period duration (GPD) with distance from glaciers. 3. We made two main predictions in relation to locally varying selection pressures: (i) populations nearest glaciers (shorter GPD and strongest time constraints) should have the fastest development rates and (ii) populations from permanent streams should be less able to respond to drying hydroperiods than populations from temporary streams. 4. All populations and both sexes accelerated development in response to late season photoperiod cues. However, only permanent stream populations showed an increase in development time with increasing GPD, suggesting that other factors were influencing populations in temporary streams. 5. Permanent stream populations showed countergradient variation (genetic and environmental influences were in opposition) in development time, and under‐compensation of growth rates resulted in a converse Bergmann cline in body size (smaller body size along gradients of declining season length). The extent of plasticity in response to hydroperiod, and the combined effects of both time constraints, differed between populations and sexes, but were not consistent among populations. 6. Taken together, our results suggest adaptive plasticity in response to season length. The lack of a predictable pattern in response to hydroperiod may be due to gene flow or weak selection. We conclude that spatially structured populations can strongly differ in phenotypic plasticity even at microgeographic scales.     

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

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

The evolution of an annual life cycle in killifish: adaptation to ephemeral aquatic environments through embryonic diapause

An annual life cycle is characterized by growth, maturity, and reproduction condensed into a single, short season favourable to development, with production of embryos (seeds, cysts, or eggs) capable of surviving harsh conditions which juveniles or adults cannot tolerate. More typically associated with plants in desert environments, or temperate‐zone insects exposed to freezing winters, the evolution of an annual life cycle in vertebrates is fairly novel. Killifish, small sexually dimorphic fishes in the Order Cyprinodontiformes, have adapted to seasonally ephemeral water bodies across much of Africa and South America through the independent evolution of an annual life history. These annual killifish produce hardy desiccation‐resistant eggs that undergo diapause (developmental arrest) and remain buried in the soil for long periods when fish have perished due to the drying of their habitat. Killifish are found in aquatic habitats that span a continuum from permanent and stable to seasonal and variable, thus providing a useful system in which to piece together the evolutionary history of this life cycle using natural comparative variation. I first review adaptations for life in ephemeral aquatic environments in killifish, with particular emphasis on the evolution of embryonic diapause. I then bring together available evidence from a variety of approaches and provide a scenario for how this annual life cycle evolved. There are a number of features within Aplocheiloidei killifish including their inhabitation of marginal or edge aquatic habitat, their small size and rapid attainment of maturity, and egg properties that make them particularly well suited to the colonization of ephemeral waters.     

Life history plasticity of Nemoura trispinosa (Plecoptera: Nemouridae) along a permanent-temporary water habitat gradient

1. The life history of the small herbivorous stonefly Nemoura trispinosa Claassen was studied in a variety of small springs in southern Ontario, Canada. Nymphs generally were able to tolerate a wide range of environmental conditions and were found in 78% of habitats sampled, although population densities differed markedly. 2. Life-cycle patterns varied from a univoltine, slow seasonal type to a univoltine, fast seasonal type with extended egg development. In one, highly stable, spring the life cycle was semivoltine. Inter-year variation was studied for 5 years in one spring and was found to be low relative to among-spring variation. 3. Differences in the life history traits of N. trispinosa populations from our spring series were most probably an expression of phenotypic plasticity rather than of genetic differentiation. 4. Maximum annual water temperature was the factor most influential on nymphal growth rate (non-linear relationship), whereas range in generation time was related to the degree of habitat permanence.