Life-history variation in contrasting habitats: flowering decisions in a clonal perennial herb (Veratrum album)

Quantifying intraspecific demographic variation provides a powerful tool for exploring the diversity and evolution of life histories. We investigate how habitat-specific demographic variation and the production of multiple offspring types affect the population dynamics and evolution of delayed reproduction in a clonal perennial herb with monocarpic ramets (white hellebore). In this species, flowering ramets produce both seeds and asexual offspring. Data on ramet demography are used to parameterize integral projection models, which allow the effects of habitat-specific demographic variation and reproductive mode on population dynamics to be quantified. We then use the evolutionarily stable strategy (ESS) approach to predict the flowering strategy-the relationship between flowering probability and size. This approach is extended to allow offspring types to have different demographies and density-dependent responses. Our results demonstrate that the evolutionarily stable flowering strategies differ substantially among habitats and are in excellent agreement with the observed strategies. Reproductive mode, however, has little effect on the ESSs. Using analytical approximations, we show that flowering decisions are predominantly determined by the asymptotic size of individuals rather than variation in survival or size-fecundity relationships. We conclude that habitat is an important aspect of the selective environment and a significant factor in predicting the ESSs.     

Life-history variation and allometry for sexual size dimorphism in Pacific salmon and trout

Allometry for sexual size dimorphism (SSD) is common in animals, but how different evolutionary processes interact to determine allometry remains unclear. Among related species SSD (male : female) typically increases with average body size, resulting in slopes of less than 1 when female size is regressed on male size: an allometric relationship formalized as 'Rensch's rule' . Empirical studies show that taxa with male-biased SSD are more likely to satisfy Rensch's rule and that a taxon's mean SSD is negatively correlated with allometric slope, implicating sexual selection on male size as an important mechanism promoting allometry for SSD. I use body length (and life-history) data from 628 (259) populations of seven species of anadromous Pacific salmon and trout (Oncorhynchus spp.) to show that in this genus life-history variation appears to regulate patterns of allometry both within and between species. Although all seven species have intraspecific allometric slopes of less than 1, contrary to expectation slope is unrelated to species' mean SSD, but is instead negatively correlated with two life-history variables: the species' mean marine age and variation in marine age. Second, because differences in marine age among species render SSD and body size uncorrelated, the interspecific slope is isometric. Together, these results provide an example of how evolutionary divergence in life history among related species can affect patterns of allometry for SSD across taxonomic scales.     

Seasonal biomass allocation in a boreal perennial grass (Phalaris arundinacea L.) under elevated temperature and CO2 with varying water regimes

The aim of this study was to analyze and model how biomass is allocated to the leaves, stems, and roots of perennial grass (reed canary grass, Phalaris arundinacea L., hereafter RCG) under elevated temperature (ET) (+approx. 3 °C) and CO₂ (approx. 700 μmol mol^?1) and with variable groundwater levels (high to low water levels) in a boreal environment. For this purpose, RCG plants were grown in environmentally controlled chambers over two growing seasons (April–September of 2009 and 2010), and the plant organ biomass (leaves, stems, and roots) was measured seven times over the entire growing season. The results showed that biomass production was mainly allocated to the leaves (LMF) and stems (SMF) early in the growing season, to the stems in the middle of the growing season, and to the roots (RMF) later in the growing season. Compared to ambient conditions, ET treatments increased LMF and SMF, and decreased RMF over the growing season under well-water conditions. Under low groundwater level, ET treatments decreased LMF and increased RMF throughout the growing season, and increased SMF in early periods and then decreased later in the growing season. CO₂ enrichment did not significantly affect the seasonal allocation pattern between plant organs. The effect of the groundwater level on biomass allocation was stronger than that of the climatic treatments. In conclusion, plant phenology controlled the seasonal course of biomass allocation in RCG and climatic treatments affected biomass allocation to each of the three plant organs, while the direction and extent of climate-related changes in biomass allocation depended on the availability of groundwater. The influence of groundwater level appeared to be crucial for the carbon gain regarding the production of RCG biomass for energy purpose and the concurrent sequestration of carbon in soils under changing climate in the mire sites used to cultivate RCG.     

Life-history variation in Crepis tectorum (Asteraceae)

Summary Populations of the monocarpic plant Crepis tectorum were grown in a series of uniform environments to test the hypothesis that weedy populations are more r-selected than populations from a more natural habitat. Weedy populations exhibited a combination of r- and K-selected traits. The relatively rapid growth, the potential for a summer annual habit, and the relatively high fecundity that characterized at least one of the two weed populations studied were considered as r-selected traits favored in habitats of unpredictable duration. However, high levels of competition from other weedy species or from the crop in arable fields may explain at least some presumably K-selected traits observed in the weedy populations, e.g. relatively large seeds and late flowering in the summer. Results indicated that stress due to abiotic factors (strong winds, desiccation and nutrient deficiency) has been a more important selective factor than r- or K-selection, in non-weedy populations from calcareous grasslands (alvars) on the Baltic islands.     

Interactive effects of lateral shade and wind on stem allometry, biomass allocation, and mechanical stability in Abutilon theophrasti (Malvaceae)

The effects of lateral shade and wind on stem allometry, whole-plant biomass allocation, and mechanical stability were examined for Abutilon theophrasti in a fully factorial glasshouse experiment. Lateral shade from neighboring plants increased stem height by 33% relative to control plants grown individually, despite a decrease in plant dry mass. Intermittent wind decreased stem height by 18% in unshaded plants, but by only 3% in shaded plants. Surprisingly, both lateral shade and wind caused decreases in stem diameter, even with diameter controlled for height, resulting in low diameter?:?height ratios in wind-treated plants relative to untreated plants. Under shade, wind-treated plants had higher root allocation than untreated plants, which allowed wind-treated shade plants to compensate for a low diameter?:?height ratio. This did not occur in the absence of shade, where stem tissue density and root allocation of wind-treated plants did not exceed that of untreated plants. Nevertheless, wind-treated plants experienced low drag relative to untreated plants due to a lower leaf area. Consequently, stem deflections of wind-treated plants did not exceed those of untreated plants at any given windspeed. Our results document a complex interaction between shade and wind on plant morphology and suggest that the nature of this interaction is generally that lateral shade acts to reduce or eliminate thigmomorphogenic responses.     

Independent effects of drought and shade on growth,biomass allocation and leaf morphology of a flammable perennial grass Tetrarrhena juncea R.Br

Knowing the abundance of different plant species provides insights into the properties of vegetation communities, such as flammability. Therefore, a fundamental goal in ecology is identifying environmental conditions affecting the abundance of plant species across landscapes. Water and light are important environmental moderators of plant growth, and by extension, abundance. In the context of understanding forest flammability, the abundance of a flammable plant species in terms of its cover or biomass can shape the flammability of the whole vegetation community. We conducted a glasshouse experiment to determine the impact of drought and shade on growth, biomass allocation and leaf morphology of forest wiregrass Tetrarrhena juncea R.Br., a rhizomatous perennial grass. When it is abundant, this species is known to contribute substantially to the flammability of eucalypt forest understories (via both ignitability and combustibility). Contrasting hypotheses in the literature predict that drought can have a weaker, stronger, or independent (uncoupled) impact on plant growth when light is limiting. We used a randomized complete block design with ten treatments from the combination of two water levels (drought, well-watered) and five light levels (100%, 80%, 60%, 40%, 20%). Drought and shade were found to have independent effects on wiregrass growth, biomass allocation, and leaf morphology, supporting the uncoupled hypothesis. Growth showed greater plasticity in response to drought, while biomass allocation and leaf morphology showed greater plasticity in response to shade. Our results suggest that wiregrass is more likely to be abundant in terms of its cover and biomass when water is not limiting. Under these conditions, the increased wiregrass abundance could create a window of increased flammability for the forest ecosystem.

     

Effect of shade on the carbon and nitrogen allocation in a perennial tropical grass, Dichanthium aristatum

Changes in sward growth above ground by shade is a controversialsubject for field experimentation because the differential allocationof carbon (C) to shoots and roots is difficult to measure. Inthis experiment the effect of three levels of irradiance (100%,56% and 33% of full sunlight) on C and nitrogen (N) allocationin Dichanthium aristatum were studied under well-watered andwell-fertilized conditions. Dry matter accumulation, weightratio per organ and N allocation indices were studied duringtwo cycles of growth, after planting and after cutting. Shadehad no effect on the C and N influxes into the whole plant,showing that the N absorption is regulated by the C assimilation.However, C, and principally N, were preferentially allocatedto the laminae under reduced irradiance. Under 100% and 56%of full sunlight, more N was allocated to the stubble component.This situation was reversed in the lowest radiation level, indicatingthat N reserves, and not exclusively C reserves, may limit theregrowth of this perennial grass when growing under high levelsof shade. The higher shoot:root ratio under shade shows thedisadvantage in the use of radiation use efficiency calculatedfrom aerial biomass data when comparing differents levels ofshade. Key words: Growth, shading, reserves, C₄ species     

Reproductive allocation of biomass and nitrogen in annual and perennial Lesquerella crops

* BACKGROUND AND AIMS: The use of perennial crops could contribute to increase agricultural sustainability. However, almost all of the major grain crops are herbaceous annuals and opportunities to replace them with more long-lived perennials have been poorly explored. This follows the presumption that the perennial life cycle is associated with a lower potential yield, due to a reduced allocation of biomass to grains. The hypothesis was tested that allocation to perpetuation organs in the perennial L. mendocina would not be directly related to a lower allocation to seeds. * METHODS: Two field experiments were carried on with the annual Lesquerella fendleri and the iteroparous perennial L. mendocina, two promising oil-seed crops for low-productivity environments, subjected to different water and nitrogen availability. * KEY RESULTS: Seed biomass allocation was similar for both species, and unresponsive to water and nitrogen availability. Greater root and vegetative shoot allocation in the perennial was counterbalanced by a lower allocation to other reproductive structures compared with the annual Lesquerella. Allometric relationships revealed that allocation differences between the annual and the perennial increased linearly with plant size. The general allocation patterns for nitrogen did not differ from those of biomass. However, nitrogen concentrations were higher in the vegetative shoot and root of L. mendocina than of L. fendleri but remained stable in seeds of both species. * CONCLUSIONS: It is concluded that vegetative organs are more hierarchically important sinks in L. mendocina than in the annual L. fendleri, but without disadvantages in seed hierarchy.     

Interpreting reproductive allometry: Individual strategies of allocation explain size-dependent reproduction in plant populations

Size-dependent or allometric relationships between reproductive and vegetative size are extremely common in plant populations. Reproductive allometry where plant size differences are due to environmental variability has been interpreted both as an adaptive strategy of plant growth and allocation, and as the product of fixed developmental constraints. Patterns of development are crucial in defining reproductive allometry but development is not fixed across individuals. For example, environmental adversity (e.g. resource impoverishment) tends to favor reproduction at relatively small sizes – an adaptive response to environmental adversity. While small individuals may have lower reproductive output than large individuals, all plants should maximize their reproductive output and relative allocation to reproduction may be constant across sizes. Thus, where individual plants within a population initiate reproduction at different sizes, no significant reproductive allometry is an appropriate null expectation. Reproductive allometry occurs in plant populations where initiating reproduction at small sizes yields relatively high or low reproductive size at final development. Both of these outcomes are common in plant populations. Our interpretation of reproductive allometry combines previous adaptive and developmental constraint interpretations, and is the first to successfully explain the range of relationships in plant populations where relative allocation has been observed to increase, decrease or remain constant will increasing plant size.     

缙云山2种禾草种群生殖配置的比较研究

利用样方法对生长于重庆缙云山不同生境条件下的2种禾草--白茅[Imperata cylindrica (L.) Beauv.]和淡竹叶(Lophatherum gracile Brongn.)的生物量配置格局进行了研究.结果表明,白茅种群在整个生长期内各构件生物量配置格局变化不大,其中对根状茎的配置比例最高,达到70.18%,有性生殖的生物量分配比例为4.51%,没有秆构件;淡竹叶种群在生长季内生物量配置格局发生较大变化,营养生长期、花期和黄熟期根构件的生物量分别占总生物量的77.33%、36.39%和48.66%,叶构件则为19.35%、45.42%和27.17%,有性生殖的生物量分配比例为11.11%.与白茅相比,淡竹叶种群秆构件明显.二者在生物量配置格局上存在显著差异,反映了二者对各自生活环境的适应特征.     

Demography and reproductive strategies of a polycarpic perennial, Trillium apetalon (Trilliaceae)

To investigate the connection between demographic strategies and reproductive strategies of a polycarpic perennial herb, Trillium apetalon Makino, we conducted three studies. First, we monitored the fate of individuals and the flowering behavior of T. apetalon for 12 years and used a transition matrix model to analyze the demography of the population. The analysis revealed that it takes a long time for individuals to go through one-leaf stage in juveniles. Elasticity analysis showed that the survival of flowering individuals was a decisive factor in the dynamics of the population. Furthermore, we found that the average remaining lifetime of flowering individuals was high relative to the other three stages. Second, to elucidate the demographic consequences of organ preformation, we investigated the development of flower buds for future years. We observed three to six flower buds per rhizome, suggesting that flower buds for the next 3–6 years were ready in advance in this plant. Third, the results of breeding experiments clarified that although this species appears to have a substantial capacity for both inbreeding and outbreeding, inbreeding plays an important role in seed production, and that crossing experiments (direct cross-pollination and self pollination) yielded similar seed-ovule ratios to those obtained from open-pollinated individuals. Our three studies suggest that the adult survival and continuous flowering strategies of T. apetalon obtained from demographic analysis are closely interlinked with breeding systems and preformation of flower buds.     

Growth, biomass allocation and photosynthesis of invasive and native Hawaiian rainforest species

Growth, biomass allocation, and photosynthetic characteristics of seedlings of five invasive non-indigenous and four native species grown under different light regimes were studied to help explain the success of invasive species in Hawaiian rainforests. Plants were grown under three greenhouse light levels representative of those found in the center and edge of gaps and in the understory of Hawaiian rainforests, and under an additional treatment with unaltered shade. Relative growth rates (RGRs) of invasive species grown in sun and partial shade were significantly higher than those for native species, averaging 0.25 and 0.17 g g⁻¹ week⁻¹, respectively, while native species averaged only 0.09 and 0.06 g g⁻¹ week⁻¹, respectively. The RGR of invasive species under the shade treatment was 40% higher than that of native species. Leaf area ratios (LARs) of sun and partial-shade-grown invasive and native species were similar but the LAR of invasive species in the shade was, on average, 20% higher than that of native species. There were no differences between invasive and native species in biomass allocation to shoots and roots, or in leaf mass per area across light environments. Light-saturated photosynthetic rates (Pmax) were higher for invasive species than for native species in all light treatments. Pmax of invasive species grown in the sun treatment, for example, ranged from 5.5 to 11.9 μmol m⁻² s⁻¹ as compared with 3.0−4.5 μmol m⁻² s⁻¹ for native species grown under similar light conditions. The slope of the linear relationship between Pmax and dark respiration was steeper for invasive than for native species, indicating that invasive species assimilate more CO₂ at a lower respiratory cost than native species. These results suggest that the invasive species may have higher growth rates than the native species as a consequence of higher photosynthetic capacities under sun and partial shade, lower dark respiration under all light treatments, and higher LARs when growing under shade conditions. Overall, invasive species appear to be better suited than native species to capturing and utilizing light resources, particularly in high-light environments such as those characterized by relatively high levels of disturbance. Received: 30 December 1997 / Accepted: 1 September 1998     

Temporal variation in reproductive allocation in a shield bug Elasmostethus interstinctus

We investigated changes in the reproductive output and the effect of female phenotype on reproductive parameters in a shield bug Elasmostethus interstinctus (L.) (Heteroptera: Acanthosomatidae) over the whole reproductive period. At the beginning and the middle of the reproductive period eggs were smaller than at the end of the period. Clutch mass and number of eggs per clutch decreased in laying sequence, first clutches being much larger than any of the later ones. Lifetime fecundity correlated positively with female size: large females produced more eggs and lived longer than small ones. Egg size did not vary with female size. Offspring survival until adulthood increased with egg weight. Individuals overwinter before reproduction, and because the nymphs from later-laid eggs have the least time to gather resources before overwintering, it may be important for later-laid eggs to be of high quality. Reproductive allocation varies during the reproductive period; females allocate resources relatively more to offspring number at the beginning of the reproductive period and more to offspring quality at the end of their life.     

Between-population variation in size-dependent reproduction and reproductive allocation in Pinguiculavulgaris (Lentibulariaceae) and its environmental correlates

Several important fitness components in herbaceous perennial plants are commonly related to plant size: flowering probability, reproductive allocation and fecundity. However, evidence for such size-dependence of fitness components is mostly anecdotal and unconnected to other life history traits. Here we report size-dependence for flowering probability and reproductive allocation in 11 populations of Pinguicula vulgaris and relate it to environmental factors. Flowering probability was size-dependent in all populations of P. vulgaris , and indicated the existence of a threshold size for reproduction. Populations at low altitudes and in wet soils showed a significantly higher threshold size for reproduction. Reproductive mass was also size-dependent in all populations. We found considerable between-population differences in the slope and the intercept of the regression between plant size and reproductive mass. This variation was weakly related to the environmental factors measured. In general, relationships between different size-dependent fitness components were low. Instead of showing a covariation of traits, in line with interpretations in terms of life history "tactics", P. vulgaris seemed to independently vary each size-dependent fitness component in each locality. In particular, no significant relationship was found between threshold size for reproduction and the slope of size-dependent reproductive allocation, as predicted by previous authors. Neither we found a significant influence of somatic cost of reproduction on size-dependent fitness components.     

Life-history theory, fertility and reproductive success in humans

According to life-history theory, any organism that maximizes fitness will face a trade-off between female fertility and offspring survivorship. This trade-off has been demonstrated in a variety of species, but explicit tests in humans have found a positive linear relationship between fitness and fertility. The failure to demonstrate a maximum beyond which additional births cease to enhance fitness is potentially at odds with the view that human fertility behaviour is currently adaptive. Here we report, to our knowledge, the first clear evidence for the predicted nonlinear relationship between female fertility and reproductive success in a human population, the Dogon of Mali, West Africa. The predicted maximum reproductive success of 4.1+/-0.3 surviving offspring was attained at a fertility of 10.5 births. Eighty-three per cent of the women achieved a lifetime fertility level (7-13 births) for which the predicted mean reproductive success was within the confidence limits (3.4 to 4.8) for reproductive success at the optimal fertility level. Child mortality, rather than fertility, was the primary determinant of fitness. Since the Dogon people are farmers, our results do not support the assumptions that: (i) contemporary foragers behave more adaptively than agriculturalists, and (ii) that adaptive fertility behaviour ceased with the Neolithic revolution some 9000 years ago. We also present a new method that avoids common biases in measures of reproductive success.     

Life-history and sex allocation in Trypoxylon (syn. Trypargilum) lactitarse (Hymenoptera; Crabronidae)

Hymenopterans have become a model for the study of factors that govern sex allocation. In 1983, Seger proposed a model to study Sphecidae wasps with a strong prediction that sex ratio for univoltine wasps should be 1 : 1 (female : male), and for partially bivoltine species it should be male-biased. The present study investigates if Trypoxylon lactitarse (Saussure, 1867) is a univoltine or a bivoltine species and if Seger's model prediction fits the pattern of sex ratio found in this species. The study was carried out at Parque Municipal das Araucárias, in the municipality of Guarapuava, state of Paraná, southern Brazil, from December 2001 to December 2004. Nests of T. lactitarse were obtained using trap-nests drilled longitudinally to a depth to 80 mm with 7.0, 10.0 and 13.0 mm opening diameter. They were placed in a very heterogeneous site filled with araucaria forests, swamps and grasslands. Trypoxylon lactitarse showed two alternative life histories: either they pupated immediately and emerged as adults later in the same season (direct development), or they entered diapause, overwintering and pupating the following spring (delayed development). Its annual sex ratios were not significantly different from 1 : 1, and both sex ratio of overwintering and sex ratio of direct-developing wasps were also not significantly different from 1 : 1, in all years of this study. By examining these results, it was possible to conclude that although T. lactitarse is a multivoltine species, with four generations per year and two alternative life histories, its sex ratio did not support Seger's model.     

Baculum variation and allometry in the muskrat (Ondatra zibethicus): a case for sexual selection

Sexual selection is a powerful force that influences the evolution of a variety of traits associated with female mate choice and male–male competition. Although other factors have been implicated, sexual selection may be particularly important in the evolution of the genitalia. Traits under sexual selection typically have high phenotypic variance and positive allometry relative to non-sexual traits. Here, we test the hypothesis that the baculum (os penis) of the muskrat (Ondatra zibethicus) is under sexual selection by examining phenotypic variance and allometry relative to non-sexual traits. Muskrats were sampled from Ontario, Canada, and a variety of traits measured. Measurements included baculum length and width, and three non-sexual traits (skull length, skull width, hind foot length). We used coefficient of variation (CV) and allometric slopes calculated using reduced major axis regression to test our hypotheses. Baculum traits had significantly higher CV’s relative to non-sexual traits. Baculum traits also showed positive allometry, whereas all non-sexual traits had negative allometric relationships. In addition, baculum width had higher CV’s and steeper allometric slopes than baculum length, indicating that, in muskrat, baculum width may be more influenced by sexual selection than baculum length. Positive allometry of the baculum is consistent with other examples of mammalian genitalia, but contrasts with negative allometry found in many insects. Other examples of positive allometry and high phenotypic variance of the baculum have suggested that females may use the baculum as an indicator of male quality. “Good genes” indicator traits may be particularly important in species that mate in an environmental context that prohibits female assessment of male quality. Muskrats mate aquatically, and thus females may be unable to properly assess males prior to copulation.     

Geographical variation in allometry in the guppy (Poecilia reticulata)

Variation in static allometry, the power relationship between character size and body size among individuals at similar developmental stages, remains poorly understood. We tested whether predation or other ecological factors could affect static allometry by comparing the allometry between the caudal fin length and the body length in adult male guppies (Poecilia reticulata) among populations from different geographical areas, exposed to different predation pressures. Neither the allometric slopes nor the allometric elevations (intercept at constant slope) changed with predation pressure. However, populations from the Northern Range in Trinidad showed allometry with similar slopes but lower intercepts than populations from the Caroni and the Oropouche drainages. Because most of these populations are exposed to predation by the prawn Macrobrachium crenulatum, we speculated that the specific selection pressures exerted by this predator generated this change in relative caudal fin size, although effects of other environmental factors could not be ruled out. This study further suggests that the allometric elevation is more variable than the allometric slope.