Seasonal timing of pseudoviviparous reproduction of Leiothrix (Eriocaulaceae) rupestrian species in South-eastern Brazil

BACKGROUND AND AIMS: Pseudovivipary is an asexual reproductive strategy. Leiothrix spiralis and L. vivipara (Eriocaulaceae) are pseudoviviparous and occur in rupestrian grasslands, a habitat that has a predominance of sandy and shallow soil, with low water retention. This study aims to investigate the seasonal variation effect of moisture availability on L. spiralis and L. vivipara pseudoviviparous reproduction, and to compare their life history attributes, on rupestrian grasslands in Southeastern Brazil. METHODS: A field study was conducted, including observations concerning pseudoviviparous reproduction and measurement of demographic variables in both L. spiralis and L. vivipara. Soil moisture measurements were also performed to study its effect on the pseudoviviparous reproduction of L. spiralis and L. vivipara. KEY RESULTS: Flower head and plantlet production in L. spiralis were highly correlated with soil moisture. All scapes split off in the drier period, indicating that this is a splitter ramet species. Plantlet mortality was positively correlated with scapes splitting off. The L. vivipara phenophases were not synchronized to the variation in soil moisture, since flower heads and plantlets were produced throughout the year. Moreover, the splitting off of scapes was not observed. In addition, plantlets were formed early, as soon as the flower heads appeared, and remained suspended. Therefore, this species was called 'canopy forming'. CONCLUSIONS: Seasonal timing of pseudoviviparous reproduction can be a vital component of the successful establishment of plantlets in L. spiralis, considering that in this species the plantlets are formed only after the flower head touches the ground. In contrast, in L. vivipara, the plantlets are formed early, without touching the ground. Moreover, L. spiralis is a splitter ramet species, while L. vivipara is a canopy-forming species. The pseudoviviparous canopy-forming strategy appears to be more advantageous than the splitter ramet strategy, because even under similar soil moisture conditions, the survival of L. vivipara plantlets was greater than that of L. spiralis.     

Dry soil determines the pseudoviviparous reproduction in Comanthera nivea (Eriocaulaceae) in the Campos Rupestres in south-eastern Brazil

Pseudovivipary is an asexual reproduction strategy correlated with extreme environments. Comanthera nivea occurs in habitats with low water retention. This study investigated the effect of soil moisture availability on C. nivea pseudoviviparous reproduction and flowering. We established four permanent plots (1 × 1 m), two in a shaded area, in which we marked 15 rosettes in each plot, and two in a full-sun area, in which we marked 10 rosettes in each plot. We made monthly observations from August 2016 to January 2017, to quantify the number of flower heads and of pseudoviviparous ramets per rosette, and measured soil moisture. Our results showed a high correlation between soil moisture and flowering. Flowering in C. nivea occurs in periods of increased soil moisture, whereas pseudoviviparous reproduction occurs in the driest period. The advantages of a pseudoviviparous canopy-forming strategy in habitats with scarcity of water in the soil are discussed.     

Genetic variation and evolutionary trade-offs for sexual and asexual reproductive modes in Allium vineale (Liliaceae)

Populations of Allium vineale commonly include individuals with very different allocation patterns to three modes of reproduction: sexual flowers, aerially produced asexual bulbils, and belowground asexual offsets. If selection is currently acting to maintain these different allocation patterns there must be a genetic basis for variation in allocation to these three reproductive modes. In addition, negative genetic correlations between reproductive traits would imply evolutionary trade-offs among reproductive strategies. We evaluated the heritability of these allocation patterns by growing 16 clones from a single population in the greenhouse at two levels of fertilization. Bulb mass and the mass and number of bulbils, offsets, and flowers were used as response variables, in addition to the proportion allocated to each reproductive mode. We found evidence of substantial heritable variation in allocation to sexual reproduction and in allocation within the two modes of asexual reproduction, indicating high sensitivity of these allocation patterns to natural selection. We also found evidence of strong negative genetic correlations between bulbil and flower traits, as well as between bulbil and offset traits, with one group of genotypes allocating greater resources to aerial asexual bulbils and the second group allocating more resources to belowground asexual offsets and aerial flowers. Phenotypic plasticity in allocation to above- vs. belowground asexual reproduction and sexual vs. asexual aerial reproduction was limited, indicating that plants are unlikely to change reproductive mode in response to nutrient availability. Together, then, we have demonstrated strong heritability for, and trade-offs in, the reproductive allocation patterns within this plant population.     

Floral variation and environmental heterogeneity in a tristylous clonal aquatic of the Pantanal wetlands of Brazil

MethodsStyle morph frequencies were sampled from 73 populations distributed across four flooding regimes differing in depth and duration. Measurements of flower size, sex-organ dimension, pollen size and pollen production were made in selected populations, and pollinator assemblages and their functional traits were recorded.ConclusionsThe similar morph structure and evenness of populations, regardless of flooding regime, suggest that sexual reproduction and clonal dispersal are sufficiently common to prevent the signature of founder events from dominating in a region. However, the pervasive occurrence of biased morph ratios in most populations suggests that many are in a non-equilibrium state. The reduced frequency of the mid-styled morph in trimorphic and dimorphic populations may be associated with the weak self-incompatibility of this morph resulting in selfing and inbreeding depression. Clonality in E. azurea and the weak self-incompatibility of the mid-styled morph may make it more vulnerable to geitonogamous selfing.     

Reproductive modes of three Ligularia weeds (Asteraceae) in grasslands in Qinghai-Tibet Plateau and their implications for grassland management

Ligularia virgaurea, L. sagitta and L. przewalskii are noxious weeds that are widely distributed in psychro-grasslands in the east of Qinghai-Tibet Plateau of China. However, little is known about reproductive modes and genetic diversity in the populations of L. virgaurea and L. sagitta under different environmental conditions, although such information is available for L. przewalskii. In this study, two experiments were conducted to examine the ex situ clonal growth of these weeds, and to determine the effects of reproductive modes on the diversity of their populations, using random amplified polymorphic DNA (RAPD) analysis. Transplanted seedlings of L. sagitta and L. virgaurea showed considerable clonal growth by forming rhizomes and ramets. The RAPD analysis showed that all individuals collected from natural populations were genetically different, indicating that no individuals originated from clonal growth. It is concluded that L. virgaurea and L. sagitta have the capacity for clonal growth, but the expression of their clonality is constrained by their growing conditions in natural grasslands. Sexual reproduction plays the major role in their population diversity in natural habitats, which requires that the control measures for these weeds should be targeted at the sexually reproducing populations.     

Life History Studies on the Genus Trillium (Liliaceae) I. Reproductive Biology of Four Japanese Species

Abstract Four Japanese Trillium (Liliaceae) species which are representative perennial herbs of the temperate deciduous forests— Trillium kamtschaticum (2x), T. apetalon (4x), T. tschonoskii (4x) and T. smallii (6x)—were studied for their reproductive characteristics, e.g., patterns of reproductive resource allocation and reproductive output.
In spite of the differences in ploidy levels (from 2x to 6x), all four species showed very similar reproductive traits. It became evident that in response to the increase in reproductive allocation to total reproductive organs (RA), the number of seeds produced per plant (P_N) clearly increased. This trend is well in accord with the relationship found in four North American species and also several temperate woodland perennial herbs which occur in closed, stable and predictable environments, and possess typical xenogamous breeding systems. Although there is no conspicuous trend between ploidy levels and P_N, one of the significant differences noted in this study was in seed weight, which ranged from 2.93 mg in diploids, to 3.42–3.45 mg in tetraploids, and to 4.47 mg in hexaploids.     

Environmental correlates of life-history variation in juvenile chinook salmon, Oncorhynchus tshawytscha (Walbaum)

Throughout its native North Pacific, the chinook salmon, Oncorhynchus tshawytscha (Walbaum), exists as twolife-history types that aredistinguished by the age at which juvenile salmon migrate to sea as smolts. 'Stream-type' chinook migrate seaward after I or more years of feeding in fresh water, whereas 'ocean-type' fish migrate to sea as newly emerged fry or after 2–3 months in fresh water. Stream-type chinook predominate in populations distant from the sea south of 56° N, and in both inland and coastal populations north of this point. By contrast, ocean-type chinook predominate in coastal populations south of 56° N, but are rare in populations in more northerly latitudes. Stream-type populations are associated with areas of low 'growth opportunity' (as measured by temperature and photoperiod regimes) and/or areas distant from the sea compared to ocean-type. Geographic variability in juvenile life history is suggested to result, in part, from environmental modulation of smolting timing via differences in growth opportunity among geo-graphic areas. In addition, differences in migration distance and temperature regime may result in selection for different sizes at migration among populations which, through differences in growth opportunity, might promote geographic variability in age at seaward migration.     

Reproductive cycles and reproductive strategies among populations of the Rose‐bellied Lizard Sceloporus variabilis (Squamata: Phrynosomatidae) from central Mexico

Species with wide distribution, generally show variations in life history characteristics, which can be attributed to environmental causes. In this study, we analyzed the reproductive cycle and reproductive characteristics from three populations (Atlapexco, San Pablo Tetlapayac, and Santa Catarina) of the lizard Sceloporus variabilis from central Mexico. The specific goal of this study was to evaluate life history characteristics such as reproductive period extent, SVL (snout‐vent length) at sexual maturity, clutch size, egg mass and volume, and RCM (relative clutch mass). The San Pablo Tetlapayac population showed a larger clutch size, RCM, egg mass, and a smaller SVL, body mass and reproductive period (January‐September), as well as egg volume than the Atlapexco and Santa Catarina populations. Reproductive cycle and reproductive characteristics were more similar between the Atlapexco and Santa Catarina populations. Differences found in the population of San Pablo Tetlapayac with respect to the Atlapexco and Santa Catarina populations could be attributed to environmental variations where lizard populations occur. Differences in the reproductive period and reproductive characteristics in each population could be the result of both historical (phylogenetic; e.g., reproductive mode) and nonhistorical (environmental; e.g., temperature, food availability) causes. This study showed that populations of the same species are under different selection pressures, and these affect the reproductive characteristics of populations. Our results also indicate that long‐term and targeted studies on predation, use and selection of food, are needed to determine the causes of these variations in populations of S. variabilis.     

The evolution of menopause in cetaceans and humans: the role of demography

Human females stop reproducing long before they die. Among other mammals, only pilot and killer whales exhibit a comparable period of post-reproductive life. The grandmother hypothesis suggests that kin selection can favour post-reproductive survival when older females help their relatives to reproduce. But although there is an evidence that grandmothers can provide such assistance, it is puzzling why menopause should have evolved only among the great apes and toothed whales. We have previously suggested (Cant & Johnstone 2008 Proc. Natl Acad. Sci. USA 105, 5332–5336 (doi:10.1073/pnas.0711911105)) that relatedness asymmetries owing to female-biased dispersal in ancestral humans would have favoured younger females in reproductive competition with older females, predisposing our species to the evolution of menopause. But this argument appears inapplicable to menopausal cetaceans, which exhibit philopatry of both sexes combined with extra-group mating. Here, we derive general formulae for ‘kinship dynamics’, the age-related changes in local relatedness that occur in long-lived social organisms as a consequence of dispersal and mortality. We show that the very different social structures of great apes and menopausal whales both give rise to an increase in local relatedness with female age, favouring late-life helping. Our analysis can therefore help to explain why, of all long-lived, social mammals, it is specifically among the great apes and toothed whales that menopause and post-reproductive helping have evolved.     

Evolution of the optimal reproductive schedule in the ant Camponotus (Colobopsis) nipponicus (wheeler): a demographic approach

1. Traits are hypothesised to optimise via natural selection. The schedule of reproduction is an important adaptive trait, but its evolution is difficult to study, as measuring parameters is usually difficult. However, the sufficient amounts of demographic data enable us to estimate these parameters. 2. Here, it is shown that the reproductive schedule of the ant Camponotus (Colobopsis) nipponicus is tuned to maximise the lifetime production of alates. 3. A colony started its reproduction 4 years after the colony founding, at which time they were far smaller than well‐developed colonies. This contradicts the prediction of the bang‐bang strategy theory. The size distribution of colonies in the study area showed that the mortality of small colonies is much higher than that of large colonies. 4. A simulation analysis suggests that the colonies that are smaller than the threshold can still achieve significant improvement in colny survival to the following year by investing all resources in colony growth instead of reproduction. A sensitivity analysis for the starting year of reproduction showed that the observed schedule maximises lifetime alate production. The demographic data suggest a stable population, which is required for optimisation through this maximisation. 5. The observed reproductive schedule must be optimised, and the breakdown of the bang‐bang theory is due to higher mortalities during the incipient stage of colonies. This study demonstrates that having enough demographic data creates a useful tool for studying the evolution of life‐history characteristics.     

Life history of the sandy beach amphipod Deshayesorchestia deshayesii (Crustacea: Talitridae) from Bizerta beach (North of Tunisia)

Population dynamics and reproductive activity of the amphipod Deshayesorchestia deshayesii were studied monthly from June 2007 to December 2008 in Bizerta beach. During this period, 6577 specimens of D. deshayesii were identified; the sex ratio was female biased, with a mean of the monthly sex ratios equal to 0.21?±?0.12. Due to the absence of embryos or eggs in the female brood pouches and juveniles in the samples, results suggested that the reproductive season extended from April to November, with a sexual resting period during winter from December to March. Brood size varied between 4 and 17 eggs and fecundity appeared to be correlated with female size. Six cohorts were identified on the first sampling date. Additionally, 12 new cohorts were detected during the study period. Life span was estimated at 5–7?months depending on the time of birth. Cohorts that appeared at the beginning of the reproductive period tended to have shorter lives than those born later in the season. These results suggest that D. deshayesii can be considered as a semi-annual species, with iteroparous females.     

Reproductive characteristics of Drosophila hibisci in the Northern Territory, Australia

Reproductive traits of Drosophila hibisci collected at 18 sites in the Northern Territory (NT) of Australia in May, 1998, as well as at two sites in north Queensland, in June, 1998, were compared to those from earlier work on a cline in ovariole number in D. hibisci along the east coast of Australia. The flies in the NT were considerably smaller, but had more ovarioles than comparably-sized flies on the east coast. Although the flies on the east coast showed an increasing number of ovarioles in populations at increasing distances from the equator, these new populations, both on the east coast and in the NT, reversed this trend, producing a generally U-shaped pattern of ovariole number with latitude among all populations. The northernmost and southernmost populations allocate more to ovariole numbers than populations in intermediate latitudes. Ovariole number is closely related to body size of females in all populations, but the regression coefficient is small at intermediate latitudes and increases at the northern and southern ends of the distribution. Egg volumes primarily varied with body size of the female (positive) and number of ovarioles per female (negatively), producing a generally inverted U-shaped pattern of egg volumes with latitude. Reproductive allocation patterns, but not thorax size or ovariole number, varied significantly in two samples taken 10 days apart at one NT site. This variation probably results from environmental differences across generations of developing larvae and is consistent with our earlier suggestion of substantial effects of the environment, primarily rainfall and temperature, on reproductive allocation in D. hibisci.     

Diversity of reproductive modes in anurans communities in the Caatinga (dryland) of northeastern Brazil

Anuran reproductive activities in xeric environments are strongly influenced by local rainfall and temperatures. Anuran species that inhabit the Caatinga biome in northeastern Brazil demonstrate numerous behavioral and reproductive strategies that avoid or minimize the negative effects of elevated temperatures and scarce water resources on egg development. Among the various anuran species found in the Caatinga region, members of the family Leptodactylidae demonstrate the greatest number of reproductive modes adapted to a terrestrial life style while most of the Hylidae species are more susceptible to variations in water availability. Laying eggs in foam nests is a common strategy among species that inhabit environments with open vegetation forms and water resources that are restricted to only short periods during the year. Data concerning anuran communities in the localities examined here indicated a greater diversity of reproductive modes in ponds with more diverse vegetation structures and longer periods of water availability indicating that the deeper the temporary ponds (permitting a longer hydroperiod) the more diverse will be the aquatic and bordering vegetation, and the richer will be the diversity of anuran species observed and the number of reproductive modes encountered there.     

Effect of reproductive modes and environmental heterogeneity in the population dynamics of a geographically widespread clonal desert cactus

The dynamics of plant populations in arid environments are largely affected by the unpredictable environmental conditions and are fine-tuned by biotic factors, such as modes of recruitment. A single species must cope with both spatial and temporal heterogeneity that trigger pulses of sexual and clonal establishment throughout its distributional range. We studied two populations of the clonal, purple prickly pear cactus, Opuntia macrocentra, in order to contrast the factors responsible for the population dynamics of a common, widely distributed species. The study sites were located in protected areas that correspond to extreme latitudinal locations for this species within the Chihuahuan Desert. We studied both populations for four consecutive years and determined the demographic consequences of environmental variability and the mode of reproduction using matrix population models, life table response experiments (LTREs), and loop and perturbation analyses. Although both populations seemed fairly stable (population growth rate, λ∼1), different demographic parameters and different life cycle routes were responsible for this stability in each population. In the southernmost population (MBR) LTRE and loop and elasticity analyses showed that stasis is the demographic process with the highest contributions to λ, followed by sexual reproduction, and clonal propagation contributed the least. The northern population (CR) had both higher elasticities and larger contributions of stasis, followed by clonal propagation and sexual recruitment. Loop analysis also showed that individuals in CR have more paths to complete a life cycle than those in MBR. As a consequence, each population differed in life history traits (e.g., size class structure, size at sexual maturity, and reproductive value). Numerical perturbation analyses showed a small effect of the seed bank on the λ of both populations, while the transition from seeds to seedlings had an important effect mainly in the northern population. Clonal propagation (higher survival and higher contributions to vital rates) seems to be more important for maintaining populations over long time periods than sexual reproduction.     

Reproductive senescence: new perspectives in the wild

According to recent empirical studies, reproductive senescence, the decline in reproductive success with increasing age, seems to be nearly ubiquitous in the wild. However, a clear understanding of the evolutionary causes and consequences of reproductive senescence is still lacking and requires new and integrative approaches. After identifying the sequential and complex nature of female reproductive senescence, we show that the relative contributions of physiological decline and alterations in the efficiency of parental care to reproductive senescence remain unknown and need to be assessed in the light of current evolutionary theories of ageing. We demonstrate that, although reproductive senescence is generally studied only from the female viewpoint, age‐specific female reproductive success strongly depends on male–female interactions. Thus, a reduction in male fertilization efficiency with increasing age has detrimental consequences for female fitness. Lastly, we call for investigations of the role of environmental conditions on reproductive senescence, which could provide salient insights into the underlying sex‐specific mechanisms of reproductive success. We suggest that embracing such directions should allow building new bridges between reproductive senescence and the study of sperm competition, parental care, mate choice and environmental conditions.     

Life History Studies on the Genus Trillium (Liliaceae) III. Reproductive Biology of Six Sessile-flowered Species Occurring in the Southeastern United States with Special Reference to Vegetative Reproduction

Abstract Life history characteristics, e.g., stage class structures of natural populations and associated reproductive characteristics, were studied for six sessile-flowered, diploid Trillium (Liliaceae) species that are restricted to the Coastal Plain region of the southeastern United States.
These sessile-flowered species reproduce primarily by vegetative propagation. Leaf area measurements were used to categorize stage class distributions of natural populations. Populations propagating mainly by vegetative reproduction had non-sigmoidal, convex-shaped population structures with most plants being in the intermediate stage classes. Low numbers of individuals in smaller stage classes and an abundance of intermediate sized individuals are due to the fact that vegetative propagules are the primary method for maintenance of local populations.
Characteristics associated with sexual reproduction were also examined. Seed output averaged about 20 seeds per plant, except for T. lancifolium which produced 9.7 seeds on the average. Seed setting rates varied between 15.61–44.15% and mean seed weight varied from 4.89 to 10.75 mg. The dominance of vegetative versus sexual reproduction may be due to the fact that all six species occur in ecologically unstable flood plain habitats.     

Association of ants with juvenile Ogyris amaryllis amaryllis Hewitson (Lepidoptera: Lycaenidae) in south-eastern Queensland

The life history of the mistletoe-feeding butterfly Ogyris amaryllis is commonly thought to involve an obligate association with ants. The distribution of eggs of subspecies O. a. meridionalis Bethune-Baker, which occurs over much of the Australian interior, is strongly associated with Iridomyrmex ants. This study examined the level and frequency of ant association exhibited by larvae and pupae of O. a. amaryllis , which occurs in coastal areas of eastern Australia. The distribution of juveniles was compared with that of ants at the scale of sites, mistletoe host trees, and larval sheltering locations. A significant positive association between larvae and ants was detected, although this association was weak and non-specific, indicating that O. a. amaryllis is facultatively myrmecophilous. Observed ant attendance of larvae was uncommon (5.6%). Available evidence suggests that either plasticity exists in the degree of myrmecophily of O. amaryllis , with different degrees of association expressed under different ecological circumstances, or different degrees of myrmecophily are characteristic of subspecific status.     

Geographic variation in life-history traits of the endemic kite skate Dipturus chilensis (Batoidea: Rajidae), along its distribution in the fjords and channels of southern Chile

Life-history traits of kite skates Dipturus chilensis were examined from two regions (c. 2286 km apart) in the sheltered fjords and channels of southern Chile. A total of 482 and 403 specimens were collected from the southern fjords (c. 42–46° S) and the fjords of Chilean Patagonia (c. 51–54° S), from September 2003 to 2004, respectively. Vertebra marginal increment analysis indicated an annual deposition of growth rings which was completed during the winter months. For each region, von Bertalanffy growth parameters showed that females attained a larger asymptotic size, L_∞, had a lower growth coefficient, K, and lived longer than males. Growth analysis indicated that D. chilensis from the Patagonian fjords had a longer life span (females: 22 v. 21 years; males 19 v. 17 years), attained a larger L_∞ (females: 150 v. 136 cm; males: 122 v. 118 cm total length, L_T) and had a lower K value (females: 0·087 v. 0·104; males: 0·110 v. 0·116) than their counterparts in the southern fjords. Comparisons with previous studies indicated that D. chilensis from both southern and Patagonian sheltered fjords had larger L_∞, and grew more slowly than their counterparts from central-southern Chile (L_∞= 119–123 cm, K= 0·123–0·127), suggesting latitudinal variations in growth. Females attained sexual maturity later than males in both regions. For both sexes, lengths at 50% maturity (L_50%) between regions were similar (females: c. 103 cm; males: c. 87 cm L_T); however, D. chilensis from Patagonia appeared to mature 1 year earlier (females: 13 v. 14 years; males: 10 v. 11 years). Specimens from Patagonia had a lower ovarian fecundity than those from the southern fjords. An increase in the proportion of mature females and males during summer, suggests that the reproductive peak occurs in this season, and no regional differences were found. The size of the egg cases increased with maternal L_T and these were longer in Patagonia. The information provided here represents the first evidence of regional variations in life-history traits for elasmobranchs in the south-eastern Pacific.