Dichogamy,gender variation and bet-hedging inPseudowintera colorata

Summary Temporal patterns of variability in the longevity of the male and female phases of individual flowers and in the gender expression of plants of a dichogamous New Zealand tree,Pseudowintera colorata (Winteraceae), were documented in field studies. Two measures for the duration of phases in a dichogamous flower are distinguished; the nominal phases based on morphological features of the flower, and the effective phases reflecting the duration of their functions. Flower and phase longevity and phenotypic gender varied considerably throughout the season and among individuals. Temporal variability in phenotypic gender was loosely synchronized among the 12 plants sampled. Three effects of an environmental factor (temperature) were noted. First, increased temperatures shortened the duration of the female phase but had no effect on the duration of the male phase. Second, pollination frequency was positively correlated with temperature. These results indirectly suggest that increased pollination may shorten the duration of the female phase. Third, average population maleness, measured as the proportion of open flowers in the population on a given day which were in the male phase, was positively correlated with temperature. It is postulated that temperature indirectly influences temporal patterns of gender expression in the population through its differential effects on the longevity of the male and female phases in individual flowers. A theoretical model of bet-hedging shows that, if the direction of an environmental effect on the proportions of the sexual phases is irreversible, selection favours asynchronous dichogamy and reduces the temporal variability as much as possible. If the direction of the response is reversible, heterodichogamy is favoured.     

The population-dynamic functions of seed dispersal

We summarize some of the population-dynamic consequences of the mosaic structure of plant populations for the evolution of seed dispersal. A fairly elaborated set of theoretical ideas exist regarding the evolution of dispersal and we have synthesized some of them in an attempt to make them more accessible to field ecologists. We consider the relationship of these general theoretical ideas to our understanding of fruit and seed dispersal.We develop three related models to describe the similarities and differences in how dispersal functions for risk reduction (bet hedging), escaping the negative consequences of crowding, and escaping high concentrations of relatives. We also briefly discuss directed dispersal as a fourth population-dynamic aspect of dispersal. Dispersal can have a risk-reducing function only when there is global (metapopulation) temporal variance in success. Dispersal to escape the negative consequences of crowding requires only spatial and local temporal environmental variation. Dispersal for escaping high concentrations of relatives requires no environmental variation, but does require genetic population structure. Directed dispersal, defined as non-random into particular patch types contingent on the expectation of local success, is always valuable when possible and represents an advantage independent the others which can occur with random dispersal.In an effort to accommodate for the differences between simple mathematical models and the behavior of complex natural fruit and seed dispersal systems we have discussed the following issues: actual patterns of patch structure and dispersal distance; the implications of plant cosexuality, perenniality, and allocation costs of dispersal structures; and the impact of the detailed nature of density dependence, breeding systems, and genetic structure. We briefly compare the population-dynamic functions of dispersal presented here with the widely cited functions of colonization, escape, and directed dispersal. Finally, we suggest how the theoretical models can be used with field data to estimate the fitness consequences of dispersal.     

Flee or fight uncertainty: plant strategies in relation to anticipated damage

In order to cope with damage, plants have evolved a number of strategies. We incorporate two of those strategies, compensatory regrowth and escaping damage in time, into a mathematical model in an attempt to outline under what circumstances one or the other of these phenotypic traits will evolve. Escaping damage in time is accomplished by flowering and setting seeds at a point of time when the risk of damage is low, whereas a compensatory capacity is made possible by activating a proportion of meristems that are left dormant. Our analysis suggests that damage that is predictable in time will favour phenotypes that flower late in the season and that have a good compensatory capacity. As damage becomes less predictable in time, a strategy that implies flowering as early as possible in the season and with no compensatory capacity at all, becomes advantageous.     

Effects of plant density and nutrient levels on fruit polymorphism in Atriplex sagittata

Atriplex sagittata is a heterocarpic species producing three types of fruits which differ in morphology and ecological properties. This study focused on variation in biomass allocation into particular fruit types under different density and fertilization levels. The reduction in total weight under stressful conditions was accompanied by a reduction of reproductive structures in terms of both total fruit production and mean fruit weight. Allocation of biomass to particular fruit types under different density-fertilization treatments showed considerable variation. The non-dormant, bracteolate fruit type (termed C) contributed 80–90% to the total fecundity and its production was not affected by density-fertilization treatments. The production of this fruit type is higher in the upper part of the maternal plant stem. Production of the very dormant ebracteate fruit type (termed A) increased under favourable conditions and was greater on the lower part of the stem, whereas more of the so-called type B fruit (covered by bracteoles and exhibiting dispersal and dormancy characteristics intermediate between types A and C) was produced under suboptimal conditions and was concentrated in the middle part of the plant body. This pattern contradicts to some extent the theory that deeply dormant and less dispersible fruits will be produced under stressful conditions and may be considered a new model for the behaviour of plants with more than two heterocarpic fruits. The type B fruit, intermediate in both morphology and ecological behaviour, provides a continuum between extreme strategies. Received: 8 February 1998 / Accepted: 19 November 1998     

Below-ground herbivory promotes soil nutrient transfer and root growth in grassland

Extremely little is known about the ecosystem-level implications of below-ground herbivory, which often represents the dominant form of consumption of primary productivity. We provide the first empirical evidence that low levels of below-ground herbivory may promote soil nutrient flux and root growth of both host plants and companion plants. Low levels of white clover ( Trifolium repens L.) root infection by clover cyst nematodes ( Heterodera trifolii Goffart) increased root growth by 141% and 219% in the host plant and the uninfected neighbouring grass ( Lolium perenne L.), respectively. Root infection increased the size of the soil microbial biomass in the root zone and the transfer of ¹⁵N from the host plant to soil and the neighbouring grass. These data suggest that low amounts of below-ground herbivory may increase the transfer of plant carbon and nitrogen below-ground, leading to increases in root growth and soil nutrient recycling in grasslands. Presumably, such interactions will influence the competitive interactions between plant species, altering plant community structure in grasslands.     

How does seed heteromorphism influence the life history stages of Atriplex sagittata (Chenopodiaceae)?

Atriplex sagittata is an annual, heterocarpic species producing three types of fruits, (i) dormant ebracteate (further termed type A), (ii) dormant bracteate (type B) and (iii) non-dormant bracteate (type C). In this study, we investigated populations grown from particular fruit types under different density regimes, and assessed their fate in particular life stages throughout a complete growing season. Differences in germination date among populations grown from seeds of the three fruit types were found, followed by differences in population density and plant size. Low population density and total biomass was recorded in populations of types A and B fruit (which are smaller and exhibit less germination), but the low density was not associated with faster growth of individual plants. Type C fruit, with bigger seeds and higher germinability, generated populations with higher densities and higher total biomass than populations generated from type A and B. There was a significant effect of plant density on dry weight of individual plants, regardless of the fruit type from which they were grown. Under both density regimes, plants grown from types A and B fruit were lighter than those grown from type C fruit. Plants grown from different fruit types produced dispersal units (fruits+bracteoles) of different mean weight which increased from A to B to C. However, the total fruit weight was not significantly different when plant size was used as a covariate. Maternal effect on seed germinability, exhibited by plants grown from the three fruit types, was not significant. However, a remarkable trend was found. Lower germination was recorded in type A and B fruits produced by plants which grew under higher densities. It is concluded that plants derived from different fruit types differ in growth characteristics and population parameters which can have pronounced effect on the life cycle and population regeneration in successive years.     

Stable germination behavior but partly changing seed–seed interactions along a steep rainfall gradient

Germination marks a critical transition in plant life that is prone to high mortality. Strong selection pressure is therefore expected to finely tune it to environmental conditions. Our study on the common Mediterranean grass Brachypodium hybridum assessed whether germination behavior changes systematically along a steep natural rainfall gradient ranging from harsh desert to rather mild mesic-Mediterranean conditions. We specifically tested hypotheses that germination behavior confers greater risk-spreading in populations from drier, unpredictable environments, and that seeds from wetter populations are better competitors.In 14 populations (spanning 114–954 mm annual rainfall) we assessed three alternative key parameters of germination in a greenhouse experiment: between-year dormancy, days to emergence within a season, and temporal spread. Addition of neighbor seeds accounted for competition as another crucial environmental factor. In six of the 14 populations, we also compared seeds originating from corresponding north (more mesic) and south (more arid) exposed hill slopes to test whether germination patterns along the large-scale rainfall gradient are paralleled at this smaller scale.B. hybridum exhibited generally high germination fractions and rapid emergence with very little temporal spread, indicating overall little risk-spreading germination. Surprisingly, none of the three parameters changed systematically with increasing aridity, neither at large scale along the rainfall gradient nor at small scale between north and south exposures. Neighbor seeds, however, mildly suppressed germination. Germination of neighbor seeds, in turn, was more strongly suppressed by B. hybridum seeds from drier populations, and this effect was stronger for forb than for grass neighbor species.Our results provide strong evidence that increased risk-spreading germination is not a universal, essential strategy to persist in increasingly dry, unpredictable environments. They also highlight that competition with neighbors occurs even at the earliest plant life stage. Since neighbor effects were species-specific, competition among seeds can affect community composition at later plant stages.     

Polygynandry inPseudotropheus zebra, a cichlid fish from Lake Malawi

Synopsis Parental care in the Malawian cichlid fishPseudotropheus zebra BB is extensive and exclusively maternal; males contribute only genetic material. The costs of searching for multiple mates (in this case risk of predation on orally incubated eggs) suggested that females should be monandrous; microsatellite genetypes of seven brooding females and their young, however, reveal extensive multiple paternity in this species, with a mean of 3.8 paternal individuals per brood. Polygynandry inP. zebra is probably not maintained by selection for genetically diverse offspring; potential explanations include avoidance of inbreeding, and bet-hedging on other male characteristics that females are unable to evaluate when selecting a mate. The observed degree of multiple paternity strongly suggests that females are free to choose mates as they will, a prerequisite of many theories positing sexual selection as a key element in Malawi chichlid evolution. It should also result in elevation of effective population sizes, and thus be antagonistic to runaway evolution of male secondary sexual characteristics, but not necessarily to other modes of sexual selection.     

Polymorphic growth in larvae of Maculinea butterflies, as an example of biennialism in myrmecophilous insects

The presence of annual and biennial individuals within the same population has been recently demonstrated in the myrmecophilous butterflies Maculinea rebeli and Maculinea alcon, which present a cuckoo strategy inside Myrmica nests, and Maculinea arion which is a predatory species. Here, we present field and laboratory data on polymorphic larval growth in two other predatory species of Maculinea: M. teleius and M. nausithous. Body mass distributions of pre-pupation larvae were bimodal in both species. These results point to the existence of larvae that develop in 1 or 2 years. We also showed that the probability of pupation depended on larval body mass. In the case of M. teleius, the critical body mass at which larvae have a 50% probability of pupation is about 80 mg. We suggest that polymorphism in Maculinea may have evolved as an adaptation to life in ant nests, a habitat which protects them from predators and provides food. However, the quality of this resource is highly variable and unpredictable. According to the bet-hedging hypothesis, if the habitat is unpredictable, females should have an advantage by producing more variable offspring. In the case of Maculinea butterflies, this may involve maintaining larvae that develop in 1 or 2 years.     

Assortative Pairing and Life History Strategy

A secondary analysis was performed on preliminary data from an ongoing cross-cultural study on assortative pairing. Independently sampled pairs of opposite-sex romantic partners and of same-sex friends rated themselves and each other on Life History (LH) strategy and mate value. Data were collected in local bars, clubs, coffeehouses, and other public places from three different cultures: Tucson, Arizona; Hermosillo, Sonora; and San José, Costa Rica. The present analysis found that slow LH individuals assortatively pair with both sexual and social partners more strongly than fast LH individuals. We interpret this phenomenon as representing (1) an adaptation for preserving coadapted genomes in slow LH strategists to maintain high copying fidelity genetic replication while producing a lower number of offspring in stable, predictable, and controllable environments and (2) a bet-hedging adaptation in fast LH strategists, favoring the genetic diversification of a higher number of offspring in unstable, unpredictable, and uncontrollable environments.