Masting, fire-stimulated flowering, and the evolutionary ecology of synchronized reproduction.

Synchronized episodic reproduction among long-lived plants shapes ecological interactions, ecosystem dynamics, and evolutionary processes worldwide. Two active scientific fields investigate the causes and consequences of such synchronized reproduction: the fields of masting and fire-stimulated flowering. While parallels between masting and fire-stimulated flowering have been previously noted, there has been little dialogue between these historically independent fields. We predict that the synthesis of these fields will facilitate new insight into the causes and consequences of synchronized reproduction. Here we briefly review parallels between masting and fire-stimulated flowering, using two case studies and a database of 1870 plant species to facilitate methodological, conceptual, geographical, taxonomic, and phylogenetic comparisons. We identify avenues for future research and describe three key opportunities associated with synthesis. First, the taxonomic and geographic complementarity of empirical studies from these historically independent fields highlights the potential to derive more general inferences about global patterns and consequences of synchronized reproduction in perennial plants. Second, masting's well developed conceptual framework for evaluating adaptive hypotheses can help guide empirical studies of fire-stimulated species and enable stronger inferences about the evolutionary ecology of fire-stimulated flowering. Third, experimental manipulation of reproductive variation in fire-stimulated species presents unique opportunities to empirically investigate foundational questions about ecological and evolutionary processes underlying synchronized reproduction. Synthesis of these fields and their complementary insights offers a unique opportunity to advance our understanding of the evolutionary ecology of synchronized reproduction in perennial plants.

Loss of Avian Intromittent Organs as a Sperm Competition Strategy: A Race to Be Last.

AbstractThe loss of the intromittent organ (IO) in the majority of birds remains unexplained. Here, I propose that IO loss results from sperm competition in the context of the unique avian sperm storage system. The first stage of fertilization is the movement of sperm through the vagina from the site of ejaculation to the sperm storage tubules (SSTs) at the uterovaginal junction. In a second stage, sperm are released from the SSTs and move through the uterus to the site of fertilization in the infundibulum. Last-male fertilization precedence can occur if sperm that arrive later at the uterovaginal junction occupy uterus-side SSTs, which then have a head start in the race to the infundibulum with each ovulation. Under such "last-in, first-out" conditions, there is strong sperm competition to arrive later at the SSTs. Consequently, the optimal male strategy would be placement of the ejaculate at the cloacal opening to allow any other sperm in the vagina to reach the SSTs first. Cloacal placement is effectively achieved by loss of the IO. The evolution of altricial development in Neoaves, the largest clade that lacks IOs, created conditions that favor IO loss. Specifically, the smaller clutch sizes and hatching asynchrony of altricial birds increase the intensity of sperm competition for fertilization of early eggs in the laying sequence and thus the selective advantage of later arrival at the SSTs. The rarity of IO loss among all animals suggests that the complex mechanism of avian fertilization creates unique conditions for sperm competition.

The ecosystem impacts of dominant species exclusion in a prairie restoration.

Dominant species often have disproportionately high abundance in restored communities compared to native remnants, which potentially could reduce the conservation value of restorations. Research is needed to determine how the abundance of dominant species in restoration plantings affects community assembly, species diversity, and ecosystem function. Most studies of dominant species in grasslands were modeled after experiments on keystone species, using the short-term experimental removal of dominants to test their functional role in ecosystems. However, the removal of established dominants constitutes a major disturbance that may influence the interpretation of their long-term functional impact. To address this, we experimentally assembled high-diversity tallgrass prairie communities that included or excluded the predicted dominant species (Andropogon gerardii and Sorghastrum nutans) from the seed mix at the time of planting, but without further manipulation of community composition. From 2013 to 2019, we measured several ecosystem functions and community dynamics in the presence or absence of dominants. Communities that included the dominant species had lower species richness, greater aboveground biomass, and reduced light availability at the soil surface. Dominant species presence also increased soil nutrient availability and rates of litter decomposition, although dominant grass litter decomposed more slowly than litter from other common species in both treatments. In the absence of the dominant grasses, communities were instead dominated by a common unplanted forb, Solidago altissima, and there was partial compensation in ecosystem functioning in these forb-dominated communities. The effects of dominant species exclusion may only be apparent in long-term studies of experimentally assembled communities that avoid the legacy effects associated with removal experiments. Furthermore, our results suggest that prairie restorations that limit or exclude the dominant grasses in seed mixes may achieve higher species diversity, increasing the conservation value of these systems.

The evolution of androgenesis.

It is well known that some species produce offspring carrying only female chromosomes by processes such as apomixis and parthenogenesis (generically termed "gynogenesis"). There are also several cases of natural reproduction by androgenesis in which diploid offspring carry nuclear chromosomes from only the male parent. We used population genetics models to investigate the conditions for invasion of rare androgenesis alleles and the consequences of their spread. Our models predict that androgenesis alleles often spread to fixation. If fixation causes the loss of females or female function in the population, population extinction occurs. Therefore, androgenesis alleles represent a new class of selfish genetic elements. Extinction is more likely in dioecious species than in hermaphrodites. Within dioecious species, extinction is more likely when androgenesis occurs via paternal apomixis (vs. fusion or doubling of haploid nuclei) and when females are the heterogametic sex (vs. male heterogamety). The apparent rarity of androgenesis compared to gynogenesis could be because androgenesis is harder to detect and more often leads to population extinction. Also, there could be greater evolutionary constraints on the origin of mutations for androgenesis. We suggest characteristics of groups in which further cases of androgenesis are more likely to be found.

Random amplified polymorphic DNA markers reveal genetic variation in the symbiotic fungus of leaf-cutting ants.

RAPD markers were used to examine the degree of genetic variation within the putatively asexual basidiomycete fungus (Lepiotaceae: provisionally named Leucoagaricus gongylophorus) associated with the leaf-cutting ant species Atta cephalotes. We analyzed fungal isolates from ant nests in two geographically distant sites, two isolates from Panama and five isolates from Trinidad. Ten decamer primers were used to amplify total DNA from these seven fungal isolates, and RAPD banding patterns were compared. Genetic similarity among isolates was determined by pair-wise comparisons of the shared number of DNA bands on an agarose gel. There was considerable genetic variation among isolates of the symbiotic fungus even within sites. Pairs of fungal isolates from the two different sites shared an average of only 36% of the bands in their RAPD profiles, while pairs from the within sites shared an average of 72% of the bands. RAPD markers may be useful for further investigation of the genetic structure of the fungal symbiont within species of leaf-cutting ants.

An exception to Darwin's syndrome: floral position, protogyny, and insect visitation in Besseya bullii (Scrophulariaceae).

Darwin pointed out that plants with vertical inflorescences are likely to be outcrossed if the inflorescence is acropetalous (flowers from the bottom up), the flowers are protandrous (pollen is dispersed before stigmas are receptive), and pollinators move upward on the inflorescence. This syndrome is common in species pollinated by bees and flies, and very few exceptions are known. We investigated flowering phenology and pollinator behavior in Besseya bullii (Scrophulariaceae) and found that it did not fit Darwin's syndrome. The vertical inflorescence was acropetalous but the flowers were distinctly protogynous, so flowers with newly receptive stigmas appeared on the inflorescence above those with dehiscing anthers. A number of small insects visited B. bullii; bees in the family Halictidae (Augochlorella striata and Dialictus spp.) were most common. When insects moved between gender phases within inflorescences, they moved up more often than down (61% versus 39% of observations, respectively) but this difference was only marginally significant. Most visits were to male-phase flowers only, and this preference was more pronounced for pollen-foraging insects than for nectar-foraging insects. B. bullii was self-compatible, so its flowering characteristics potentially could result in considerable self-pollination. However, an average of 38% of the lowermost flowers opened before any pollen was available on the same inflorescence; these "solo females" had a high probability of outcrossing (though fruit set was relatively low in the bottom portion of the inflorescence). Upper flowers may also be outcrossed because downward insect movement was not uncommon. Therefore protogyny in B. bullii may not necessarily lead to more selfing than would protandry.

Characteristics of pollen production in a population of New Zealand snow-tussock grass (Chionochloa pallens Zotov).

Intraspecific variation in pollen production and pollen size were studied within a population of Chionochloa pallens, an alpine snow-tussock grass of New Zealand. I sampled plants from two sites at the altitudinal extremes (1620 and 1070 m) of the distribution of C. pallens on Mt. Hutt, South Island. There was a highly skewed distribution of pollen production among plants; the largest 10% of plants produced over a third of the pollen. In contrast to some other species, pollen production in C. pallens can be accurately estimated by simple measures such as tussock area and number of flowering shoots. Other components of yield (number of spikelets per shoot, number of florets per spikelet, and average anther length) varied significantly among individuals, but these characteristics had a small impact on total pollen yield. There was continuous variation in pollen size among individuals; between the plants with the largest and the smallest pollen grains, there was a 21 % difference in pollen diameter and a 76% difference in pollen volume. Interplant differences in pollen size have been found in a number of plants, although the biological significance of this variation is unknown. There were few differences in pollen characteristics between sites. Plants at the low-altitude site had slightly greater pollen yield because total plant size was greater.

SEX ALLOCATION AND OUTCROSSING RATE: A TEST OF THEORETICAL PREDICTIONS USING BROMEGRASSES (BROMUS).

Predictions of sex-allocation theory were tested by comparisons among hermaphroditic bromegrass (Bromus) species that differed in outcrossing rate. Relative maternal and paternal investment were calculated using both the ratio of pollen to seed production, and absolute allocations in units of energy, nitrogen, phosphorus, potassium, magnesium, and calcium. Outcrossing rate had a large effect on sex allocation; species having greater outcrossing rates had relatively more paternal reproductive effort. Bromus inermis was obligately outcrossing, and nearly half of its reproductive effort was devoted to pollen production. Three partially outcrossed species, B. kalmii, B. ciliatus, and B. latiglumis, invested between 5% and 11% of reproductive effort in pollen production. Paternal investment was less than 2% in the selling species B. tectorum. Estimates of sex allocation were relatively unaffected by the resource currency used in calculation. The differences among species in sex allocation were mostly due to differences in anther size and seed set.>.

Intrapopulation gender variation in common ragweed (Asteraceae: Ambrosia artemisiifolia L.), a monoecious, annual herb.

We measured variation in gender among individuals within populations of ragweed (Ambrosia artemisiifolia) in an abandoned old-field and in the greenhouse. There was great variability in sex expression, from all-female to approximately 78% male. Plants differed significantly in gender in different locations within the field. Plants in an area abandoned from agriculture one year previously were more male than plants in a nearby area abandoned four years previously. In the greenhouse, soil moisture treatments and levels of attack by spittlebugs (Philaenus spumarius) did not affect gender. Height was positively correlated with relative maleness in both populations. Plants with greater shoot weight were relatively more male in the greenhouse, but not in the field. The gender variation we observed either has a genetic basis or is controlled by environmental variables other than those we investigated.