Characterization of polymorphic microsatellites for the invasive grass Microstegium vimineum (Poaceae)

? Premise of the study: Microsatellite markers were developed for the invasive plant Microstegium vimineum (Poaceae) to assess its population structure and to facilitate tracking of invasion expansion. ? Methods and Results: Using 454 sequencing, 11 polymorphic and six monomorphic microsatellite primer sets were developed for M. vimineum. The primer sets were tested on individuals sampled from six populations in the United States and China. The polymorphic primers amplified di-, tri-, and tetranucleotide repeats with three to 10 alleles per locus. ? Conclusions: These markers will be useful for a variety of applications including tracking of invasion dynamics and population genetics studies.     

Limits to local spatial spread in a highly invasive annual grass (Microstegium vimineum)

The potential roles of seed and microsite limitation in local spatial spread of the invasive grass Microstegium vimineum were experimentally investigated in a woodland and open lawn in central New Jersey, USA. Plots (30 × 30 cm) in three sites previously unoccupied by M. vimineum for at least 8 years (woodland interior, woodland edge, and open lawn) were sown with ~262 M. vimineum seeds in early spring 2008. Seedling emergence, density, summer growth and autumn reproduction were compared to plots in a nearby control population where natural recruitment occurred. Seedling emergence was greatest in the open lawn (54% of seeds sown) where plants showed the greatest growth and reproduction due to high light availability. Seedling emergence was lowest in the woodland interior (24%) and edge sites (9%), and growth and reproduction were greatly reduced there (relative to the control). Plots in the open lawn supported a consistently high density of M. vimineum (>1,000 plants per m²) through the growing period (April to October). The number of seeds in both cleistogamous and chasmogamous spikelets was correlated with shoot dry mass and thus, total seed production was greatest in the sunny open lawn where plants were largest, despite high density there. Across all sites, plants in plots at the highest densities produced the most seeds. Total seed production correlated with levels of light, but not soil moisture. Both seed availability and microsite limitation may reduce the probability of establishment of new M. vimineum populations into previously unoccupied sites. Intraspecific density does not negatively affect survival or reproduction. Light and soil moisture can be limiting abiotic factors in some areas, but poor, natural seed dispersal limits the distribution of this invasive species on a local scale.     

Population differentiation in the tiller architecture of Microstegium vimineum (Poaceae) in relation to habitat

Grass tillers grow by addition of modular units known as phytomers. Differences in phytomer organ size produce subindividual variation with potential adaptive value. Here, patterns in organ mass along tillers in the invasive annual Microstegium vimineum are related to habitat and tiller architecture. In an earlier study, seed families were collected from two populations: one from a sunny, woodland edge and the other from a shady understory in New Jersey, USA. Plants from these seeds were grown in a greenhouse. Phytomers along primary tillers were divided into culms, leaves, and cleistogamous or chasmogamous spikelets and seeds, dried and weighed. These data were used to examine the quantitative genetics of subindividual variation among families and populations, and the relationship of tiller fitness (based on total seed mass) to the mean or subindividual variance of phytomer traits. Phytomer position along a tiller was the major determinant of organ mass. Leaf mass increased from basal to upper nodes; cleistogamous reproductive mass decreased from upper to lower nodes. Phytomer organs were heaviest in the population from the sunny habitat. Family explained < 18% of variation in organ mass. Tiller fitness was positively correlated with mean culm, and leaf mass, but negatively correlated with coefficients of variation. Field‐collected tillers showed evidence of selection for increased leaf mass. Subindividual variation in M. vimineum is mostly due to phytomer position along a tiller and the prevailing light environment. Differentiation between sunny and shady populations suggests selection favors heavier phytomer leaves and culms, especially in the shady understory where this species is most invasive.     

Water-induced stress influences the relative investment in cleistogamous and chasmogamous flowers of an invasive grass,Microstegium vimineum (Poaceae)

Background: Global climate change has the potential to shape evolutionary trajectories of invasive species via many routes, including through changes in mating systems. Many cleistogamous (CL) plants adjust investment in CL (selfed) vs. chasmogamous (CH, potentially outcrossed) progeny across environmental gradients. However, the details of such adjustments are lacking for highly invasive plant species.

Aims: We used a highly invasive grass, Microstegium vimineum, as a model for understanding how changes in water-induced stress (including potential associated changes in soil nutrient availability) might affect mating systems and thus evolutionary change in invasive species. We predicted that plants would respond to increased water-induced stress through a relative reduction in investment in CL vs. CH reproduction (i.e., a decrease in the CL:CH ratio).

Methods: Under greenhouse conditions, we measured fecundity (number of inflorescences and florets per plant) as well as relative investment in CL vs. CH florets (CL:CH ratios for number of inflorescences, florets per inflorescence, overall florets) in response to three watering treatments approximating mesic (low) to inundated (high) conditions.

Results: Plant biomass was significantly lower in high-watering treatment relative to intermediate and low treatments, indicating that the high-water condition was stressful. Contrary to expectations, stressed plants significantly increased relative investment in CL reproduction, a pattern associated with decreased inflorescence number and increased numbers of CL florets per inflorescence.

Conclusions: We conclude that changes in water-induced stress could strongly influence realised rates of outcrossing in this invasive plant, leading to mating system evolution, and altered invasiveness.     

Per capita community-level effects of an invasive grass, Microstegium vimineum, on vegetation in mesic forests in northern Mississippi (USA)

Quantifying per capita impacts of invasive species on resident communities requires integrating regression analyses with experiments under natural conditions. Using multivariate and univariate approaches, I regressed the abundance of 105 resident species of groundcover plants and tree seedlings against the abundance and height of an invasive grass, Microstegium vimineum, within 117 plots in four mesic floodplain forests in Mississippi (USA). Microstegium vimineum was most productive (i.e., tallest and most abundant) in canopy gaps in floodplains, and a significant amount of variation in resident species composition was directly explained by canopy gaps and stand age. The relatively small (but statistically significant) percentage of variation in resident species composition (1.8%) explained by M. vimineum in the multivariate analysis was attributable to significant relationships with a few common species. Most of these were negative relationships with shady mesic forest indicators. Most positive relationships were with infrequent disturbance indicators and with species with growth phenologies that differed from that of M. vimineum. Results of field competition experiments with the three most common species to show significant relationships with M. vimineum revealed asymmetric competitive effects of M. vimineum on Chasmanthium laxum and positive responses of Quercus alba seedlings and Leersia virginica adults to the removal of M. vimineum in one growing season. Results of this study suggest that negative per capita community-level effects of M. vimineum are likely to be greater in shady forests than in open floodplain forests due to the relative paucity of vulnerable species in the latter.     

BIODIVERSITY RESEARCH: Population expansion in an invasive grass,Microstegium vimineum: a test of the channelled diffusion model

Aim The greatest biodiversity impact of non‐native plant species is caused by rapid expansion of colonist populations. Unfortunately, invasion has rarely been documented in real time at a population scale, and demographic mechanisms of invasion remain unclear. Our goal is to describe real‐time expansion of populations, using channelled diffusion as a null model. Location The study examined three populations of the invasive annual grass Microstegium vimineum in mature second‐growth forests of south‐eastern Ohio and nearby West Virginia, USA. Methods Distributions were recorded in belt transects perpendicular to population edges over a period of 3 years. A second group of belt transects documented spread along five types of potential movement corridor. Observed changes in distribution were compared with predictions from a diffusion model. A seed‐sowing experiment tested seed availability, microsite quality and proximity to potential movement corridors as factors controlling population spread. Results Population boundaries showed little change over the study period. Colonization was limited by propagule availability over distances as little as 0.25 m, and to a lesser extent by litter cover. Populations did not advance along several potential movement corridors including unpaved roads, off‐road vehicle trails and footpaths. Advance was observed along deer trails and stream courses but did not conform to the wave‐form distribution predicted by diffusion theory. During the study, seeds were moved out of experimental plots by sheet flow and minor flooding events along small streams. Main conclusion At a population level, invasion is driven by processes that are episodic in time and non‐random in space – probably a common condition in non‐native plant species. Spatially realistic models are likely to be more useful than diffusive models in managing invasions at these scales.     

Relative growth rate and biomass allocation in 20 Aegilops (Poaceae) species

This paper analyses relationships between relative growth rate ( rgr ), seed mass, biomass allocation, photosynthetic rate and other plant traits as well as habitat factors (rainfall and altitude) in 20 wild species of Aegilops L. and one closely related species of Amblyopyrum (Jaub. & Spach) Eig., which differ in ploidy level (diploid, tetraploid and hexaploid). The plants were grown hydroponically for 20 d in a growth chamber. The relationships between parameters were calculated either using the phylogenetic information (phylogenetically independent contrasts, PIC) or without using the phylogenetic information (trait values of taxa, TIP). The results using the two approaches were very similar, but there were a few exceptions in which the results were different (e.g. rgr vs. seed mass). Specific leaf area ( sla ) was positively correlated with leaf area ratio ( lar ) and negatively correlated with net assimilation rate ( nar ), which together resulted in the absence of a correlation between sla and rgr . Leaf photosynthetic rates (expressed on a mass or area basis) showed no correlation with rgr . rgr was positively correlated with the stem mass ratio and negatively with root mass ratio. Species with a lower d. wt percentage have a higher rgr . Aegilops species from locations with higher annual rainfall invested less biomass in roots and more in shoots (leaves and stems) and had a higher rgr . Diploid species had a lower seed mass and initial mass than the hybrids (tetraploid and hexaploid species), but there was no correlation of rgr with ploidy level. Polyploid species, which have higher seed mass, occur at a higher altitude than diploid species. Our results show that variation in rgr in Aegilops and Amblyopyrum spp. is associated mainly with variation in biomass allocation (proportion of biomass in stems and roots) and d. wt percentage, and not with variation in sla , leaf photosynthetic rates or seed mass.     

Arbuscular mycorrhizal fungi enhance P uptake and alter plant morphology in the invasive plant Microstegium vimineum

Invasive plant species can interact with native soil microbes in ways that change how they use nutrients and allocate biomass. To examine whether Microstegium vimineum form symbiotic associations with arbuscular mycorrhizal fungi (AMF) and whether AMF mediate nutrient acquisition and growth of the plant, we conducted a field survey in Raleigh, NC and Hangzhou, China and two experiments in growth chambers. This is the first report that M. vimineum is mycorrhizal, with colonization rates of 47 and 21 % in its native and invaded range, respectively. In the growth chamber, addition of an AMF inoculum mixture significantly promoted M. vimineum biomass accumulation in both field and sterilized soils, particularly after 64 days of growth. Arbuscular mycorrhizal fungi also increased plant phosphorous (P) uptake but did not consistently affect total plant nitrogen (N) acquisition, leading to decreases in plant N:P ratios. More interestingly, AMF significantly altered plant morphology, increasing the number of stolons and aerial roots per individual (59 and 723 %), aerial roots per gram aboveground biomass (374 %) and aerial roots per stolon (404 %). Our results suggest that mycorrhizal enhancement of plant growth by stimulating tillering may serve as another mechanism by which M. vimineum can quickly take over new territory. Future studies on invasive plant-microbial interactions are needed to understand the mechanisms through which microbes contribute to the competitive ability of invasive plants.     

Nitrogen deposition and herbivory affect biomass production and allocation in an annual plant

If environmental conditions vary, plasticity in life-history traits is predicted. A recent model indicates that males and females should differ in life-history traits, because sexes differ in optimal attributes depending on species ecology. In this study we test the impact of two biotic factors in combination (presence/absence of predators and low/high food level) on gender specific life-history traits in the damselfly Coenagrion puella (Odonata). Results show that predator presence and low food density decreased activity in both sexes. Additionally, individuals with less food grew more slowly, emerged later, remained smaller and had a higher mortality. At low food densities, however, and in contrast to former investigations, individuals from treatments with predator presence were the same size or larger than individuals without predators. Gender had a strong impact on larval activity and life-history traits and sexes differed in development. Females were less active and took longer to complete development, but emerged at a larger size, weight and fat content. This study highlights the importance of gender specific approaches in life-history research.     

Plant community and phylogenetic shifts in acid seep springs over 49 years following Microstegium vimineum invasion

Plant Ecology - Community composition and diversity can change following exotic invasion. We tested the extent to which acid seep spring communities in southern Illinois had changed in composition...     

Inflorescence development in a high-altitude annual Mexican teosinte (Poaceae)

Some have postulated that highland Mexican maize was derived from an ancient high-altitude teosinte and that later introgression between the two taxa occurred. We used scanning electron microscopy to examine the inflorescence development in both the tassel and ear of a high-altitude Toluca teosinte. One of the most interesting observations was the presence of atypical multiranked orthostiches in the central spike of some male Toluca teosinte inflorescences. Most tassels exhibited a central spike with a pure, four-ranked, tetrastichous phyllotaxy or an intermediate (distichous/tetrastichous) phyllotaxy. A few A(1) tassels had a more typical distichous (two-ranked) central spike. Most ears showed the two-rank condition expected for teosintes. However, three ears displayed an intermediate (distichous/tristichous or distichous/ tetrastichous) phyllotaxy and one ear was tetrastichous. Our analysis of spikelet and floret development in all Toluca inflorescences revealed a pattern similar to that in landrace and U.S. maize, as well as to their close relatives, the teosintes. We suggest that this investigation may reveal inflorescence development in a natural maize-teosinte hybrid. This study further supports our hypothesis that both maleness and femaleness in the Zea inflorescences are derived from a common developmental pathway and underpins a proposal that andropogonoid grasses share a common pattern of inflorescence development.     

Photosynthetic responses of Microstegium vimineum (Trin.) A. Camus, a shade-tolerant, C4 grass, to variable light environments

Microstegium vimineum (Trin.) A. Camus, a shade-tolerant C₄ grass, has spread throughout the eastern United States since its introduction in 1919. This species invades disturbed understory habitats along streambanks and surrounding mesic forests, and has become a major pest in areas such as Great Smoky Mountains National Park. The focus of this study was to characterize the photosynthetic induction responses of M. vimineum, specifically its ability to utilize low light and sunflecks, two factors that may be critical to invasive abilities and survival in the understory. In addition, we were curious about the ability of a grass with the C₄ photosynthetic pathway to respond to sunflecks. Plants were grown under 25% and 50% ambient sunlight, and photosynthetic responses to both steady-state and variable light were determined. Plants grown in both 25% and 50% ambient sun became 90% light saturated between 750–850 μmol m⁻² s⁻¹; however, plants grown in 50% ambient sun had significantly higher maximum steady-state photosynthetic rates (16.09 ± 1.37 μmol m⁻² s⁻¹ vs. 12.71 ± 1.18 μmol m⁻² s⁻¹). Both groups of plants induced to 50% of the steady-state rate in 3–5 min, while it took 10–13 min to reach 90% of maximum rates, under both flashing and steady-state light. For both groups of plants, stomatal conductance during induction reached maximum rates in 6–7 min, after which rates decreased slightly. Upon return to low light, rates of induction loss and stomatal closure were very rapid in both groups of plants, but were more rapid in those grown in high light. Rapid induction and the ability to induce under flashing light may enable this species to invade and dominate mesic understory habitats, while rapid induction loss due to stomatal closure may prevent excess water loss when low light constrains photosynthesis. The C₄ pathway itself does not appear to present an insurmountable barrier to the ability of this grass species to respond to sunflecks in an understory environment. Received: 21 February 1997 / Accepted: 10 October 1997     

Comparative leaf anatomy of the annual Muhlenbergia (Poaceae)

A leaf anatomical survey was conducted to gather critical data for interpreting relationships among 32 species of annual Muhlenbergia . Phenetic analyses, utilizing 14 anatomical characters, support recognition of four major species groups. These four major species groups are not concordant with previously reported groups. Muhlenbergia brandegei , known only from three islands in Baja California Sur, is believed to be more closely related to M. appressa, M. microsperma , and M. tenuifolia , rather than, M. biloba , as suggested in prior publication. Muhlenbergia diversiglumis is morphologically similar to species of Pereilema and does not show close affinities with other annual species of Muhlenbergia . Anatomically, M. ciliata, M. pectinata , and M. tenella are very similar and form the most distinctive group among the annual Muhlenbergia .     

Taxonomic studies in some annual genera of the Triticeae (Poaceae)

The small annual genera Crithopsis, Heteranthelium, Henrardia and Amblyopyrum are revised, maps showing the distribution of the species are presented and the known chromosome numbers are confirmed. The phylogenetic relationships of each genus are discussed. Results of intergeneric hybridizations including the four genera are reported.     

Altered allocation to roots and shoots in the endophyte‐infected seedlings of Puccinellia distans (Poaceae)

Endophytes play an important role in ecological and evolutionary processes in plants and have marked economic value. Seed‐transmitted fungal endophytes are conventionally regarded as mutualistic symbionts, but their fitness consequences for the offspring of the host are not clear. Puccinellia distans infected with the fungus Epichloë typhina (E+) produces seeds that are several times smaller than normal (E?). This observation suggests that the E+ seedlings face a developmental disadvantage. Our growth chamber experiments compared the germination rates of the small E+ and large E? seeds of P. distans and examined the biomass allocation of seedlings to roots and shoots. The E+ seedlings germinated more slowly and maintained shorter shoots and a smaller root biomass for 30–50 days after sowing. Despite this disadvantage, the E+ plants more quickly increased their total size, attaining a larger shoot and whole‐plant biomass. The shoot:root biomass ratio increased more rapidly through time in the E+ seedlings, attaining a value nine times higher in the E+ than the E? group 50 days after sowing. Such differences between the E+ and E? seedlings were not explained by the growth allometry between shoots and roots. The seedlings of P. distans infected with the Epichloë endophyte were initially handicapped by their postponed emergence, but this disadvantage was quickly overcome by their superior growth capacity. The decrease in the relative allocation to roots may indicate that endophytes increase the performance of roots as resource‐acquiring organs and/or reduce the role of roots in protection against herbivores.     

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.     

Influence of nutrient availability on the mechanisms of tolerance to herbivory in an annual grass, Avena barbata (Poaceae)

Tolerance, or the capacity of a genotype to survive and reproduce following herbivore damage, varies widely across the plant kingdom. One proximate cause of this variation is resource availability, which can influence tolerance through mechanisms such as growth rate and photosynthesis. We examined the effect of high and low soil nutrient levels on the relationship between tolerance and two of its underlying mechanisms, biomass regrowth and photosynthetic upregulation, among genotypes of the Mediterranean annual grass Avena barbata. Although defoliated plants did not reach the same biomass as controls, biomass regrowth was higher at high nutrients. However, increased seed abortion at high nutrients caused tolerance to be the same in both nutrient treatments. Increased seed abortion also uncoupled biomass regrowth from tolerance at high nutrients. We found no evidence for photosynthetic upregulation in defoliated compared to control plants in either nutrient treatment. However, tolerance was positively correlated with predefoliation photosynthetic efficiency at high nutrients. Thus, constitutive photosynthetic efficiency may be a better predictor of tolerance than photosynthetic responses following herbivory in A. barbata. More generally, our results highlight the possibility that the mechanisms of tolerance can differ across resource environments even if tolerance is the same.     

Pleurovularia, a new genus of hyphomycetes proposed for a parasite on leaves of Microstegium sp. (Poaceae)

The hyphomycete Ovularia polliniae, formerly excluded from Ramularia and allied genera and tentatively considered as related to Beniowskia sphaeroidea, was recently collected on leaves of the grass Microstegium sp. in Taiwan. Based on studies with light and transmission electron microscopy, O. polliniae was redescribed. Using type specimens, the taxonomic status of both fungal species was reevaluated. In O. polliniae, the conidiophores emerge directly through the outer cell wall of the host epidermis. They are either sparsely branched at the base or simple and composed of verruculose intercalary and terminal conidiogenous cells that produce conidia through minute, slightly darkened scars. The conidia are solitary, one-celled, obovoid, hyaline, smooth, and often have a large vacuole. By the combination of these characteristics, this fungus differs from similar species of Beniowskia, Ramularia, and Ramulariopsis. The new genus Pleurovularia and the new combination Pleurovularia polliniae are proposed to accommodate this parasite on Microstegium. Received: May 24, 2001 / Accepted: September 3, 2001     

A new Pharus (Poaceae) endemic to Ecuador

The new species Pharus ecuadoricus , endemic to the Pacific lowlands of Ecuador, is described, illustrated, and compared with its closest relative, P. parvifolius.