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.     

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...     

Leaf litter and understory canopy shade limit the establishment, growth and reproduction of Microstegium vimineum

The invasive grass Microstegium vimineum is widely distributed in closed-canopy forests, but often is patchily distributed under uniform canopy conditions. We hypothesized that the occurrence of patches of invasion may be related to two interacting factors, the presence of dense understory shrub layers and the presence of thick litter layers on the forest floor. Seeds of M. vimineum were sown in plots located under or distant from Lindera benzoin (spicebush) shrubs, and with or without litter manipulations (none, half of natural amount, naturally occurring amount, double the natural amount) in a mature forest in central New Jersey, USA. Populations were monitored for germination and survivorship, as well as growth and fecundity of surviving plants. Neither shrub-associated shade nor litter depth affected seed germination, but both factors affected survivorship, growth and reproduction. The presence of shade from the shrubs reduced survivorship and seed set. Seeds germinating on top of the litter layer also experience higher mortality than seeds germinating under litter and in contact with soil. These results suggest that the interacting effects of shade from understory strata and deep litter layers may limit the spread of M. vimineum. The loss of shrub layers due to intense deer browse and other factors may thus accelerate the spread of this highly invasive species.     

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.     

Variation in species richness and species pool size across a pH gradient in forests of the southern Blue Ridge Mountains

Pärtel (Ecology 83: 2361–2366, 2002) andEwald (Folia Geobot. 38: 357–366, 2003) suggest that the relationship between local species density and soil pH is determined by regional species pool size, which in turn reflects the relative abundance of soil types during the evolutionary history of the flora. Ewald observed that calcareous sites in Central Europe have higher species density and larger species pools than acidic sites, and argues that this is the consequence of a Pleistocene bottleneck for acidophiles.The flora of the southern Blue Ridge Mountains USA has always been associated with primarily acidic soils. We used vegetation and soil data from 3328 100 m² southern Blue Ridge forest plots to examine the generality of the Pärtel-Ewald hypothesis. The Blue Ridge flora with less than 20% of species confined to sites above pH 4.7 contrasts dramatically with that of Europe. However, regional species pool size increases with pH. Genus- and family-level pools increase with pH, suggesting an ancient origin for this pattern. Mean species density is also strongly positively correlated with soil pH. Thus, both regional species pool size and plot species density of southern Blue Ridge forests fail to conform to predictions derived from the work of Pärtel and Ewald.The increase in species pool with increase in pH exhibited by southern Blue Ridge forests appears to reflect broad species distributions and tolerance for high pH conditions among species that grow predominantly under acid conditions. We conclude that richness on higher pH sites is a consequence of generally more favorable conditions for plant growth and/or establishment. Ewald may be correct in asserting that the abundance of high pH specialists in Central Europe is an historical artifact, but our data contradict the assertion of Pärtel and Ewald that evolution of a flora in a primarily low pH environment will necessarily translate into a negative correlation between pH and species density in local vegetation.     

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.     

秦岭山地天然油松群落主要植物种群生态位特征

应用Levins、Hurlbert生态位宽度及Pianka生态位重叠研究方法,对秦岭山地天然油松群落内20种主要乔木及26种主要灌木的种群生态位特征进行了分析。结果表明:(1)油松、锐齿槲栎、华山松、漆树为乔木层的优势种群,而榛子、卫矛、忍冬、荚蒾为灌木层的优势种群。(2)乔木层中上述优势种群具有较大的生态位宽度,构成了该群落的主体;灌木层中荚蒾、胡颓子、卫矛、榛子等种群的生态位宽度较大。(3)乔木层主要种群间生态位重叠相对较少,其中冬瓜杨-尖叶四照花、盐肤木-山杨、小叶杨-白桦、小叶杨-少脉椴等种对生态位重叠值较高;灌木层主要种群间生态位重叠较多,其中卫矛-陕西卫矛、绣线菊-忍冬、粉背黄栌-黑刺菝葜、高山杜鹃-中华柳等种对生态位重叠较高。(4)群落内优势种群生态位宽度较大,与其他种群间的生态位重叠值相对较高,但生态位重叠与生态位宽度之间没有绝对的正相关关系。     

Phytomass structure of natural plant communities on spodosols in southern Venezuela: the Bana woodland

Bana, or Low Amazon Caatinga is an evergreen sclerophyllous woodland. It occurs on bleached quartz sands in the lowlands of SW Venezuela, where it occupies relatively small ‘islands’ amidst Tall Amazon Caatinga which is exclusively developed on tropaquods. There is an outer vegetation belt about 20 m in width in which trees over 10 m in height occur (Tall Bana); its structure and floristic composition resemble Tall Amazon Caatinga. Low Bana (maximum tree height usually below 5 m) follows next. The central part is occupied by Open Bana in which even lower trees are very widely spaced. Destructive phytomass sampling was carried out for chemical analyses in seven plots along a 150 m line across the zonation. The total dry matter of living plants including roots of Tall Bana (30–32 kg/m²) compares rather well with 41 kg/m² in Tall Amazon Caatinga. This is only 9–14 kg/m² in Low Bana, and 4–6 kg/m² in Open Bana. The average root % of total phytomass increases from 41% in Tall Bana to 63% in Low Bana, and is 88% in Open Bana. Average total dry dead above-ground phytomass (including standing trees and stumps) declines from 1 kg/m² in Tall Bana to 0.2 kg/m² in Open Bana. An accumulation of dead matter in Low and Open Bana, relative to the above-ground phytomass of living plants, is noted and this contrasts with the general absence of raw humus in the soil. Eighty-two species of woody plants (dbh≥1 cm) were recorded on the total plot area (640 m²); 90% of the species are also known to occur in Tall Amazon Caatinga. The species number declines from 59 in Tall Bana to 18 in Open Bana. Mesophylls sensu strictu dominate in Tall Bana, while notophylls are dominant in Low and Open Bana. Herbaceous species are less numerous: most of them belong to the Araceae, Bromeliaceae, Orchidaceae, Droseraceae, Eriocaulaceae and Xyridaceae.     

Regeneration dynamics of Tsuga canadensis in the southern Appalachian Mountains, USA

Tsuga canadensis is often considered a foundation species because of its influence on the biological and ecological processes of upland forest systems. With hemlock woolly adelgid (Adelges tsugae; HWA)-induced mortality of T. canadensis in southern Appalachian upland forests, there is a crucial need to understand the regeneration dynamics of this tree species. This research was conducted within mature T. canadensis forests of Great Smoky Mountains National Park (GSMNP) prior to widespread HWA infestation. Tsuga canadensis regeneration abundance is lower than has been reported in more northern locales, but varies widely among sites in the southern Appalachian Mountains. Ordination results suggest landscape-scale patterns of regeneration abundance that are influenced by gradients in understory Rhododendron maximum cover, soil sand and nutrient content. Regression models reinforce the observed gradients in regeneration abundance. Sites with abundant T. canadensis regeneration have less R. maximum cover, soil sand and litter depth, but more soil nutrient availability. Tsuga canadensis seedlings and saplings tend to be spatially aggregated at the 14 × 21 m plot scale. Sites with spatially aggregated seedlings and saplings have greater canopy openness and more species with gap affinity. This research elucidates associations between successful T. canadensis regeneration and site factors such as soil nutrient availability and canopy dynamics. More forest surveys are required to understand the conditions necessary for successful T. canadensis regeneration. Land managers involved in forest restoration efforts may have to utilize appropriate silvicultural techniques conjoined with R. maximum removal operations to regenerate T. canadensis in southern Appalachian upland forests.     

Dynamics of plant communities in the Lomas of southern Peru

A study of the floristic composition of the plant communities of the Lomas was carried out in the Coastal Range of Arequipa, Peru. Data on the seasonal and elevational variation of those communities was obtained by monthly censusing a 20-meter circle sampling stations established at 100 m intervals from 300 to 1 000 m of elevation. The 80 recorded plants belong to 60 genera and to 20 families. Dominating life forms are Therophytes and Chamaephytes. Four vegetational zones (xeric, subxeric, mesic, and submesic), are separated by significant differences in the number of plants of each zone. Considering the entire Lomas transect, no significant statistical differences among the seasons were found. The floristic relationship among the altitudinal samples was found to be a reflection of a moisture pattern. The persistence in the Lomas of a mesophytic flora could be the result of a continuous adaptation to the fogginess and reduced solar radiation of this particular zone in a desert area.This study was supported by the Universidad Nacional de San Agustin, Arequipa, Perú, and by the Universidad de Los Andes through the Consejo de Desarrollo Cientifico y Humanistico (Proycet C-142-79). Amelia Díaz de Paseual, William E. Duellman, Evaristo López, Luis Malaret, Mario Ricardi and Antonio Vivas made helpful comments and criticisms. Field trips were shared with José Dávila, Evaristo López and Alberto Nuñez. The identification of plants was checked by Ramón Ferreyra and Oscar Tovar of the Museo Javier Prado, Lima, Perú, and Alwing Gentry of the Missouri Botanical Garden. I am grateful to all these institutions and people.     

Impact on plant communities by white-tailed deer in Mississippi,USA

Background: The abundance of white-tailed deer (Odocoileus virginianus) in the eastern United States has escalated during the twentieth century, potentially impacting plant communities.

Methods: We measured understorey plant cover and biomass five years after excluding deer from mature forests of three ecological regions in Mississippi, USA. We extended the significance of P values to 0.10 to detect developing impacts.

Results: Deer impacts were limited and varied by ecological region. We recorded 151 species in cover transects. Consistent exclosure treatment effects were detected in two regions where there was greater cover of two deer forages and less cover of three non-forages. Species richness was greater in exclosures in one region, but otherwise species richness and diversity indices did not differ. We recorded 127 species in biomass quadrats. Exclosure treatment effects on biomass were inconsistent. Out of five species with significant differences, three had more biomass in controls, including two deer forages. Except for greater total biomass in controls of one region, there were no differences by growth form or total vegetation for canopy coverage or biomass. Ordination of community canopy cover demonstrated similarity of paired exclosure and controls.

Conclusions: Exclosure treatment effects on canopy cover and species richness in two regions indicated limited negative impacts from deer foraging. A time frame of more than five years may be required for exclusion to allow recovery of vegetation, even with relatively open canopies and a long growing season.     

Experimental manipulation of natural plant communities

Attempts to elucidate the factors controlling the structure of plant communities have relied increasingly on field experiments. This is a powerful approach for testing theoretical predictions that offers important advantages over observational and comparative studies. However, field experiments suffer from intrinsic difficulties as well as more-easily remediable limitations. Recent progress has been made by new approaches including the use of multifactor experiments, and the development and dissemination of better statistical tools.     

Large herbivores influence the composition and diversity of shrub-steppe communities in the Rocky Mountains, USA

It is widely believed that wild and domestic herbivores have modified the structure and composition of arid and semi-arid plant communities of western North America, but these beliefs have rarely been tested in long-term, well-replicated studies. We examined the effects of removing large herbivores from semi-arid shrublands for 40–50 years using 17 fenced exclosures in western Colorado, USA. Shrub cover was greater (F=5.87, P=0.0020) and cover (F=3.01, P=0.0601) and frequency (F=3.89, P=0.0211) of forbs was less inside the exclosures (protected) relative to grazed plots. However, we found no significant effects (minimum P=0.18) of protection from grazing on cover or frequency of grasses, biotic crusts, or bare soil. Although mean species richness and diversity were similar between treatments, protected areas had much higher dominance by fewer species, primarily sagebrush. Exclusion of herbivores changed the relationship between species richness and evenness. Consistent with theoretical expectations, species evenness was positively correlated with richness in protected plots (r ²=0.54). However, contrary to theory, evenness and richness were inversely related in grazed plots (r ² _adjacent=0.72, r ² _distant=0.84). We suggest that these differences resulted because grazing acts as a stressor promoting facilitative relationships between plant species that might compete for resources in the absence of grazing. We conclude that exclusion of grazing in the sites we studied caused minor changes in cover and diversity of herbaceous plants, but caused a clear increase in the cover of shrubs. Importantly, the exclusion of ungulates changed the relationship between evenness and richness.     

Diversity and productivity of plant communities across the Inland Northwest,USA

No definitive explanation for the form of the relationship between species diversity and ecosystem productivity exists nor is there agreement on the mechanisms linking diversity and productivity across scales. Here, we examine changes in the form of the diversity–productivity relationship within and across the plant communities at three observational scales: plots, alliances, and physiognomic vegetation types (PVTs). Vascular plant richness data are from 4,760 20 m² vegetation field plots. Productivity estimates in grams carbon per square meter are from annual net primary productivity (ANPP) models. Analyses with generalized linear models confirm scale dependence in the species diversity–productivity relationship. At the plot focus, the observed diversity–productivity relationship was weak. When plot data were aggregated to a focus of vegetation alliances, a hump-shaped relationship was observed. Species turnover among plots cannot explain the observed hump-shaped relationship at the alliance focus because we used mean plot richness across plots as our index of species richness for alliances and PVTs. The sorting of alliances along the productivity gradient appears to follow regional patterns of moisture availability, with alliances that occupy dry environments occurring within the increasing phase of the hump-shaped pattern, alliances that occupy mesic to hydric environments occurring near the top or in the decreasing phase of the curve, and alliances that occupy the wettest environments having the fewest species and the highest ANPP. This pattern is consistent with the intermediate productivity theory but appears to be inconsistent with the predictions of water–energy theory.     

Phytomass structure of natural plant communities on spodosols in southern Venezuela: The tall Amazon Caatinga forest

The phytomass structure of the evergreen lowland forest vegetation (Tall Amazon Caatinga) supported by tropaquods near San Carlos de Rio Negro, Federal Amazon Territory of Venezuela was studied in 13 10 m× 10 m plots. The plots were laid out subjectively to cover a low topographical gradient along which the forest on tropaquod is grading into a low woodland (Bana, or Low Amazon Caatinga). The phytomass was estimated by destructive sampling. The total living phytomass (dry matter) varies between 199 t/ha in one plot including a natural gap, to 822 t/ha in a plot located near a blackwater creek draining the 10 ha study site in which the vegetation was surveyed. Dead aboveground phytomass (dry matter) varies between 2 and 37 t/ha. The average leaf area index is 5.1. Leaves are sclerophyllous. The mesophyll leaf size class is dominant. Compared with other Amazonian rainforests the Tall Amazon Caatinga is lower in aboveground phytomass and wood volume, respectively, but its root average proportion is 2.4 times greater. One hundred and thirty species (dbh≥1 cm) were recorded in all plots. 14–45 species per plot composed the aboveground phytomass. 20 species each had a relative frequency of 50 or more percent. 7 of these species plus 10 less frequent ones each represented 10 or more percent of the basal area in at least 1 plot. Sixteen species each contributed 10 or more percent to the aboveground phytomass, in at least 1 plot. There is only 1 species (Micranda sprucei, Euphorbiaceae) which was observed in all plots. Its contribution to basal area and aboveground phytomass is considerable. M. sprucei is also the dominant species of the top canopy layer. Eperua leucantha, Caesalpiniaceae, is subdominant.     

Seasonal occupancy of sympatric larger carnivores in the southern San Andres Mountains,south-central New Mexico,USA

We investigated factors facilitating coexistence of pumas (Puma concolor), coyotes (Canis latrans), and bobcats (Lynx rufus) in the arid San Andres Mountains of south-central New Mexico, during the season (winter and spring, prior to the annual monsoon) of greatest resource stress. We established a camera-trapping grid in the San Andres, 2007–2011, and modeled occupancy of the three carnivores as a function of habitat, prey, and presence of the other carnivore species. Species interaction factors were >1.3 for each pair of carnivores, and the presence of the other carnivore species never significantly influenced occupancy of any other carnivore. Similarly, occupancy of the San Andres landscape was positively correlated among all carnivores. Occupancy of pumas was most influenced by proximity of water; coyote occupancy was influenced by terrain ruggedness and presence of medium (primarily lagomorph) prey, and bobcat occupancy was influence primarily small prey and proximity to water. The three carnivores also did not show temporal partitioning in use of habitats. Rather than segregation driven by competition, predation, or despotism, our results appeared to reflect preferences for differing habitat characteristics between ambush and cursorial predators and preferred habitats for travel.     

Structural characteristics and canopy dynamics of Tsuga canadensis in forests of the southern Appalachian Mountains, USA

Tsuga canadensis (L.) Carr. forests of the southern Appalachian Mountains are currently facing imminent decline induced by a nonnative insect pest, the hemlock woolly adelgid (Adelges tsugae Annand). To effectively manage these forest systems now and in the future, land managers need baseline data on forest structure and dynamics prior to large-scale Tsuga canadensis mortality. Most of our knowledge concerning the dynamics of Tsuga canadensis forests comes from more northern locations such as the Great Lakes region and New England and, therefore, may not pertain to the ecological systems found within the southern Appalachian Mountains. We examined the structure and canopy dynamics of four Tsuga canadensis forest stands within the Cataloochee watershed, in the far eastern part of Great Smoky Mountains National Park (GSMNP). We characterized the environmental settings and vertical forest layers, as well as the diameter and age-structures of each Tsuga canadensis forest stand. These environmental and structural data showed that there were indeed differences between forest stands with and without successful Tsuga canadensis regeneration. The two forest stands exhibiting successful Tsuga canadensis regeneration were located above 1,000 m in elevation on well-drained, moderately steep slopes and had the greatest canopy openness. Structural data from these two forest stands indicated a history of more continuous Tsuga canadensis regeneration. We also constructed disturbance chronologies detailing the history of canopy response to disturbance events and related these to Tsuga canadensis regeneration within each forest stand. Student t-tests adjusted for unequal variances indicated significant differences in the number of release events per tree between forest stands with and without successful Tsuga canadensis regeneration. While forest stands with successful Tsuga canadensis regeneration were more frequently disturbed by minor to major canopy disturbances, events of moderate intensity were found to be most significant in terms of regeneration. These data will be of value to land managers maintaining stands of Tsuga canadensis where treatment for hemlock woolly adelgid infestation has been successful. In areas where treatment is impractical or unsuccessful, land managers will be able to use these data to restore Tsuga canadensis forests after the wave of hemlock woolly adelgid induced mortality has passed. As of August 2008, Joshua A. Kincaid will be a member of the Environmental Studies program at Shenandoah University in Winchester, Virginia, USA