Performance of two Picea abies (L.) Karst. stands at different stages of decline

Summary The annual replacement of tillers of Agropyron desertorum (Fisch. ex Link) Schult., a grazing-tolerant, Eurasian tussock grass, was examined in the field following cattle grazing. Heavy grazing before internode (culm) elongation seldom affected tiller replacement. Heavy grazing during or after internode elongation, which elevates apical meristems, increased overwinter mortality of fall-produced tillers and reduced the number and heights of these replacement tillers. Unexpectedly, tussocks grazed twice within the spring growing season tended to have lower overwinter tiller mortality, greater tiller replacement, and larger replacement tillers than tussocks grazed only once in late spring. These responses of twice-grazed tussocks, however, were still less than those of ungrazed tussocks or tussocks grazed moderately in early spring. The presence of ungrazed tillers on partially grazed tussoks did not increase the replacement of associated grazed tillers relative to tillers on uniformly grazed plants. This result indicates that resource sharing among tillers, if present, is short-lived or ecologically unimportant in this species. Although A. desertorum is considered grazing-tolerant, tiller replacement on heavily grazed tussocks, particularly those grazed during or after internode elongation when apical meristems were removed, was usually inadequate for tussock maintenance. These observations at the tiller (ramet) level of organization in individual tussocks (genet) may explain the often noted reduction in stand (population) longevity with consistent heavy grazing.     

Spatial arrangement of tiller replacement in Agropyron desertorum following grazing

Summary The spatial arrangement of tiller replacement was assessed on grazed and ungrazed tussocks of Agropyron desertorum (Fisch. ex Link) Schult. for three annual cycles. Frequency distributions of the number of replacement tillers per single progenitor were also determined. Tiller replacement was usually greater on the perimeter of tussocks than within the core, with or without grazing. Replacement was inversely related to grazing intensity, both on the perimeter and within the core of tussocks. Heights of replacement tillers on the perimeter or within the core seldom differed. Furthermore, grazing seldom affected the number of replacement tillers per progenitor. Greater tillering on the perimeter than within the core indicates that the tussocks were expanding. Apparently, grazing neither enhances tussock expansion and subsequent disintegration, nor does it necessarily lead to patches of tillers (multiple tillering per progenitor) within tussocks of A. desertorum.     

Effect of competition on stable carbon isotope ratios of two tussock grass species

Summary Previous studies have shown that plant carbon isotope composition varies when plants experience differences in water and nutrient availability. However, none have addressed the effect of root interactions, including competition for these soil resources, on carbon isotope ratios. We studied the effect of interspecific root interactions on the productivity and carbon isotope ratios of two Great Basin tussock grass species (Agropyron desertorum and Pseudoroegneria spicata). We compared grasses grown in mixture with sagebrush (Artemisia tridentara) to grasses in similar mixtures but where root interactions with sagebrush were limited by fiberglass partitions. During both years of the study, tussocks growing in competition with sagebrush produced tissue with more negative ¹³C values than grasses experiencing limited root interaction with sagebrush. The magnitude of this difference (0.5 to 0.9%) is similar to that found in other studies when soil fertility and moisture availability were altered.     

Characteristics of successful competitors: an evaluation of potential growth rate in two cold desert tussock grasses

Summary Within the first few weeks after seedling emergence, Agropyron desertorum, a more competitive tussock grass, had a much higher mean relative growth rate (RGR) than Agropyron spicatum, a very similar, but less competitive species. However, beyond the early seedling stage, the two grasses had a remarkably similar whole-plant RGR in hydroponic culture and aboveground RGR in glasshouse soil, if root temperatures were above approximately 12°C. At soil temperatures between 5 and 12°C, A. desertorum exhibited a 66% greater aboveground RGR than A. spicatum (P<0.05). Both species responded similarly to warming soil temperatures. In the field, however, tiller growth rates were generally similar. Neither species showed marked tiller elongation until a couple of weeks after snowmelt, by which time soil temperatures, at least to a depth of 10 cm, were above 12°C for a significant portion of the day. Aboveground biomass accumulation over a three-year period indicated that both grasses had similar potential growth rates whereas Artemisia tridentata ssp. vaseyana, a common neighbor planted in the same plots, had a much greater potential growth rate. The greater competitive ability of adult A. desertorum, as compared to A. spicatum, cannot be attributed to appreciable differences in potential growth rates.     

Growth and carbon allocation of Agropyron desertorum following autumn defoliation

Summary Growth and carbon allocation of a cool season tussock grass, Agropyron desertorum, following defoliation of newly initiated tillers in the autumn of 1988 and 1989 were investigated. Tiller density and mortality, reproductive shoot density, root density, biomass, individual tiller weight, carbon allocation, and soil water depletion were used to evaluate the response of A. desertorum to autumn grazing. Tiller recruitment was lower in the autumn-defoliated treatment in both years compared with the control because of the cessation of tiller development following autumn defoliation. Autumn defoliation also significantly reduced the movement of ¹³C to the roots in 1988 but not in 1989. Soils were cooler and drier in 1989. Other plant growth measurements and soil water depletion rates were not different between treatments. Autumn defoliation in 1988 did not influence tiller recruitment in the following autumn. Two consecutive years of autumn defoliation did not affect tiller overwinter mortality or peak standing crop in 1990.     

Morphological Analysis of Tillering in Agropyron spicatum and Agropyron desertorum

The caespitose grasses Agropyron spicatum and Agropyron desertorumexhibit a striking difference in tillering response followingexperimental clipping treatment, with plants of A. desertorumproducing up to 18 times more tillers. The two species are similarin many aspects of their phenology and physiology. Previousexamination of current photosynthate production and levels ofstored carbohydrates indicate only slight differences betweenthe species. The possible role of three anatomical/morphologicalconstraints in controlling tillering was examined. No evidencefor such constraints was found. A basal cluster of buds is presenton the parent tillers. The mean bud number per tiller was similarfor both species and the range (3–9) was identical. Nearlyall of the bud apical meristems appeared anatomically viablethroughout the growing season and vascular development occurredto within 250 to 490 µm of the various bud apices of bothspecies. Both normal fall tillers and summer tillers producedunder clipping treatment originated from the largest, most distalbuds of the basal cluster of buds. However, precocious, morphologicallydistinctive, second-order tillers occasionally grew out fromthe smaller, most basal buds of some elongating fall tillers. Agropyron spicatum, Agropyron desertorum, bluebunch wheatgrass, crested wheatgrass, bud, tiller, tillering ability, meristematic potential, vascular development, regrowth     

Comparative demography of co-occurring introduced and native tussock grasses: persistence and potential expansion

Summary Demographic characteristics associated with the maintenance and growth of populations, such as seed dynamics, seedling emergence, survival, and tiller dynamics were examined for two tussock grasses, the native Agropyron spicatum and the introduced Agropyron desertorum in a 30-month field study. The introduced grass was aerially sown onto a native grassland site. Seed production of the introduced grass was greater than the native grass in both above- and below-average precipitation years. Seeds of A. spicatum were dispersed when they mature, while A. desertorum retained some seeds in inflorescences, and dispersed them slowly throughout the year. This seed retention allowed some seeds of the introduced grass to escape peak periods of seed predation during the summer and allowed seeds to be deposited constantly into the seed bank. Carryover of seeds in the seed bank beyond one year occurred in the introduced grass but not in the native species. For both species, seedling emergence occurred in both autumn or spring. Survival rates for A. desertorum were higher than A. spicatum when seedlings emerged between November and March. Survival rates of cohorts emerging before November favored A. spicatum whereas survival rates did not differ between species for cohorts emerging after March. Individuals of both species emerging after April were unable to survive the summer drought. Demographic factors associated with seeds of A. desertorum seemed to favor the maintenance and spread of this introduced grass into native stands formerly dominated by A. spicatum.     

Growth and tillering patterns within tussocks of Eriophorum vaginatum

Growth patterns were investigated for Eriophorum vaginatum tussocks from disturbed and undisturbed tussock tundra at two sites in Alaska. Total basal area of tussocks decreased with increased cryoturbation but mean basal area per tussock did not. Flowering was observed in tussocks of significantly smaller size on disturbed compared to undisturbed tundra. For tussocks with < 10% cover by shrubs and moss, number of tillers per tussock was linearly related to tussock diameter in most disturbed and undisturbed sites. Exceptions occurred in an area that had been bladed with a bulldozer 7 yr before our survey where tiller number increased as the square of diameter and in an area with much frost activity where tiller number was not emulated with diameter. The ratio of daughter tillers to adult tillers decreased with diameter in disturbed tundra, whereas the trend was less pronounced in undisturbed tundra.
Microsuccession in undisturbed tussock tundra was investigated by sampling tussocks with different amounts of shrub and moss cover for number of daughter tillers per adult tiller, weight per tiller, percent nitrogen, and percent phosphorus. Small tussocks without cover by other species and large, partially covered tussocks were not significantly different by any measures, but tussocks that were almost completely buried had significantly (p < 0.05) lower values of tillering index, weight per tiller, and percent phosphorus.     

Potential and constraints for grasses to cope with spatially heterogeneous radiation environments

Placing plant organs into upper canopy layers or gaps is considered advantageous for avoiding neighbours. Current research only covers the lack of selective branching in response to heterogeneous radiation in dicots. Due to some unique shade avoidance regulatory pathways and clonal structure, directional response may occur in grasses. I used Festuca rubra L., a grass with two types of branches: intravaginal tillers developed from young buds close to the primary shoot, and extravaginal tillers developed from older buds and placed on rhizomes in a certain distance from the mother tussocks. Tussocks grown from vegetativelly multiplied initial tillers were exposed to three radiation regimes in two green-house experiments: full sunlight, full shading simulating canopy and heterogeneous radiation. Interaction of the treatments with orientation of the initial tillers relative to their mother tillers, which may constrain morphology of the developing tussocks, were studied. Even though shading decreased the number of intravaginal tillers, heterogeneous radiation did not influence direction of their outgrowth. However, shading activated dormant buds at the tussock base and thus stimulated extravaginal tillering, with a tendency to develop more extravaginal tillers towards the shaded sites.     

Morphological Constraints of Shoot Demography of a Clonal Plant: Extra- and Intravaginal Tillers of Festuca rubra

Abstract Festuca rubra forms tillers in two different ways: extravaginally and intravaginally. Demography of these two tiller types was observed in seventeen selected tussocks of Festuca rubra s.s. over four growing seasons. Extravaginal tillers were bigger at birth and on the average produced twice as many daughter tillers per tiller. In general, the natality and mortality of extravaginal tillers were less regular than that of intravaginal tillers. Overall tillering rate per tiller was correlated with the density of the surrounding vegetation; mortality, natality and tiller life span were not. High density of the surrounding vegetation did not result in increased formation of extravaginal tillers. The proportion of the extravaginal tillers was not correlated with the density of the F. rubra tussocks. There is no evidence for foraging by extravaginal tillers, but they do act as founders of small clusters of tillers.     

Negative Plant–Soil Feedback and Positive Species Interaction in a Herbaceous Plant Community

Increasing evidence shows that facilitative interaction and negative plant–soil feedback are driving factors of plant population dynamics and community processes. We studied the intensity and the relative impact of negative feedback on clonal growth and seed germination of Scirpus holoschoenus, a ‘ring’ forming sedge dominant in grazed grassland, and the consequences for species coexistence. The structure of aboveground tussocks was described. A Lithium tracer assessed belowground distribution of functional roots. Seed rain and seedling emergence were compared for different positions in relation to Scirpus tussocks. Soil bioassays were used to compare growth on soil taken from inside and outside Scirpus tussocks of four coexisting species (Mentha acquatica, Pulicaria dysenterica, Scirpus holoschoenus and Dittrichia viscosa). We also compared plant performance of dominant plant species inside and outside Scirpus tussocks in the field. The ‘ring’ shaped tussocks of S. holoschoenus were generated by centrifugal rhizome development. Roots were functional and abundant under the tillers and extending outside the tussocks. The large roots mats that were present in the inner tussock zone were almost all dead. Seedling emergence and growth both showed a strong negative feedback of Scirpus in the inner tussock zone. Scirpus clonal development strongly reduced grass biomass. In the degenerated tussock zone, Pulicaria and Mentha mortality was lower, and biomass of individual plants and seed production were higher. This positive indirect interaction could be related to species-specific affinity to soil conditions generated by Scirpus, and interspecific competitive release in the degenerated tussock zone. We conclude that Scirpus negative feedback affects its seedling emergence and growth contributing to the development of the degenerated inner tussock zone. Moreover, this enhances species coexistence through facilitative interaction because the colonization of the inner tussock zone is highly species-specific.     

Rapid recovery of kohekohe (

Soil conditions, vegetation features and soil fauna were recorded in montane tall tussock grassland dominated by narrow- leaved snow tussock Chionochloa rigida ssp. rigida up to 30 months after a spring fire. Burning reduced the stature of tussocks and the size and density of tillers in the first growing season. After two growing seasons, tussock canopy development and tiller size remained below those found in the unburnt grassland nearby. New tillers and tussocks established following the prolific fire-induced flowering one year after burning. After the fire and sheep grazing, intertussock cover became progressively dominated by introduced grasses and herbs. While soil pH, moisture content, bulk density, surface litter and total nematodes showed significant treatment (burning) effects, these properties also showed significant year-to-year variation. The greatest increase in any nematode group was in Paratylenchus, a distinctive genus widespread in tussock grasslands and apparently responsive to environmental fluctuation and root development; its population was 100x and 29x greater in the burned area than in the control area 16 and 30 months after burning. Subject to detailed testing, populations of mites and collembola may provide relatively simple indicators of recovery of ecosystem function of such grasslands after burning.     

Avian predation at a Southern Rockhopper Penguin colony on Staten Island,Argentina

We studied predation risk in relation to nest location and subcolony size in Southern Rockhopper Penguins (Eudyptes chrysocome chrysocome) during the chick-rearing period. Striated Caracaras (Phalcoboenus australis), the main predator, preferentially attacked from tussock grasses which are found in the periphery of all subcolonies (peripheral tussocks) and often scattered within them (central tussocks). The greatest numbers of predation and attempted predation events were observed on nests in the periphery of the subcolony next to peripheral tussocks, and on those nests next to central tussocks. Central tussocks offer Striated Caracaras an additional “edge” area from which to prey, much in the same way as do the peripheral tussocks. Predation rate per individual was not correlated with subcolony size possibly due to the presence of central tussocks which, by creating an extra edge area, change the subcolony shape. There is a suggestion (P = 0.06) of increased probability of nest success with subcolony size.     

The effects of the spatial pattern of defoliation on regrowth of a tussock grass

Summary The spatial pattern of foliage removal from a tussock grass can influence regrowth through effects on daily carbon gain (CER_d). This field study examined the extent to which tussock photosynthetic responses to different defoliation patterns were associated with changes in whole-canopy attributes (e.g., foliage age structure, canopy light microclimate). During the spring growing season, 60% of the green foliage area was removed from individual Agropyron desertorum tussocks with scissors in different spatial patterns. These patterns represented extremes of defoliation patterns that might be inflicted by natural herbivores. Tussock photosynthesis (per unit foliage area) at high light (2000 mol photons m^–2 s^–1 between 400 and 700 nm; P₂₀₀₀) increased following clipping with all defoliation patterns. The increases in P₂₀₀₀ were greater when leaves were removed from low in the tussock (older leaves) than if leaves high in the canopy (younger leaves) were removed. These relative changes of P₂₀₀₀ among clipping patterns paralleled the responses of CER_d and regrowth from an earlier study. Furthermore, the changes in P₂₀₀₀ corresponded with increases in the proportion of foliage within the tussocks that was directly illuminated at midday. The greater photosynthesis of tussocks after lower-leaf removal was directly related to a higher proportion of younger foliage and a smaller fraction of foliage shaded within the tussock. In a dense canopy, such as these grass tussocks, the influence of defoliation on whole-canopy attributes may be of primary importance to whole-plant photosynthetic responses.     

Interactions between tussock grass (Poa spp.) and Eucalyptus pauciflora seedlings near treeline in South-Eastern Australia

Summary Observations of tussock grasses (mainly Poa species) and tree seedlings (Eucalyptus pauciflora) growing near treeline suggested that the tussocks may physically protect and compete with the tree seedlings. An analysis of nearest neighbour data from sites burnt in 1972–3 showed the there is a minimum separation between seedlings and tussocks, indicating that competition is taking place. Correlation and principal components analyses show that seedlings growing close to tussocks tend to be taller, with fewer stems and leaves than those growing further away. This same trend from tall, few-stemmed individuals to shorter multistemmed individuals also occurs with an increase in altitude. It is concluded that the trend in habit is related to exposure to environmental extremes. The concept of the regeneration niche and its application to E. pauciflora seedlings is discussed.     

The timing and degree of root proliferation in fertile-soil microsites for three cold-desert perennials

Summary Root proliferation in nutrient-rich soil patches is an important mechanism facilitating nutrient capture by plants. Although the phenomenon of root proliferation is well documented, the specific timing of this proliferation has not been investigated. We studied the timing and degree of root proliferation for three perennial species common to the Great Basin region of North America: a shrub, Artemisia tridentata, a native tussock grass, Agropyron spicatum, and an introduced tussock grass, Agropyron desertorum. One day after we applied nutrient solution to small soil patches, the mean relative growth rate of Agropyron desertorum roots in these soil patches was two to four times greater than for roots of the same plants in soil patches reated with distilled water. Most of the increased root growth came from thin, laterally branching roots within the patches. This rapid and striking root proliferation by Agropyron desertorum occurred in response to N-P-K enrichment as well as to P or N enrichment alone. A less competitive bunchgrass, Agrophyron spicatum, showed no tendency to proliferate roots in enriched soil patches during these two-week experiments. The shrub Artemisia tridentata proliferated roots within one day of initial solution injection in the N-enrichment experiment, but root proliferation of this species was more gradual and less consistent in the N-P-K and P-enrichment experiments, respectively. The ability of Agropyron desertorum to proliferate roots rapidly may partly explain both its general competitive success and its superior ability to exploit soil nutrients compared to Agropyron spicatum in Great Basin rangelands of North America.     

Establishing Native Grasses in a Big Sagebrush–Dominated Site: An Intermediate Restoration Step

Many semiarid rangelands in the Great Basin, U.S.A., are shifting dominance to woody species as a consequence of land degradation including intense livestock grazing and fire suppression. Whereas past rehabilitation efforts in Big sagebrush (Artemisia tridentata) steppes removed the shrub and added introduced forage grasses to successfully shift communities from shrublands to grasslands, current consensus is that native species should be included in restoration projects and that retention of some woody plants is desirable. We examined the potential for interseeding grasses into dense shrub communities as a precursor to thinning shrubs and releasing grasses from shrub interference. We compared seedling establishment of the native grass, Bluebunch wheatgrass (Pseudoroegneria spicata), with that of the Eurasia grass, Crested wheatgrass (Agropyron desertorum), in dense Ar. tridentata stands. Shrubs may play an important role as nurse plants for seedling establishment (reduced solar radiation, “island of fertility” effect) but result in highly contrasting light environments and root interference for seedlings. In experimental plots, we examined effects of Ar. tridentata shade levels (0, 40, 70, and 90% reduction of solar radiation) and initial root exclusion (present/absent) on the establishment and growth of P. spicata and Ag. desertorum seedlings. With this design we evaluated the interference effects of Ar. tridentata on the two grasses and identified the most beneficial microsites for grass restoration in Ar. tridentata–dominated communities. We predicted seedling survival and growth to be greater under moderate shade (40% reduction) and limited root competition than under no or strong shade conditions (0 and 90%) and unrestricted root interactions. Fifty to 85% of the P. spicata and Ag. desertorum seedlings survived the dry summer months of 1995 and 1996 and the intervening winter. Neither shading nor root exclusion from Ar. tridentata affected final seedling survival of either species. Seedling biomass of both grass species was negatively affected by initial root interactions with Ar. tridentata. However, the analysis of seedling biomass variability (coefficient of variation) indicated that in all shade and root‐exclusion treatments, some seedlings of both species developed to large individuals to survive in Ar. tridentata–dominated rangelands. Thus, the use of interseeding techniques shows promise for restoring herbaceous species in dense Ar. tridentata stands and should be given further consideration when shrub retention is an important consideration.     

Post‐fire shrub recruitment in a semi‐arid grassland: the role of microsites

Question: (1) Which factors regulate post‐fire recruitment and spread of the shrub Senecio bracteolatus in Patagonian grasslands? (2) What is the role of the grass Stipa speciosa on S. bracteolatus establishment in the post‐fire succession? Location: Northwest Patagonia, Argentina. Methods: We studied the effect of fire on S. bracteolatus recruitment and density by comparing these variables between burned and unburned grasslands. In burned areas, we compared abiotic characteristics and seedling establishment under the canopy of grasses (S. speciosa) and in gaps (inter‐tussock areas). Post‐fire interactions between S. bracteolatus seedlings and S. speciosa were studied using field and greenhouse experiments. Results: Density of S. bracteolatus was higher in burned than in unburned areas. In burned sites, seedlings were more abundant under tussock grasses, whereas juveniles were more abundant in gaps. Tussocks generated more attenuated micro‐environmental conditions than gaps during stressful summers. Gaps were more abundant in burned sites, while “under tussock” microsites were more frequent in unburned sites. In burned areas, tussocks allowed higher establishment of seedlings (facilitation), but gaps allowed more seedling growth and higher persistence of juveniles. Conclusions: Fire promoted S. bracteolatus recruitment in Patagonian grasslands by increasing the availability of favourable gap microsites. Grass protection for shrub seedlings became negative with time, probably due to competition with grasses. Gaps led to better performance and persistence of shrub plants. Six years after fire, higher shrub recruitment and adult density (observed as a trend) in burned grassland provides an opportunity for potential S. bracteolatus invasion.     

Species interactions at the level of fine roots in the field: influence of soil nutrient heterogeneity and plant size

Interference at the level of fine roots in the field was studied by detailed examination of fine root distribution in small soil patches. To capture roots as they occur in natural three-dimensional soil space, we used a freezing and slicing technique for microscale root mapping. The location of individual roots intersecting a sliced soil core surface was digitized and the identity of shrub and grass roots was established by a chemical technique. Soil patches were created midway between the shrub, Artemisia tridentata, and one of two tussock grasses, Pseudoroegneria spicata or Agropyron desertorum. Some soil patches were enriched with nutrients and others given only deionized water (control); in addition, patches were located between plants of different size combination (large shrubs with small tussock grasses and small shrubs with large tussock grasses). The abundance of shrub and grass roots sharing soil patches and the inter-root distances of individual fine roots were measured. Total average rooting density in patches varied among these different treatment combinations by only a factor of 2, but the proportion of shrub and grass roots in the patches varied sixfold. For the shrub, the species of grass roots sharing the patches had a pronounced influence on shrub root density; shrub roots were more abundant if the patch was shared with Pseudoroegneria roots than if shared with Agropyron roots. The relative size of plants whose roots shared the soil patches also influenced the proportion of shrub and grass roots; larger plants were able to place more roots in the patches than were the smaller plants. In the nutrient-enriched patches, these influences of grass species and size combination were amplified. At the millimeter- to centimeter-scale within patches, shrub and grass roots tended to segregate, i.e., avoid each other, based on nearest-neighbor distances. At this scale, there was no indication that the species-specific interactions were the result of resource competition, since there were no obvious patterns between the proportion of shrub and grass roots of the two species combinations with microsite nutrient concentrations. Other potential mechanisms are discussed. Interference at the fine-root level, and its species-specific character, is likely an influential component of competitive success, but one that is not easily assessed.     

Effect of gibberellic acid treatment,and nutrient supply through detached tillers,upon haploid frequency in barley

Summary Haploids from Hordeum vulgare (2n = 14) X H. Bulbosum (2n = 14) crosses result after fertilization from the subsequent elimination of bulbosum chromosomes during early embryo development. Seed set from the cross is high but embryo culture is necessary to obtain seedlings. Application of gibberellin A₃ (GA₃) to pollinated florets was effective for increasing the frequency of haploid seedlings produced on both nutrient-fed detached tillers and intact plants. GA₃ increased both seed set and embryo yield. The number of cells per embryo during its transition to the haploid state was increased two to three times following GA₃ treatments. Enhanced embryo and endosperm development was attributed to increased mitotic activity. The number of visibly differentiated embryos was doubled to about 35 % of the cultured embryos after GA₃ was applied to detached tillers in nutrient solution. About 70 % of the resulting haploid plants developed from the visibly differentiated embryos. The detached tiller technique offers a convenient method of culturing haploids from field-grown plants.