Initial Defoliation Research Articles

Mineral nutrients improve phosphonate effectiveness against cork oak root disease

Phosphonate fungicides are the most efficient chemicals for the control of oomycete root diseases which are causing significant economic and ecological losses in Mediterranean climate-type forests and natural ecosystems worldwide. However, a more environmental-friendly disease management demanded by society requires the optimization of chemicals application. Thereby, the main goal of this study was to develop a new resistant inducer (RI) formulation based on potassium phosphonate (PP) combined with mineral nutrients efficient against the root rot caused by Phytophthora cinnamomi and Pythium spiculum on cork oak that improve potassium phosphonate effectiveness. The potential phytotoxicity of five RI formulations composed of PP at five concentrations (25, 40, 50, 100 and 200 g/l) and each including 2.15 g/l ZnCl2 (Zn) and 0.25 g/l CaCl2 (Ca), were tested on 1-year old Quercus suber seedlings. Two weeks after treatments, only seedlings treated with the RIs containing the two highest doses of PP (100 and 200 g/l) showed symptoms of toxicity. RI with 40 g/l of H3PO3 (RI40), as well as its compounds individually and in combination two by two (PP (40 g/l), Ca, Zn, PP + Ca, PP + Zn, Ca + Zn), were tested against root disease development induced by P. cinnamomi and Py. spiculum on cork oak seedlings. For both oomycetes, seedlings submitted to every treatment with PP exhibited a significantly lower severity of root symptoms than plants artificially inoculated and treated with water. Furthermore, plants treated with RI40 did not differ to the plants growing in uninfested soil, improving PP alone effect. These results were validated by a field experiment where 60 mature cork oaks in three different defoliation classes (asymptomatic, slight, and moderate defoliation) were trunk injected with RI40 or water (controls). Three years after treatments, root disease development was significantly lower in cork oaks with slight and moderate initial defoliation level and treated with RI40 in comparison with water-treated controls. No effect was detected over both pathogens’ dynamics in soil. Results strongly support that PP effectiveness in controlling Q. suber root disease caused by P. cinnamomi and Py. spiculum is improved by its combination with Zn and Ca, allowing to reduce the PP application rates according to current society demands and the European Plant Health Legislation.

Oil spill from the Era: Mangroves taking eons to recover

Mangroves are highly susceptible to oil exposure. Depending on the severity, oil exposure can result in initial defoliation and eventual recovery through to mass mortality and complete loss of habitat. Some aspects of the impact of oil on mangroves and their recovery are well studied, but the focus has been on short-term responses, and the understanding of the longer-term trajectory of mangrove recovery from oiling is very limited. Here, we combine field results from sampling in the two years following a significant oiling event, with analysis of canopy cover in aerial images from before the event to 26 years afterwards. Approximately 100 ha of a monospecific stand of Avicenna marina mangroves were oiled as a result of a spill from the Era tanker in Spencer Gulf in southern Australia in September 1992. While lightly oiled trees made a full recovery, trees in heavily oiled areas experienced mass defoliation and ultimately mortality within several months of the oiling event. An analysis of aerial images indicated that there was no recovery in heavily oiled areas for 10 years following the oiling event. Between 10 and 25 years, seedling establishment and growth saw canopy cover increase to 35% of pre-oiling cover within heavily oiled areas. Predictive modelling estimates that complete recovery of mangroves to pre-oiling cover will take 55 years (median prediction in 2047). Our findings indicate that although mangroves can recover following a heavy oiling event, the rate of recovery can be slow, with full recovery in the order of half a century, much longer than has previously been anticipated.

Fosetyl-aluminium injection controls root rot disease affecting Quercus suber in southern Spain

In Spain, natural and managed cork oak woodlands are severely affected by decline caused by the root rot oomycetes Phytophthora cinnamomi and Pythium spiculum. The need for environmental-friendly disease management makes the use of resistance inducers a suitable option. As the use of phosphite is prohibited in Spain, trunk injections of fosetyl-aluminium were tested in a seminatural cork oak woodland (dehesa). Sixty cork oaks in three different defoliation classes (asymptomatic, slight, and moderate defoliation) were selected for fosetyl- aluminium or control (water) trunk injections, and periodically checked for defoliation and presence of both pathogens in roots and rhizosphere soil. Three years after treatments, fosetyl-aluminium injection significantly decreased defoliation in treated cork oaks in comparison with water-injected controls, regardless of the initial defoliation class considered. Changes in inoculum density in the rhizosphere or presence of the two pathogens in roots were not significantly dependent on treatment. However, a trend to a lower presence of pathogens in roots of treated oaks was observed with increasing soil inoculum densities. Fosetyl-aluminium, a phosphonate fungicide registered in the European Union, was experimentally proved to protect cork oaks against root rot caused by P. cinnamomi and Py. spiculum, even exhibiting a therapeutic effect on preexisting infections. This effective measure can therefore be used to fight against this highly destructive disease in seminatural woodlands and in protected natural ecosystems.

Trunk injection of fosetyl‐aluminium controls the root disease caused by Phytophthora cinnamomi on Quercus ilex woodlands

In Spain, Quercus open woodlands are animal ranching systems of organic production seriously threatened by the exotic pathogen Phytophthora cinnamomi. The root disease it causes kills thousands of oaks annually. Effective disease management needs to integrate different techniques, and the use of a resistance inducer such as fosetyl‐Al can play a key role, because the use of potassium phosphite is prohibited in Spain. In a woodland where the pathogen recently arrived, 60 holm oaks in three different defoliation classes (asymptomatic, slight and moderate defoliation) were selected for trunk injection with pressurised capsules containing 4% of commercial fosetyl‐Al or water (controls). Holm oaks were checked periodically for defoliation and presence of the pathogen in roots and rhizosphere soil. Three years after treatments, defoliation was significantly lower in oaks treated with fosetyl‐Al, which even increased canopy cover, in comparison with control oaks, independent of the initial defoliation class considered. Chlamydospore density in rhizosphere soil, as well as the presence of the pathogen into the roots, was not significantly influenced by fosetyl‐Al treatments, although a trend to a lower presence of P. cinnamomi in roots was observed in treated oaks at every soil inoculum density detected. This study has shown that fosetyl‐Al, a phosphonate registered as a fungicide in the European Union, provides protection to holm oaks against P. cinnamomi, even exhibiting a therapeutic effect on pre‐existing infections. Consequently, this effective measure should be considered as part of the integrated approach to control this highly destructive pathogen in holm oak woodlands.

Phenology and Abundance of Northern Tamarisk Beetle, Diorhabda carinulata Affecting Defoliation of Tamarix

Timing and spatial dynamics of tamarisk (Tamarix spp. L.) defoliation by the biological control agent Diorhabda carinulata (Desbrochers) were evaluated. Relative abundance of D. carinulata and the phenology of tamarisk along the San Juan and Colorado rivers were recorded in 2011–2012. D. carinulata began reproducing in the spring when temperatures were >15°C. Variation in spring temperature-rise affected the timing of development of larvae of the first summer generation and initial defoliation of tamarisk at each site. Shortening day lengths in mid- to late-summer cued D. carinulata to enter reproductive diapause resulting in cessation of defoliation. The critical day length for inducing reproductive diapause was 33–47 minutes shorter than that of populations of D. carinulata released into North America in 2001. Variation in spring temperature-rise combined with timing of shortening day length resulted in differences in D. carinulata voltinism per site. During the active season, larvae were less likely to establish in areas where defoliation was >70%. Lack of reestablishment of larvae led to temporary loss of D. carinulata from the locations and allowed tamarisks to sprout new canopies. Defoliation of tamarisk was dictated by environmental cues and abundance of D. carinulata, and in turn large amounts of defoliation negatively affected abundance of D. carinulata.

Carbon allocation to growth and storage in two evergreen species of contrasting successional status.

A prevailing hypothesis in forest succession is that shade-tolerant species grow more slowly than shade-intolerant species, across light conditions, because they prioritize carbon (C) allocation to storage. We examined this hypothesis in a confamilial pair of species, including one of the fastest-growing tree species in the world (Eucalyptus globulus) and a shade-tolerant, slow-growing species (Luma apiculata). Seedlings were subjected to one out of four combinations of light (high vs. low) and initial defoliation (90% defoliated vs. nondefoliated) for four months. Growth, C storage concentration in different organs, leaf shedding, and lateral shoot formation were measured at the end of the experiment. Eucalyptus globulus grew faster than L. apiculata in high light, but not in low light. Both species had lower C storage concentration in low than in high light, but similar C storage concentrations in each light condition. Defoliation had no effect on C storage, except in the case of the old leaves of both species, which showed lower C storage levels in response to defoliation. Across treatments, leaf shedding was 96% higher in E. globulus than in L. apiculata while, in contrast, lateral shoot formation was 87% higher in L. apiculata. In low light, E. globulus prioritized C storage instead of growth, whereas L. apiculata prioritized growth and lateral branching. Our results suggest that shade tolerance depends on efficient light capture rather than C conservation traits.

Timing and frequency are the critical factors affecting the impact of defoliation on long term growth of plantation eucalypts

Insect defoliation of plantation Eucalyptus sp. is a ubiquitous problem, not only in their native Australia but also in many other countries where Australian defoliators have invaded the introduced eucalypt plantations. Although eucalypts are very resilient to defoliation, their growth suffers and reduces the economic benefits of the resource. An artificial defoliation trial investigated the critical factors relating to patterns of insect defoliation that affect the long-term growth of the plantation eucalypt, E. nitens, in Tasmania, Australia. Current season’s adult-phase foliage was removed manually from two- to three-year-old trees to test four factors: severity (50% or 100% of current adult foliage, equivalent to 11% or 25% of total foliage); disbudding following defoliation (with or without disbudding), timing of defoliation (early or late in the summer) and frequency (for one or two consecutive years). Growth parameters of the trees were measured annually for four years after the initial defoliation, and again thirteen years later, before harvest. The most significant factors affecting the growth of the trees were timing and frequency of defoliation; severity of defoliation and disbudding did not have significant effects over the long term. Trees that received either light or heavy defoliation late in the season for two consecutive years were at least 17% smaller in diameter and MAI in diameter was reduced by at least 21% compared to untreated trees over one rotation. This means they would need to be grown for three to four more years to reach the same stand volume as undefoliated trees at harvest. This would have serious cost implications for plantation managers. To prevent these economic losses, an integrated pest management system should focus on protecting eucalypts from defoliation of 50% or more of current season’s adult foliage late in the summer, and in particular, preventing defoliation from occurring in concurrent years. In addition, the continual decline in growth rates of defoliated trees relative to undefoliated trees beyond the initial four years of measurement also suggests that defoliation impacts predicted by models based on short term studies may need to be treated conservatively.

Biological control reduces growth, and alters water relations of the saltcedar tree (Tamarix spp.) in western Nevada, USA

We monitored the impacts of a biological control agent, the saltcedar leaf beetle (Diorhabda carinulata), on the saltcedar tree (Tamarix spp.) at two sites (Humboldt and Walker rivers) in Nevada, USA. At the Humboldt site trees that had experienced three to four defoliation events had more negative water potentials and lower foliar Δ13C than trees farther from the release site that had experienced only one defoliation event. We established paired trees (exposed to D. carinulata and sprayed with insecticide) at both sites and monitored impacts. Beetles reduced stem growth during the first year of defoliation at both sites but not in the second year at the Humboldt site. Defoliation did not affect midday water potentials, or leaf gas exchange during the first two years of defoliation of paired trees at either site. Furthermore there was no difference in foliar Δ13C in either year at the Humboldt site but defoliation during the first year lead to higher foliar Δ13C at the Walker site. These results suggest that initial defoliation by D. carinulata reduces growth but not water relations of saltcedar. However, repeated defoliation, potentially acting through reduced root growth, leads to an overall reduction in the water status of this invader.

Tree Mortality in Mixed Pine–Hardwood Stands Defoliated by the European Gypsy Moth (Lymantria dispar L.)

Abstract Defoliation by the European gypsy moth (Lymantria dispar L.) and subsequent tree mortality have been well documented in the northeastern United States. In this study we evaluate tree mortality after initial defoliation in mixed pine–hardwood stands in the southeastern United States as the range of European gypsy moth expands. In 1993, 46 mature pine–oak and pine–sweetgum stands were selected on the coastal plain in Maryland and Virginia along the advancing gypsy moth front. Initial stand conditions, defoliation, and tree mortality after a single defoliation outbreak were monitored for 4 years. Susceptible oaks and sweetgum were defoliated at the greatest intensities. After defoliation, mortality in attacked stands was 4.4 m2/ha higher in pine–oak stands and 1.7 m2/ha higher in pine–sweetgum stands, representing 13 and 3% of the total basal area, respectively. Mortality of pines was observed mainly among suppressed and intermediate trees. Logistic regressions were derived to correlate individual tree mortality to stand and tree factors associated with gypsy moth attack. On the basis of these models, susceptible understory species with degraded crowns in pine–oak stands were at the greatest risk of mortality. Dominant overstory pines in pine–hardwood stands appear to be at low risk.

Comparative defoliation tolerance of temperate perennial grasses

AbstractThe defoliation tolerance of cultivars of four temperate perennial pasture grasses, perennial ryegrass (Lolium perenne, cv. Yatsyn1), phalaris (Phalaris aquatica cv. Australian), tall fescue (Festuca arundinaceae cv. Demeter) and cocksfoot (Dactylis glomerata cv. Porto), was determined under controlled conditions over a period of 12 weeks. Undefoliated plants were compared with defoliated plants, where only half of one leaf was left intact at the initial defoliation, and leaf regrowth was harvested every 3–4 d. The growth responses measured were plant tiller number, dry weight, relative leaf regrowth rate, root:shoot ratio, sheath:stem ratio and specific leaf weight. All species showed morphological adaptations that potentially increased their ability to tolerate defoliation (e.g. increased allocation to shoot at the expense of roots and lower specific leaf weights) but cocksfoot was found to be the most defoliation‐tolerant and perennial ryegrass the least. The adaptation that favoured cocksfoot most strongly was high sheath:stem ratio, which, it is proposed, allowed it to maintain photosynthesis and a level of carbon supply sufficient to support regrowth throughout the experiment. The strategy of perennial ryegrass which favours leaf growth and leads to rapid leaf turnover rates made it particularly susceptible to defoliation under the conditions of this experiment. This highlights the likely importance of defoliation‐avoidance responses in explaining the well‐known grazing resistance of this species. Phalaris and tall fescue showed responses that were intermediate between the other two species. The importance of defoliation‐avoidance mechanisms and implications for grazing management are discussed.

Performance of the green spruce aphid, Elatobium abietinum (Walker) on previously defoliated Sitka spruce

Abstract 1 The green spruce aphid, Elatobium abietinum, is an important defoliator of Sitka spruce in the U.K. However, it is usual for years in which high E. abietinum populations occur to be followed by a year with low aphid densities. The possibility that the performance of E. abietinum is reduced on previously infested Sitka spruce, and that this is the cause of year‐to‐year fluctuations in population density, was investigated by comparing population development and the growth rate of individual aphids on experimentally defoliated trees.2 Separate experiments were performed to determine whether aphid performance was reduced either in the autumn immediately after defoliation in the spring, or was reduced in the spring of the next year. Different rates of initial defoliation on trees used to test aphid performance were created by artificially infesting the trees with aphids in the spring before the experiments, and varying the time of infestation.3 Population development and the mean relative growth rate (MRGR) of individual aphids on previously defoliated and undefoliated Sitka spruce did not differ significantly in the spring of the next year. No differences were observed in the nutrient content of the 1‐year‐old needles of previously defoliated or undefoliated trees at this time.4 In the autumn and winter immediately after spring defoliation, aphid MRGR was significantly higher on trees that had been heavily defoliated earlier in the season compared with trees that had been lightly defoliated. However, the difference in MRGR decreased over the winter period. Nitrogen, phosphorous and potassium concentrations were 9.4–12.2% higher, at the beginning of the autumn, in the current year needles of heavily defoliated trees than in the current year needles of lightly defoliated trees.5 The experiments indicate that high populations of E. abietinum in the spring do not induce any defensive mechanisms in Sitka spruce that adversely affect subsequent generations of the aphid. By contrast, the results suggest that high spring densities of the aphid improve the nutritional quality of the current year's foliage for autumn generations.

Seasonal Dynamics and Defoliation Impact on Herbage Yield in Aspen Boreal Habitats of Alberta

Within the aspen boreal ecosystems, little information exists on the seasonal dynamics of available herbage and the effects of varying defoliation regimes on accumulated herbage growth and associated opportunities for animal production. We examined seasonal changes in herbage phytomass in conjunction with defoliation treatments in Bromus inermis-Poa pratensis grasslands in central Alberta. Changes in herbage pools were examined by sampling at five monthly intervals from April to September 1997 and 1998, inclusive. Vegetation was also subjected to a factorial experiment with an initial defoliation in late-May, June or July, at heights of 2.5, 7.5, or 15 cm, and repeated at 3-, 6- or 9-week intervals until the end of September. Green herbage, standing dead and fallen litter increased from spring to summer and decreased from summer to fall. Average growing conditions resulted in a peak phytomass of 350 g m-2, and varied by year. Weathering losses of green herbage, standing dead and fallen litter over winter were 34%, 52% and 51%, respectively. Dry matter losses of total herbage (all three pools) over winter were 58% of 1997 summer green phytomass. Initial timing, height, and frequency of clipping all affected accumulated herbage yield (P < 0.001). The greatest accumulated herbage yield was from clipping initiated in May. Light clipping resulted in less phytomass accumulation relative to moderate and heavy clipping. Clipping frequencies of six and nine weeks resulted in similar phytomass removal, but were greater than herbage removal associated with three weeks frequency. The interaction between clipping date and frequency of clipping demonstrates the importance of temporal rest and sensitivity of forage plants to defoliation, and lends support to the use of rotational grazing systems.

Defoliation Regime Effects on Accumulated Season‐long Herbage Yield and Quality in Boreal Grassland

AbstractWithin boreal grasslands, little information exists on the effects of initial defoliation date, frequency, and intensity on accumulated herbage yield and quality. We investigated the effects of initial defoliation in May, June or July, at heights of 5, 10 or 15 cm, and repeated at 2‐, 4‐ or 6‐week intervals throughout the growing season. Harvested material was combined with year‐end residual biomass, and examined for herbage removed, crude protein (CP), crude protein yield (CPY) and neutral detergent fibre (NDF). Compared to single defoliated check plots, total, grass and forb dry matter (DM) yields were lower under repeated defoliation by 25, 38 and 17 %, respectively. The majority of total herbage produced was harvested in the spring clipping. Total, grass and forb DM yields peaked under moderate (10 cm) clipping. Total DM and grass biomass were maximized with long (6 week) recovery periods. In contrast, forb biomass was greatest with May defoliation followed by a 4‐week interval. While maximum grass CPY was found under 10‐cm defoliation, forb CPY peaked with early and moderate to intensive defoliation. These results indicate that season‐long herbage biomass, along with CPY, can be maximized in boreal grasslands through controlled defoliation.

Effect of defoliation height and redefoliation interval on regrowth and survival of perennial ryegrass (Lolium perenne) in subtropical dairy pastures

A cut plot study was undertaken on the subtropical north coast of New South Wales, Australia, to determine the effect of defoliation height and redefoliation interval on dry matter yield and persistence of perennial ryegrass (Lolium perenne L.) pastures. The pasture was established on 7 April 1998 and plots were irrigated to replace evapotranspiration loss. The study was a completely randomised block design with plots of 2 by 1 m and treatments replicated 3 times. In winter (commencing 13 July) plots were defoliated to 20, 50 or 120 mm stubble height and either not redefoliated or redefoliated at 3, 6 or 3 and 6 days after initial defoliation. In spring (commencing 28 October) plots were redefoliated as for winter but only to 50 mm stubble height. After imposition of the redefoliation treatments, the plots were allowed to regrow until the non-redefoliated treatments had regrown 3 new leaves per tiller (subsequently referred to as a regrowth cycle) and then again defoliated (regrowth cycle 1). Plots cut in winter were then halved with one half (A plots) continuing to be subject to the redefoliation treatment for 4 more regrowth cycles until regrowth cycle 1 in spring was completed on 24 November, while the other half (B plots) were a carryover comparison of redefoliation treatment in regrowth cycle 1. Both A and B plots continued to be subjected to the same defoliation height treatments as imposed in regrowth cycle 1. From 24 November to 30 March 1999, plots were defoliated at 50 mm height each time 3 new leaves per tiller had regrown.Plots defoliated to 20 or 50 mm height during regrowth cycle 1 in winter yielded 21% more dry matter than plots cut to 120 mm height while redefoliation at 6 or 3 and at 6 days produced 14% less dry matter than plots not redefoliated or redefoliated at 3 days. Continued redefoliation at 6 days (comparison within A plots) reduced dry matter yield by 63% compared with no redefoliation or redefoliation at 3 days, but only in plots defoliated to 20 or 50 mm height.Plant density in the autumn (March 1999) of the year after establishment was positively related to defoliation height over regrowth cycles 1–5 of the previous year (35, 55 or 77 plants/m2 for plots defoliated at 20, 50 or 120�mm, respectively). Plant density of plots not redefoliated or redefoliated at 3 days over regrowth cycles 1–5 was 63% higher (70 plants/m2) than for the other treatment combination (43 plants/m2) at P = 0.07 level of significance.Plants cut to 20 or 50 mm stubble height at the commencement of regrowth cycle 1 in winter had a stubble water-soluble carbohydrate content of 5.2%, decreasing to 2.3% at day 6 post-defoliation. The water-soluble carbohydrate content of plants cut to 120 mm were initially higher at 8% and fell to only 6.4% by day 6.The redefoliation treatments imposed in this study were designed to simulate the regrazing of regrowth shoots in an extended grazing bout at various defoliation heights. The results confirm the negative effects of redefoliation, at 6 days in the winter to spring period, on both dry matter yield and plant survival over the subsequent summer in the subtropics. In contrast to winter, redefoliation in late spring had no effect on dry matter yield or plant density. The results also indicate a compromise between the benefits of more lax grazing for persistence and harder grazing for pasture utilisation.

Temporal patterns of nitrogen leakage from mid‐Appalachian forested watersheds: Role of insect defoliation

Fluxes of dissolved nitrogen (N) as nitrate from forested watersheds in the mid‐Appalachian region have important water quality ramifications for small acid‐sensitive streams and for downstream receiving waters such as the Chesapeake Bay. Previous studies of N leakage have suggested that annual dissolved N fluxes from small watersheds can vary by several orders of magnitude and may be increasing as second‐growth forests gradually become N saturated from the accrual of atmospheric N loadings. In this study, we examined the temporal (intra‐annual and interannual) variability in dissolved nitrate fluxes from five small (area < 15 km2) forested watersheds in the mid‐Appalachian region from 1988 to 1995. At all sites, nitrate concentrations were observed to increase dramatically during storm flow events, with nitric acid contributing significantly to depressions in pH and acid‐neutralizing capacity; annual nitrate fluxes were dominated by high‐discharge periods. Interannually, the fluxes at each site varied by 1–2 orders of magnitude, but the patterns of N leakage displayed considerable synchrony with outbreaks of gypsy moth caterpillar defoliation that began in the late 1980s and early 1990s in this region. N leakage from forested watersheds apparently lagged the initial defoliation by several months to perhaps a year or more. Defoliation outbreaks by the gypsy moth caterpillar (or other herbivorous pests) thus provide an alternative explanation of N leakage from forest ecosystems. Poorly documented insect defoliations, rather than premature N saturation of intact forest ecosystems, need to be considered as a possible explanation of N leakage from forested watersheds in the mid‐Appalachian region and elsewhere.

Relationship between environmental factors, effects of fungicide applications, and premature defoliation of black locust trees (Robinia pseudoacacia L.)

From 1989 through to and including 1994 extensive premature defoliation of black locust ( Robinia pseudoacacia L.) was observed in a progeny test plantation in Greene County, in the Georgia Piedmont. During July 1991 and 1992, all observed trees suffered greater than 50% defoliation. Leaflets of black locust showing symptoms of leaf spotting began abscising as early as mid-June following wet springs. Onset and progress of defoliation was followed on 50 trees in two plots in 1993 and 1994. In both years, sample trees were cut at the base and allowed to regrow from sprouts. A protectant fungicide, chlorothalonil (Daconil WDG 1.5 g/l water) was applied to run-off weekly to one-half of the growing sprouts beginning 17 April through 22 August 1994 by backpack sprayer. Air temperature, relative humidity, rainfall, and leaf wetness were monitored. Abscission of unprotected leaflets in 1993, a dry, hot year, began in mid-July and averaged 36% by late August; abscission of unprotected leaflets in 1994, a wet, cool year, began in mid-June and averaged 80% by late August; whereas, abscission of protected leaflets in 1994 began in late June and averaged 70% by late August. The Gompertz model described the increase of leaflet abscission with time. The estimated rate of leaflet abscission for unprotected leaflets in 1993 was significantly slower ( k = 0.056/wk) compared to the rate of abscission of unprotected leaflets in 1994 ( k =0.070/wk). Estimated rates of leaflet abscission were not significantly different between protected ( k = 0.063/wk) and unprotected trees ( k =0.070/wk) in 1994. Cumulative hours of leaf wetness of 600 h were associated with initial leaf spotting and defoliation in 1993 and in 1994. Delay in leaflet abscission of protected trees indicates that fungal infection may have contributed to premature defoliation of black locust with onset related to moisture conditions favourable for infection.

Seedling White Clover Response to Defoliation

Defoliation during the early stages of seedling development can influence growth of white clover (Trifolium repens L.). Our objective was to determine the response of cultivars with diverse leaf sizes to defoliation across a range of seedling growth stages. ‘Regal’ (large‐leaf), ‘Louisiana S‐1’ (medium‐large‐leaf), ‘Grasslands Huia’ (medium‐smallleaf), and ‘Aberystwyth S184’ (small‐leaf) were grown in the greenhouse and were initially clipped to remove all leaf (except cotyledons) and petiole tissue (1.0‐cm stubble) when seedlings attained a unifoliolate leaf or one, two, four, and eight trifoliolate leaves. Seedlings were subsequently clipped in the same manner every 7 or 28 d for the next 28 d. Plant response to defoliation was assessed 7, 14, 21, and 28 d after the last clipping. Seedling shoot dry weight of all cultivars increased linearly in response to initial leaf stage when subsequent clipping occurred every 7 d after the initial defoliation. When subsequent clipping was delayed 28 d, defoliation after the four‐trifoliolate stage did not influence shoot weight of any cultivar except Regal. Despite fewer numbers of leaves after regrowth, seedling leaf area of Regal was always greater than that of the other cultivars. Number of stolon branches of Regal and Louisiana S‐1 increased in response to leaf stage at initial defoliation (0.15 branches stage−1), while that of Grasslands Huia and Aberystwyth S184 was relatively unchanged after the four‐ to six‐trifoliolate stage. Permitting white clover seedlings to attain advanced leaf stages before initial defoliation and increasing the time interval before subsequent defoliation will enhance seedling growth and potential survival.

Delay of harvest effects on forage yield and regrowth in spring and winter cereal mixtures

Spring-planted mixtures of spring and winter cereals extend the grazing season and maximize dry matter yield if the initial defoliation is delayed until the milk stage of kernel development of the spring cereal component. However, fall regrowth may be less than spring-planted winter cereal monocrops. Research was conducted at Lacombe, Alberta to determine the effect of time of initial cut and winter cereal species on initial yield, regrowth yield and factors affecting regrowth (tiller density, water-soluble carbohydrate (WSC) and etiolated regrowth immediately post-cutting) of the winter cereal component of spring-seeded monocrops of fall rye (Secale cereale L.), winter triticale (× Triticosecale Wittmack) and winter wheat (Triticum aestivum L.) and in binary mixtures with spring barley (Hordeum vulgare L.). Treatments were planted in early May and harvested initially when the barley reached the boot (B), heads emerged (H), H + 2 wk, H + 4 wk and H + 6 wk stages. Three weeks after the initial cut a regrowth harvest was conducted. Initial yields of both mixtures and monocrops increased until approximately H + 4 wk and H + 6 wk respectively, with no differences due to species in the mixture. The effect of crop stage at initial harvest on regrowth was much larger than the species effect. Monocrop regrowth decreased almost linearly with delay of defoliation, while that of the mixture (winter and spring components combined) declined at a faster rate. Tiller density, WSC and etiolated regrowth also followed decreasing trends with time of initial cut in the monocrops and mixtures. These trends were due directly or indirectly to very low light intensities in the lower levels of the canopies of both cropping systems after H + 2 wk. Both initial and regrowth yields of the winter cereal component of mixtures involving rye and triticale were generally superior to those involving wheat. The ability of the winter cereal component to regrow in mixtures may be related to plant size and therefore ability to compete during initial growth. In general, the initial harvest of the mixture should occur no later than H + 2 wk of the spring cereal component in order to allow sufficient time for recovery of the winter cereal component after harvest. Otherwise, a spring-seeded winter cereal would be a better alternative for fall pasture. Key words: Delayed harvest, spring and winter cereals, forage yield, regrowth

Impacts of Defoliation on Tiller Production and Survival in Northern Wheatgrass

Although northern wheatgrass (Agropyron dasystachyum (Hook.) Scribn.) is a dominant or co-dominant species that decreases under grazing in northern Mixed Prairie, little is known about its response to herbage removal at different times during the growing season. The objective of this research was to determine the effects of defoliation on the tiller production and survival of this native perennial on a clayey range site in mixed prairie in south-central Saskatchewan. Vegetation was subjected to a factorial experiment with an initial defoliation in early-May, June, July, or August and repeated at 2- or 6-week intervals until mid-September in the same plots for 3 years. An undefoliated control was also included. On average defoliation enhanced tillering (71%) and survival relative to the control, and tiller recruitment was greatest during June and September 1989. Generally tiller survival decreased as the date of emergence in the growing season was delayed. Numbers of tillers emerging was positively correlated with soil water (r = 0.77). Some tillers of northern wheatgrass lived 5 years. The 2- and 6-week intervals of defoliation had little influence on tiller survival, but initiating defoliation near the time of tiller emergence reduced survival whereas delaying defoliation until August increased their survival. Increased tillering may be an adaptive feature enabling northern wheatgrass to tolerate defoliation by re-establishing lost photosynthetic area and maintaining or even increasing basal area. Thus, once released from grazing it may rapidly increase phytomass production in a relatively short time. Delaying grazing until August will maximize tiller survival of northern wheatgrass.

Defoliation of a Northern Wheatgrass Community: Above- and Belowground Phytomass Productivity

A defoliation study was conducted on a fair condition, clayey range site that is potentially dominated by northern wheatgrass (Agropyron dasystachyum (Hook.) Scribn.) in south-central Saskatchewan. Vegetation was subjected to a factorial experiment with an initial defoliation in early-May, June, July, or August and repeated at 2- or 6-week intervals until mid-September in the same plots for 3 years. An undefoliated control was also included. Herbage removed, residual live, dead, total and root phytomass were measured. Defoliation reduced all yield components, with the exception of herbage removed. Residual live grass was reduced 37, 57, and 46%, respectively, in first second, and third years; the sedge and forb components of live residual phytomass generally were not affected by defoliation. Compared to control, dead phytomass was reduced 77% in the first year, 67% in the second, and 52% in the third year across treatments. Total herbage yield across defoliation treatments ranged from 68 to 93% of control Total live phytomass (herbage removed + residual live phytomass) in defoliated plots equaled control Herbage removal was greatest when initially defoliated in early July and thereafter at 2-week intervals. When defoliated at 6-week intervals residual live and dead phytomass were generally greater than when herbage was removed biweekly. Yields were higher when the first defoliation was delayed and repeated at 6-week intervals. Generally, root phytomass was not different among defoliation treatments, but total belowground phytomass was reduced 30% in the 0-30-cm depth after 3 year of defoliation. This northern mixed prairie ecosystem is sensitive to herbage removal. Maximum forage yield can be obtained if grazing is deferred until after peak growth in July.