Initial regenerative processes ofDistylium racemosum andPersea thunbergii in an evergreen broad-leaved forest

We studied the initial regenerative processes ofDistylium racemosum andPersea thunbergii, major canopy species, in an old-growth evergreen broad-leaved forest, Kagoshima, southwestern Japan. Although the two species coexisted, the patterns of seed production, seedling emergence, and sapling survival were different between them. During the observation (1989–1991),P. thunbergii produced large numbers of seeds in alternate years, whileD. racemosum had no mast year. The density of established seedlings was high and increased with mass seed production forP. thunbergii, but was fairly low forD. racemosum. The mortality of seedlings and saplings ofP. thunbergii was higher than that ofD. racemosum. In a closed stand, the sapling bank was maintained by dense seedling supply forP. thunbergii, and by extremely low mortality of individuals germinated before the beginning of observation forD. racemosum. Since saplings ofD. racemosum showed continuous height growth beneath the closed canopy, the possibility of recruitment into the upper layer seems to be high. The strategy ofP. thunbergii might be to wait for suitable conditions favoring sapling growth on various site by means of frequent and large seed productivity and wide dispersal of seeds. Thus both species might be able to coexist through environmental heterogeneity.     

Comparative studies on acorn and seedling dynamics of fourQuercus species in an evergreen broad-leaved forest

The dynamics and regeneration probabilities ofQuercus acuta, Q. gilva, Q. salicina, andQ. sessilifolia were studied in a 0.48 ha plot in an old-growth evergreen broad-leaved forest, Kagoshima prefecture, southwestern Japan. The canopy was well closed and had only one small gap. Seventeen tree species occurred in the canopy. The fourQuercus species ranked second to fifth in basal area. Excluding the trees originated from sprouts,Q. acuta and Q. gilva lacked trees at the subcanopy layer, andQ. salicina andQ. sessilifolia had some trees at the same layer. For eachQuercus species, the amount of fallen acorns fluctuated from year to year. The number of current year seedlings was positively correlated with the number of fallen acorns in the previous year. A disproportion between the spatial distribution of fallen acorns and that of established seedlings has been shown. The establishment of seedlings was frequently observed where no acorns had fallen in the previous year. Saplings of four species survived for several years and formed sapling banks under the closed canopy. During the three years of the study, the mortality ofQ. salicina andQ. sessilifolia saplings was lower than that ofQ. acuta andQ. gilva. However, the differences in the average annual height growth of saplings between four species were not significant, and the most of them did not grow much. These results suggest that the saplings of the fourQuercus, especiallyQ. gilva andQ. acuta, have no chance to recruit to the canopy or subcanopy layer within a closed stand, and that some changes in the environment caused by gap formation are required for their recruitment.     

Limits to tree species invasion in pampean grassland and forest plant communities

Factors limiting tree invasion in the Inland Pampas of Argentina were studied by monitoring the establishment of four alien tree species in remnant grassland and cultivated forest stands. We tested whether disturbances facilitated tree seedling recruitment and survival once seeds of invaders were made available by hand sowing. Seed addition to grassland failed to produce seedlings of two study species, Ligustrum lucidum and Ulmus pumila, but did result in abundant recruitment of Gleditsia triacanthos and Prosopis caldenia. While emergence was sparse in intact grassland, seedling densities were significantly increased by canopy and soil disturbances. Longer-term surveys showed that only Gleditsia became successfully established in disturbed grassland. These results support the hypothesis that interference from herbaceous vegetation may play a significant role in slowing down tree invasion, whereas disturbances create microsites that can be exploited by invasive woody plants. Seed sowing in a Ligustrum forest promoted the emergence of all four study species in understorey and treefall gap conditions. Litter removal had species-specific effects on emergence and early seedling growth, but had little impact on survivorship. Seedlings emerging under the closed forest canopy died within a few months. In the treefall gap, recruits of Gleditsia and Prosopis survived the first year, but did not survive in the longer term after natural gap closure. The forest community thus appeared less susceptible to colonization by alien trees than the grassland. We conclude that tree invasion in this system is strongly limited by the availability of recruitment microsites and biotic interactions, as well as by dispersal from existing propagule sources.     

The spatial pattern and canopy-understory association of trees in a cool temperate,mixed forest in western Japan

The spatial distribution pattern of trees and the association between canopy and understory individuals were examined with reference to the distribution of tree crowns in a cool temperate, mixed forest in Ohdaigahara, western Japan. Line transect and contact sampling methods were used to examine the pattern over various spatial scales. These methods are useful to detect patterns over a large study area. The dominance ofChamaecyparis obtusa on steep slopes forming large patches suggested that the distribution of this species is a consequence of landslides. UnderstoryFagus crenata showed a clumped distribution, and the relative coverage of this species was larger in canopy gaps than under a closed canopy. Understory individuals ofAbies homolepis showed a positive association with canopy trees ofF. crenata but a negative association with conspecific canopy individuals. These patterns suggested thatF. crenata regenerates in canopy gaps and is replaced byA. homolepis. The dynamics of these two species are consistent with the process of gap dynamics. The effects of both small- and large-scale disturbance must be evaluated to understand the mechanisms of patch formation and the coexistence of forest tree species.     

Regeneration in subalpine coniferous forests

The regeneration process of a subalpine coniferous forest, a mixed forest ofTsuga diversifolia (dominant species),Abies veitchii, Abies mariessi, andPicea jezoensis var.hondoensis, was studied on the basis of annual ring data. The age class distribution was discontinuous and four age groups occurred in the study plot (30m×30m). The canopy layer was a mosaic of patches (83.8–133.7 m² patch area), which had different mean ages. The recruitment of canopy trees was carried out only by advance regeneration in the plot. The diameter growth ofAbies andPicea exceeded diameter growth ofTsuga in the gap.Abies lived for 200–300 years and their trunks were susceptible to heart rot.Picea lived for 300–400 years andTsuga for more than 400 years. The regeneration process derived from the analysis of the plot consisted of three phases leading to the development of a even-aged patch; (1) the establishment of saplings before a gap opening, (2) the opening of a gap in the canopy and repair of the canopy by advance regenerated saplings dominated by rapid growth species,Abies andPicea, and (3) the dying off of canopy trees as each species reached the end of its life-span, resulting in pure patches of long-livedTsuga.     

Establishment and growth of mangrove seedling in mangrove forests of southern Thailand

The demography and growth of mangrove seedlings in southern Thailand were studied for an 18-month period. Plots for measurement of seedlings were established using gradients of light and surface soil texture. The number of established propagules peaked from April to August, the peak forRhizophora being earlier than that forBruguiera. Under the mangrove forest canopy, the survivorship of seedlings established during May to December 1982 was 33% in June 1983. Light conditions and soil texture did not strongly affect initial seedling establishment, and the relationship between species seed size and water depth at a particular site influenced that were able to become established there. The elongation growth was large during the rainy season and in open sites. The growth ofR. apiculata Blume,B. parviflora Wight & Arn. andB. cylindrica Blume in open sites was more than ten times that of seedlings established in the shade. It has noted that light conditions affected the growth of young trees (>1 year old) to a greater degree than that of current-year seedlings.     

Seedling establishment ofPolygonum cuspidatum andPolygonum weyrichii var.alpinum at high altitudes of Mt Fuji

Seedling establishment ofPolygonum cuspidatum (Seib. et Zucc.) andPolygonum weyrichii (F. Schmit) var.alpinum (Maxim.), dominant early-successional species in a volcanic gravel area on Mt Fuji, Japan, was compared for current-year seedlings at 2500 m a.s.l. The rate of survival in the first winter after germination at 2500 m was 2% and 24% forP. cuspidatum andP. weyrichii seedlings, respectively, and correlated with the upper distribution limit of each (2500–2600 m forP. cuspidatum; 3200–3300 m forP. weyrichii). The critical size for survival was 12 mg dry weight for overwintering current-year seedlings of both species. At 2500 m, the percentage of seedlings that attained the critical size for survival at the end of the first growing season was 19% and 85% forP. cuspidatum andP. weyrichii, respectively. Growth curves indicated that the larger-seeded characteristic ofP. weyrichii contributed to larger end-of-season size and resulted in higher rate of seedling survival at 2500 m, compared with smaller-seededP. cuspidatum. Current-year seedlings ofP. weyrichii, at least up to 3250 m, attained the first year annual growth critical for overwintering and therefore for successful seedling establishment, due to the large-seeded characteristic. At the top (3770 m) beyond the upper distribution limit ofP. weyrichii, most seedlings raised artificially failed to attain the critical size for survival in the first growing period due to the reduced growing period. Failure of seedling establishment would thus appear to restrict the distribution of the species at altitudes higher than 3300 m.     

林隙对香樟和枫香幼苗早期生长阶段功能性状的影响

探究林隙对不同需光性树种早期生长特征和功能性状的影响,对揭示林隙微生境影响次生林内幼苗更新机制具有重要意义。以亚热带次生林中耐荫常绿树种香樟和阳性落叶树种枫香幼苗为试验对象,研究大林隙(D/H介于1.5—2.0)、中林隙(D/H介于1.0—1.5)和小林隙(D/H介于0.5—1.0)对不同需光树种幼苗早期(1—3年生)生长特征和功能性状的影响。结果表明：(1)林隙大小对两种幼苗的生长均有显著影响。其中,中林隙可显著促进香樟2—3年生幼苗的生长,大林隙对枫香1—3年生幼苗的生长均具有显著促进作用。(2)对林隙环境因子与幼苗功能性状的关系进行冗余分析表明,香樟幼苗功能性状的变化与林隙土壤有机质含量、水解性氮含量、酸碱度和有效磷含量密切相关,而枫香幼苗功能性状则主要受林隙土壤酸碱度、有机质含量、水解性氮含量、土壤含水率、冠层透光率和土壤有效磷含量的影响。(3)维持较高的根重比、细根比根长、叶碳氮比和叶碳磷比是幼苗应对林隙环境影响的重要生理生态调节机制。     

Spatial pattern of seedlings 1 year after fire in a Mediterranean pine forest

Summary The spatial distribution of seedlings of the dominant perennial plant species (Pinus halepensis, Cistus salviifolius, Rhus coriaria) and may annual species was studied after a wild fire in an eastern Mediterranean pine forest. The spatial distribution of all seedlings is affected by the location of the old burned pine trees. Seedling density of Pinus and Cistus is higher at a distance from the burned pine canopy and lower near the burned pine trunk. It is also higher beneath small burned pine trees than under big ones. Rhus seedling density is higher under big burned pine trees and also near the burned trunks. Seedlings of Pinus, Cistus and Rhus growing under the burned canopy of big pine trees tend to be taller than seedlings under small ones or outside the burned canopy. Most annual species germinate and establish themselves outside the burned canopies, and only a few annual species are found beneath them. It is suggested that variation in the heat of the fire, in the amount of ash between burned pine trees of different sizes, and in the distance from the burned canopy are responsible for the observed pattern of seedling distribution. The possible ecological significance of the spatial pattern of seedlings distribution and their differential growth rate are discussed.     

The regeneration of pioneer tree species under browsing pressure of Sika deer in an evergreen oak forest

The effects of Sika deer (Cervus nippon) browsing on the regeneration of pioneer species were studied in relation to canopy gaps in a warm temperate evergreen oak forest in Kasugayama, Nara City. Four study sites, three in canopy gaps and one under a closed canopy, were selected and each divided into fenced and unfenced plots. Under the closed canopy, seedlings of all the pioneer species died irrespective of browsing pressure. However, in the canopy gap sites, seedlings of the pioneer species could establish and grow well. The seedling survival ratio in the fenced plots in the canopy gaps was >60% 1 year after germination. However, in the unfenced plots, only <20% of the seedlings survived 1 year, with all dying within 3 years after germination. Thus, the regeneration of pioneer trees in this forest was strongly inhibited by deer browsing. Successful regeneration of a pioneer,Zanthoxylum ailanthoides, occurred for several years even after two major wind disturbances during the past 90 years. This may be due to less browsing pressure from the deer.     

An interactive effect of simultaneous death of dwarf bamboo, canopy gap, and predatory rodents on beech regeneration

To clarify the interactive effect of the simultaneous death of dwarf bamboo (Sasa kurilensis), forest canopy gap formation, and seed predators on beech (Fagus crenata) regeneration, we analyzed beech demography from seed fall until the end of the first growing season of seedlings in an old-growth forest near Lake Towada, northern Japan. The simultaneous death of S. kurilensis took place in 1995. We established four types of sampling site differing in forest canopy conditions (closed or gap) and Sasa status (dead or alive). Beech seed survival and emergence ratio were both highest in gaps with dead Sasa (gap-dead), because rate of predation was lowest. Seedling survival during the first growing season was also highest in the gap-dead treatment, because of less predation and less damping off. As a result, even though density of seed fall was lowest in the gap-dead treatment, the living seedling density there was highest at the end of the first growing season. Predation, which caused the greatest mortality during the seed and seedling stages, was significantly lower at both sites in gaps and sites with dead Sasa. This was probably due to changes in the behavior of rodents in response to the structure of the forest canopy and undergrowth. Both the death of Sasa and canopy gap formation allowed seedlings to avoid damping off because of the high light availability. The indirect effect of the simultaneous death of Sasa and canopy gap formation in reducing predation contributed more to beech regeneration than their direct effect in increasing light for the seedlings.     

Hydrellia spp. [Diptera: Ephydridae] attackingHydrilla verticillata in south India

The leaf-mining fliesHydrellia sp. andH. pakistanae Deonier were observed attacking the aquatic weedHydrilla verticillata (L.f.) L.C. (Hydrocharitaceae) in south India. Their life histories were very similar. The duration of the immature stages ofH. pakistanae was 25 to 29 days. The number of leaves ofH. verticillata mined by a single larva varied from 8 to 12. In the laboratory, adults feeding on 50% aqueous solution of honey lived for 6 to 21 days and laid an average of 26 eggs. Several overlapping generations occurred in the field except during the period of drought. BothHydrellia sp. andH. pakistanae were heavily parasitized by the braconidsAdemon sp. nr.decrescens Nees andChaenusa sp.      

林窗对川西周公山柳杉人工林林下物种多样性的影响

采用典型样地法,以川西周公山柳杉人工林5种不同大小的林窗为研究对象,以林下非林窗为对照,研究了不同大小的林窗对柳杉人工林物种多样性的影响,同时分析了不同梯度林窗下林窗中心、林窗边缘、及林下群落的物种组成、物种多样性的变化情况。结果表明:(1)在所调查的18个样地231个样方中共记录到维管束植物141种,隶属于76科113属;随着林窗面积的增大,群落各层次的物种数呈现出先升高后降低的趋势,灌木层物种数在各林窗梯度上表现为林缘林下林窗中心,草本层物种数在各林窗梯度上表现为林缘林窗中心林下。(2)不同林窗优势种及其重要值不同,即在小林窗内,优势种为柳杉和野桐,其重要值之和高达0.292 3;在大林窗内,杉木及亮叶桦为群落优势物种,群落内出现大量其更新幼苗。(3)不同大小的林窗表现为灌木层物种丰富度指数(D)、Shannon-Wienner指数(H)、和Pielou均匀度指数(Jsw)值在400~450 m2面积的大林窗内达到一个均优水平,草本层物种的多样性在面积为100~150m2的小林窗内达到较高水平;不同梯度的林窗各层次群落D、H值整体表现为林缘林窗中心林下。研究认为:林窗的存在会改变群落物种组成,提高群落物种多样性水平,并且大林窗(400~450m2)更利于柳杉人工林林下树种更新及物种多样性的提高。     

Survivorship and growth of seedlings of four dipterocarp species in a tropical rainforest of Sarawak,East Malaysia

Survivorship and growth of seedlings of four dipterocarp species (Dipterocarpus actangulus, D. globosus, Dryobalanops aromatica, Dryobalanops lanceolata) were studied for 2.5 years in a mixed dipterocarp forest in Sarawak, East Malaysia. Predispersal seed predation rates were larger forD. globosus (75%) thanD. lanceolata (27–34%) andD. aromatica (18–26%). Less than 20% of the twoDryobalanops seeds were damaged by vertebrates after seed dispersal. During the period from seed dispersal to the time when the seedlings had shed cotyledons, more dispersed seeds died in the twoDipterocarpus (ca 90%) than the twoDryobalanops (ca 60–70%). The major mortality factors during this period were uprooted and seed/seedling predation by insects or vertebrates. After the seedlings shed cotyledons, all species showed constant mortality rates of 34, 15–16, 17 and 6%/year forD. actangulus, D. lanceolata, D. aromatica andD. globosus, respectively, in the forest understorey. Mortality was lower in less shaded conditions than in more shaded ones forD. aromatica andD. actangulus, but not significantly different forD. lanceolata andD. globosus. A majority of dead seedlings were killed by fallen branches or were found standing with wilted leaves, probably due to water stress. No significant correlation was found between seed/seedling mortality and distance from mother trees or the initial density of seeds/seedings for all species. The mean leaf production was positively correlated with the estimated diffuse light factor of their habitats for each species.     

Gap formation, microsite variation and the conifer seedling occurrence in a subalpine old-growth forest, central Japan

To evaluate the effects of canopy gaps and forest floor microsites (soil, fallen logs, root-mounds, buttresses and stumps) on regeneration of subalpine forests, the gap regeneration and seedling occurrence of conifers (Abies mariesii, Abies veitchii, Picea jezoensis var. hondoensis and Tsuga diversifolia) were studied in two stands of a subalpine old-growth forest, central Japan. The percentage of gap area to total surveyed area was 11.2–11.3% in the stands. Gap regeneration was not common for P. jezoensis var. hondoensis and T. diversifolia. In contrast, gap regeneration by advanced regeneration was relatively common for Abies. Seedling occurrence of P. jezoensis var. hondoensis and T. diversifolia was restricted on elevated surfaces such as stumps and root-mounds, while Abies seedlings could occur on soil as well as on elevated surfaces. Rotten stumps were the most favorable microsites for conifer seedling occurrence, which covered small area in the forest floor. Although canopy gaps were not always favorable for seedling occurrence, all conifer seedlings were larger under canopy gaps than under closed canopy. Canopy gaps and forest floor microsites clearly affected seedling occurrence and growth of conifers. This suggests that regeneration of conifers is related to the difference of growth advantage under canopy gaps and favorable microsites for seedling occurrence.     

Importance of seed size for the establishment of seedlings of five deciduous broad-leaved tree species

Effects of seed size and phenology on the establishment of five deciduous broad-leaved tree species were examined in deciduous woodland. Treatments included absence and presence of litter in the forest understory, a small gap, and a large gap. Seedling emergence of large-seeded speciesQuercus mongolica var.grosseserrata andAcer mono was not reduced by accumulation of litter in the forest understory, but was promoted in the large gap where litter was less. Seedling emergence of small-seeded species,Alnus hirsuta, Cercidiphyllum japonicum andBetula platyphylla var.japonica, was reduced by the litter in almost all of the sites. Seedlings of large-seeded species avoid shade stress phenologically by unfolding all of their large leaves in a short period before canopy closure in the forest understory. These species had little mortality after seedling emergence. In contrast, small-seeded species have a longer duration of leaf emergence, shorter leaf longevity, and rapid leaf turnover in all the sites. These seedlings attained similar height to those of the large-seeded species at the end of the second year in the large gap, but survival and height growth rate decreased after canopy closure in the forest understory. We suggest that the importance of seed size in determining seedling establishment largely depends on the relationships between seasonal changes of environmental conditions and phenological traits of seedlings, which are related to seed size.Abbreviations Ah Alnus hirsuta - Am Acer mono - Cj Cercidiphyllum japonicum - Bp Betula platyphylla var.japonica - Qm Quercus mongolica var.grosseserrata     

Variation with soil depth,topographic position and host species in the capacity of soils from an Australian locale to nodulateCasuarina andAllocasuarina seedlings

Sandy alluvial soils in a floodplain supporting a native stand ofCasuarina cunninghamiana Miq. produced about three times as many nodulated seedlings and more than twice as many nodules per nodulated seedling on roots of baitedCasuarina spp. than did clay loam red earth soils from the adjacent valley slope. Moist and well-aerated subsurficial alluvial sands had the greatest nodulation capacity of all the soils sampled. For all topographic positions, soil samples from depths greater than 20 cm promoted 76% more nodulated Casuarina seedlings than samples from the surficial 20 cm.Seedlings of three provenances ofC. cunninghamiana, together with seedlings ofC. glauca Sieb. ex Spreng.,C. cristata F. Muell ex Miq. andC. obesa Miq. developed significantly more nodules per pot and nodules per nodulated seedling in soils from this locale than seedlings of twoCasuarina equisetifolia Forst. provenances. Seedlings of two provenances ofAllocasuarina torulosa (Ait.) L. Johnson had fewer than 1% nodulated seedlings, a significantly lower level by far than that ofCasuarina seedlings.A. torulosa provenances also had significantly fewer nodulated seedlings per pot and nodules per nodulated seedling than all Casuarina hosts excepting one poorly-nodulated provenance ofC. equisetifolia.Nodulated seedlings of allCasuarina species had the capacity to fix atmospheric N₂, as indicated by acetylene-reduction capability. The presence of yellow cladodes and low rates of acetylene reduction per plant forC. cristata Miq. suggest that this association was poorly effective.     

Facilitation and Inhibition of Seedlings of an Invasive Tree (Acer platanoides) by Different Tree Species in a Mountain Ecosystem

Facilitation is known to be an important process structuring natural plant communities. However, much less is known about its role in facilitating the invasion of ecosystems by non-native plant species. In this study we evaluated the effects of invasive (Acer platanoides) and native (Pseudotsuga menziesii) forest types on the performance of A. platanoides seedlings, and related these effects to structural and functional properties associated with the two forest types, in a native P. menziesii forest that is being invaded by A. platanoides. Acer platanoidesseedlings had higher densities, recruitment, and survival, and experienced less photoinhibition and water stress when beneath conspecific canopies than in the adjacent P. menziesii forest. Soil moisture and canopy cover were greater in the invaded patch than the native forest. There was no difference in soil fertility or understory light levels between locations. These demographic (i.e. seedling survival), physiological, and environmental differences appeared to be due to the effects of A. platanoides and P. menziesii trees. Thus, Acer trees appear to produce a more mesic environment by modifying the structure and phenology of the forest canopy and by altering the timing of transpirational water loss relative to P. menziesii. Environmental modification by invaders that lead to positive effects on conspecifics may help us to understand the dramatic success and lag periods of some invasive species     

Spatial patterns of tree seedling establishment and their relationship to environmental variables in a cold-temperate deciduous forest of eastern North America

Patterns of seedling recruitment may have persistent effects on population and community processes. Assuming seed availability is not limiting, the environmental sieve (i.e., the suite of factors influencing seed germination and seedling emergence and survival) determines how many seedlings establish and, most importantly, where they do so. In this study, we identify the spatial structure of some resources and abiotic conditions known to be significant for tree seedling emergence and survival and determine how these environmental factors influence the establishment of Fagus grandifolia, Acer saccharum, Fraxinus americana, and Ostrya virginiana in a deciduous forest of southern Québec (Canada). We expect an increase from Fagus, through Acer and Fraxinus, to Ostrya in the control of environmental variables on seedling emergence and survival, because of differences in the seed size of these species. Density of newly-emerged seedlings of all four species showed positive spatial autocorrelation at distances of up to ca. 10 m. Environmental variables were also structured at the same spatial scale, except for soil moisture. Acer seedling emergence pattern was positively correlated to photosynthetic photon flux density (PPFD), and the pattern of Fraxinus to soil N and moisture. Seedling survival was not spatially autocorrelated for any of the four species, although it was positively density-dependent in Acer and Fagus. In only Ostrya was seedling survival correlated (positively) to one of the environmental variables studied, i.e., PPFD. Overall, environmental variables were spatially less heterogeneous than seedling emergence and survival. Either seed availability was not saturating or factors not considered here, such as competition and predation (the intensity of which often varies with resources and/or abiotic conditions), modified the influence that the physical environment had on patterns of seedling establishment. Our prediction of a greater environmental control on seedling emergence and survival in small-seed species was not totally confirmed.     

Inter-relationship between shoot weight,severity of root rot and survival rate of subterranean clover inoculated with certain pathogenic fungi

Summary Plant survival rate, root disease index and fresh shoot weight of subterranean clover seedlings inoculated with fungal pathogens (Fusarium avenaceum, F. oxysporum, Phoma medicaginis, Pythium irregulare andRhizoctonia solani, both singly and in combinations) were generally inter-correlated over a wide range of temperature (10, 15, 20 and 25°C) and moisture conditions (45, 65% water holding capacity and flooding). There was a negative correlation between root disease index and shoot weight for all treatments exceptF. avenaceum + P. irregulare. Root disease index and seedling survival rate were negatively correlated except forF. oxygsporum, Phoma medicaginis, P. irregulare andF. oxysporum + F. avenaceum. However, a good positive correlation was found between the survival rate and shoot weight for all treatments with the exception ofPhoma medicaginis.