Dynamics of resprouting in the lignotuberous shrub Banksia oblongifolia

The resprouting response of different sized Banksia oblongifolia lignotubers (genets) was followed in two field experiments. In the first, the density and speed of resprouting, and the growth in length of the leading shoot from each lignotuber in response to fire and to the time elapsed since the last fire was monitored for 18 months after fire and clipping treatments. In the second, sizes of bud banks were estimated by repeatedly clipping new shoots from individual lignotubers. Density of resprouting (shoots dm^?2 lignotuber) decreased with increasing lignotuber size, and the length of the leading shoot increased. The direct effect of fire was to reduce shoot density by about 75%. The speed of resprouting (time taken by a cohort of shoots to reach 50% of their peak density) was similar after fire and clipping, but leading shoots grew significantly longer after fire. The elapsed time since lignotubers were last burnt did not influence their density of resprouting, but it did influence the speed of resprouting. Shoots from clipped lignotubers that had burnt 3 years earlier took about 90 days to each 50% of their peak density while shoots on lignotubers last burnt 5 and 17 years earlier took about 40 days. Death of shoots was unrelated to crowding in any stand. More lignotubers from the oldest unburnt stand were grazed by herbivores. The number of buds converted into shoots after successive clippings was surprisingly small; for most lignotubers this reserve was less than three times the size of their standing crop of shoots. In general, the smaller lignotubers carried a higher proportion of dormant buds in relation to their standing crop of shoots. About 30% of buds remained dormant after the first clipping and about 10% after the second and third clippings. Evidence suggests that buds are replaced within 6 months of fire. No lignotubers survived four clippings over 15 months.     

Persistence of resprouting species after fire in natural and post‐mine restored shrublands in southwestern Australia

Questions: Is post‐fire persistence of resprouting species lower in restored sites, and is survival related to lignotuber size? Location: Southwestern Australia, Eneabba, 300 km north of Perth. Methods: Post‐fire persistence of 10 lignotuberous shrub species was compared between three sites restored 8–24 years ago after mineral‐sand mining and three surrounding natural shrubland sites (8–24 years since previous fire). Results: Overall persistence of species was 11–93% in restored sites (mean 52%) and 79–100% in natural sites (mean 96%). Persistence increased with time since rehabilitation for five species with <25% of individuals in three species surviving in the youngest stand. For equivalent crown size, average lignotuber circumferences were 50% smaller at restored sites and this probably accounted for their higher post‐fire mortality. Apart from differences in the age of plants, restored sites had lower soil penetrability than natural sites, which may have restricted rootstock development. A tradeoff favoring a higher crown volume to lignotuber size ratio was apparent in nine of the ten species with greater crown volumes (by 37%) and smaller lignotubers (by 36%) in restored sites. Two resprouting species for which crown seed store was quantified had much higher fecundity in restored sites. Conclusions: Fires reduced resprouter persistence in restored sites owing to poor development/insufficient size of lignotubers. Further management after fires is required, including application of resprouter seeds/seedlings on restored topsoil, transplanting adult resprouters (where viable) from natural areas ahead of the mining front. Low intensity/patchy fires are recommended on long unburnt sites. Resprouter survival would have likely been much greater in the first place if a deeper sandy soil profile was rehabilitated, thereby providing a more suitable medium for lignotuber development.     

Comparative physiology of burned and unburned Rhus laurina after chaparral wildfire

Summary Laurel Sumac (Rhus laurina) is a dominant member of the coastal chaparral community of southern California that survives periodic burning by wildfires by resprouting from a lignotuber (root crown). We investigated the physiological basis for resprouting by comparing shoot elongation, leaf nitrogen content, tissue water status, leaf conductance to water vapor diffusion, and photosynthetic rates of post-fire R. laurina to those of adjacent unburned shrubs. Resprouts had higher rates of shoot elongation, leaf conductance, and photosynthesis than mature, unburned shrubs. Leaf nitrogen contents were elevated in burned shrubs even though their leaves developed interveinal chlorosis. A comparison of soil water potential to predawn water potential indicated that roots of R. laurina remain active below 2 m during the first summer drought after wildfire. Our results support the hypothesis that lignotubers not only contain dormant buds that develop into aerial shoots after wildfire but they also supply nutrient resources that enhance shoot elongation. Because R. laurina is relatively sensitive to drought, yet very successful in its rapid recovery after fire, maintaining an active root system after shoot removal may be the primary function of the massive lignotuber formed by this species.     

Continuous basal sprouting from a lignotuber:Arbutus unedo L. andErica arborea L., as woody Mediterranean examples

Summary The two dominant species of the Corsican mattoral,Arbutus unedo L. andErica arborea L., can produce abundant sprouts from the lignotuber not only immediately after fire but also more or less continuously in the absence of major disturbance. The lignotuber appears to be more important during the early stages of development; the result is an increase in the number of sprouts during the 25 years following the establishment of the individuals. Later the lignotuber seems to lose the ability to ensure the development of new basal sprouts. A hypothesis is that the presence of a lignotuber is related to the growth form.Arbutus unedo andErica arborea show behaviour intermediate between acrotony and basitony, as the shoots show acrotony, and continuous sprouting is characteristic of basitonic species. The fact that sprouting from the lignotuber is not necessarily a result of fire suggests that the relation between fire and vegetation in the Mediterranean region should be reconsidered.     

Hydraulic flow characteristics in the lignotuberous mallee Eucalyptus behriana F. Muell. in the field

Abstract The lignotuberous mallee Eucalyptus behriana F. Muell, had much lower predawn leaf water potentials (not higher than – 1.2MPa) than other eucalypts (as high as – 0.2MPa), even after extended rain. This led to the expectation that the lignotuber of E. behriana might have specific hydraulic characteristics. Keeping the soil around partially defoliated mallces for several days underwater did not raise the water status above the maximum leaf water potential observed under natural conditions. Digging a plant out and placing its roots in water after removal of the soil rapidly increased the water status to a level consistant with other eucalypts. This indicated that the major impedance to water uptake was a component of the soil rather than in the roots or in the lignotuber. Some of the individual mallces had only two major stems or branches. One stem or branch was kept covered throughout the experiments to prevent transpiration. The other stem was subjected to a variety of different conditions in order to modify water loss from it. The transpiring branch affected the water status of the non-transpiring plant parts. Hydraulic resistances in the shoot and root/lignotuber were determined from differences in the leaf water potential of covered and uncovered branches, at high water flow rates through the plant. Resistances in branches, including the liquid phase component of the leaf, were significantly larger than in root or lignotuber. The total plant hydraulic resistance of E. behriana was similar to that of other eucalypts, such as E. pauciflora Sieb. ex Spreng. or E. delegatensis R. T. Bak., even though its growth form was different and its natural leaf water potentials were much lower. An osmotic adjustment at the leaf level was observed in the mallee, keeping its bulk leaf turgor in the same range as compared to the other eucalypt species.     

Lignotuber size of Erica australis and its relationship with soil resources

Abstract. It has been proposed that resprouting after a disturbance would be favoured under conditions of low soil resource availability. In lignotuberous plants of Mediterranean‐type areas, successful resprouting after disturbances such as fire depends on the size of the lignotuber, but little is known about the role of soil resources in determining the relative size of this organ. In this work we tested whether the relative size of the lignotuber in the resprouting shrub Erica australis is related to the availability of soil resources. At each of 13 different sites in Spain, 10 plants were chosen and the sizes of their various parts (above‐ and below‐ground) measured. Additionally, at each site we evaluated soil fertility, foliar N and P concentrations and plant water potential in the middle of the summer. The relationships between lignotuber dimensions and the various plant parts were assessed. In all cases, significant differences between sites were found indicating consistent differences in lignotuber size across the range of plant sizes. This was particularly the case when lignotuber dimensions were expressed as a function of foliar biomass. Lignotuber dimensions relative to foliar biomass were positively correlated with soil pH and negatively with plant water potentials in midsummer. There were, however, no clear relationships between lignotuber relative dimensions and soil N and P contents or other measures of site productivity. In summary, lignotubers were not smaller at more productive sites, in fact they were relatively larger at sites where soils were less acidic, but where plant water conditions were less favourable. These findings contradict predictions made based on current theory on the role of soil fertility allocation to resprouting.     

Phosphorus accumulation by lignotubers of jarrah (Eucalyptus marginata Donn ex Sm.) seedlings grown in a range of soils

Summary Jarrah seedlings were grown in six virgin Western Australian soils for up to 27 months. Lignotubers were produced in all soils and formed 10–16% of plant dry weight. The phosphorus concentration in the lignotuber (250–800 g g^–1) was nearly twice that in the stem and roots. The lignotuber contained 10–30% of total plant phosphorus and like the leaves was a sink for phosphorus. In one lateritic soil the phosphorus concentrations of lignotuber and stem barks were similar. However, in the same plants the concentration of phosphorus in the lignotuber wood was five times the phosphorus concentration in stem wood. Hence both lignotuber bark and sap wood in young jarrah seedlings are storage sites for phosphorus. X-ray probe analysis showed that wood phosphorus was associated with the ray parenchyma. Unlike phosphorus, nitrogen did not accumulate in the lignotuber and the concentrations of nitrogen were similar for roots, lignotubers and stems.     

Mathematical inference of the underground clonal growth of Epipactis helleborine (L.) Crantz (Orchidaceae, Neottieae)

Epipactis helleborine (L.) Crantz (Orchidaceae, Neottieae) can spread by sexual or vegetative propagation. The choice of strategy likely depends on the environmental conditions. The rhizome is the organ of vegetative reproduction; hence, it is crucial to understand its development. Unfortunately, it is hardly possible to investigate rhizome morphology directly, since E. helleborine is a protected species in most European countries. The goal of our investigation was to infer the growth patterns of underground parts of an orchid population from long-term annual observations of its aboveground shoots. We implemented the Minimum Spanning Tree method to determine a likely set of underground connections between shoots and to simulate the annual growth of new rhizomes. Furthermore, we modelled the spatial distribution of shoots with a density kernel estimator to compare the density gradients with the direction of growth of the rhizomes. Observed shoot numbers fluctuated between 72 and 183 from year to year. Our results suggest that (1) vegetative reproduction prevails in the studied population, (2) the population consists of about a dozen clones with a diameter of up to 6 m, (3) rhizomes produce up to five new shoots at one branch end per year, (4) rhizomes develop in the direction of decreasing population density, and (5) nodes of rhizomes may produce new offshoots after up to 7 years of dormancy.     

川西北高寒沙区切断根茎对赖草和沙生苔草克隆生长的影响

为了研究高原亚寒带沙化生境中切断根茎对克隆植物基株扩展能力和分株定居能力的影响,在川西北若尔盖高原沙化区内,对根茎禾草赖草和沙生苔草进行了以切断根茎为处理的野外实验。结果表明,赖草和沙生苔草基株的幼小部分(观测单元)地上枝总长度增量、主根茎长度增量和根茎总长度增量显著减少,而对根茎数增量、主根茎节增量和根茎节总数增量影响不显著;赖草观测单元地上枝数增量显著减少,而对沙生苔草地上枝数增量无影响;赖草地上枝与根茎的相关性质发生逆转。这表明．在高原亚寒带半湿润沙化生境中．克隆整合效应显著促进基株幼小部分地上枝和根茎的伸长生长,但对新生根茎的产生和根茎节分化没有影响;切断根茎处理导致赖草、沙生苔草生殖生长与营养生长间竞争加剧,同时使赖草地上部分与地下部分间竞争加剧;观测单元在缺少与基株(或上级株系)的克隆整合作用时,赖草受到的影响大于沙生苔草。     

Quantifying above- and below-ground growth responses of the western Australian oil mallee,Eucalyptus kochii subsp. plenissima,to contrasting decapitation regimes

Resprouting in the oil mallee, Eucalyptus kochii Maiden & Blakely subsp. plenissima Gardner (Brooker), involves generation of new shoots from preformed meristematic foci on the lignotuber. Numbers of such foci escalated from 200 per lignotuber in trees aged 1 year to 3,000 on 4- to 5-year-old trees. Removal of shoot biomass by decapitation 5 cm above ground in summer (February) or spring (October) resulted in initiation of 140-170 new shoots, but approx. 400 shoots were induced to form if crops of new shoots were successively removed until sprouting ceased and rootstocks senesced. Initially, the new shoot biomass of regenerating coppices increased slowly and the root biomass failed to increase appreciably until 1.7-2.5 years after cutting. Newly cut trees showed loss of fine root biomass, and structural roots failed to secondarily thicken to the extent shown by uncut trees. After 2 years, the biomass of shoots of coppiced plants was only one-third that of uncut control trees and shoot:root dry mass ratios of coppiced plants were still low (1.5-2.0) compared with those of the controls (average ratio of 3.1). Spring cutting promoted quicker and greater biomass recovery than summer cutting. Starch in below-ground biomass fell quickly following decapitation and remained low for a 12-18 month period. Utilization of starch reserves in naturally regenerating coppices was estimated to provide only a small proportion of the dry matter accumulated in new shoots. Results are discussed in relation to their impact on coppicing ability of the species under natural conditions or when successively coppiced for shoot biomass production.     

伊乐藻和黑藻断枝根和芽的发生及生长研究

为了解外来种伊乐藻的无性繁殖力、评价其生态安全性,采用插植方式比较研究了伊乐藻(Elodea nuttallii)和本土种黑藻(Hydrilla verticillata)两种沉水植物不同节数(1至4节)和不同节位断枝的不定根和新芽的发生及生长情况。通过室内4周的3次平行实验,结果表明:两者顶芽段均有形成不定根继而形成新植株的能力,而顶芽以下茎段只有本身具有腋芽的断枝才有形成新芽和不定根的能力。两者具相同节数的断枝形成不定根的百分率及根、芽长度,以具顶芽断枝的均明显高于不具顶芽断枝的,具顶芽四节断枝的不定根生成率最高达到90%以上。不具顶芽断枝形成新芽和不定根的百分率及长度随着断枝节数的增加均呈显著递增趋势,每类断枝的发芽率显著大于其生根率;伊乐藻和黑藻枝条一般分别每7节和5节具有一个腋芽,只有具腋芽断枝才能存活,因此,对不具顶芽断枝,7节和5节分别是其形成新苗所需的最短断枝长度。根和芽的长度随节位的下降大致呈递增的趋势。但是节数对形成根、芽的影响显著大于节位的影响。具顶芽断枝的顶芽的增长量和具顶芽4节断枝的生物量增量伊乐藻的高于黑藻,其余指标伊乐藻均显著低于黑藻。伊乐藻断枝的繁殖力总体上低于黑藻。       

Mortality of resprouting chaparral shrubs after a fire and during a record drought: physiological mechanisms and demographic consequences

We examined postfire regeneration of chaparral shrubs during an intense drought. This study focused on the demography and physiology of shrub species that resprout from a basal lignotuber following fire. We found significant levels of resprout mortality when intense drought occurred in the year following fire during the period of shrub recovery. Three of the seven sampled resprouting species had the greatest or near greatest levels of mortality ever recorded when compared to previous studies. Most shrub mortality occurred during the drought after individuals had resprouted (i.e. individuals survived fire, resprouted and then subsequently died). Physiological measurements of species with high mortality suggested that resprout stems were highly embolized and xylem hydraulic conductivities were close to zero during the peak of the drought. In addition, lignotubers of two of the three species experiencing high mortality were depleted of starch. Population densities of most shrub species declined after the drought compared with their prefire levels, with the exception of one drought tolerant obligate seeding species. Resprouting shrub species may deplete their carbohydrate reserves during the resprouting process, making them particularly vulnerable to drought because of the need to transpire water to acquire the CO₂ that is used to supply energy to a large respiring root system. Drought appears to interact with fire by altering postfire shrub recovery and altering species abundances and composition of chaparral communities.     

Root starch storage and allocation patterns in seeder and resprouter seedlings of two Cape Erica (Ericaceae) species

Post-fire sprouting of dormant buds in resprouter plants is facilitated by stored carbohydrate reserves, with starch being the critical reserve. Starch is mainly stored in xylem parenchyma ray tissue of woody underground organs, such as burls, lignotubers, and roots. We carried out a comparative analysis of the pattern of starch storage and the proportion of parenchymatic ray tissue in the upper root or cotyledonary region of seedlings from seeder and resprouter forms within two Cape Erica (Ericaceae) species: E. coccinea L. and E. calycina L., which were raised in the greenhouse under controlled irrigation. We also explored the root-to-shoot allocation patterns of seeder and resprouter seedlings in these two species. Resprouter seedlings of both species showed higher relative amounts of upper-root starch and upper-root storage tissue as well as a higher root-to-shoot allocation than their seeder counterparts. Pronounced swelling of the upper root region suggests ontogenetic development of a lignotuber in the resprouter forms of the two Erica species. The distinct allocation of starch in roots seems to be genetically determined and would account for the apparent differences in the root-to-shoot allocation patterns between both regeneration forms from the early seedling stage.     

Rhizome development ofPhragmites australis in a reed community

Variations in the height, shoot density, biomasses of above- and below-ground parts and rhizome distributions ofPhragmites australis were investigated along a line-transect in a reed community at Yufutsu Mire, Hokkaido. Relationships of performance of the reed plants to soil conditions and species compositions were also examined. Three types of rhizome development were recognized in reed plants; (1) the central part of the reed community, characterized by well developed rhizomes and dense aerial shoots, (2) the intermediate part, characterized by development of rhizomes along both the peat and surface layers and very dense aerial shoots, and (3) the marginal part, characterized by development of rhizomes only along the peat layer and sparse aerial shoots. Observation showed that rhizomes in the surface layer actively produced aerial shoots, whereas rhizomes in the peat layer contributed to the spreading of their distribution range. With the growth of rhizomes, organic debris originating from dead rhizomes and roots accumulated in the mineral soil to promote organic soil formation. In dense parts of the reed stand, species composition was poor because of the shading and litter accumulation by reed plants. The effects of microtopography and water level on the establishment of reed seedlings were also considered.     

Biology of the bamboo Chusquea culeou (Poaceae: Bambusoideae) in southern Argentina

Over a period of 7 years the biology and phenotypic variability of Chusquea culeou were studied at 5 locations in cool temperate forests of southern Argentina. Excavated rhizomes had an average of 1.1 successful rhizome buds, and an average of 2.1 years elapsed between successive generations of rhizomes. Rhizome buds usually develop within the first four years after a rhizome forms. Height, volume and weight of a culm can be calculated from its diameter 1 m above the ground. Culm size, length of foliage leaf blades, and pattern of secondary branching differed among study sites. Dead culms were numerous and commonly remained erect for more than 7 years after dying. New culm shoots appear in spring and reach full size within a few months. Shoots can grow more than 9 cm/day. Less than half of the shoots survived a year; most were killed by moth larvae. Multiple primary branch buds emerge through the culm leaf sheaths in the second spring. The mean number of branch buds at mid-culm nodes varied between 34.8 and 81.5, and the mean number of primary branches was between 22.8 and 40.8. Number and length of branches, and number and length of foliage leaf blades at each node is related to the position of the node on a culm. Most branches grow about 3 cm and produce 1 to 3 foliage leaves annually. Foliage leaf blades generally live 2 years or more; few survive 6 years. Relative lengths of foliage leaf blades and their spacing along a branch permit recognition of annual cohorts.Both gregarious and sporadic flowering have been reported, and every year a few isolated plants flower and die. Length of the life cycle is unknown. Seedlings require up to 15 years to produce culms of mature size. Foliage branches may live more than 23 years, and culms may survive 33 years. Extensive loss of new shoots to predation suggests that gregarious flowering may be driven by a need to escape parasitism. C. culeou clumps expand slowly. Average annual rate of increase of the number of live culms in a clump was 4.6%. Methods of seed dispersal are undocumented. A dense stand of Chusquea culeou had an estimated phytomass of 179 tons/hectare (dry weight), 28% of which was underground. Net annual production was about 16 t/ha dry weight.     

Floral determination in adventitious shoot meristems regenerated from leaves of day-neutral tobacco

The developmental patterns of adventitious shoots regenerated from cultured leaf tissue and the petioles of leaf cuttings of flowering day-neutral tobacco plants were assessed. Leaf tissue, removed from leaves within the inflorescence, was cultured on hormone-free medium and produced a limited number of adventitious shoots. These shoots produced the same number of nodes that would be produced by a seedling meristem. Most leaf-cutting derived shoots produced the same number of nodes that would be produced by a seedling meristem prior to flowering, while a few shoots produced very few nodes and flowered. Some plants appeared to be intermediate in response. Results indicated that the developmental state or the developmental response of all petiole cells is not the same. The influence of position and the potential reversibility in developmental fate of cell populations within leaves is discussed in the context of existing theory.     

Sprouting in temperate trees: A morphological and ecological review

Sprouting in trees, which results in the production of secondary trunks, is an induced response to injury or to a dramatic change in surrounding environmental conditions. This article reviews the forestry and ecology literature to produce an integrated view of the role of sprouting in both disturbed habitats and closed-canopy forests. Sprouting is a universal attribute of temperate angiosperm trees through the sapling stage of development but is much less common among gymnosperms. Four basic types of sprout morphologies are described: collar sprouts from the base of the trunk, sprouts from specialized underground stems (lignotubers and rhizomes), sprouts from roots, and opportunistic sprouts from layered branches. In a survey of 68 species of trees native to northeastern North America, 41% were found to retain the ability to sprout from the collar into adulthood; 26% sprout from branch layers under natural conditions; and 25% have the capacity to form root suckers. Sprouting in seedlings promotes their survival under a variety of stressful conditions, including suppression by canopy trees, herbivory, site exposure, and desiccation. In contrast, sprouting in mature trees extends the life span of the individual following damage and, in the case of root-suckering species, promotes the colonization of new ground. Although the sprouting of mature trees is more conspicuous than the sprouting of seedlings, its ecological significance is not as great. As a broad generalization, species that grow in stressful sites or sites with frequent disturbances are likely to sprout more vigorously and to retain the sprouting ability longer than are species that grow in less stressful sites or sites with less frequent disturbance. Near the limits of a species’ altitudinal or latitudinal range, the production of basal sprouts, root suckers, rhizomes, and/or branch layers allows trees to spread into adjacent areas, thereby circumventing the difficulties associated with seedling establishment.     

Dietary use of wild plant resources in the Sikkim Himalaya, India

The edible wild plants are greatly valued throughout the Himalayan region and serve as an important source of food for indigenous communities. This paper describes the botanical richness, elevational distribution and dietary use of the edible wild plant resources from the Sikkim Himalaya (Eastern Himalaya), many with promising potential. A total of 190 wild plant species have been screened from the Sikkim Himalaya, this derived from 143 genera and 78 families and accounting for nearly 15% of total edible wild plants resources of India. Of the total, 65% were edible for their fruits, 22% for leaves/shoots, 7% for flowers and 3% for roots/ rhizomes. Nearly 91 wild edible species were recorded from low-hills, 70 from mid-hills and 28 species from high-hill areas. Within Sikkim state, the North and East districts represent maximum diversity of edible wild plants due to the wilderness and inaccessibility to most of the habitats. An average rural family annually consumes nearly 8 types of edible wild plants, and a few species provide over five meals in a season. Selected plants also form a source of earning to a few families that sell them in local markets. It is suggested that the high diversity of edible plants needs to be conserved for future use. Some species may be grown in traditional agroforestry systems and on marginal lands of otherwise low agricultural value. Such measures may help protect wild plant resources in their natural habitats.     

Effects of Severing Rhizome on Clonal Growth in Rhizomatous Grass Species Psammochloa villosa and Leymus secalinus

A field experiment was conducted in Inner Mongolian dune to investigate the contribution of clonal integration to genet expansion and ramet establishment in clonal plants in semi-arid inland dune. In the experiment, the developing parts (plant observing unit) of two rhizomatous grass species Psammochloa villosa (Trin.) Bor and Leymus secalinus (Georgi.) Tzvel. were subjected to the treatment of severing rhizomes. The 7 characters relevant to the elonal growth of the plant observing units were investigated such as total length of shoots, number of shoots, number of rhizomes, length of main rhizome, total length of rhizomes, number of main rhizome nodes and total number of rhizome nodes. The results showed that the increments in those plant characters were reduced by the treatment in P. villosa, while no effect of the treatment was observed in L. secalinus. The results suggest an interspecific difference in clonal integration and in its contribution to clonal growth. In P. villosa, clonal integration and clonal growth offer great contributions to the survival of the plants in the sandy grassland characterized where water shortage, strong transevaporation, infertility, habitat patchiness and frequent disturbance were most characteristic.     

Root biomass,root distribution and the fine-root growth dynamics of Quercus coccifera L. in the garrigue of southern France

Quercus coccifera L., the characteristic scrub oak of the garrigue, covers more than 100,000 ha in southern France alone. Precipitation in this area averages 900 mm/year and summer rains are not rare. A total belowground biomass of 7.2 kg/m², including rhizomes and lignotubers, was harvested. Roots were concentrated in the uppermost 50 cm of the soil. It was hypothesized that low winter temperatures inhibit active fine-root growth. This hypothesis was tested by means of fine-root extractions of soil samples from 0–50 cm depth from November 1987 to June 1988. Although the fine-root analysis could not be extended into late summer and fall, the data supported the hypothesis. Ratios of live/dead fine roots reached their minimum at 0.2–0.3 from December to April. They increased to 1.0–1.2 during late spring and early summer. Initiation of fine-root growth in early April was synchronous with bud break. Starch contents of roots, rhizomes, and lignotubers fluctuated from 4.3% in January to 8.3% in April. The starch stored in belowground organs of Q. coccifera in a closed canopy stand amounted to about 500 g/m² in April. This amount declined to 400 g with bud burst and fine-root growth initiation.