An experimental test of limits to tree establishment in Arctic tundra

1 Five treeline species had low seed germination rates and low survivorship and growth of seedlings when transplanted into Alaskan tundra. Seed germination of all species increased with experimental warming, suggesting that the present treeline may in part result from unsuccessful recruitment under cold conditions.
2 Growth, biomass and survivorship of seedlings of treeline species transplanted into tundra were largely unaffected by experimental warming. However, transplanted seedlings of three species ( Betula papyrifera , Picea glauca and Populus tremuloides ) grew more when below‐ground competition with the extant community was reduced. All three measures of transplant performance were greater in shrub tundra than in the less productive tussock or heath tundra. Establishment of trees in tundra may thus be prevented by low resource availability and competition.
3 Two species ( Alnus crispa and Populus balsamifera ) had low seed germination and survivorship of germinated seeds; transplants of these species did not respond to the manipulations and lost biomass following transplanting into tundra. Isolated populations of these two species north of the present treeline in arctic Alaska probably became established during mid‐Holocene warming rather than in recent times.
4 Of all the species studied here, Picea glauca was the most likely to invade intact upland tundra. Its seeds had the highest germination rates and it was the only species whose seedlings survived subsequently. Furthermore, transplanted seedlings of Picea glauca had relatively high survivorship and positive growth in tundra, especially in treatments that increased air temperature or nutrient availability, two factors likely to increase with climate warming.     

Studies in the ecology of coastal heath in New South Wales

Effects of heating dry seed were investigated in Banksia serratifolia and B. ericifolia. Seeds were inviable after 7 min exposure at 150°C or higher temperatures but remained viable after exposure at 100°C and lower temperatures. Imbibed seeds show no dormancy, and seeds survive fire insulated from heat in the thick woody follicles which may persist unopened for several years on the parent plant but open following fire. Effects of nutritional supplements were observed in the seed germination and seedling survival and growth of Banksia serratifolia, B. aspleniifolia and B. ericifolia. Germination with radicle and hypocotyls extension was not significantly affected by a wide range of mineral nutrient regimes. In pot experiments using a heath soil seedling survival and growth were adversely affected by moderate additions of phosphate with surviving seedlings showing toxicity symptoms and poor development of proteoid roots. Moderate supplements of potassium had no adverse effect, and in B. serratifolia and B. ericifolia lessened the adverse effects of phosphate addition. The slow growth of the seedlings accords with the normal low nutrient availability in heath soils but seedlings show little capacity for rapid exploitation of any temporary increase of available nutrients after fire.     

Seed dispersal, seedling establishment and gap partitioning among tropical pioneer trees

1 We examined the abundance and distribution patterns of pioneer seeds in the soil seed bank, and of pioneer seedlings in 53 recently formed gaps, in a 50‐ha forest dynamics plot on Barro Colorado Island (BCI), Panama. The aim was to assess the importance of dispersal limitation (failure of seeds to arrive at all sites suitable for their germination) and establishment limitation (failure of seeds having reached a site to germinate successfully and establish as seedlings) in determining patterns of gap occupancy.
2 The abundance of seeds in the soil seed bank was strongly negatively correlated with seed size, but was not correlated with the abundance of reproductive‐sized adult trees in the plot. In contrast, the abundance of pioneer seedlings > 10 cm height in natural gaps was strongly correlated with adult abundance, but was not correlated with seed size.
3 Seedlings were non‐randomly distributed among gaps, but seedling abundance was not directly related to gap size, and there was no evidence of partitioning of the light environment of gaps by small seedlings. Large differences in growth and mortality rates among species were observed after 1 year, and this may result in the gap size partitioning previously found in saplings of the same species.
4 Seedlings of most species, particularly those with large seeds, were relatively more abundant than expected in gaps close to their conspecific adults. Proximity to reproductives, and by inference dispersal limitation, therefore exerts some effect on seedling distribution. None the less, large differences between seed and seedling abundances for some species, and low seedling occupancy rates in some gaps close to adult conspecifics, suggest that seedling emergence probabilities and species‐specific establishment requirements may also be important determinants of local abundance.     

Demography of a non-sprouting and resprouting Hakea species (Proteaceae) in fire-prone Eucalyptus woodlands of southeastern Australia in relation to stand age,drought and disease

Population size-structures, seed production, canopy seed storage (serotiny), and recruitment were investigated in relation to fire, drought and disease for a pair of co-occurring resprouting and non-sprouting shrub species from the genus Hakea (Proteaceae) in fire-prone Eucalyptus woodlands in western Victoria, Australia. The non-sprouter species, Hakea decurrens, showed faster height growth, higher seed production and higher seed viability than the resprouter, Hakea rostrata. Population size structures in stands up to 24 years since last fire showed no evidence of inter-fire recruitment for either species. Following a fire in 1990 in a mixed species stand 15–20 years old, the estimated number of viable seeds released from canopy-stored seed banks was approximately equal for both species. However, the rate of seedling establishment in the first year was about 10 times higher, and seedling suvivorship over the first 5 years was seven times higher, for the non-sprouter. Seedlings of Hakea decurrens and resprouts of Hakea rostrata began to produce seeds within three years of the last fire, while the few surviving seedlings of Hakea rostrata showed no evidence of reproductive maturity after six years. Inter-fire recruitment was recorded for the non-sprouter, Hakea decurrens, in the oldest stand (burned in 1967) between 24 and 28 years since last fire. This was associated with an increased rate of seed release from serotinous fruits due to the onset of high rates of adult plant mortality. High adult mortality and increased seed release correlated with increasing stand age, the occurrence of severe drought, and the likely presence of Phytophthora cinnamomi, a fungal pathogen which damages the root system, reducing water and nutrient uptake. There were no new recruits for the resprouter Hakea rostrata in this stand, but old plants continued to resprout from basal lignotubers and no mortality was observed. While recruitment of strongly serotinous shrub species is commonly described as being restricted to the immediate post-fire period, the present study illustrates that other events (e.g., senescence, drought, disease) can lead to recruitment of serotinous non-sprouters and may be important in the maintenance of populations during unusually long periods without fire.     

Exudation of carboxylates in Australian Proteaceae: chemical composition

Roots of a wide range of plant species exude carboxylates, such as citrate, into the rhizosphere. In the present study, seedlings of a range of Australian Banksia, Hakea and Dryandra species (Proteaceae) were assayed for their exudation of carboxylates. All of these species (Hakea prostrata, Hakea undulata, Hakea petiolaris, Hakea baxteri, Banksia grandis, Banksia prionotes, Banksia occidentalis and Dryandra sessilis) form cluster roots when grown in nutrient solution with a low phosphate concentration. Exudation of carboxylates was studied for cluster roots and non‐cluster roots separately, and for the entire root system. Cluster roots of these Proteaceae exuded malate, malonate, lactate, acetate, maleate, citrate, fumarate, cis‐ and trans‐aconitate. The relative contributions of each of these carboxylates differed between species. Malate, malonate, lactate, citrate and trans‐aconitate, however, were invariably present in large proportions of total carboxylate exudation. Non‐cluster roots of H. prostrata exuded a spectrum of carboxylates (mainly malonate, lactate and citrate), which differed somewhat from the exudation pattern of cluster roots (mainly malate, malonate, lactate and citrate). The rate of exudation for cluster roots of the seven species was approximately 1·6 nmol g^?1 FM s^?1, which is considerably higher than that reported for a variety of crop and native species that do or do not form cluster roots. Contrary to what occurs in the cluster roots of Lupinus albus, which release carboxylates accompanied by protons so that the rhizosphere is acidified, the present Proteaceae exude the carboxylates as anions without concomitant proton release. The role of carboxylates in the mobilization of phosphate and other nutrients from soil is discussed.     

Seedling growth of four fire-following Mediterranean plant species deprived of single mineral nutrients

1. Anthyllis vulneraria, Cistus creticus, Hippocrepis unisiliquosa and Pinus brutia are frequent post-fire colonizers whose seedlings are subjected to the mineral conditions imposed by ash in the early stages of growth. We test the hypothesis that the effective internal supply of mineral elements in the seeds of these species may complement the external availability of specific nutrients.
2. Newly germinated seedlings were grown in nutrient solutions, each deficient in one of the following elements: nitrogen, phosphorus or potassium. Control treatments consisted of full nutrients and distilled water. Seedling growth was monitored over 12weeks. The final dry masses of the seedlings were taken as a measure of the availability of each element from the seeds' own stored reserves.
3. For the two legume species nitrogen was the most limiting element followed by phosphorus. Potassium deficiency had no effect on the final legume dry biomass. For Cistus, the order was nitrogen followed by potassium, while phosphorus deprivation had no effect on growth during the 12weeks of the experiment. Deprivation of single minerals had no significant effect on seedling growth of Pinus over the same period. The two legumes and Cistus when denied nitrogen had relatively greater relative root ratios than control plants.
4. The relative seedling requirements of each species for early external supplies of the various nutrients for establishment in the field are discussed with reference to the adaptation of the four species to post-fire regeneration.     

Evaluation of Restoration Techniques for the Succulent Karoo, South Africa

Abstract Possible constraints on the passive recovery of bare areas in the Karoo, a semiarid region in South Africa, include inadequate supply of seed, availability of suitable microsites for plant establishment, altered soil properties, and the truncation of key soil biotic processes. Here we investigate the possibility of initiating the restoration of bare areas by soil surface treatments with gypsum (CaSO₄) and/or organic mulch. We also apply an exogenous seed source to test the hypothesis that seed availability limits autogenic recovery. Both gypsum and mulch improved rain water infiltration, gypsum more so than mulch, and both treatments resulted in significantly higher numbers of reseeded seedlings compared with controls. Gypsum also improved the survival of the cohorts of seedlings of the larger seeded Tripteris sinuata. Tripteris showed the highest number of seedlings (maximum count of 150 seedlings/1,000 viable seeds sown) and surviving plants of the three reseeded species, which included two small‐seeded species, Ruschia spinosa and Chaetobromus dregeanus. Throughout the study period significantly higher plant volumes of naturally seeded annuals and perennials were recorded in the gypsum and/or mulch treatments compared with the controls. Germination and emergence of reseeded and naturally seeded plants appears to be determined by the availability of cool season (autumn to spring ) soil moisture, whereas follow‐up rainfall during this time is important for plant survival. Mulching of bare areas in the Succulent Karoo has the potential to re‐create vegetated areas that will further capture and conserve water, soil, and nutrients. Gypsum also showed positive results but might not be a cost‐effective option because of transport costs to these remote arid areas.     

Root:shoot ratio in developing seedlings: How seedlings change their allocation in response to seed mass and ambient nutrient supply

Root:shoot (R:S) biomass partitioning is one of the keys to the plants' ability to compensate for limiting resources in the environment and thus to survive and succeed in competition. In adult plants, it can vary in response to many factors, such as nutrient availability in the soil or reserves in the roots from the previous season. The question remains whether, at the interspecific level, reserves in seeds can affect seedlings' R:S ratio in a similar way. Proper allocation to resource‐acquiring organs is enormously important for seedlings and is likely to determine their survival and further success. Therefore, we investigated the effect of seed mass on seedling R:S biomass partitioning and its interaction with nutrient supply in the substrate. We measured seedling biomass partitioning under two different nutrient treatments after 2, 4, 6, and 12 weeks for seventeen species differing in seed mass and covering. We used phylogenetically informed analysis to determine the independent influence of seed mass on seedling biomass partitioning. We found consistently lower R:S ratios in seedlings with higher seed mass. Expectedly, R:S was also lower with higher substrate nutrient supply, but substrate nutrient supply had a bigger effect on R:S ratio for species with higher seed mass. These findings point to the importance of seed reserves for the usage of soil resources. Generally, R:S ratio decreased over time and, similarly to the effect of substrate nutrients, R:S ratio decreased faster for large‐seeded species. We show that the seed mass determines the allocation patterns into new resource‐acquiring organs during seedling development. Large‐seeded species are more flexible in soil nutrient use. It is likely that faster development of shoots provides large‐seeded species with the key advantage in asymmetric above‐ground competition, and that this could constitute one of the selective factors for optimum seed mass.     

Nutrient availability and limitation in traditionally mown and in abandoned limestone grasslands: a bioassay experiment

Management is necessary for the conservation of limestone grasslands. However, the traditional management of hay-making every year in July is no longer profitable for farmers. Hence many species-rich grasslands have become abandoned. The aim of this study is (a) to investigate the consequences of abandonment (as compared with annual mowing) on the availability of soil nutrients, and (b) to determine the type of nutrient limitation. The soil was taken from a long-term experimental site set up 22 years ago in northern Switzerland. The availability of soil N and P was assessed in a bioassay where soil from mown and unmown plots was supplied with different nutrient solutions or deionised water as control. Seedlings of Galium mollugo s.str. L. and Raphanus sativus ssp. oleiferus (DC) Metzg. were used as phytometer plants. Their growth in the control treatment was limited by N and P almost to the same extent, indicating that the availability of both elements was very low in this soil. P limited plant growth slightly more, but was overcome in the case of Raphanus by a high P content in the seeds. The N and P availability responded differently to management. The availability of N was slightly higher in soil from the abandoned plots, whereas the P availability did not differ significantly. Accumulation of nitrogen in the soil after abandonment did presumably not have any decisive effect on the vegetation because the amount was small and because the vegetation seemed primarily P-limited. The direct effect of mowing or abandonment on plants is therefore likely to be much more important for species composition than the minor changes in soil nutrients.     

Effects of supplementary feeding on growth and reproduction of three carnivorous plant species in a subarctic environment

1 The effect of supplementary feeding on growth and reproduction of three carnivorous plants species was investigated over a 6‐year period. Pinguicula alpina , P. villosa and P. vulgaris populations growing at two altitudes in a subarctic environment were fed with fruit flies ( Drosophila melanogaster ).
2 Fed plants increased in size relative to control plants during the first years. Subsequently a stable size difference between feeding levels was established. The weight of the over‐wintering part was higher in fed plants than in control plants.
3 The flowering frequency (i.e. proportion of plants in a population carrying flowers) was also higher in the fed plants. The proportion of flowering plants increased in the feeding treatment compared to control plants during more‐or‐less the whole experimental feeding period (5–6 years). Seed production also increased slowly in response to feeding.
4 No feeding effect on nitrogen and phosphorus concentrations in seeds and over‐wintering parts was found.
5 In most characteristics the high‐altitude populations were less responsive than populations growing at low altitude. In the high‐altitude population reproduction failed repeatedly, suggesting that seed output may be more dependent on abiotic factors than on resource availability in cold environments.
6 Pinguicula villosa and P. vulgaris used the new resources the same summer for increased leaf rosette growth and current reproduction, showing these species to be income breeders, while the third species ( P. alpina ) allocated a larger proportion to storage and future reproduction, characteristic of a capital breeder.     

Hypericum cumulicola demography in unoccupied and occupied Florida scrub patches with different time‐since‐fire

1 Metapopulation models predict that unoccupied, but suitable, patches will exist for species subject to extinction and colonization dynamics. We compared the demographic responses of Hypericum cumulicola , a rare herbaceous species almost entirely restricted to Florida rosemary scrub, when transplanted to occupied or unoccupied patches.
2 Seedlings were transplanted and seeds buried into Florida rosemary scrub patches differing in time since last fire, and in the presence or absence of H. cumulicola. We used a replicated, factorial design to place the transplants and seeds in the field, and monitored their performance for 18 months.
3 Neither time‐since‐fire nor prior H. cumulicola site occupancy affected survival of transplants. Only time‐since‐fire affected growth. Time‐since‐fire, H. cumulicola occupancy, and their interaction affected reproductive effort, but these effects were not consistent between years.
4 Flowering and seed production led to subsequent seedling recruitment near transplants, mainly in recently burned sites. Genetic screening of transplants and seedlings showed that transplants in occupied sites could have crossed with nearby resident plants, but that offspring in sites previously unoccupied were likely to have been parented only by nearby transplants.
5 Seeds buried, and later exhumed, germinated after 1 or 2 years of burial, demonstrating a persistent soil seed bank from which populations could recover after fire. Neither time‐since‐fire nor H. cumulicola occupancy affected seed dormancy or germination.
6 Similar demography in unoccupied and occupied patches suggests that the patchy pattern of site occupancy by H. cumulicola is probably due to limited dispersal and periodic extinction, especially associated with long fire‐free intervals. Conservation measures need to protect unoccupied patches to allow metapopulation dynamics and persistence.     

Fertilization affects the establishment ability of species differing in seed mass via direct nutrient addition and indirect competition effects

Fertilization causes species loss and species dominance changes in plant communities worldwide. However, it still remains unclear how fertilization acts upon species functional traits, e.g. seed mass. Seed mass is a key trait of the regeneration strategy of plants, which influences a range of processes during the seedling establishment phase. Fertilization may select upon seed mass, either directly by increased nutrient availability or indirectly by increased competition. Since previous research has mainly analyzed the indirect effects of fertilization, we disentangled the direct and indirect effects to examine how nutrient availability and competition influence the seed mass relationships on four key components during seedling establishment: seedling emergence, time of seedling emergence, seedling survival and seedling growth. We conducted a common garden experiment with 22 dry grassland species with a two‐way full factorial design that simulated additional nutrient supply and increased competition. While we found no evidence that fertilization either directly by additional nutrient supply or indirectly by increased competition alters the relationship between seed mass and (time of) seedling emergence, we revealed that large seed mass is beneficial under nutrient‐poor conditions (seedlings have greater chances of survival, particularly in nutrient‐poor soils) as well as under competition (large‐seeded species produced larger seedlings, which suffered less from competition than small‐seeded species). Based on these findings, we argue that both factors, i.e. nutrient availability and competition intensity, ought to be considered to understand how fertilization influences seedling establishment and species composition with respect to seed mass in natural communities. We propose a simple conceptual model, in which seed mass in natural communities is determined by competition intensity and nutrient availability. Here, we hypothesize that seed mass shows a U‐shaped pattern along gradients of soil fertility, which may explain the contrasting soil fertility‐seed mass relationships found in the recent literature.     

Post‐fire seed predation: Does distance to unburnt vegetation matter?

Human‐induced changes to fire regimes result in smaller, more patchy fires in many peri‐urban areas, with a concomitant increase in potential edge effects. In sclerophyll vegetation, many structurally dominant serotinous plants rely on the immediate post‐fire environment for recruitment. However, there is little information about how fire attributes affect seed predation or recruitment for these species. We examined the influence of distance to unburnt vegetation on post‐dispersal seed predation for five serotinous species from sclerophyll vegetation in the Sydney region, south‐eastern Australia; Banksia serrata L.f., Banksia spinulosa Sm. var. spinulosa, Hakea gibbosa (Sm.) Cav., Hakea teretifolia (Salisb.) Britten (all Proteaceae) and Allocasuarina distyla (Vent.) L. Johnson (Casuarinaceae). We used cafeteria trials and differential exclusion of vertebrates and invertebrates to test whether rates of seed removal for these five species differed among (i) unburnt, (ii) burnt‐edge (approx. 10 m from unburnt vegetation) and (iii) burnt‐interior (approx. 100 m from unburnt vegetation) locations. When all animals had access to seeds, seeds were removed at lower rates from burnt‐interior areas than from other locations. Vertebrates (small mammals) showed this pattern markedly the first time the experiment was run, but in a repeat trial this effect disappeared. Rate of seed removal by invertebrates differed among plant species but we did not detect any such differences for removal by vertebrates. Overall rates of seed removal also differed significantly between the two fires studied. Our results indicate that small mammal seed predation can be substantial for large‐seeded serotinous shrubs, and that differences in the perimeter: area ratio, severity or size of a fire are likely to affect seed predation.     

Sources of nutrients to rooted submerged macrophytes growing in a nutrient-rich stream

1. The relative contribution of roots and leaves to nutrient uptake by submerged stream macrophytes was tested in experiments where plants were grown in an outdoor flow-channel system. Water was supplied from a nutrient-rich stream with inorganic nitrogen and phosphorus concentrations typical of Danish streams.
2. Four submerged macrophyte species were tested, Elodea canadensis , Callitriche cophocarpa , Ranunculus aquatilis and Potamogeton crispus, and all species were able to satisfy their demand for mineral nutrients by leaf nutrient uptake alone. This was evident from manipulative experiments showing that removal of the roots had no negative impact on the relative growth rate of the plants. Further, the organic N and P concentrations of the plant tissue was constant with time for the de-rooted plants.
3. Enrichment of water and/or sediment had no effect on the relative growth rate of two species, E. canadensis and C. cophocarpa , indicating that in situ nutrient availability was sufficient to cover the needs for growth. Despite the lack of a response in growth rate, a reduced root/shoot biomass ratio was observed with nutrient enrichment of water and/or sediment, and an increased tissue-P concentration in response to open-water enrichment.
4. The open-water nutrient concentrations of the stream in which the experiments were performed are in the upper part of the range found for Danish farmland streams (the majority of Danish streams). Still, however, the negligible effect of nutrient enrichment on the growth of submerged macrophytes observed suggests that mineral nutrient availability might play a minor role in controlling macrophyte growth in most Danish streams.     

Nutrient limitations and their implications on the effects of atmospheric deposition in coastal dunes; lime‐poor and lime‐rich sites in the Netherlands

1 A survey of plant and soil parameters was carried out in dry dune grasslands along the Dutch coast in the lime‐ and iron‐poor Wadden district and initially lime‐ and iron‐rich Renodunaal district, in order to detect differences in nutrient availability related to soil characteristics and potential sensitivity to atmospheric deposition of nitrogen.
2 Plant biomass and phosphorus pools in the shoot were higher in the Wadden district. The low foliar nitrogen concentrations and nitrogen/phosphorus ratios in the Wadden district suggested nitrogen‐limitation, while in the Renodunaal district there appeared to be a balanced supply of both nitrogen and phosphorus.
3 Soil pH, soil organic matter, soil nitrogen and phosphorus concentrations and total amounts were generally higher in the Renodunaal district. In both districts mineral phosphorus decreased with acidification and phosphorus oxalate (iron and aluminium bound) increased.
4 In the Wadden district iron is present primarily in iron–organic matter complexes, which leads to reversible binding of phosphorus. In the Renodunaal district large amounts of iron (hydr)oxides occur and at high pH may contribute to reversible phosphorus‐sorption, but at low pH this probably leads to immobilization of phosphorus.
5 While pools of soil phosphorus are low in the Wadden district, the phosphorus availability may be relatively high due to the comparatively loose nature of phosphorus‐sorption. As a result the area may be nitrogen‐limited and grass‐encroachment may thus have resulted from atmospheric deposition of nitrogen.
6 In the Renodunaal district, atmospheric deposition probably only accelerates grass‐encroachment, because deposition of acid and nitrogen increases the availability of both nitrogen and phosphorus and maintains the 'co‐limitation'.     

Impact of Douglas-fir and Scots pine seedlings on plagioclase weathering under acidic conditions

The weathering of soil minerals in forest ecosystems increases nutrient availability for the trees. The rate of such weathering and its relative contribution to forest tree nutrition, is a major issue when evaluating present and potential forest stand productivity and sustainability. The current paper examines the weathering rate of plagioclase with and without Douglas-fir or Scots pine seedlings, in a laboratory experiment at pH 3–4 and 25 °C. All nutrients, with the exception of Ca, were supplied in sufficient amounts in a nutrient solution. The objective of the experiment was to evaluate the potential of trees to mobilise Ca from the mineral plagioclase that contained 12% of Ca. Amounts of nutrients supplied in the nutrient solution, amounts accumulated in the living tissue of the seedlings and amounts leached from the experimental vessels, were measured. A weathering balance, accounting for leached + accumulated − supplied amounts, was established. Bio-induced weathering, defined as the weathering increase in the presence of trees, relative to the weathering rate without trees (geochemical weathering; control vessels), under the present experimental conditions, explained on average, 40% of total weathering (biological + geochemical). These conditions appeared more beneficial to Scots pine (higher relative growth rate, higher Ca incorporation) than to Douglas-fir.     

Shifts in seed size across experimental nitrogen enrichment and plant density gradients

We tested the hypotheses that increased soil resource availability selects for larger seeded plants by: (1) increasing community seed density and seedling competition, (2) increasing light competition, which favours larger seeded species if their additional seed resources are allocated to shoots. We also tested the hypothesis that plants respond to increased nutrient availability by producing larger seeds. In a controlled environment experiment, we grew monocultures of five species of co-occurring annual plant species at two N addition rates and measured seed size and the number of seeds produced. In a second experiment, we sowed seed of the five species together, manipulated nitrogen (N) addition rate and community seed density and measured species performance. In the first experiment, elevated N addition increased seed size, but only in larger seeded species. In the second experiment, high N addition increased community seed production. However, contrary to our hypothesis small seeded species were selected under high seed density, possibly because they germinated and grew more rapidly thus enabling them to pre-empt limiting resources. Larger seeded species were favoured by high N addition, as hypothesised. However, our data suggest that this was due to their seed reserves boosting survival below the denser canopies of fertile conditions rather than via competitive advantages. Our results point to a largely overlooked role of recruitment in determining community response to fertilisation. Although our results may not generalise to all plant communities they suggest that seed traits play a role in community response to both the direct effect of N addition and the stimulation of seed density caused by increased productivity. These results are also consistent with the view that the advantage of large seed size is not a competitive one, but greater survival in the face of environmental hazards.     

Small-scale spatial heterogeneity in the vertical distribution of soil nutrients has limited effects on the growth and development of Prosopis glandulosa seedlings

We examined the above- and below-ground responses of seedlings of the woody shrub Prosopis glandulosa to the spatial heterogeneity of soil nutrients within the root zone. We performed a microcosm experiment where seedlings were grown with different combinations of nutrients [nitrogen (N), phosphorus (P), and both combined (NP)] and under different levels of nutrient heterogeneity (nutrients supplied as patches located in the bottom and/or upper portion of rooting zone versus homogeneous distribution). Seedling morphology and biomass did not show a strong response to changes in nutrient ion or spatial heterogeneity. Height, number of leaves, and specific leaf area did not vary significantly between treatments. The number of leaves, foliar biomass, stem biomass and biomass allocation to stems of seedlings showed more responsiveness to the addition of N and NP than to the addition of P. The spatial heterogeneity of nutrients affected the diameter, root biomass and leaf N content. Seedlings had higher diameter and root biomass when the nutrients were homogeneously distributed as compared to their placement as patches in the bottom of the microcosms. Their leaf N concentration increased in those treatments where the nutrient patch was located in the lower half as compared to the upper half of the microcosms. Root foraging responses to nutrient patches varied with their location. Significant root proliferation was observed when patches of N, P and NP were located in the upper portion of the rooting zone; when they were located in the lower portion such a response was observed only for P. Despite our findings that Prosopis seedlings have a low overall responsiveness to small-scale vertical differences in soil nutrient heterogeneity, our results suggest that these differences may modify the growth dynamics of the secondary roots of this ecologically important dryland species during the early stages of its development.     

Savanna soil fertility limits growth but not survival of tropical forest tree seedlings

Background and Aims

Cerradão (Brazilian woodland savannas) and seasonally dry forests (SDF) from southeastern Brazil occur under the same climate but are remarkably distinct in species composition. The objective of this study was to evaluate the role of soil origin in the initial growth and distribution of SDF and Cerradão species.

Methods

We conducted a greenhouse experiment growing Cerradão and SDF tree seedlings over their soil and the soil of the contrasting vegetation type. We evaluated soil nutrient availability and seedling survivorship, growth and leaf functional traits.

Results

Despite the higher nutrient availability in SDF soils, soil origin did not affect seedling survivorship. The three SDF species demonstrated home-soil advantage, enhanced growth with increasing soil nutrient availability and had higher growth rates than Cerradão species, even on Cerradão soils. Growth of Cerradão seedlings was not higher on Cerradão soil and, overall, was not positively correlated with soil nutrient availability.

Conclusions

SDF species are fast-growing species while Cerradão trees tend to be slow-growing species. Although savanna soil reduces growth of forest species, our findings suggest that soil chemical attributes, alone, does not exclude the occurrence of SDF seedlings in Cerradão and vice-versa.     

Nutrient and Carbon Additions to the Microbial Soil Community and its Impact on Tree Seedlings in a Boreal Spruce Forest

We have added glucose and nutrients to manipulate soil microbial activity and nutrient availability in a boreal spruce forest to study the performance of birch and spruce seedlings in relation to the soil microbial community. The proportion of aboveground biomass in the seedlings was largest in plots amended with extra nutrients, while ectomycorrhizal (ECM) colonisation was low in these plots. ECM appeared beneficial for growth of both species, but only at low levels of colonisation (<25% ECM colonised root-tips). The soil microbial biomass, as determined by total PLFA, was largest in plots treated with glucose and there was a significant negative relationship between birch seedling size and levels of total PLFA in soil. This could be taken to suggest that poor seedling growth was due to nutrient limitation caused by microbial assimilation. However, the treatment response of the birch seedlings was generally weak, and spruce often showed no response at all to the addition of nutrients and glucose. The most consistent parameter for the variation in plant performance, as well as for the microbial soil community, was the block-effect. This suggests a strong spatial structure in the soil microbial community, and that this structure was robust with respect to our treatments even though they continued over a 3-year period.