Conspecific versus heterospecific litter effects on seedling establishment

Plant litter is an important determinant of seed germination and seedling establishment. Positive effects of litter have received considerable attention, but few studies have explicitly tested whether seedlings are more facilitated by conspecific litter compared to heterospecific litter. In order to contrast conspecific and heterospecific facilitative effects on seedling establishment, we used Anthriscus sylvestris, Angelica sylvestris, Pimpinella saxifraga and different combinations of their seeds and litter seedbeds as a model system. Although litter had a significant species-specific effect on seedling emergence, we found no evidence of strictly conspecific facilitation. Anthriscus sylvestris displayed a positive response to all types of litter. In contrast, there was a clear negative effect of conspecific litter in Pimpinella saxifraga. Activated carbon did not modify the negative effect, indicating that chemical compounds were not the cause. Our study suggests a high level of idiosyncrasy in response to litter at the species level.     

Changing leaf litter feedbacks on plant production across contrasting sub-arctic peatland species and growth forms

Plant species and growth forms differ widely in litter chemistry, which affects decay and may have important consequences for plant growth via e.g. the release of nutrients and growth-inhibitory compounds. We investigated the overall short-term (9.5 months) and medium-term (21.5 months) feedback effects of leaf litter quality and quantity on plant production, and tested whether growth forms can be used to generalise differences among litter species. Leaf litter effects of 21 sub-arctic vascular peatland species on Poa alpina test plants changed clearly with time. Across all growth forms, litter initially reduced plant biomass compared with untreated plants, particularly litters with a high decomposition rate or low initial lignin/P ratio. In the second year, however, litter effects were neutral or positive, and related to initial litter N concentration (positive), C/N, polyphenol/N and polyphenol/P ratios (all negative), but not to decomposability. Differences in effect size among several litter species were large, while differences in response to increasing litter quantities were not significant or of similar magnitude to differences in response to three contrasting litter species. Growth forms did not differ in initial litter effects, but second-year plant production showed a trend (P < 0.10) for differences in response to litters of different growth forms: evergreen shrubs < graminoids or deciduous shrubs < forbs. While long-persisting negative litter effects were predominant across all growth forms, our data indicate that even within nutrient-constrained ecosystems such as northern peatlands, vascular plant species, and possibly growth forms, differ in litter feedbacks to plant growth. Differences in the composition of undisturbed plant communities or species shifts induced by external disturbance, such as climate change, may therefore feedback strongly to plant biomass production and probably nutrient cycling rates in northern peatlands. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Negative and positive interactions among plants: effects of competitors and litter on seedling emergence and growth of forest and grassland species

Living plant neighbours, but also their dead aboveground remains (i.e. litter), may individually exert negative or positive effects on plant recruitment. Although living plants and litter co‐occur in most ecosystems, few studies have addressed their combined effects, and conclusions are ambivalent. Therefore, we examined the response in terms of seedling emergence and growth of herbaceous grassland and forest species to different litter types and amounts and the presence of competitors. We conducted a pot experiment testing the effects of litter type (grass, oak), litter amount (low, medium, high) and interspecific competition (presence or absence of four Festuca arundinacea individuals) on seedling emergence and biomass of four congeneric pairs of hemicryptophytes from two habitat types (woodland, grassland). Interactions between litter and competition were weak. Litter presence increased competitor biomass. It also had positive effects on seedling emergence at low litter amounts and negative effects at high litter amounts, while competition had no effect on seedling emergence. Seedling biomass was negatively affected by the presence of competitors, and this effect was stronger in combination with high amounts of litter. Litter affected seedling emergence while competition determined the biomass of the emerged individuals, both affecting early stages of seedling recruitment. High litter accumulation also reduced seedling biomass, but this effect seemed to be additive to competitor effects. This suggests that live and dead plant mass can affect species recruitment in natural systems, but the mechanisms by which they operate and their timing differ.     

Response of Litter Decomposition to Simulated N Deposition in Disturbed, Rehabilitated and Mature Forests in Subtropical China

The response of decomposition of litter for the dominant tree species in disturbed (pine), rehabilitated (pine and broadleaf mixed) and mature (monsoon evergreen broadleaf) forests in subtropical China to simulated N deposition was studied to address the following hypothesis: (1) litter decomposition is faster in mature forest (high soil N availability) than in rehabilitated/disturbed forests (low soil N availability); (2) litter decomposition is stimulated by N addition in rehabilitated and disturbed forests due to their low soil N availability; (3) N addition has little effect on litter decomposition in mature forest due to its high soil N availability. The litterbag method (a total of 2880 litterbags) and N treatments: Control-no N addition, Low-N: −5 g N m⁻² y⁻¹, Medium-N: −10 g N m⁻² y⁻¹, and High-N: −15 g N m⁻² y⁻¹, were employed to evaluate decomposition_. Results indicated that mature forest, which has likely been N saturated due to both long-term high N deposition in the region and the age of the ecosystem, had the highest litter decomposition rate, and exhibited no significant positive and even some negative response to nitrogen additions. However, both disturbed and rehabilitated forests, which are still N limited due to previous land use history, exhibited slower litter decomposition rates with significant positive effects from nitrogen additions. These results suggest that litter decomposition and its responses to N addition in subtropical forests of China vary depending on the nitrogen status of the ecosystem.     

Effects of kelp (Macrocystis integrifolia) on soil chemical properties and crop response

In 1981 a two-year field plot experiment was established to assess the effects of quantities (0, 7.5, 15, 30, 60 and 120 t ha⁻¹) of fresh kelp (Macrocystis integrifolia) on crop growth and nutritional response and chemical properties of a fine-textured soil. Soil was analyzed for NO₃−N, NH₄−N, electrical conductivity, pH, Cl and exchangeable cations (K, Mg, Ca, Mn and Na). The plots were planted to beans (Phaseolus vulgaris) in the first year and peas (Pisum sativum) in the second year. Marketable bean yields increased in the first year with kelp applications up to 60 t ha⁻¹, with yields, emergence and flowering being reduced by the 120 t ha⁻¹ application. Soluble salts (EC) and Cl concentrations in the soil eight days after application increased linearly and sharply with increasing quantities of kelp. Increased K concentration and moisture content, characteristics of plants growing in a salt-stressed soil environment, were measured. A subsequent companion greenhouse experiment confirmed that the reduced bean emergence and growth with 120 t ha⁻¹ applications of kelp were primarily due to soluble salts. The only growth effects upon peas in the second year was a slight reduction in leaf plus stem yields with increasing applications of kelp.     

Leaf litter of <Emphasis Type="Italic">Kageneckia angustifolia</Emphasis> D. Don (Rosaceae) inhibits seed germination in sclerophyllous montane woodlands of central Chile

Leaf litter accumulation can have either positive, negative or neutral effects on seed germination and seedling recruitment. In montane woodlands of the Mediterranean zone of central Chile, large amounts of leaf litter accumulate beneath the crowns of the summer semi-deciduous tree Kageneckia angustifolia and no regeneration of this or other plant species has been observed beneath this tree throughout the year. In a sample plot of 5000 m² we selected ten K. angustifolia trees and measured (1) leaf litter accumulation beneath and outside canopy; (2) the effects of time elapsed since burial on viability of K. angustifolia seeds with and without a leaf litter cover; (3) field seed germination with presence or absence of leaf litter and (4) the possible chemical effects of K. angustifolia leaf litter leachates on seed germination of its own seeds and of other two co-occurring native shrubs species (Guindilia trinervis and Solanum ligustrinum). Our results show that a considerable accumulation of leaf litter occurred beneath K. angustifolia, and litter negatively affected seed viability and germination of this species in the field. Under laboratory conditions, K. angustifolia leaf litter leachates inhibited seed germination of its own seeds and of the two native shrub species. Chemical effects are likely involved in the negative effects of leaf litter on the recruitment of K. angustifolia in the montane sclerophyllous woodland of central Chile.     

Response of nutrient dynamics of decomposing pine (Pinus massoniana) needles to simulated N deposition in a disturbed and a rehabilitated forest in tropical China

The effects of simulated N deposition on changes in mass, C, N and P of decomposing pine (Pinus massoniana) needles in a disturbed and a rehabilitated forest in tropical China were studied during a 24-month period. The objective of the study was to test the hypothesis that litter decomposition in a disturbed forest is more sensitive to N deposition rate than litter decomposition in a rehabilitated forest due to the relatively low nutrient status in the former as a result of constant human disturbance (harvesting understory and litter). The litterbag method and N treatments (control, no N addition; low-N, 5 g N m⁻² year⁻¹; medium-N, 10 g N m⁻² year⁻¹) were employed to evaluate decomposition. The results revealed that N addition increased (positive effect) mass loss rate and C release rate but suppressed (negative effect) the release rate of N and P from decomposing needles in both disturbed and rehabilitated forests. The enhanced needle decomposition rate by N addition was significantly related to the reduction in the C/N ratio in decomposing needles. However, N availability is not the sole factor limiting needle decomposition in both disturbed and rehabilitated forests. The positive effect was more sensitive to the N addition rate in the rehabilitated forest than in the disturbed forest, however the reverse was true for the negative effect. These results suggest that nutrient status could be one of the important factors in controlling the response of litter decomposition and its nutrient release to elevated N deposition in reforested ecosystems in the study region.     

Litter pool sizes,decomposition, and nitrogen dynamics in <Emphasis Type="Italic">Spartina alterniflora</Emphasis>-invaded and native coastal marshlands of the Yangtze Estuary

Past studies have focused primarily on the effects of invasive plants on litter decomposition at soil surfaces. In natural ecosystems, however, considerable amounts of litter may be at aerial and belowground positions. This study was designed to examine the effects of Spartina alterniflora invasion on the pool sizes and decomposition of aerial, surficial, and belowground litter in coastal marshlands, the Yangtze Estuary, which were originally occupied by two native species, Scirpus mariqueter and Phragmites australis. We collected aerial and surficial litter of the three species once a month and belowground litter once every 2 months. We used the litterbag method to quantify litter decomposition at the aerial, surficial and belowground positions for the three species. Yearly averaged litter mass in the Spartina stands was 1.99 kg m⁻²; this was 250 and 22.8% higher than that in the Scirpus (0.57 kg m⁻²) and Phragmites (1.62 kg m⁻²) stands, respectively. The litter in the Spartina stands was primarily distributed in the air (45%) and belowground (48%), while Scirpus and Phragmites litter was mainly allocated to belowground positions (85 and 59%, respectively). The averaged decomposition rates of aerial, surficial, and belowground litter were 0.82, 1.83, and 1.27 year⁻¹ for Spartina, respectively; these were 52, 62 and 69% of those for Scirpus litter at corresponding positions and 158, 144 and 78% of those for Phragmites litter, respectively. The differences in decomposition rates between Spartina and the two native species were largely due to differences in litter quality among the three species, particularly for the belowground litter. The absolute amount of nitrogen increased during the decomposition of Spartina stem, sheath and root litter, while the amount of nitrogen in Scirpus and Phragmites litter declined during decomposition for all tissue types. Our results suggest that Spartina invasion altered the carbon and nitrogen cycling in the coastal marshlands of China.     

Litter production, leaf litter decomposition and nutrient return in Cunninghamia lanceolata plantations in south China: effect of planting conifers with broadleaved species

This study compared litter production, litter decomposition and nutrient return in pure and mixed species plantations. Dry weight and N, P, K, Ca, Mg quantities in the litterfall were measured in one pure Cunninghamia lanceolata plantation (PC) and two mixed-species plantations of C. lanceolata with Alnus cremastogyne (MCA) and Kalopanax septemlobus (MCK) in subtropical China. Covering 6 years of observations, mean annual litter production of MCA (4.97 Mg·ha⁻¹) and MCK (3.97 Mg·ha⁻¹) was significantly higher than that of PC (3.46 Mg·ha⁻¹). Broadleaved trees contributed 42% of the total litter production in MCA and 31% in MCK. Introduction of broadleaved tree species had no significant effect on litterfall pattern. Total litterfall was greatest in the dry season from November to March. Nutrient returns to the forest floor through leaf litter were significantly higher in both MCA and MCK than in PC (P < 0.05). The amounts of N, K, and Mg returned to the forest floor through leaf litter were highest in the MCA, and P and Ca returns were highest in the MCK. Percent contribution of broadleaf litter to total nutrient returns ranged from 41.7% to 86.9% in MCA and from 49.3% to 74.8% in MCK. The decomposition rate of individual leaf litter increased in the order: C. lanceolata < K. septemlobus < A. cremastogyne. Litter mixing had a positive effect on decomposition rate of the more recalcitrant litter and promoted nutrient return. Relative to mass loss of A. cremastogyne decomposing alone, higher mass loss of the mixture of C. lanceolata and A. cremastogyne was observed after 330 days of decomposition. These results indicate that mixed plantations of different tree species have advantages over monospecific plantations with regards to nutrient fluxes and these advantages have relevance to restoration of degraded sites. Responsible Editor: Alfonso Escudero.     

Contrasting litter effects on old field tree germination and emergence

Because the fate of seeds is critical to understanding the invasion of old fields by trees, and plant litter is an important component defining the old field microsite of dispersed seeds, I investigated the effects of litter type (Solidago spp./goldenrod,Quercus spp./oak, mixed) and litter amount (100–800 g/m²) on tree seed germination and seedling emergence. I found that at all densities bothSolidago andQuercus litter greatly reduced emergence of the small-seeded, bird-dispersed species,Juniperus virginiana andCornus florida. For one of the large-seeded, mammal-dispersed species,Carya tomentosa, high densitySolidago litter and high density mixed litter treatments reduced emergence. For the other large-seeded species,Quercus rubra, the high density mixed litter treatment and all levels ofSolidago litter reduced emergence.Quercus seedlings emerged twice as often as the other three species in control pots without litter.Carya emerged before the other species but the high density oak treatment delayed the expansion of its cotyledons. My results suggest that litter may contribute to the slow rate of tree invasion and the low probability of tree establishment in old fields. However, old field litter studies taken together point to the difficulty in drawing general conclusions about any net effect of litter on old field tree establishment.     

Invasion of <Emphasis Type="Italic">Spartina alterniflora</Emphasis> Enhanced Ecosystem Carbon and Nitrogen Stocks in the Yangtze Estuary,China

Whether plant invasion increases ecosystem carbon (C) stocks is controversial largely due to the lack of knowledge about differences in ecophysiological properties between invasive and native species. We conducted a field experiment in which we measured ecophysiological properties to explore the response of the ecosystem C stocks to the invasion of Spartina alterniflora (Spartina) in wetlands dominated by native Scirpus mariqueter (Scirpus) and Phragmites australis (Phragmites) in the Yangtze Estuary, China. We measured growing season length, leaf area index (LAI), net photosynthetic rate (Pn), root biomass, net primary production (NPP), litter quality and litter decomposition, plant and soil C and nitrogen (N) stocks in ecosystems dominated by the three species. Our results showed that Spartina had a longer growing season, higher LAI, higher Pn, and greater root biomass than Scirpus and Phragmites. Net primary production (NPP) was 2.16 kg C m⁻² y⁻¹ in Spartina ecosystems, which was, on average, 1.44 and 0.47 kg C m⁻² y⁻¹ greater than that in Scirpus and Phragmites ecosystems, respectively. The litter decomposition rate, particularly the belowground decomposition rate, was lower for Spartina than Scirpus and Phragmites due to the lower litter quality of Spartina. The ecosystem C stock (20.94 kg m⁻²) for Spartina was greater than that for Scirpus (17.07 kg m⁻²), Phragmites (19.51 kg m⁻²) and the mudflats (15.12 kg m⁻²). Additionally, Spartina ecosystems had a significantly greater N stock (698.8 g m⁻²) than Scirpus (597.1 g m⁻²), Phragmites ecosystems (578.2 g m⁻²) and the mudflats (375.1 g m⁻²). Our results suggest that Spartina invasion altered ecophysiological processes, resulted in changes in NPP and litter decomposition, and ultimately led to enhanced ecosystem C and N stocks in the invaded ecosystems in comparison to the ecosystems with native species.     

杉木凋落物覆盖对自身幼苗出土和早期生长的影响

凋落物可通过物理和化学作用显著影响幼苗出土和早期生长,进而影响天然更新.杉木是中国南方重要的造林树种,但存在着天然更新障碍,其原因可能是林下较厚的凋落物层阻碍了杉木幼苗出土和早期生长.本试验利用覆盖自然和塑料凋落物来研究凋落物对杉木幼苗出土和早期生长的影响,并检验其影响是物理作用还是化学作用.本试验设置2种凋落物类型(自然和塑料凋落物)和4个覆盖厚度(对照,0 g·m^-2;浅层,200 g·m^-2;中层,400 g·m^-2;深层,800 g·m^-2).结果表明: 与对照(0 g·m^-2)相比,浅层(200 g·m^-2)凋落物覆盖对出苗率有促进作用但不显著,深层(800 g·m^-2)凋落物覆盖对出苗率和存活率有显著抑制作用.随着凋落物覆盖厚度的增加,幼苗根长不断减小,而茎长逐渐增加.凋落物浅层覆盖下杉木幼苗的根生物量、叶生物量和总生物量均最大,深层覆盖下最小.幼苗的根冠比随着凋落物覆盖厚度的增加而不断减小.与对照相比,凋落物覆盖下幼苗光合与非光合组织生物量比均有所增加.相同覆盖厚度下,自然和塑料凋落物对杉木幼苗出土和早期生长的影响均无显著差异,表明短期内凋落物覆盖对幼苗出土和早期生长的影响主要是物理作用.随着凋落物覆盖厚度的增加,杉木幼苗出土和早期生长表现为先促进后抑制,且杉木幼苗为了穿过厚厚的凋落物层倾向于把资源分配给地上部分.本研究结果为凋落物是影响杉木幼苗建植和天然更新的一个重要生态因子提供了试验证据.     

Effect of grass litter on seedling recruitment of the critically endangered Cistus heterophyllus in Spain

Cistus heterophyllus Desf. is an Iberian-north African endemic plant species with only one wild population left in Spain. A sowing experiment under controlled conditions was designed to assess both the seedling emergence rate and the growth fitness of C. heterophyllus on a litter layer of Brachypodium retusum (Pers.) Beau., which forms dense carpets around the remnant population of C. heterophyllus. In order to test the physical effects of B. retusum litter on Cistus seedling recruitment, six sowing treatments in pots were compared that combined three doses of litter (0, 0.6 and 1.1 kg m⁻²) and different physical locations of the seeds (above, below the litter layer or within a litter–peat mixture). B. retusum litter interfered with the seedling recruitment of C. heterophyllus, negatively affecting both seedling emergence and their early growth. Such an effect was more severe when the seeds were located above a 0.6 kg m⁻² litter layer, although seedlings from seeds sown under the litter showed a growth reduction ranging from 58% to 100% compared to the no-litter treatment. A significant negative effect of the litter–peat mixture on the number of leaves, leaf size and number of branches of seedlings was also inferred. These data provide experimental evidence on the serious threat that B. retusum litter represents for the conservation and regeneration of the last population of C. heterophyllus in Spain.     

CONTRASTING GERMINATION AND SEEDLING GROWTH OF BETULA ALLEGHANIENSIS AND RHUS TYPHINA SUBJECTED TO VARIOUS AMOUNTS AND TYPES OF PLANT LITTER

Several characteristics of the environment, such as plant litter, may interact with species characteristics, such as seed size, to determine patterns of plant establishment and thereby influence abundance and spatial location of plants long after the factor has itself disappeared. To determine if litter might differentially affect seedling establishment of Betula alleghaniensis Britton and Rhus typhina L., two woody species with different sized seeds, we conducted two greenhouse experiments. We documented emergence, morphology, and growth patterns of seedlings covered by three amounts each of Fagus grandifolia Ehrh. leaves or Tsuga canadensis L. needles. All amounts of litter significantly reduced and delayed final Betula alleghaniensis seedling emergence, and the resultant seedlings had reduced root: shoot ratios, longer hypocotyls, and were less robust. Litter did not affect Rhus typhina emergence but did alter biomass allocation: stem, leaf, and root biomass of Rhus seedlings was reduced by greater amounts of litter, and by leaf litter relative to needle litter. We attribute the different responses of Betula and Rhus to differences in seed size (Betula mean seed size = 1.0 mg; Rhus mean seed size = 8.5 mg) and germination cues (Betula requires light, Rhus requires chemical or heat scarification), and suggest that differential response to litter may contribute to coexistence among these sympatric species.     

The Sesquiterpene Caulerpenyne from Caulerpa spp. is a Lipoxygenase Inhibitor

Many algae contain secondary metabolites with the potential to gain importance as pharmaceutically active secretions. Caulerpa racemosa var. cylindracea and Caulerpa prolifera are very abundant on the Mediterranean coastlines. The methanolic extracts of C. racemosa and C. prolifera were tested for inhibitory effects on soybean lipoxygenase. The extract of C. prolifera showed potent inhibitory effect in a lipoxygenase enzyme activity assay. HPLC comparison revealed that C. racemosa extract contained less caulerpenyne, the major secondary metabolite of both algae. In accordance with these findings, purified caulerpenyne inhibited lipoxygenase with an IC₅₀ of 5.1 μM. The enzyme kinetic studies indicated that both K _M and V _max decreased from 0.041 to 0.019 mM and 312.5 to 151.5 U mL⁻¹ in the presence of 5 μM caulerpenyne, revealing an un-competitive type of inhibition kinetics. The major secondary metabolite of Caulerpa species, caulerpenyne, is thus a novel lipoxygenase inhibitor that can be easily obtained in high quantities from the abundant algae.     

The influence of a neotropical herbivore (<Emphasis Type="Italic">Lamponius portoricensis</Emphasis>) on nutrient cycling and soil processes

The role of phytophagous insects in ecosystem nutrient cycling remains poorly understood. By altering the flow of litterfall nutrients from the canopy to the forest floor, herbivores may influence key ecosystem processes. We manipulated levels of herbivory in a lower montane tropical rainforest of Puerto Rico using the common herbivore, Lamponius portoricensis (Phasmatidea), on a prevalent understory plant, Piper glabrescens (Piperaceae), and measured the effects on nutrient input to the forest floor and on rates of litter decomposition. Four treatment levels of herbivory generated a full range of leaf area removal, from plants experiencing no herbivory to plants that were completely defoliated (>4,000 cm² leaf area removed during the 76-day study duration). A significant (P<0.05) positive regression was found between all measures of herbivory (total leaf area removed, greenfall production, and frass-related inputs) and the concentration of NO ₃ ⁻ in ion exchange resin bags located in the litter layer. No significant relationship was found between any of the herbivory components and resin bag concentrations of NH ₄ ⁺ or PO ₄ ⁻ . Rates of litter decay were significantly affected by frass-related herbivore inputs. A marginally significant negative relationship was also found between the litter mass remaining at 47 days and total leaf area removed. This study demonstrated a modest, but direct relationship between herbivory and both litter decomposition and NO ₃ ⁻ transfer to the forest floor. These results suggest that insect herbivores can influence forest floor nutrient dynamics and thus merit further consideration in discussions on ecosystem nutrient dynamics.     

An isotopic method for testing the influence of leaf litter quality on carbon fluxes during decomposition

During microbial breakdown of leaf litter a fraction of the C lost by the litter is not released to the atmosphere as CO₂ but remains in the soil as microbial byproducts. The amount of this fraction and the factors influencing its size are not yet clearly known. We performed a laboratory experiment to quantify the flow of C from decaying litter into the soil, by means of stable C isotopes, and tested its dependence on litter chemical properties. Three sets of ¹³C-depleted leaf litter (Liquidambar styraciflua L., Cercis canadensis L. and Pinus taeda L.) were incubated in the laboratory in jars containing ¹³C-enriched soil (i.e. formed C4 vegetation). Four jars containing soil only were used as a control. Litter chemical properties were measured using thermogravimetry (Tg) and pyrolysis–gas chromatography/mass spectrometry–combustion interface–isotope ratio mass spectrometry (Py–GC/MS–C–IRMS). The respiration rates and the δ¹³C of the respired CO₂ were measured at regular intervals. After 8 months of incubation, soils incubated with both L. styraciflua and C. canadensis showed a significant change in δ¹³C (δ¹³C_final = −20.2 ± 0.4‰ and −19.5 ± 0.5‰, respectively) with respect to the initial value (δ¹³C_initial = −17.7 ± 0.3‰); the same did not hold for soil incubated with P. taeda (δ¹³C_final:−18.1 ± 0.5‰). The percentages of litter-derived C in soil over the total C loss were not statistically different from one litter species to another. This suggests that there is no dependence of the percentage of C input into the soil (over the total C loss) on litter quality and that the fractional loss of leaf litter C is dependent only on the microbial assimilation efficiency. The percentage of litter-derived C in soil was estimated to be 13 ± 3% of total C loss.     

Species-specific effects of woody litter on seedling emergence and growth of herbaceous plants

The effect of litter on seedling establishment can influence species richness in plant communities. The effect of litter depends on amount, and also on litter type, but relatively little is known about the species-specific effects of litter. We conducted a factorial greenhouse experiment to examine the effect of litter type, using two woody species that commonly co-occur in boreonemoral forest--evergreen spruce (Picea abies), deciduous hazel (Corylus avellana), and a mixture of the two species--and litter amount--shallow (4 mm), deep (12 mm) and leachate--on seedling emergence and biomass of three understorey species. The effect of litter amount on seedling emergence was highly dependent on litter type; while spruce needle litter had a significant negative effect that increased with depth, seedling emergence in the presence of hazel broadleaf litter did not differ from control pots containing no litter. Mixed litter of both species also had a negative effect on seedling emergence that was intermediate compared to the single-species treatments. Spruce litter had a marginally positive (shallow) or neutral effect (deep) on seedling biomass, while hazel and mixed litter treatments had significant positive effects on biomass that increased with depth. We found non-additive effects of litter mixtures on seedling biomass indicating that high quality hazel litter can reduce the negative effects of spruce. Hazel litter does not inhibit seedling emergence; it increases seedling growth, and creates better conditions for seedling growth in mixtures by reducing the suppressive effect of spruce litter, having a positive effect on understorey species richness.     

Effects of understory removal, N fertilization, and litter layer removal on soil N cycling in a 13-year-old white spruce plantation infested with Canada bluejoint grass

Canada bluejoint grass [Calamagrostis canadensis (Michx.) Beauv., referred to as bluejoint below] is a competitive understory species widely distributed in the boreal region in North America and builds up a thick litter layer that alters the soil surface microclimate in heavily infested sites. This study examined the effects of understory removal, N fertilization, and litter layer removal on litter decomposition, soil microbial biomass N (MBN), and net N mineralization and nitrification rates in LFH (the sum of organic horizons of litter, partially decomposed litter and humus on the soil surface) and mineral soil (0–10 cm) in a 13-year-old white spruce [Picea glauca (Moench.) Voss] plantation infested with bluejoint in Alberta, Canada. Removal of the understory vegetation and the litter layer together significantly increased soil temperature at 10 cm below the mineral soil surface by 1.7 and 1.3°C in summer 2003 and 2004, respectively, resulting in increased net N mineralization (by 1.09 and 0.14 mg N kg⁻¹ day⁻¹ in LFH and mineral soil, respectively, in 2004) and net nitrification rates (by 0.10 and 0.20 mg N kg⁻¹ day⁻¹ in LFH and mineral soil, respectively, in 2004). When the understory vegetation was intact, nitrification might have been limited by NH₄ ⁺ availability due to competition for N from bluejoint and other understory species. Litter layer removal increased litter decomposition rate (percentage mass loss per month) from 2.6 to 3.0% after 15 months of incubation. Nitrogen fertilization did not show consistent effects on soil MBN, but increased net N mineralization and nitrification rates as well as available N concentrations in the soil. Clearly, understory removal combined with N fertilization was most effective in increasing rates of litter decomposition, net N mineralization and nitrification, and soil N availability. The management of understory vegetation dominated by bluejoint in the boreal region should consider the strong effects of understory competition and the accumulated litter layer on soil N cycling and the implications for forest management.     

Invasion by a Perennial Herb Increases Decomposition Rate and Alters Nutrient Availability in Warm Temperate Lowland Forest Remnants

We determined the impact of the invasive herb, Tradescantia fluminensis Vell., on litter decomposition and nutrient availability in a remnant of New Zealand lowland podocarp–broadleaf forest. Using litter bags, we found that litter beneath mats of Tradescantia decomposed at almost twice the rate of litter placed outside the mat. Values of k (decomposition quotient) were 9.44±0.42 yrs for litter placed beneath Tradescantia and 5.42±0.42 yrs for litter placed in native, non-Tradescantia plots. The impact of Tradescantia on decomposition was evident through the smaller forest floor mass in Tradescantia plots (2.65±1.05 t ha⁻¹) compared with non-Tradescantia plots (5.05±1.05 t ha⁻¹), despite similar quantities of annual leaf litterfall into Tradescantia plots (6.85±0.85 t ha⁻¹ yr⁻¹) and non-Tradescantia plots (7.45±1.05 t ha⁻¹ yr⁻¹). Moreover, there was increased plant nitrate available, as captured on resin bags, in Tradescantia plots (25.77 ± 8.32 cmol(−)/kg resin) compared with non-Tradescantia plots (9.55±3.72 cmol(−)/kg resin). Finally, the annual nutrient uptake by Tradescantia represented a large proportion of nutrients in litterfall (41% N, 61% P, 23% Ca, 46% Mg and 83% K), exceeded the nutrient content of the forest floor (except Ca), but was a small proportion of the topsoil nutrient pools. Taken together, our results show that Tradescantia increases litter decomposition and alters nutrient availability, effects that could influence the long-term viability of the majority of podocarp–broadleaf forest remnants affected with Tradescantia in New Zealand. These impacts are likely mostly due to Tradescantia's vegetation structure (i.e., tall, dense mats) and associated microclimate, compared with native ground covers. This revised version was published online in July 2006 with corrections to the Cover Date.