Natural-enemies affect the seed and litter fall dynamics of Melaleuca quinquenervia in the wetlands,and influence long-term species diversity in leaf-litter

Wetlands Ecology and Management - The exotic tree Melaleuca quinquenervia (melaleuca) has invaded and formed monotypic forest stands in ecologically sensitive wetlands of Florida. We hypothesized...     

Decline in exotic tree density facilitates increased plant diversity: the experience from Melaleuca quinquenervia invaded wetlands

The Australian tree Melaleuca quinquenervia (melaleuca) formed dense monocultural forests several decades after invading parts of Florida and the Caribbean islands. These dominant forests have displaced native vegetation in sensitive wetland systems. We hypothesized that native plant diversity would increase following recent reductions in density of mature melaleuca stands in south Florida. We therefore examined data on changes in melaleuca densities and plant species diversity derived from permanent plots that were monitored from 1997 to 2005. These plots were located within mature melaleuca stands in nonflooded and seasonally-flooded habitats. Two host-specific biological control agents of melaleuca, Oxyops vitiosa and Boreioglycaspis melaleucae, were introduced during 1997 and 2002, respectively. Also, an adventive rust fungus Puccinia psidii and lobate-lac scale Paratachardina pesudolobata became abundant during the latter part of the study period. Overall melaleuca density declines in current study coincided with two to four fold increases in plant species diversity. The greatest declines in melaleuca density as well as the greatest increases in family importance values and species diversity indices occurred in nonflooded as compared to seasonally-flooded habitats. Most pioneer plant species in study sites belonged to Asteraceae, Cyperaceae, Poaceae, and Ulmaceae. The rapid reduction in melaleuca density and canopy cover during the study period may be attributed to self-thinning accelerated by the negative impact of natural enemies. Densities of other woody plants, particularly Myrica and Myrsine, which were sparsely represented in the understory by a few suppressed individuals also declined during the same period, possibly due to infestation by the generalist lac-scale. These findings indicate that natural-enemy accelerated self-thinning of melaleuca densities is positively influencing the native plant diversity and facilitating the partial rehabilitation of degraded habitats.     

<Emphasis Type="Italic">Melaleuca quinquenervia</Emphasis> dominated forests in Florida: analyses of natural-enemy impacts on stand dynamics

Melaleuca quinquenervia (melaleuca) is a native of Australia but has become an invasive plant in Florida, USA. We conducted a long-term demographic study of melaleuca in three sections (central, transitional, and peripheral) of monoculture stands located in Florida, and quantified absolute density, diameter at breast height and basal area of trees by section at three sites. Additionally, we monitored the impacts of natural enemy (insects and fungi) on melaleuca populations which became apparent after 2001. Both absolute density and basal area, from before (1997–2001) and after noticeable natural-enemy impact (2002–2005), were compared. Prior to the natural-enemy impact, absolute density of melaleuca trees declined primarily due to self-thinning and associated losses of small trees, but diameter at breast height increased, as did the basal area. Later during the period when natural enemies prevailed, absolute density declined at a significantly greater rate across all sections but was highest at the periphery. The decrease in mean absolute density and basal area/ha of melaleuca during the natural-enemy impacted period coincided with the increased incidence of the populations of plant-feeding insects and fungi. The mean diameter at breast height continued to increase in all sections of the stands throughout the study period. An increasing trend in basal area prior to natural-enemy impact was reversed after increase in natural-enemy abundance and noticeable impact in all three sections of the stands. These findings lend support to a growing body of literature that implicates natural enemies as increasingly important density-independent regulators of M. quinquenervia populations. The U.S. Government's right to retain a non-exclusive, royalty-free license in and to any copyright is acknowledged     

Impacts of the exotic,nitrogen-fixing black locust (Robinia pseudoacacia) on nitrogen-cycling in a pine–oak ecosystem

We investigated the influence of the exotic nitrogen-fixing black locust (Robinia pseudoacacia) on nitrogen cycling in a pitch pine (Pinus rigida) −scrub oak (Quercus ilicifolia, Q. prinoides) ecosystem. Within paired pine-oak and adjacent black locust stands that were the result of a 20-35 year-old invasion, we evaluated soil nutrient contents, soil nitrogen transformation rates, and annual litterfall biomass and nitrogen concentrations. In the A horizon, black locust soils had 1.3-3.2 times greater nitrogen concentration relative to soils within pine-oak stands. Black locust soils also had elevated levels of P and Ca, net nitrification rates and total net N-mineralization rates. Net nitrification rates were 25-120 times greater in black locust than in pine-oak stands. Elevated net N-mineralization rates in black locust stands were associated with an abundance of high nitrogen, low lignin leaf litter, with 86 kg N ha^–1 yr^–1 in leaf litter returned compared with 19 kg N ha^–1 yr^–1 in pine-oak stands. This difference resulted from a two-fold greater litterfall mass combined with increased litter nitrogen concentration in black locust stands (1.1% and 2.6% N for scrub oak and black locust litter, respectively). Thus, black locust supplements soil nitrogen pools, increases nitrogen return in litterfall, and enhances soil nitrogen mineralization rates when it invades nutrient poor, pine-oak ecosystems. This revised version was published online in June 2006 with corrections to the Cover Date.     

High litterfall in old-growth and secondary upper montane forest of Costa Rica

Tropical upper montane forests usually comprise trees of small stature with a relatively low aboveground productivity. In contrast to this rule, in the Cordillera de Talamanca (Costa Rica), tall trees (>35 m in height and more than 60 cm in diameter) are characteristic for the upper montane old-growth oak forests which are growing at an altitude of almost 3,000 m close to the alpine timberline. For these exceptional forests, productivity data are not yet available. In this study, we analyzed litterfall and its components (tree leaves, litter of epiphytic vascular and non-vascular plants, mistletoes, twigs and other canopy debris) in three forest stands belonging to different successional stages and related seasonal changes in litterfall to micrometeorological variables. The studied stands were early-successional forest (10–15-year-old), mid-successional forest (40-year-old), and old-growth forest. The stands are dominated by Quercus copeyensis and are located at 2,900-m altitude. Total litterfall was highest in the mid-successional forest (1,720 g m⁻² y⁻¹), and reached 1,288 g m⁻² y⁻¹ in the old-growth forest and 934 g m⁻² y⁻¹ in the early-successional forest. Litter mass was dominated by leaves in all stages (56–84% of total litterfall). In the old-growth forest, however, twigs and small canopy debris particles (33%), epiphytes (6%), and mistletoes (5%) also contributed substantially to litter mass. Leaf litterfall showed a clear seasonal pattern with a negative correlation to monthly precipitation and highest values in the dry season (January–April). However, the strongest correlation existed with minimum air temperature (negative), probably because temperatures already dropped at the end of the rainy season, when precipitation had not yet declined and leaf shedding already increased. In contrast, litterfall of epiphyte mass, and twigs and other debris was mostly dependent on occasional strong winds. We conclude that the upper montane oak forests of the Cordillera de Talamanca are exceptional with respect to the large tree size and the relatively high productivity as indicated by litterfall. Litter mass was especially high in the mid-successional and old-growth forests, where the observed annual totals are among the highest recorded for tropical forests so far.     

Litterfall dynamics in a mixed conifer-angiosperm forest in northern New Zealand

Litterfall in a mixed conifer-angiosperm temperate forest in northern New Zealand was traced for 5 years to determine the patterns of litter production and turnover for conifer and angiosperm components of the forest. Basal area and above-ground biomass was shared approximately equally between conifer (mostly Agathis australis; New Zealand kauri) and angiosperm species (plus tree ferns). The five-year mean annual litterfall, excluding macro-litter, was 7.76± 0.39(SEM) t ha^?1 and ranged from 6.77±0.70 t ha^?1 in 1983–4 to 8.79±1.00 t ha^?1 in 1987–8. Mean monthly litterfall showed a strong seasonal pattern with low rates in winter and early spring, increasing to a peak in early autumn. There were major differences in the nature and timing of litterfall between the conifer and angiosperm fractions. Angiosperm leaf litter reached a maximum in early summer, while conifer litterfall showed highest rates for leaves, twigs and cone scales in late summer-autumn. Conifer reproductive structures (strobili and cone scales) contributed from 13 to 21% of total litterfall, a value high relative to other temperate forests. However, conifer leaf turnover was low relative to that for the angiosperms. Size of the microlitter store was 16.16±1.97 t ha^?1 prior to conifer cone fall, and 18.70±2.02 t ha^?1 following it, and conifer litter made up 76–78% of the total litter store. The estimated mean annual decomposition constant, k, was 0.39 overall, 0.33 for conifer leaf litter and 0.71 for angiosperm leaf litter, values which agree well with previously published rates for decomposition in this forest stand. Differences in the costs of biomass production and rates of turnover, as measured by litterfall and decomposition, may help to explain the functional coexistence of conifers and angiosperms in mixed forests.     

Field colonization, population growth, and dispersal of Boreioglycaspis melaleucae Moore, a biological control agent of the invasive tree Melaleuca quinquenervia (Cav.) Blake

Invasion of native plant communities by the Australian paperbark tree (“melaleuca”), Melaleuca quinquenervia, complicates restoration of the Florida Everglades. Biological control, within the context of a comprehensive management program, offers a means to suppress regeneration of melaleuca after removal of existing trees and a mechanism to forestall reinvasion. To meet this need, a biological control program commenced in 1997 upon the release of an Australian weevil (Oxyops vitiosa [Pascoe] [Coleoptera: Curculionidae]). Release of a second biological control agent, the melaleuca psyllid (Boreioglycaspis melaleucae Moore), followed in February 2002 at field sites containing mixed age-class melaleuca stands or coppicing stumps. Each site was inoculated with 7000–10,000 adult psyllids, with one exception where 2000 nymphs were released on seedlings the following December. Psyllid populations established everywhere irrespective of colony source, site conditions, or the quantity released, although numbers released and, to a lesser degree, colony age influenced the numbers of colonies produced. Quantity included in the release was the major determinant of the resultant number of colonies, although the duration of their tenure in quarantine culture may have also influenced this. One site, comprised mainly of coppicing stumps, contained 3.3 million psyllids per ha within 3 months after release. Less than 1% of the coppices at a similar site harbored psyllid colonies 2 months after release (May 2002), but this rose to 75% in October then to 100% by December. The census population exceeded 715,000 adults and nearly 11 million nymphs by late January 2003. Psyllid populations dispersed 2.2–10.0 km/year, with the slower rates in dense, continuous melaleuca stands and faster rates in fragmented stands. Over 1 million psyllids had been redistributed to 100 locations as of December 2005. This species now occurs throughout much of the range of melaleuca in south Florida due to natural range expansion as well as anthropogenic dissemination.     

Field colonization,population growth,and dispersal of Boreioglycaspis melaleucae Moore,a biological control agent of the invasive tree Melaleuca quinquenervia (Cav.) Blake

Invasion of native plant communities by the Australian paperbark tree (“melaleuca”), Melaleuca quinquenervia, complicates restoration of the Florida Everglades. Biological control, within the context of a comprehensive management program, offers a means to suppress regeneration of melaleuca after removal of existing trees and a mechanism to forestall reinvasion. To meet this need, a biological control program commenced in 1997 upon the release of an Australian weevil (Oxyops vitiosa [Pascoe] [Coleoptera: Curculionidae]). Release of a second biological control agent, the melaleuca psyllid (Boreioglycaspis melaleucae Moore), followed in February 2002 at field sites containing mixed age-class melaleuca stands or coppicing stumps. Each site was inoculated with 7000–10,000 adult psyllids, with one exception where 2000 nymphs were released on seedlings the following December. Psyllid populations established everywhere irrespective of colony source, site conditions, or the quantity released, although numbers released and, to a lesser degree, colony age influenced the numbers of colonies produced. Quantity included in the release was the major determinant of the resultant number of colonies, although the duration of their tenure in quarantine culture may have also influenced this. One site, comprised mainly of coppicing stumps, contained 3.3 million psyllids per ha within 3 months after release. Less than 1% of the coppices at a similar site harbored psyllid colonies 2 months after release (May 2002), but this rose to 75% in October then to 100% by December. The census population exceeded 715,000 adults and nearly 11 million nymphs by late January 2003. Psyllid populations dispersed 2.2–10.0 km/year, with the slower rates in dense, continuous melaleuca stands and faster rates in fragmented stands. Over 1 million psyllids had been redistributed to 100 locations as of December 2005. This species now occurs throughout much of the range of melaleuca in south Florida due to natural range expansion as well as anthropogenic dissemination.     

Ground beetle diversity in ancient woodland remnants in north-western Germany (Coleoptera, Carabidae)

We investigated ground beetle communities (Coleoptera, Carabidae) in ancient woodland remnants in north-western Lower Saxony, Germany. A total of 90 pitfall traps was exposed in a stratified design in 10 stands of mature oak–beech and oak–hornbeam forests in the year 2003. Overall, 47 species (10,676 individuals) were recorded. Among these were the two relict species Carabus glabratus and Abax parallelus, and 14 further eurytopic forest species. Eleven species exhibited a high frequency and were found in all of the ten stands. Multiple linear regressions showed several significant relationships at two scales for species richness of different groups of carabids and for several of the measured environmental factors. Forest area, litter depth, amount of dead wood, distance to forest edge, and soil moisture were found to be key factors determining species richness. Furthermore, recent disturbance by logging reduced the number of forest species. According to direct gradient analyses soil moisture and litter depth have greatest influence on species communities of both, forest species and widespread species. Habitat suitability models for the two recorded relict species, A. paralellus and C. glabratus, were developed using logistic regression. The presence of A. parallelus in the mature ancient woodland remnants depends mainly on higher values of soil moisture, whereas for C. glabratus none of the measured environmental variables appeared to be key factors. Implications for the conservation of carabid assemblages in mature ancient woodlands include the advice to spread out logging over long periods of time and over various woodlands in order to keep the stand disturbance at a long-term low level. Variation in logging practices may help to conserve diverse structures. Afforestation with non-native tree species should be avoided in the managed ancient woodlands. Finally, especially the preservation of a high soil moisture seems to be important to conserve typical carabid communities.     

Effects of forest-pasture edge on C,N and P associated with <Emphasis Type="Italic">Caesalpinia eriostachys</Emphasis>, a dominant tree species in a tropical deciduous forest in Mexico

Forest fragmentation in tropical ecosystems can alter nutrient cycling in diverse ways. We have analysed the effects of the forest-pasture edge on nutrient soil dynamics in a tropical deciduous forest (TDF) in Mexico. In two remnant forest fragments, both larger than 10 ha, litterfall, litter and soil samples associated to the tree Caesalpinia eriostachys were collected at five distances from the pasture edge into the inner forest (10 m in the pasture and 0–10, 30–40, 70–80 and 100–110 m towards the forest interior). We measured the concentrations of carbon (C), nitrogen (N) and phosphorus (P) in litterfall, surface litter and soil, and soil microbial C (C_mic) and microbial N (N_mic). Soil nutrient concentrations and C_mic and N_mic were lower in the pasture soils than in the forest soil samples. Total C and N pools, and C_mic and N_mic in the pasture were lower than in the forest. In contrast, net N immobilization and the increase in N_mic from rain to dry season increased from the edge to the inner forest. Soil P concentration was lower in the pasture and at the first distance class in the forest margin (0–10 m) than in the sites located further into the forest, while litter P concentration had the inverse pattern. Litterfall P was also reduced near the edge and increased towards the forest interior. As a consequence, litterfall C:P and N:P ratios decreased from the edge to the inner forest. These results suggest that the forest–pasture edge disrupts P dynamics within the first 10 m in the forest. Thus, plants' use of nutrients and productivity could be altered in the edge of fragmented forests.     

Relationship of litterfall to basal area and climatic variables in cool temperate forests of southern Tasmania

Components of litter accession were measured for 2 years in two re growth eucalypt stands, a Nothofagus mixed forest, and a tall shrubland in the temperate forests of southern Tasmania. Total annual litterfall (t ha^-1) ranged from 4.77 to 5.64 in the regrowth eucalypt stands and 4.06 to 4.94 and 1.95 to 2.17 in the Nothofagus mixed forest and tall shrubland, respectively. Significant correlations were found between annual litterfall of individual tree species and their respective basal area measured at 1.3 m height. The seasonal patterns of litterfall were most closely related to mean maximum temperatures. However, fall of non-leaf material was also related to gale force winds associated with the spring equinox.     

Regional‐Scale Variation in Litter Production and Seasonality in Tropical Dry Forests of Southern Mexico1

Highly seasonal rainfall creates a pulse of litterfall in the southern Yucatan peninsula region, with cascading effects on the timing of essential nutrient fluxes, microbial dynamics, and vegetation growth. I investigated whether forest age or a regional environmental gradient related to rainfall has a greater effect on patterns of litterfall in this increasingly human‐dominated landscape. Litterfall was sampled in 10–13 stands in each of three locations spanning a rainfall gradient of ca 900–1400 mm/yr. Litter was collected monthly from November 1998 through January 2000 in mature forests and in secondary forests aged 2–25 yr. Despite a substantial precipitation gradient, age was the only significant predictor of annual litter mass. Two‐ to five‐yr‐old forests produced significantly less litter than 12–25‐yr‐old secondary forests (4.6 vs. 6.2 Mg/ha/yr), but the difference between older secondary forests and mature forests (9 percent) was not significant. Litter production increased with rainfall, but not significantly so. The pattern of litterfall was similar across locations and age classes, with a peak during late March or early April. However, litterfall seasonality was most pronounced in the old secondary and mature forests. Litterfall was more evenly distributed throughout the year in forests under 10 yr old. Seasonality of litterfall was also less pronounced at the wettest site, with less disparity between peak litterfall and off‐peak months. Seasonality was not related to soil texture. Forest age and rainfall are important drivers of litterfall dynamics; however, both litter mass and degree of seasonality depended more strongly on forest age. Thus, the impact of land‐use change on litter nutrient cycling is as great, if not greater, than the constraint imposed by the major natural environmental factor affecting tropical dry forests.     

Invasion of a West Everglades wetland by Melaleuca quinquenervia countered by classical biological control

The population dynamics of Melaleuca quinquenervia were monitored over a 5-year period in a cypress-pine wetland while subjected to two levels of herbivory. The trees had been recruited during 1998–1999 after a destructive crown fire. Half of 26 experimental plots were sprayed every 4–6 weeks with an insecticide to reduce herbivory by the biological control agents Oxyops vitiosa and Boreioglycaspis melaleucae. After only 1-year melaleuca density increased by 26% in sprayed plots and by 7% in unsprayed plots. However, over the entire 5-year period melaleuca density increased in sprayed plots by 0.1% while decreasing by 47.9% in unsprayed plots when compared to initial densities. Annual mortality of melaleuca never exceeded 6% in any year in sprayed plots but ranged from 11% to 25% in unsprayed plots. There was a significant year by treatment interaction indicating the importance of the environment on tree mortality. Limited seed production occurred on sprayed trees but never on unsprayed trees. Mean tree height increased by 19.6% in sprayed plots while declining by 30.6% in unsprayed plots. Coverage by native vegetation did not increase with decreasing melaleuca density. This is the first study with controls that quantifies the population level regulation of melaleuca by introduced biological control agents and corroborates other correlative studies that documented significant changes in melaleuca communities after the introduction and establishment of biological control agents.     

Litterfall of tropical forested wetlands of Veracruz in the coastal floodplains of the Gulf of Mexico

Canopy productivity of five seasonally flooded forests in the central area of the Gulf of Mexico was estimated by measuring the litterfall. Productivity was estimated on a monthly basis from November 2005 to October 2006, and values ranged between 9 and 15 t ha⁻¹ yr⁻¹. A total of 57 plant species were recorded based on the litter collected. The dominant species (Pachira aquatica, Annona glabra, Hippocratea celastroides and Dalbergia brownei) were the principal litterfall producers. The contribution of two life forms was assessed: trees (including trees, shrubs and palms) and lianas (climbers, lianas and creepers). Lianas were found to be extremely productive and represented between 8 and 62% of the total litterfall at the sampled locations. Leaves contributed most to the litter, followed by branches and reproductive structures. Fruit and seed fall coincided with the rainy season, and accounted for 50–90% of the production and shedding during the flooded season. Species that released seeds during this time had their seeds dispersed by water. Flower production occurred during the dry season (March–June). Species richness did not explain litterfall productivity. The litterfall productivity of these forests is similar to that of mangrove ecosystems.     

Litterfall dynamics in carbonate and deltaic mangrove ecosystems in the Gulf of Mexico

From 1996 to 2002, we measured litterfall, standing litter crop, and litter turnover rates in scrub, basin, fringe and riverine forests in two contrasting mangrove ecosystems: a carbonate-dominated system in the Southeastern Everglades and a terrigenous-dominated system in Laguna de Terminos (LT), Mexico. We hypothesized that litter dynamics is driven by latitude, geomorphology, hydrology, soil fertility and soil salinity stress. There were significant temporal patterns in LT with litterfall rates higher during the rainy season (2.4 g m⁻² day⁻¹) than during the dry season (1.8 g m⁻² day⁻¹). Total annual litterfall was significantly higher in the riverine forest (12.8 Mg ha⁻² year⁻¹) than in the fringe and basin forests (9.7 and 5.2 Mg ha⁻² year⁻¹, respectively). In Southeastern Everglades, total annual litterfall was also significantly higher during the rainy season than during the dry season. Spatially, the scrub forest had the lowest annual litterfall (2.5 Mg ha⁻² year⁻¹), while the fringe and basin had the highest (9.1 and 6.5 Mg ha⁻² year⁻¹, respectively). In LT, annual standing litter crop was 3.3 Mg ha⁻¹ in the fringe and 2.2 Mg ha⁻¹ in the basin. Litter turnover rates were significantly higher in the fringe mangrove forest (4.1 year⁻¹) relative to the basin forests (2.2 year⁻¹). At Southeastern Everglades there were significant differences in annual standing litter crop: 1.9, 3.3 and 4.5 Mg ha⁻¹ at scrub, basin and fringe mangrove sites, respectively. Furthermore, turnover rates were similar at both basin and fringe mangrove types (2.1 and 2.0 year⁻¹, respectively) but significantly higher than scrub mangrove forest (1.3 year⁻¹). These findings suggest that litter export is important in regulating litter turnover rates in frequently flooded riverine and fringe forests, while in infrequently flooded basin forests, in situ litter decomposition controls litter turnover rates.     

Conversion of secondary broadleaved forest into Chinese fir plantation alters litter production and potential nutrient returns

Litter production, litter standing crop, and potential nutrient return via litterfall to soil were studied during a 4-year period (January 2004–December 2007) in a Chinese fir (Cunninghamia lanceolata (Lamb.) Hook) plantation and a secondary broadleaved forest in Hunan Province in subtropical China. Mean annual litterfall in the sampling sites varied from 358 g m⁻² in the pure plantation to 669 g m⁻² in the secondary broadleaved forest. Total litterfall followed a bimodal distribution pattern for both forests. Amount of litterfall was also related to the air temperature in both forests. During the period under this study, annual variation in the total litterfall in the pure plantation was significantly higher than that in the secondary broadleaved forest. Litterfall was markedly seasonal in the both forests. Leaf proportions of litterfall in the pure plantation and secondary broadleaved forest were 58.1 and 61.7%, respectively. Total potential nutrient returns to the soil through litterfall in the pure plantation were only 46.2% of those in the secondary broadleaved forest. Total litter standing crop was 913 and 807 g m⁻² in the pure plantation and secondary broadleaved forest, respectively. Our results confirm that conversion from a secondary broadleaved forest into a pure coniferous plantation changes the functioning of the litter system.     

Leaf litter decomposition in temperate deciduous forest stands with a decreasing fraction of beech (Fagus sylvatica)

We hypothesised that the decomposition rates of leaf litter will increase along a gradient of decreasing fraction of the European beech (Fagus sylvatica) and increasing tree species diversity in the generally beech-dominated Central European temperate deciduous forests due to an increase in litter quality. We studied the decomposition of leaf litter including its lignin fraction in monospecific (pure beech) stands and in stands with up to five tree genera (Acer spp., Carpinus betulus, Fagus sylvatica, Fraxinus excelsior, Tilia spp.) using a litterbag approach. Litter and lignin decomposition was more rapid in stand-representative litter from multispecific stands than in litter from pure beech stands. Except for beech litter, the decomposition rates of species-specific tree litter did not differ significantly among the stand types, but were most rapid in Fraxinus excelsior and slowest in beech in an interspecific comparison. Pairwise comparisons of the decomposition of beech litter with litter of the other tree species (except for Acer platanoides) revealed a “home field advantage” of up to 20% (more rapid litter decomposition in stands with a high fraction of its own species than in stands with a different tree species composition). Decomposition of stand-representative litter mixtures displayed additive characteristics, not significantly more rapid than predicted by the decomposition of litter from the individual tree species. Leaf litter decomposition rates were positively correlated with the initial N and Ca concentrations of the litter, and negatively with the initial C:N, C:P and lignin:N ratios. The results support our hypothesis that the overall decomposition rates are mainly influenced by the chemical composition of the individual litter species. Thus, the fraction of individual tree species in the species composition seems to be more important for the litter decomposition rates than tree species diversity itself.     

Nutrient cycling in Pinus sylvestris stands in eastern Finland

Nutrient cycling within three Pinus sylvestris stands was studied in eastern Finland. The aim of the study was to determine annual fluxes and distribution of N, P, K, Ca, Mg, Zn, Fe, B, and Al in the research stands. Special emphasis was put on determining the importance of different fluxes, especially the internal cycle within the trees in satisfying the tree nutrient requirements for biomass production. The following nutrient fluxes were included, input; free precipitation and throughfall, output; percolation through soil profile, biological cycle; nutrient uptake from soil, retranslocation within trees, return to soil in litterfall, release by litter decomposition. The distribution of nutrients was determined in above- and belowground tree compartments, in ground and field vegetation, and in soil.The nitrogen use efficiencies were 181, 211 and 191 g of tree aboveground dry matter produced per g of N supplied by uptake and retranslocation in the sapling, pole stage and mature stands, respectively. Field vegetation was more efficient in nitrogen use than trees. Stand belowground/aboveground and fine root/coarse root biomass ratios decreased with tree age. With only slightly higher fine root biomass, almost three times more nitrogen had to be taken-up from soil for biomass production in the mature stand than in the sapling stand.The annual input-output balances of most nutrients were positive; throughfall contained more nutrients than was lost in mineral soil leachate. The sulphate flux contributed to the leaching of cations, especially magnesium, from soil in the mature stand.Retranslocation supplied 17–42% of the annual N, P and K requirements for tree aboveground biomass production. Precipitation and throughfall were important in transferring K and Mg, and also N in the sapling stand. Litterfall was an important pathway for N, Ca, Mg and micro nutrients, especially in the oldest stands.     

Litter production,nutrient recycling and litter accumulation inPinus caribaea Morelet var.hondurensis stands in the northern Guniea savanna of Nigeria

Summary The seasonal pattern and quantity of litterfall were studied during a two-year period in two unthinned stands ofPinus caribaea Morelet var. hondurensis Barr. and Golf. in Nigeria. Although pine needles were cast continuously throughout the year, the peak period of litterfall occurred in the dry months of March and April. Mean values of annual litterfall were 3068 and 3665 kg/ha in the two stands aged 7–9 and 9–11 years respectively. Nutrient returns in litterfall in the stands had mean values of 15.0, 0.6, 17.3, 18.2 and 6.3 kg/ha of N, P, K, Ca and Mg respectively. Comparatively low amounts of N and P returned in litterfall were attributed to soil deficiencies of the two elements.Measurements of ground litter showed considerable dry matter accumulation (11378 kg/ha) in the litter layers. Estimates of litter decomposition rate and recycling time showed that it would take 3 to 4 years for the organic matter in annual litterfall to decompose completely as contrasted to about 2 to 5 months often reported under mixed nautral savanna vegetation in the same climatic environment. Similar estimates of nutrient recycling time also showed that between 2 to 4 years were required to mineralize nutrient elements in the annual litterfall; the relative mobilities of the elements were in the order K>Mg>P>NCa.     

Reactions of the herb and moss layer,tree saplings and the shrub layer to tree deaths in forests of the Wielkopolska National Park (Western Poland)

This study examines the impact of canopy and canopy gaps on the development of lower forest layers in five protected phytocoenoses of oak-hornbeam habitats (natural and regeneration stands) and oak-pine habitats in the Wielkopolska National Park (WPN). In the studied forests the most common form of dead trees are those which are uprooted (45–59%), while the most frequently dying tree is pine (40–88%). The total area of gaps in relation to the studied forest area ranges from 329 to 2356 m²/ha.