A Tropical Freshwater Wetland: III. Direct Use Values and Other Goods and Services

Understanding the economic value of a wetland as well as how people perceive its role in providing goods and services can provide insight into the wetland's actual role and the kinds of policies needed to ensure sustainable use. On the island of Kosrae, Federated States of Micronesia, freshwater forested wetlands dominated by Terminalia carolinensis (hereafter called Terminalia) are often found just upslope from mangrove forests, which appear to be hydrologically connected to them. Many of these Terminalia forests have been converted into agroforests. A survey of 10% of the households on Kosrae showed that 89% owned some Terminalia land. Most grew taro, bananas, and sugar cane, either in or immediately adjacent to Terminalia forests. Most owned canoes constructed of Terminalia logs, and nearly half had harvested trees from these forests during the past year: 64% to clear land for agricultural purposes, 36% for building canoes, and 31% for other uses. Terminalia forests provided over $3.1 million worth of goods to Kosraeans, primarily from agricultural production. Approximately 2/3 of those surveyed understood that Terminalia grows best in a wetland setting. Most thought that Terminalia forests provide erosion protection and improve water quality. However, very few were cognizant of the ecological links between Terminalia and mangrove forests. Kosraeans attached little importance to the fact that Terminalia is endemic to the eastern Caroline Islands. If human dependence on these wetlands increases, the integrity of Terminalia forests, as well as adjacent mangrove forests, could be at risk.     

Host and habitat preferences of polypore fungi in Micronesian tropical flooded forests

The distribution and ecological impacts of plant-associated fungi is determined in large part by their degree of specificity for particular host species or environmental conditions. Here we evaluate the host and habitat preferences among the Aphyllophorales, a guild of wood-decay basidiomycete fungi usually considered to be host generalists. We determined the patterns of host association in three well-defined, floristically distinct, tropical wetlands — freshwater forested wetlands, saltwater mangrove forests, and peatlands with scattered trees — on the islands of Kosrae and Pohnpei in the Federated States of Micronesia. Of 33 fungal species, 20 were locally rare. Of the 11 species sufficiently common to evaluate habitat specificity, nine showed significant habitat preferences. Of eight species common enough to evaluate within-habitat host specificity, six showed strong host preferences. All except one of the nine habitat-specialized fungi showed either statistically significant host specificity or strong numerical biases toward single host species. Our results suggest that host preferences may be important in shaping the assemblages of wood-decay fungi, and that the effect of environment on the distribution of susceptible plant species, rather on the fungi themselves, may ultimately drive the apparent habitat specificity of many fungi.     

Environmental Effects of Canopy Gap Formation in High-Rainfall Mangrove Forests1

This study investigated the importance of gap formation in mangrove swamps on the island of Kosrae, Federated States of Micronesia, in order to understand better both natural processes of forest development and the effects of harvesting trees for firewood in these wetlands. Measurements were concentrated in seven plots located near four rivers: three in fringe zones and four in basin zones. Each plot was a cluster of five points and covered an area of ca 1.3 ha. From every point in each of the seven plots, the nearest canopy gap ≥10 m² was located; 25 of the 35 gaps were formed by harvesting. Porewater salinity was significantly higher under the canopy in fringe mangrove forests than in basin mangrove forests. Although gaps were small (mean gap size = 158 m²; median gap size = 92 m²), soil temperatures were significantly higher in gaps of both zones. Soil redox potential was significantly lower and porewater salinity significantly higher in the gaps than under the canopy in the basin zone only. Higher porewater salinity may be attributed to high evaporation rates from the soil and high transpiration rates from trees surrounding gaps. There were significantly more seedlings in gaps than under the canopy only in the fringe zones. Although gap formation alters the soil environment of Kosraean mangrove swamps, high freshwater input may buffer these effects in basin mangrove swamps by reducing porewater salinity. Current harvesting rates do not appear to be changing canopy species composition, but large gaps, especially in mangrove forests in more arid areas, may lead to major changes.     

Distribution of spore-positive and spore-negative nodules ofAlnus incana ssp.rugosa in Maine,USA

The distribution of spore-positive (sp+) and spore-negative (sp−) root nodules ofAlnus incana ssp.rugosa (DuRoi) Clausen (speckled alder) was examined at 29 sites with a wide range of environmental conditions in Maine, USA. These included: pH 3.4 to 7.0, soil texture ranging from coarse gravel to clay to organic soils, elevation from 3 to 591 m and latitude 43 to 47°N. Habitat types included disturbed areas, streamsides, swamps and old fields. Sp (−) nodules were substantially more common, making up 76% of all nodules, whereas only 24% were sp (+). Sp (−) nodules often occurred in pure stands and predominated at disturbed sites with mineral soils at the surface and in old fields and swamps with pH>4.0 Sp (+) nodules were nearly always found in mixture with sp (−) nodules. They occurred primarily at streamside and lakeshore sites where they made up 40% of the nodules and at sites with pH<4.0 regardless of habitat type. It is suggested that sp (−) strains ofFrankia may be maintained at a site by saprophytic growth in soil and thus nodulate newly established hosts, whereas sp (+) strains may be maintained primarily by spore production within nodules and thus depend on extended presence of the host.     

Tracking Human Disturbance in Mangroves: Estimating Harvest Rates on a Micronesian Island

Disturbance is an integral component in mangrove forest dynamics, influencing forest structure, composition, and function. The impacts of human disturbance, however, threaten mangrove forests throughout the world. Small-scale wood harvesting on the small Pacific island of Kosrae, Federated States of Micronesia, provided an instructive scenario for exploring the dynamics of human disturbance. Natural disturbances on the island are rare, but the growing island population harvests mangrove trees for firewood and construction materials, placing pressure on the forest. In order to determine recent harvest rates, we estimated gap ages by developing a time scale for mangrove wood decomposition and by quantifying growth rates for Rhizophora apiculata and Bruguiera gymnorhiza seedlings. Stump and log decomposition patterns were useful in aging gaps, although some patterns were more reliable than others. Seedlings of both species added approximately 5 nodes/year depending on light conditions. The island-wide harvest rate was 10% over the last 10 years, but the rates varied widely among different parts of the island. Rhizophora apiculata has been harvested preferentially, and a dearth of young trees where harvesting has been heaviest portends a decline of this highly desired species in the forest. Socio-economic data substantiated some but not all of the trends we observed. Even on a small island, local differences in both natural and anthropogenic factors are important to understanding forest dynamics.     

Gap regeneration in primary evergreen broad-leaved forests with or without a major canopy tree,Distylium racemosum,southwestern Japan: A comparative analysis

Gap regeneration was studied in a typical primary evergreen broad-leaved forest withoutDistylium racemosum, at the Kasugayama Forest Reserve, southwestern Japan and the results were compared with those from other primary evergreen broad-leaved forests in southwestern Japan, whereD. racemosum was the dominant species. Several common types of gap regeneration behavior were recognized among the major tree species and forests with or withoutD. racemosum consisted of three typical regeneration guilds which could be detected in the principal component analysis.Castanopsis cuspidata frequently regenerated in gaps from saplings recruited before gap formation in the forest withoutD. racemosum, although elsewhere, in forests withD. racemosum, it lacked advanced regeneration and regenerated in gaps from saplings recruited after gap formation. Some evergreenQuercus had their regenerations in gaps of the forest withoutD. racemosum, although elsewhere, in forests withD. racemosum, evergreenQuercus might not regenerate. The results indicate that tree species may change their regeneration behavior depending on the presence or absence of another key dominant species. This suggests that the presence and the dominance of a potential competitor induces shifts in the regeneration niche of other coexisting tree species.     

Forest/environment relationships in Yosemite National Park,California, USA

Forest compositional patterns in Yosemite National Park, California, were related to environmental factors through numerical classification of forest types, arrangement of forest types along elevational and topographic gradients, and development of regression models relating basal area of common tree species to environmental variables. The eight forest types are differentiated primarily by elevation zone and secondarily by topographic setting. Lower montane forests (1200–1900 m) were divided into the Abies concolor/Calocedrus type occurring primarily on mesic sites and the Pinus ponderosa/Calocedrus type predominantly on xeric sites. Upper montane forests (1900–2500 m) included the Abies concolor/Abies magnifica type on mesic sites, the Abies magnifica/Pinus type on somewhat more xeric sites, and Juniperus occidentalis/Pinus jeffreyi woodlands on granitic domes. Subalpine forests (2500–3300 m) embraced three types: Tsuga mertensiana/Pinus forests on mesic sites, monotypic Pinus contorta forests on drier sites, and Pinus albicaulis/Pinus contorta groves at treeline. Regression models consistently included elevation and soil magnesium content as explanatory variables of species basal area totals. The two Abies spp. were negatively correlated with soil magnesium levels, whereas other montane species (e.g. Calocedrus decurrens, Pinus lambertiana, and Pinus ponderosa) exhibited positive correlation with soil magnesium. Topography and soil physical properties were only infrequently incorporated into species regression models.Abbreviations DBH= diameter at breast height (1.4 m) - DCA= detrended correspondence analysis - TWINSPAN= two-way indicator species analysis     

Distribution of spore-positive and spore-negative nodules in stands ofAlnus glutinosa andAlnus incana in Finland

Summary The distribution of spore positive (Sp⁺) and spore negative (Sp⁻) nodules on the two native alder species (A. incana andA. glutinosa) in Finland was investigated. Nodules were collected throughout the country from different ecosystems (forests, swamps, lake- sea- and riversides, old pastures and fields as well as from alder plantations). OnA. incana Sp⁺ nodules predominated, whereas onA. glutinosa the vast majority of the nodules were of the Sp⁻ type. Sp⁺ nodules onA. glutinosa were found only at sites where the two alder species grew close together. This distribution pattern indicates an association of nodule type with alder species, the reasons for which are discussed. Indications of saprophytic growth in the Sp⁻ strain were also found.     

Organic Matter Dynamics on the Forest Floor of a Micronesian Mangrove Forest: An Investigation of Species Composition Shifts1

Species composition shifts in mangrove forests may alter organic matter dynamics. The purpose of this study was to predict the effect of species replacements among mangrove trees on organic matter dynamics in a mangrove forest on the island of Kosrae, Federated States of Micronesia. We were particularly interested in elements of the carbon cycle that affect peat accumulation rates, organic matter exports to the estuary and coral reef systems, and soil microbiology. We compared organic matter production and decomposition rates among three mangrove species that commonly grow in similar hydrogeomorphic settings: Rhizophora apiculata BL, which is selectively harvested; Bruguiera gymnorrhiza, which may gradually replace Rhizophora; and Sonneratia alba, which is producing few mature fruits. Sonneratia had significantly higher rates of root production (estimated with ingrowth chambers) than Bruguiera or Rhizophora. Sonneratia foliage had significantly faster decomposition rates and significantly lower lignin:nitrogen ratios than Bruguiera foliage. Live root mass was positively correlated with ingrowth and soil carbon, although soil carbon and ingrowth were not significantly correlated with each other. Humic acid concentrations were significantly higher in Sonneratia rhizospheres than in either Bruguiera or Rhizophora rhizospheres and were positively correlated with root ingrowth. The species changes taking place on Kosrae are likely to result in lower rates of root production and foliage decomposition, and more refractory carbon pools in soil.     

湖南栎类天然次生林幼树更新特征及影响因子

以湖南典型栎类天然次生林为研究对象,基于51块样地的调查数据,采用k-means聚类分析划分林分类型,研究湖南不同栎类天然次生林幼树更新特征,分析了湖南不同栎类天然次生林幼树更新指标(幼树密度、幼树平均地径、平均高以及平均冠幅)与环境因子、林分因子的相关性,旨在阐明环境因子、林分因子对幼树更新的影响,以期为湖南不同栎类天然次生林的恢复与经营管理提供理论依据。结果表明:(1)利用聚类分析可将研究区内栎类天然次生林划分为5个类型,包括甜槠锥栗混交林(CC)、亮叶水青冈多脉青冈混交林(FC)、石栎樟树混交林(LC)、枹栎甜槠混交林(QC)、青冈栎混交林(CG)。(2)不同类型栎类天然次生林更新幼树优势种分化明显,物种丰富度差异显著(P0.05)。5种不同栎类次生林幼树密度均未超过500株/hm~2,更新情况较差;幼树数量差异显著(P0.05),为亮叶水青冈多脉青冈混交林石栎樟树混交林青冈栎混交林枹栎甜槠混交林甜槠锥栗混交林;生长情况差异显著(P0.05),为青冈栎混交林亮叶水青冈多脉青冈混交林枹栎甜槠混交林甜槠锥栗混交林石栎樟树混交林。(3)相关分析结果显示,不同类型次生林幼树更新的主要影响因子存在差异。甜槠锥栗混交林中幼树密度与腐殖质厚度呈显著负相关(P0.05);幼树平均高与灌木盖度呈显著正相关(P0.05);幼树平均地径与草本盖度、灌木盖度呈显著正相关(P0.05)。亮叶水青冈多脉青冈混交林中幼树密度与海拔、腐殖质厚度、枯落物厚度呈显著正相关(P0.05),与草本盖度呈极显著正相关(P0.01);幼树平均地径与郁闭度呈显著负相关(P0.05);幼树平均高、幼树平均冠幅与坡位呈显著正相关(P0.05)。石栎樟树混交林中幼树密度与坡向、土壤厚度呈显著正相关(P0.05),其余因子对幼树生长无显著影响。枹栎甜槠混交林中幼树密度与郁闭度、乔木密度呈极显著正相关(P0.01),与坡位呈显著负相关(P0.05);幼树平均冠幅与坡度呈显著负相关(P0.05)。青冈栎混交林中幼树平均地径与土壤厚度呈显著正相关(P0.05),与乔木密度呈极显著正相关(P0.01);幼树平均冠幅与灌木盖度呈显著正相关(P0.05)。     

Importance of fragmentation-tolerant species as seed dispersers in disturbed landscapes

Forest fragmentation can negatively affect plants if animal seed-dispersers become locally extinct in fragments. We conducted a 2-year experiment to evaluate the importance of tree squirrels (Sciurus) as seed dispersers for Quercus, Carya, and Juglans, and to assess dispersal consequences in patches where fragmentation-sensitive eastern gray squirrels (Sciurus carolinensis) are absent. We accounted for fates of ∼15,700 seeds from five tree species in four exclosure treatments at 18 fragments during a high (2003–2004) and low seed (2004–2005) year. Two treatments excluded Sciurus to mimic disperser loss. We sampled nut-tree seedling density at 259 sites across eight watersheds, half of which were too fragmented to support S. carolinensis, but supported fragmentation-tolerant fox squirrels (Sciurus niger). Autumn-to-spring seed survival was low (∼1%) for all species during low seed production. During high seed production, survival was higher for Juglans nigra (20%) and Carya ovata (16%) than for three Quercus species (∼4% for Quercus palustris and Quercus rubra in two exclosure types; ∼1% for Quercus alba in all treatments). Survival of J. nigra, C. ovata, and Q. rubra was ≥2.1–7.7 times higher for seeds in exclosures that Sciurus could access. Seed displacement distance was higher in the low seed than the seed-rich year, but the proportion of seeds surviving to greater distances was higher in seed-rich years for all seed types except Q. rubra. This affirms the importance of masting to seed survival and dispersal, but also suggests an advantage to trees of producing seed in non-mast years. Seedling densities were comparable in watersheds with and without S. carolinensis. These results demonstrate the importance of tree squirrels as dispersers of nut-bearing trees, but suggest that fragmentation may not disrupt dispersal of certain species if losing S. carolinensis from disturbed landscapes is compensated for by fragmentation-tolerant fox squirrels (S. niger).     

Genetically Distinct Populations in an Asian Soldier-Producing Aphid, Pseudoregma bambucicola (Homoptera: Aphididae), Identified by DNA Fingerprinting and Molecular Phylogenetic Analysis

To estimate genetic structure of a soldier-producing aphid, Pseudoregma bambucicola, samples from natural populations throughout southeastern Asia were analyzed by a DNA fingerprinting technique. We unexpectedly found that P. bambucicola comprises two geographic groups, the northern group and the southern group, which are genetically distinct from each other but morphologically almost indistinguishable. Molecular phylogenetic and statistical analyses based on mitochondrial ribosomal DNA sequences demonstrated that the northern and southern groups of P. bambucicola are not closely related but constitute distinct lineages in the genus Pseudoregma. Detailed morphological reexamination revealed that the two groups could be distinguished by the number of setae on the 8th abdominal tergite of 1st instar nymphs and soldiers. From these results, it was suggested that P. bambucicola should be divided into two species. The northern group from Japan, Taiwan, Hong Kong, and northern Vietnam retains the name P. bambucicola, whereas we suggest that the name P. carolinensis (R. Takahashi, 1941, Tenthredo 3, 208–220) should be used for the southern group from Thailand, Malay Peninsula, Java, Irian Jaya, and Micronesia. The morphological resemblance between P. bambucicola and P. carolinensis might be due to shared ancestral characters of the genus Pseudoregma.     

Differentiation of perennial and annual types due to habitat conditions in the wild riceOryza perennis

Populations of the common wild rice of AsiaO. rufipogon Griff. (=Oryza perennis Moench) were studied with regard to interrelations between life-history traits and habitat conditions. They showed a perennial-annual continuum and differed in reproductive allocation and many other traits. Perennial populations were found in deep swamps, while annual populations were in shallower, temporary swamps which were parched in the dry season. The perennial and annual types tended to be in association with other perennial and annual plants, respectively, suggesting their niche differentiation. Intermediate perennial-annual populations were in communities with a high species diversity. In one population, plants growing on the periphery of the swamp were of annual and those in deeper center were of perennial type. Another population seemed to be differentiated into an annual and an intermediate type in accordance with different degrees of habitat disturbance. The small genetic distances found between the sub-populations suggested their differentiation within a gene pool.Contribution from National Institute of Genetics, Japan, No. 1415.     

Nymphal survival and habitat distribution of Ixodes scapularis and Amblyomma americanum ticks (Acari: Ixodidae) on Fire Island, New York, USA

The distribution and survival of Ixodes scapularis and Amblyomma americanum were studied in deciduous and coniferous wooded habitats and in open habitats on Fire Island, New York, USA. The survival of nymphal I. scapularis in field enclosures was greater in forests than in open habitats, suggesting that greater survival contributes to the higher tick population in the woods. The nymphs of each species were more common in deciduous thickets (predominantly Aronia arbutifolia and Vaccinium corynbosum) than in coniferous woods (mostly Pinus rigida) in most but not all years. Larval I. scapularis were more common in coniferous sites in 1994, while the same ticks, as nymphs, were more common in deciduous sites in 1995. The survival of the nymphs was not consistently greater in either the deciduous or coniferous woods. Therefore, factors other than nymphal survival (e.g. larval overwintering survival and tick movement on hosts) probably influenced the relative nymph abundance in different forest types. Overall, the survival of A. americanum was far higher than that of I. scapularis.     

Occurrence and importance of mycorrhizae in aquatic trees of New South Wales,Australia

Vesicular arbuscular mycorrhizal (VAM) infection was found in KOH-cleared and lactophenolblue-stained roots of Salix babylonica, Melaleuca quinquenervia and Casuarina cunninghamiana. These are all trees growing on creeks and river banks, in stationary or slowly flowing fresh or brackish waters in swamps, creeks, drains and channels, and in seepage areas of New South Wales, Australia. Larger and older roots lacked VAM infection in the inner cortex, probably due to suberisation of cells, and the endophyte was restricted to the epidermal layers. Spores and sporocarps of the VAM fungi Glomus fasciculatus, G. mosseae, Sclerocystis rubiformis, Gigaspora margarita and an unidentified Scutellospora sp. were wet sieved and decanted from aquatic sediments and soils. The presence of similar VAM fungal spores in the aquatic sediments and terrestrial soil suggests that they probably enter the aquatic sediments through run off from the land ecosystem. All three plants formed vesicular arbuscular (VA) mycorrhizae almost exclusively in the marshy, periodically inundated soils, but the same plant species formed endo-/ ectomycorrhizae when growing in soil with higher redox potentials (E _h). Salix and Melaleuca tree roots possessed both VAmycorrhizae and ectomycorrhizae. VAM roots of Casuarina were equipped with both N-fixing Frankia nodules and proteoid roots. VAM endophytes did not invade nodular cortical tissues, suggesting the presence of an exclusion mechanism which needs further study. The highest VAM infection was found in nodulated specimens. Free-floating roots growing in water close to the banks were non-mycorrhizal but were mycorrhizal in the bottom-rooting state. VAM spore number and mycorrhizal infection seem to be associated with redox-potential, i.e. lower at sites such as swamps, water or sediments with lower E _h values than in terrestrial soils with higher E _h values. A relationship between soil moisture gradient and VAM infection pattern became apparent from the study of a C. cunninghamiana transect on a creek embankment, i.e. typical vesicles and arbuscules were found in roots from drier soils, there was a lack of arbuscules in relatively wet soils but large lipid-filled intracellular vesicles were present, and typical vesicles and arbuscules were absent in flooded creek beds where roots were associated with coenocytic intercellular hyphae with abundant lipid droplets. The importance of VA mycorrhiza, ectomycorrhizae, N-fixing root nodules and proteoid roots at the land-water interface is discussed with reference to the use of these trees as pioneering species for stabilising river and stream banks, reducing erosion, windbreaking, and as a long-term and inexpensive means of achieving biological control of aquatic weeds by shading waterways.     

北京地区油松人工林不同演替类型空间结构对林下植被及土壤的影响

以北京地区油松(Pinus tabuliformis)人工林不同演替类型林分为研究对象,研究油松纯林、油松-栓皮栎(Quercus variabilis)混交林和栓皮栎纯林三种不同演替类型林分的空间结构、林下植被和土壤水分的变化规律及其相互作用关系。结果表明：(1)林分水平及垂直空间结构、草本层物种多样性、更新幼树生长、土壤持水和透气性能等指标在三种不同演替类型林分间差异显著(P<0.05),林分空间结构参数中的角尺度、林层指数和开敞度显著影响了各类型林分的灌草多样性,混交度、林层指数和大小比数显著影响了更新幼树的生长,混交度和林层指数显著影响了土壤水分的变化(P<0.05)。(2)松栎混交林灌草生物量、天然更新幼树的生长以及土壤水分物理状况均好于纯林,并主要受林分混交度和林层指数的共同作用。(3)各演替类型林分内均存在栓皮栎更新幼树,混交林栓皮栎更新幼树数量最多、长势最好,对林地资源的竞争最为激烈。因此,可以通过调整林分空间结构实现种间关系及林地资源的调控,以充分发挥森林生态系统的各项功能与价值。     

Successional trends and apparent <Emphasis Type="Italic">Acer saccharum</Emphasis> regeneration failure in an oak-hickory forest in central Missouri,USA

Permanent vegetation plots established in 1968 at the Baskett Research and Education Area (BREA) were remeasured in 2004/2005 to assess trends in canopy composition, vegetation–environment relationships, and regeneration patterns. Quercus spp. [particularly Quercus alba (white oak)] remained dominant in BREA forests. However, Nonmetric Multidimensional Scaling Ordination of a dataset composed of tree importance values from both samples indicated the two most common community types (Dry Ridge and Slope; Mesic Slope) were shifted away from Quercus spp. abundance toward greater importance of Acer saccharum (sugar maple) in ordination space. Plots in glade-like environments shifted toward greater Juniperus virginiana (eastern redcedar) importance. Environmental analysis indicated that soil properties (pH, organic matter, Ca, Mg, and K) and slope position were highly correlated with ordination axes. Saplings of Quercus spp. exhibited greatly reduced abundance in the most recent sample and the reduction in Quercus spp. density was highest in plots with greatest canopy cover. While Acer saccharum saplings were still quite abundant in BREA forests, smaller sapling size classes were becoming notably depleted, reflecting growth of a cohort of plants established before the 1968 sample toward and into tree-sized individuals. Seedlings surveys emphasized that A. saccharum seedlings have become very rare in BREA forests, while Quercus spp. seedling densities are higher, but likely below those needed for eventual adequate recruitment to the canopy. Although a number of mechanisms to explain apparent A. saccharum regeneration failure can be developed, herbivory, recent climate trends (warming late-winter conditions) and increased litter layer depth are the most plausible explanations for the observed scarcity of A. saccharum seedlings. Although increasing importance of A. saccharum in the canopy of BREA forests appears inevitable in coming decades, longer-term trends are unclear given the shortage of seedling regeneration.     

文峪河上游华北落叶松林的种子雨、种子库与幼苗更新

华北落叶松林下更新不良,为探究其制约因素,开展了山西省文峪河上游5个华北落叶松林分的种子雨、土壤种子库和幼苗更新的研究。结果表明:(1)华北落叶松种子主要集中于9—10月散落。2011年为华北落叶松种子丰年:种子产量高,种子雨密度达(961.93±377.40)粒/m2;种子质量高,完整种子占(89.31±16.13)%。2012年为种子平年,种子产量低,种子雨密度为(252.73±115.12)粒/m2。华北落叶松种子雨主要源于毗邻树木,华北落叶松纯林和落叶松云杉林的种子雨密度显著高于其他3个针阔混交林。(2)土壤种子库主要由上年种子雨组成,2012年4月的土壤种子库密度为(695.18±297.23)粒/m2,完整种子占(59.73±9.56)%。种子自然萌发前,约(78.98±24.76)粒/m2具发芽力,基本可满足更新需要。但种子活力保持期少于2 a,只能形成短期持久土壤种子库。(3)华北落叶松更新不良,种子年后仍难以实现幼苗建成,当年生幼苗的出现频度平均为1.6%,且林下难以存活。幼苗发生与种子储量关联性不强,种源条件不是制约华北落叶松更新的主要因素。