Takuya Kajimoto

Estimation of root biomass based on excavation of individual root systems in a primary dipterocarp forest in Pasoh Forest Reserve, Peninsular Malaysia.

Precise estimation of root biomass is important for understanding carbon stocks and dynamics in tropical rain forests. However, limited information is available on individual root masses, especially large trees. We excavated 121 root systems of various species (78) and sizes (up to 116 cm in dbh), and estimated both above- and below-ground biomass in a lowland primary dipterocarp forest in the Pasoh Forest Reserve, Peninsular Malaysia. A tree census was conducted in four research plots (each 0.2 ha) and stand-level biomass was estimated. We examined relationships between tree size parameters and masses of coarse roots (roots ≥ 5 mm in diameter) and derived a dbh-based allometric equation. The amounts of coarse roots that were lost during excavation were corrected. Coarse-root biomass before and after correction for lost roots was estimated to be 63.8 and 82.7 Mg ha -1 , indicating that significant amounts of roots (23%) were lost during the sampling. We also estimated the biomass of small root (<5 mm) by applying pipe-model theory. The estimate, 13.3 Mg ha -1 , was similar to another estimate of small roots, 16.4 Mg ha -1 , which was obtained directly by the soil-pit sampling method. Total below-ground (BGB) and above-ground biomass (AGB) was estimated to be 95.9 and 536 Mg ha -1 , respectively. The biomass-partitioning ratio (BGB/AGB) was about 0.18. In conclusion, the dbh-based allometric equation for coarse roots developed in this study, which kept good linearity even including the data of larger trees, might be useful for evaluating below-ground carbon stocks in other stands of similar forest (old-growth dipterocarp) in South-East Asia.

Root system development of Larix gmelinii trees affected by micro-scale conditions of permafrost soils in central Siberia

Spatial distributions of root systems of Larix gmelinii (Rupr.) Rupr. trees were examined in two stands in central Siberia: an even-aged stand (ca. 100 yrs-old) and a mature, uneven-aged (240–280 yrs-old) stand. Five larch trees of different sizes were sampled by excavating coarse roots (diameter > 5 mm) in each stand. Dimensions and ages of all first-order lateral roots were measured. Micro-scale conditions of soil temperature and soil water suction (each 10 cm deep) were also examined in relation to earth hummock topography (mound vs. trough) and/or ground floor vegetation types (moss vs. lichens). All larch trees developed superficial root systems, consisting of the aborted short tap root (10–40 cm in soil depth) and some well-spread lateral roots (n = 4 – 13). The root network of each tree was asymmetric, and its rooting area reached about four times the crown projection area. Lateral roots generally expanded into the upper soil layers of the mounds where summer soil temperature was 1–6 °C higher than inside nearby troughs. Chronological analysis indicated that lateral root expansion started successively from lower to upper parts of each aborted tap root, and some lateral roots occurred simultaneously at several decades after tree establishment. The process of root system development was likely to be primarily linked with post-fire dynamics of rhizosphere environment of the permafrost soils.

Effects of Snowfall Fluctuation on Tree Growth and Establishment of Subalpine Abies mariesii near Upper Forest-limit of Mt. Yumori, Northern Japan

We examined effects of annual snowfall fluctuation on tree growth and establishment in a mature stand (ca. 200 yr old) of Japanese subalpine Abies mariesii Mast, near upper forest-limits at leeward site. Relatively tall and aged fir trees (>5 m, >100 yr old) retained various scars of past mechanical damage due to snow pressures, e.g., multiple-, tip dieback- and broken-stems, and canopy anomaly. Ring-width pattern analysis showed that these deformed trees mostly experienced abrupt growth reduction. Some events of such abrupt growth reduction occurred simultaneously just after winters with heavy snowfall (e.g., mid-1960s), indicating that the A. mariesii trees often suffered from snow damage due to settlement force of excess snowpack. Particularly, in snowy winters, mechanical breakage of branches was likely to occur intensively at the height of 1 to 3 m above the maximum snowpack level of normal snowfall winters (ca. 3 m). On the other hand, some aged, but suppressed, trees sharply enhanced their growth rates simultaneously after such period of the intensive snow damage. This indicated that the snow damage upon the relatively tall trees sometimes facilitated regeneration of nearby shaded-trees by creating small-scale gaps. The snow-induced disturbance events may have occurred at about 10-yr intervals during the last half of 20th century. Our findings suggested that population dynamics of A. mariesii near its upper forest-limit, especially located at leeward and snowy site, was closely affected by the repetitive events of snow damage following snowy winters: the event primarily acted as a major limiting factor for canopy development of individuals, and sometimes triggered growth of suppressed-trees.

Individual-based measurement and analysis of root system development: Case studies for Larix gmelinii trees growing on the permafrost region in Siberia

We present results of individual-based root system measurement and analysis applied for Larix gmelinii trees growing on the continuous permafrost region of central Siberia. The data of root excavation taken from the three stands were used for the analyses; young (26 years old), mature (105 years old), and uneven-aged over-mature stand (220 years old). In this article, we highlight two topics: (1) factors affecting spatio-temporal pattern of root system development, and (2) interactions between aboveground (i.e., crown) and belowground (i.e., root) competition. For the first topic, the detailed observation of lateral roots was applied to one sample tree of the overmature stand. The tree constructed a superficial (<30 cm in depth) and rather asymmetric root system, and each lateral root expanded mainly into elevated mounds rather than depressed troughs. This indicated that spatial development of an individual root system was largely affected by microtopography (i.e., earth hummocks). For these lateral roots, elongation growth curves were reconstructed using annual-ring data, and annual growth rates and patterns were compared among them. The comparison suggested that temporal root system development is associated with differences in carbon allocation among the lateral roots. For the second topic, we examined relationships between individual crown projection area (CA) and horizontal rooting area (RA) for the sample trees of each stand. RA was almost equal to CA in the young stand, while RA was much larger (three or four times) than CA in the mature and overmature stands. Two measures of stand-level space occupation, crown area index (aboveground: CAI; sum of CAs per unit land area) and rooting area index (belowground: RAI; sum of RAs), were estimated in each stand. The estimates of RAI (1.3–1.8 m2 m−2) exceeded unity in all stands. In contrast, CAI exceeded unity (1.3 m2 m−2) only in the young stand, and was much smaller (<0.3 m2 m−2) in the two older stands. These between-stand differences in RAI–CAI relationships suggest that intertree competition for both aboveground and belowground spaces occurred in the young stand, but only belowground competition still occurred in the two older stands. Based on this finding, we hypothesized that competition below the ground may become predominant as a stand ages in L. gmelinii forests. Methodological limitations of our analysis are also discussed, especially for the analysis using the two indices of space occupation (CAI, RAI).

Effects of Snow Avalanche Disturbance on Regeneration of Subalpine Abies mariesii Forest, Northern Japan

Abstract We examined the effects of snow avalanche on stand structure and the subsequent regeneration of subalpine Abies mariesii Mast forest in northern Japan, and discussed whether gap formation after avalanche disturbance leads to immediate seedling establishment or not. Tree size and age, tree-ring chronology, and seedling density were compared between, on, and around a relatively large avalanche path, which was created in the mid-1980s within Hachimantai National Park. On the avalanche path, only smaller and younger trees (height <5 m, 50–100 yr old) escaped the mechanical damage of avalanche by leaning into the remaining snowpack, and old canopy trees were mostly killed by stem breakage. Tree size structure of the avalanche path, including both living and dead individuals, was relatively similar to that of nearby undisturbed forest, indicating that a mature stand (>ca. 200 yr old) had been previously developed on the avalanche slope without large-scale disturbances in the past. Most of the surviving, younger trees showed abrupt growth release during the few years (1987–1989) after the avalanche event. However, densities of both post-avalanche (<15 yr old) and pre-avalanche (≥15 yr old) seedlings were much lower on the avalanche path than in the nearby forest, especially at microsites covered with dense dwarf bamboo (Sasa kurilensis) shrub. The pre-avalanche seedlings on the avalanche path sharply increased annual height growth rates after the mid-1980s avalanche, but such positive growth response did not continue longer than 6 to 7 yr. These findings suggested that gap formation following the infrequent, large-scale avalanche disturbance did not necessarily lead to immediate seedling recruitment and/or further growth release of pre-avalanche seedlings. The constraint of seedling establishment was primarily explained by the lack of potential seed supply, and shading effect of the dwarf bamboo bush. Consequently, post-avalanche regeneration of the subalpine fir forest was likely to depend on the smaller individuals (2–5 m in height) that were able to avoid both the mechanical damage of avalanche and mortality from shading by the dwarf bamboo.

Winter climate change in plant–soil systems: summary of recent findings and future perspectives

The winter climate is changing in many parts of the world, and it is predicted that winter climate change will modify the structure and function of plant–soil systems. An understanding of these changes and their consequences in terrestrial ecosystems requires knowledge of the linkage between above- and below-ground components as well as the species interactions found in plant–soil systems, which have important implications for biogeochemical cycles. However, winter climate-change studies have focused on only a part of the ecosystem or ecological process. We summarize here recent findings related to the effects of winter climate and its changes on soil nitrogen (N) dynamics, greenhouse gas (N2O) emissions from the soil, N use by individual plants, vegetation development, and interactions between vegetation and pollinators to generate an integrative understanding of the response of the plant–soil system to winter climate change. This review indicates that the net effects on plants, soil microbes, pollinators, and the associated biogeochemical cycles are balanced among several processes and are highly variable depending on the context, such as the target species/functional group, original winter condition of the habitat, and type of climate change. The consequences of winter climate change for species interactions among plants, associated animals, and biogeochemical cycles are largely unknown. For further research, a large-scale comparative study to measure ecosystem-level functions is important, especially in less-cold ecosystems.

Long-term growth analyses of Japanese cedar trees in a plantation: neighborhood competition and persistence of initial growth deviations

The individual growth of tree diameter at breast height (dbh) is analyzed in an even-aged plantation of Cryptomeria japonica from stand age of 45 to 94 years, to examine how the growth of individual trees has been affected by the changes in spacing resulting from thinning operations. At any age, a significant proportion (0.37–0.46) of the variation in dbh growth during a 5–11-year period was explained by dbh at the beginning of the period, probably due to greater leaf mass of larger trees. Next, either one-sided or two-sided competition was added to the model, by calculating the basal area (BA) of neighboring trees around each tree within a given radius or BA for trees having larger dbh than the focal tree within the radius. After preliminary analyses, a radius of 8 m was selected as the critical range for tree competition. Although both types of competition explained a significant proportion (0.09–0.43) of growth variation, one-sided competition was not significant at ages greater than 54 years. Based on the model at 45 years of age, the initial deviation of growth rate for each tree from the predicted rate was calculated and added to the models as a third variable. This raised the coefficient of determination up to 0.50–0.74. These findings have practical significance for forest plantation management, particularly for controlling the growth of standing trees via thinning, to produce high-quality timber in the future.

Factors affecting seedling recruitment and survivorship of the Japanese subalpine stone pine, Pinus pumila, after seed dispersal by nutcrackers

To clarify the factors affecting the seedling establishment of the subalpine stone pine (Pinus pumila Regl.) from nutcracker-cached seeds, subsequently emerged seedlings were examined in two plots on Mt Akitakomagadake, northern Japan. The survivorship of older seedlings (2–20 years old) was also monitored during six seasons at nine different sites. In one plot with 18 caches, seed germination occurred between June and mid-July. During this period, nutcracker’s retrieved the stored pine seeds from 16 caches and ate some seeds immediately. Simultaneously, this nutcracker behavior caused mechanical damage to newly emerging seedlings (e.g. uprooting and tearing off cotyledons). Such initial loss to in situ harvesting and mechanical damage accounted for 75% of the total seeds remaining in the caches. The number of established current seedlings sharply declined during the first 2 years, and the survival rate was 4.4% over four winters. Two major mortality factors were identified: uprooting by frost-heave soil disturbance in the spring of the second year and standing death by drought or other physiological stresses in early summer. In another plot with 13 caches, survivorship of newly emerged seedlings was also low (4.2%), but mortality was mostly due to summer drought, indicating that the frost-heave event was a site-specific disturbance factor. For older seedlings, survival rates reached approximately 90% even after six seasons and summer drought stress was a major mortality factor. My findings suggest that seedling recruitment of P. pumila was largely limited by both nutcracker disturbance and external disturbance (and/or stresses) at an early stage. However, after the critical first few years, pine seedlings were highly likely to survive and grow to the sapling stage despite the harsh environment of the high mountains.

Mechanical Damage on Abies mariesii Trees Buried below the Snowpack

Abstract On a gentle leeward slope in a snowy forest limit in northern Honshu Island, Japan, mechanical damage by snow settlement and creep on Abies mariesii trees buried below the snowpack was examined to detect signs of the snow-damage effect on future survival and crown development. Damage types were recorded based on direct observation of crowns in 1996, a year of high snow accumulation exceeding 4.5 m, and 1997, a year of moderate snow accumulation. Of 153 trees examined, 63% were damaged in 1996 and 15% were damaged in 1997. The most destructive damage type was breakage of stems ≥5 cm in diameter, which occurred on eight trees in 1996 and three in 1997, resulting in foliage loss and death of some trees. The prevalent damage type was branch tearing at branch-stem junctions primarily within a height range of 4–6 m, which occurred on 171 branches in 1996 and 5 in 1997. Under snowy and windy conditions, stem breakage and branch tearing, caused by forces active within restricted layers of the snowpack, may reduce the future survival and crown development of A. mariesii buried below the snowpack in years of heavy snowfall.

Biomass and Productivity of Siberian Larch Forest Ecosystems

Boreal forests are expected to affect global carbon balance significantly due to their large areas (e.g., Goulden et al. 1998; Schulze et al. 1999; Chapin et al. 2000; Jarvis et al. 2001). Biomass and productivity of boreal forests have been examined mainly in the evergreen taiga established in the regions of nonpermafrost or discontinuous permafrost, e.g., interior Alaska (Van Cleve et al. 1983, 1986; Vogt et al. 1996), northern Canada (Gower et al. 2001; Bhatti et al. 2002; Bond-Lamberty et al. 2002), and northern Europe (Schulze 2000). In contrast, comparable studies are considerably limited in the boreal forests of Siberia, where three Larix species are growing separately in different locations and soils: each of two similar species, Larix gmelinii (Rupr.) Rupr. and L. cajanderi Mayr, dominates in the continuous permafrost regions in Central and Northeastern Siberia, respectively, while Larix sibirica L. grows mainly in the nonpermafrost regions of Western or Mountainous southern Siberia (Abaimov 1995; also see Chaps. 1, 3 and 4).