Distribution of different mycorrhizal classes on Mount Koma, northern Japan

To investigate the role of mycorrhizae in nutrient-poor primary successional volcanic ecosystems, we surveyed mycorrhizal frequencies on the volcano Mount Koma (42°04N, 140°42E, 1,140 m elevation) in northern Japan. After the 1929 eruptions, plant community development started at the base of the volcano. Ammonia and nitrate levels, along with plant cover, decreased with increasing elevation, whereas phosphorus did not. In total, 305 individuals of 56 seed plant species were investigated in three elevational zones (550–600 m, 650–700 m, and 750–800 m). Five mycorrhizal classes were classified based on morphological traits: ecto- (ECM), arbuscular (AM), arbutoid, ericoid, and orchid mycorrhiza. All plant species were mycorrhizal to at least some extent, with most widespread tree species being heavily ectomycorrhizal. In addition, of 16 tree species collected in all three zones, 6 differed in the frequencies of ECM on roots between elevational zones, and 3 of these 6 species increased in frequency with increasing elevation. These results suggest that ECM colonization in some tree species is related to establishment in nutrient-poor habitats. All species of Ericaceae and Pyrolaceae had ericoid mycorrhizae, and an Orchidaceae species had orchid mycorrhizae. Herbaceous species, except for the low mycorrhizal frequency of Carex oxyandra and two Polygonaceae species, and ericoid and orchid mycorrhizal species, were generally AM. Of herbaceous species, Anaphalis margaritacea var. angustior increased AM frequency and decreased ECM frequency with increasing elevation, and Hieracium umbellatum increased ECM frequency. In total, the establishment of herbaceous species was not sufficiently explained by AM colonization on roots. Tree individuals developed 2–3 classes of mycorrhizae more than herbs at each elevational zone. We conclude that the symbiosis between seed plants and mycorrhizae, ECM in particular, greatly influences plant community structures on Mount Koma. Not only a single mycorrhizal class, but combinations of mycorrhizal classes should be studied to clarify effects on plant community dynamics.     

Plant community dynamics on the volcano Mount Koma, northern Japan, after the 1996 eruption

A small eruption occurred in the early spring of 1996 on the summit of Mount Koma, northern Japan, 67 years after the 1929 catastrophic eruption. To identify damage on the plant communities and recovery patterns, we established 400 50 × 50 cm permanent plots in four locations along the gradient of the thickness of tephra produced in 1996: non-disturbed (N), weakly disturbed (W), middle disturbed (M), and heavily disturbed (H). Annual monitoring was conducted from 1996 to 2001. Gullies and rills formed in M and H, indicating that the ground surface movements were more intense there. Mean vascular plant species richness increased from H to N, but did not increase from 1996 to 2001 in any location. In an area where tephra thickness was 10 cm, the recovery was mostly conducted by the species that could have survived during the eruption. A shrub,Salix reinii, was the leading species in most sites throughout most years. Large perennial herbs,Polygonum weyrichii, P. sachalinense andMiscanthus sinensis were common in the disturbed areas, in particular in M and H. Those three species develop large underground organs and enlarge clonally, suggesting that the tolerance to ground surface stability is the most important trait for the recovery of those species.Carex oxyandra established not only by vegetative regeneration but also by seedling regeneration. A short forb,Campanula lasiocarpa, could establish only in W by seedling regeneration. Mosses and lichens were predominant in plots in N and W, but less represented and not greatly increasing their cover in M and H up to 2001. We concluded that the community recovery was delayed in all the disturbed areas, mostly due to low seedling establishment. Only a few specific species established by vegetative reproduction. Disturbance gradients such as thickness of tephra and/or ground surface stability have determined plant community structure and development.     

Differential establishment and survival of species in deciduous and evergreen shrub patches and on bare ground,Mt. Koma,Hokkaido, Japan

The effects of patches formed by a deciduous shrub (Salix reinii) and an evergreen shrub (Gaultheria miqueliana) on cohabitants were examined on an active volcano Mt. Koma, northern Japan, using bareground as a comparative baseline (control). Species richness increased with increasing the areas of three habitats (Salix patches, Gaultheria patches and bareground), but significantly slowed in Gaultheria patches where the shoot densities of cohabitants also decreased. Calamagrostis hakonensis and Spiranthes sinensis were 2–3 times more likely to establish in Salix patches than in bareground, Anaphalis margaritacea var. angustior and Campanula lasiocarpa were 4–5 times more likely to establish on bareground than in Salix patches. Cohabitants was fewer with Gaultheria. Light intensities of the two patch types were ca 1/3 of that on bareground. Light intensity on Gaultheria was ca 88% of that on Salix patch and decreased with increasing patch size, because the differences in stem and leaf architectures and phenology between the two patch-forming species. Moisture and total nitrogen in volcanic deposits were 2–3 times higher in the patches than in bareground. In the patches, low light intensity restricted seed germination, in particular, for the plants characteristic of bareground, e.g., Anaphalis, while high soil moisture increased the survival rates of Calamagrostis. In Gaultheria patches, despite high moisture, the reduced light intensity influenced seed germination and inhibited the establishment of most species. Seedbanks estimated by a lab-experiment indicated that Salix patches trapped more seeds. As a consequence, Salix patches act as facilitator, in particular, for Calamagrostis, by improving moisture and nutrients status of volcanic deposits, moderating light intensity and seed trapping. Deciduous shrub patches, such as Salix reinii, have an improving role in accelerating rates of succession in harsh environments, while evergreen shrub patches slow plant establishment. Nomenclature: Ohwi and Kitagawa (1983)     

Natural regeneration patterns of the introduced larch, Larix kaempferi (Pinaceae), on the volcano Mount Koma, northern Japan

The 1929 eruption of Mount Koma (1140 m in altitude) completely deforested the vegetation on the summit area of the volcano. An alien species of larch, Larix kaempferi (=L. leptolepis) was planted on the lower slopes of the volcano between 1953 and 1963, but since then has become abundant on the summit. To determine the regeneration patterns of the larch, we measured stem densities on aerial photographs taken in 1963, 1973 and 1994, and in the field during 1996. Larix kaempferi stems with crown diameters > 2 m were mapped on aerial photographs divided into 1083 100 m×100 m grids. Stem densities on the summit increased from 0.8/ha in 1963, to 14.1/ha in 1994. Willows and birches also established there but did not grow as large as the larch. All these species are wind dispersed, and larch densities were higher in areas closer to the plantations, suggesting that wind intensity and direction determined seed migration. Four environmental factors—slope gradient, direction, elevation, and distance from the plantations in each grid—were correlated with larch stem densities. Multiple regression showed that establishment patterns of L. kaempferi in the early stages were mainly related to distance from seed source (the plantations). Later, geographical disturbances and/or physiological stresses became more important. Density effects on tree invasion have been weak until now. We concluded that revegetation primarily depends on the chance of seed immigration, and that the larches may be an earlier successional species than any other native tree species.     

Influence of a non-native invasive tree on primary succession at Mt. Koma, Hokkaido, Japan

The slopes of Mt. Koma in Japan are undergoing primary succession as a result of a 1929 eruption. Understory vegetation below a non-native invasive tree species, Larix kaempferi, a native tree, Betula ermanii, and in the open were compared to determine if the non-native tree species was influencing species composition. Larix canopies are significantly larger than Betula canopies. Vegetation under Larix canopies had significantly greater richness and diversity than vegetation in the open, vegetation under Betula was intermediate but was significantly greater than the open in diversity. Vegetation cover was highest under Betula and significantly lower in the open. Larix canopy size was positively correlated with size and number of Salix reinii shrubs. Betula canopy size was positively correlated with size but not with number of Salix reinii shrubs. Species assemblages in the three sites are slightly different as shown by DCA. Due to the limited species pool on Mt. Koma the greatest possible extent of differences between the three microsites is not large. At this point Larix certainly appears to be accelerating succession for the non-tree species. If Larix persists on the slopes then succession would be permanently deflected towards a Larix forest. This would be a case of succession being deflected towards dominance by the introduced species. This revised version was published online in June 2006 with corrections to the Cover Date.     

Distribution of plants in relation to microsites on recent volcanic substrates on Mount Koma,Hokkaido, Japan

On Mount Komas volcanically devastated summit six common plant species colonized 12 microsite types differentially. Flat sites covered 65%, rills and gullies 16% and biotic sites 13% of the study area. Most species preferred sites near rocks and avoided flat and biotic microsites. Polygonum and Salix preferred gully bottoms and edges. Polygonum and Salix seedlings avoided flat areas and Polygonum seedlings strongly colonized gullies. Carex oxyandra seedlings preferred Salix patches. The distributions of seedlings and adult plants were correlated. Biotic microsites supported the largest Carex individuals, rills supported the largest Agrostis individuals and the largest Salix individuals were in gully bottoms and flats.     

Origin of plants recovering on the volcano Usu,northern Japan,since the eruptions of 1977 and 1978

The vegetation near the summit of the volcano Usu was destroyed during eruptions in 1977 and 1978 by 1–3 m thick layers of volcanic deposits. Thereafter, the vegetation gradually recovered and by 1984 134 plant species were recognized of which 95 species had established by vegetative reproduction, 18 by seed immigration from elsewhere, 5 by artificial introduction for soil erosion control and 17 species from viable seeds buried in the former topsoil. The summit area was still covered by thick layers of volcanic ash, which were however rapidly eroded by rains. Vegetatively reproducing plants such as Petasites japonicus var. giganteus and Polygonum sachalense contribute more effectively to the revegetation process than plants of other types of origin.     

Tree regeneration patterns on Mount Usu, northern Japan, since the 1977–78 eruptions

Although the summit of Mount Usu was deforested by the 1977–78 eruptions, vegetative regeneration on the caldera rim was rapid due to the erosion of thick volcanic deposits by snow and rain. To obtain the mechanisms underlying regeneration patterns after the eruptions, we monitored the growth of permanently-marked stems from 1983 to 1990. Regeneration was from resprouting-branches buried in the volcanic deposits on the caldera rim, while on the crater basin, where thick volcanic deposits accumulated, regeneration was from seedlings. The seedling regeneration lagged approximately 3 years behind vegetative regeneration. Stem densities averaged 14,000 ha^-1 in the vegetatively-regenerated community on the caldera rim, and 28,000 ha^-1 in the seedling regeneration on the crater basin. Populus maximowiczii accounted for ca. 75% of total stems on the caldera rim, while P. maximowiczii accounted for ca. 30% on the crater basin where Salix integra and Betula platyphylla var. japonica were also common. In both stands, immigration and mortality rates were very low. The growth of vegetatively regenerated stems expressed as stem height and diameter was significantly faster than that of stems grown from seedlings. Herbivory damage on the terminal shoots of tall stems was restricted on the caldera rim and was restricted for B. platyphylla var. japonica on the crater basin, perhaps due to fast growth supporting herbivore avoidance or low palatability. Height growth was restricted when neighbors established in close proximity, especially in the seedling-regenerated forest. The results suggest that vegetative regeneration is rapid due to three mechanisms: 1) faster plant growth; 2) herbivore avoidance; and 3) decreased interference by neighboring.     

An experimental test of the effects of food resources and hydraulic refuge on patch colonization by stream macroinvertebrates during spates

1. The passive or active movement of organisms between habitat patches plays important roles in achieving ecosystem resilience to disturbance and long-term control of population levels. However, causal mechanisms of disturbance-induced movements of mobile biota across heterogeneous habitat patches at a relatively short time-scale are little understood. 2. We experimentally tested the effects of food resource values on macroinvertebrate colonization of hydraulic refugia from spates in a second-order creek. Experimental cages were colonized by macroinvertebrates with combinations of resource types (natural or polyester leaves), and extent of exposure to stream flow (exposed to or sheltered from current); one half of each set was collected before and after a spate. This experiment was repeated over three spates of varying magnitude and seasonal contexts. 3. Pre-spate colonization was consistently greater for the cages with natural leaves relative to artificial leaves regardless of the extent of flow exposure. Two autumn spates with relatively low and stable antecedent flow conditions caused large movements of organic matter and macroinvertebrates across the stream, showing community-level accumulations into hydraulically sheltered patches independent of food treatment. The smallest spate with high and variable antecedent flows during winter resulted in negligible responses, which we interpret to be a result of depletion of easily transportable organic matter and organisms. 4. Two detritivorous taxa, the mayfly Paraleptophlebia spp. and stonefly Despaxia augusta (Banks) were the most responsive to autumn spates, and had disproportionately higher colonization rates of cages when provided with natural leaves during the largest autumn spate. Preferential settlement in food-enriched hydraulic refugia was attributable to taxon-specific mobility related to efficient acquisition of detritus resource, whose availability varies spatially and temporally. 5. Our findings suggest (1) detritivorous macroinvertebrate colonization of hydraulic refugia can be influenced by hydraulic controls as well as food resource value, and (2) pre-spate environmental conditions in terms of resource distribution and availability may pre-condition organisms' susceptibility to spates and also affect refugium usage, at least in food-limited, detritus-based stream systems.     

Effects of resource distribution, patch spacing, and preharvest information on foraging decisions of northern bobwhites

We investigated patch assessment by northern bobwhites (Collinusvirginianus) in an experimental arena where the distributionof resources in patches, preharvest information about thesepatches, and spacing of patches varied. We found that preharvestinformation about patch quality and a bimodal distribution ofpatch rewards allowed birds to selectively exploit patches highin resources. In contrast, uniform distribution of patch qualitiesand lack of preharvest information caused birds to forage nonselectivelyamong patches. Birds distinguished among patches of differentquality when these patches were spaced 13 m apart, but failedto react to patch quality differences when patches were 0 or3 m apart We also found a strong effect of the level of patchdepletion on foraging decisions: as resources in die arena becamescarce, birds increasingly foraged selectively in die most profitablepatches. Foraging decisions of bobwhites are biased by die waythey experience and memorize a spatially and temporally variableenvironment. The relative cost of this cognitive bias (i.e.,lost opportunity) is nonlinearty related to die mean resourcedensity in die environment and to die difference between thismean density and die resource density in die exploited patch.Cognitive bias should be considered when evaluating patch assessmentcapabilities of foragers in complex environments.     

Woody plant establishment during the early stages of volcanic succession on Mount Usu, northern Japan

The establishment patterns of woody plants were investigated on the volcano Usu, 9 years after the 1977–1978 eruptions. The former vegetation was covered by a 1–3 m thick volcanic deposit. Trees producing wind-dispersed seeds capable of long distance dispersal, such as Populus maximowiczii, Betula platyphylla var. japonica, Salix hultenii var. angustifolia, and Salix sachalinensis, were dominant. Most trees were only 2–4 years old in 1986, suggesting that chances for seedling establishment were restricted. The tree heights did not differ significantly among the species, while lengths of annual shoots differed due to herbivore preferences. Trees were established at higher densities on gravel-dominated pumice surfaces than on fine-textured surfaces. Tree density was not greatly affected by the nutrient content of deposits. From 1987 to 1990, tree height increments did not differ between the gravel and non-gravel areas. Ground surface texture is an important factor in determining seedling establishment in the early stages of volcanic succession, and nutrient status is unimportant.     

Species turnover and diversity during early stages of vegetation recovery on the volcano Usu,northern Japan

To study the rate of revegetation during succession on the volcano Usu, northern Japan, vegetation structure and species composition were monitored from 1984 to 1988 in permanent quadrats near the summit of the volcano, which had been almost completely deforested by 1 to 3 m thick volcanic deposits in 1977–78. Analyses of vegetation structure included species richness, species diversity (IT), evenness (J'); year-to-year changes were quantified using Community Coefficient (CC), and Percentage Similarity (PS). While species turnover rates fluctuated and vegetation cover gradually increased, species richness, diversity and evenness did not fluctuate much. The diversity parameters showed slightly different levels for three habitat types distinguished. CC values comparing subsequent years with the starting year 1984 suggest only minor changes in qualitative species composition; PS values decreased more rapidly, indicating larger changes in species cover. A few well-rooted perennial plants were predominant, therefore, PS decline resulted from dominance-controlled community structure. PS-value detected not only habitat difference, but also the rate of the volcanic succession, suggesting that PS is most effective to evaluate successional pace.     

The role of refugia and dispersal in primary succession on Mount St. Helens,Washington

Abstract. An intense lateral blast devastated Mount St. Helens in 1980, but forest understory species survived in some north‐slope ‘refugia’. We explored the effects of refugia on colonization of barren pumice in 1997 and 1998, 18 yr after the eruption. The seed rain of 23 colonizers came mostly from populations that had previously established in refugia. Parachutists had small, vagile seeds, parasailors had winged seeds, and tumblers were blown along the ground. The latter two groups are heavier and dispersed more slowly, but are more likely to survive. The proportion of the vegetation represented by wind‐dispersed species increased with distance from refugia. Parachutist's density declined with time and proximity to refugia. As vegetation adjacent to refugia developed, populations of parasailors and tumblers expanded, foreshadowing their dominance in more remote pumice. Refugia played a critical role in determining the rate and course of succession by providing fertile islands that permitted pioneers and dry meadow species to establish near barren pumice. Species that survived in refugia played a negligible role in colonization. This study showed that when refugia contrast sharply with new substrates, they accelerate recovery by facilitating the invasion of pioneer species.     

Influence of sulfur-oxidizing bacteria on the budget of sulfate in Yugama crater lake, Kusatsu-Shirane volcano, Japan

Sulfur-oxidizing bacteria, Thiobacillus thiooxidans, werefound in a highly acidic (pH = 11.5) crater lake, Yugama,seasonally flowing streams and soil in the catchment area of thecrater. Thiobacillus ferrooxidans was also found in some ofthe streams but not in the lake itself. The lake water containsaqueous carbon dioxide, hydrogen sulfide, polythionates andelemental sulfur in suspension which are the substrates for thegrowth of the sulfur-oxidizing bacteria as no organic compoundsexcept for the microorganisms themselves were detected. Thebacteria isolated from the Yugama water preferred polythionatesin the following order: S₄O₆ ²> S₅O₆ ²> S₆O₆ ²On the other hand,H₂S was more rapidly consumed by the bacteria thanpolythionates which were followed by elemental sulfur. In thecase of test-tube incubation, the optimum pH of the solution forgrowth of the bacteria was between 1.0 and 1.5, and forcultivation in growth medium plates between 2.5 and 3.5. Thebacteria hardly proliferated at pH 0.5 or below. In accordancewith these characteristics of the bacteria, numbers of thebacteria in the surface Yugama crater lake water were at minimum(< a few cells/mL) in February and at maximum (10⁶ cell/mL)in August. The bacterial activity changed in accordance with thesurface lake water temperature, but not necessarily with thevariations in H₂S and polythionates concentrations of the lakewater. Based on the variation in sulfur isotope ratios of sulfateand experimentally determined rate of oxidation of H₂S in thelake water, the sulfate production rate by the bacteria in thecatchment area and the lake were estimated to 9.5 and 8.4g/m²/day, respectively, during the period from 1988 to 1990when the volcanic activity at Yugama was at minimum. Also stream,hydrothermal, H₂S-oxidated SO² ₄-inputs and outputs byseepage and precipitation have been calculated as 4.1, 32, 0.56,36, and 1.2 ton/day, respectively.     

Elevational gradients of small mammal diversity on the northern slopes of Mt. Qilian, China

Aim Small mammal species richness and relative abundance vary along elevational gradients, but there are different patterns that exist. This study reports the patterns of distribution and abundance of small mammals along the broader elevational gradient of Mt. Qilian range. Location The study was conducted in the Mt. Qilian range, north‐western China, from June to August 2001. Methods Removal trapping was conducted using a standardized technique at 7 sites ranging between 1600 and 3900 m elevation within three transects. Correlation, regression and graphical analyses were used to evaluate the diversity patterns along this elevational gradient. Results In total, 586 individuals representing 18 nonvolant small mammal species were collected during 20 160 trap nights. Species composition was different among the three transects with 6 (33%) of the species found only within one transect. Elevational distribution and relative abundance of small rodents showed substantial spatial variation, with only 2 species showing nonsignificant capture frequencies across elevations. Despite these variations, some general patterns of elevational distribution emerged: humped‐shape relationships between species diversity and elevation were noted in all three transects with diversity peaks at middle elevations. In addition, relative abundance was negatively correlated with elevation. Conclusions Results indicate that maximum richness and diversity of nonvolant small mammals occurred at mid‐elevations where several types of plants reached their maximum diversity and primary productivity, and where rainfall and humidity reached a maximum. It is demonstrated that the mid‐elevation bulge is a general feature of at least a large portion of the biota on the Mt. Qilian range.     

The effects of patch richness on con-specific interference in the parasitoid Ibalia leucospoides (Hymenoptera: Ibaliidae)

Abstract We experimentally studied con-specific interference in Ibalia leucospoides, a parasitoid of the woodwasp Sirex noctilio, on pine logs containing variable numbers of woodwasp larvae. Competiton occurred when two different-sized female parasitoids foraged on a host patch, and consequently the small female always abandoned the patch. Regardless of host density, con-specific presence did not affect the attack rate, the number of hosts attacked, nor patch residence time by the winner, when compared to a control female that foraged alone. In contrast, con-specific presence reduced patch time and the number of hosts attacked by the loser. Finally patch time (by both) as well as number of hosts attacked (by the winner only) increased with host density per patch. Our results suggest that con-specific presence has different consequences for different-sized females during patch exploitation.     

Effects of environmental parameters and irrigation on the turgor pressure of banana plants measured using the non‐invasive,online monitoring leaf patch clamp pressure probe

Turgor pressure provides a sensitive indicator for irrigation scheduling. Leaf turgor pressure of Musa acuminate was measured by using the so‐called leaf patch clamp pressure probe, i.e. by application of an external, magnetically generated and constantly retained clamp pressure to a leaf patch and determination of the attenuated output pressure P_p that is highly correlated with the turgor pressure. Real‐time recording of P_p values was made using wireless telemetric transmitters, which send the data to a receiver base station where data are logged and transferred to a GPRS modem linked to an Internet server. Probes functioned over several months under field and laboratory conditions without damage to the leaf patch. Measurements showed that the magnetic‐based probe could monitor very sensitively changes in turgor pressure induced by changes in microclimate (temperature, relative humidity, irradiation and wind) and irrigation. Irrigation effects could clearly be distinguished from environmental effects. Interestingly, oscillations in stomatal aperture, which occurred frequently below turgor pressures of 100 kPa towards noon at high transpiration or at high wind speed, were reflected in the P_p values. The period of pressure oscillations was comparable with the period of oscillations in transpiration and photosynthesis. Multiple probe readings on individual leaves and/or on several leaves over the entire height of the plants further emphasised the great impact of this non‐invasive turgor pressure sensor system for elucidating the dynamics of short‐ and long‐distance water transport in higher plants.     

Effects of water stress on the gas exchange,the activities of some enzymes of carbon and nitrogen metabolism,and on the pool sizes of some organic acids in maize leaves

Water-stressed maize (Zea mays L.) leaves showed a large decrease in leaf conductance during photosynthesis. Net CO₂ uptake and evaporation declined fast at mild stress (=–0.6 to –1.0 MPa) and slower at more severe stress (=–1.0 to -1.2 MPa), whereas the CO₂ concentration in the intercellular spaces (C_i) did not drop to the CO₂ compensation point. The activities of the enzymes of photosynthetic carbon metabolism tested in this study dropped by approx. 30% at =-1.2 MPa. Glutamine synthetase activity was unaffected by water stress, whereas the activity of nitrate reductase was almost completely inhibited. The decline of enzyme activities in relation to was correlated with a concomitant decrease in the content of total soluble protein of the stressed leaves. The total leaf pools of malate, pyruvate and oxaloacetate decreased almost linearly in relation to , thus obviously contradicting the almost constant C_i. In comparison to the controls (=0.6 MPa) the content of citrate and isocitrate increaed markedly at =-0.9 MPa and decreased again at =-1.2 MPa.Abbreviations PCR photosynthetic carbon reduction cycle - PCO photosynthetic carbon oxidation cycle - PEP phosphoenolypyruvate - RuBP ribulose-1,5-bisphosphate