Gap formation and species diversity in Japanese beech forests: a test of the intermediate disturbance hypothesis on a geographic scale

To evaluate whether the intermediate-disturbance hypothesis applies on regional scales, the relationship between the species diversity and gap formation regime of beech forests was examined. The mean gap size and the variation of gap sizes showed no correlation with species diversity. The mean windstorm interval varied widely, but geographical trends, such as latitudinal gradient, were not observed. However, locations that sustained an intermediate frequency of disturbance had the highest species diversity. Although a latitudinal gradient of disturbance was not apparent, the intermediate-disturbance hypothesis was partly supported on a geographic scale. The most predictable model for species diversity was a multiple regression model composed of two factors, the windstorm interval and the cumulative temperature of the growing season. The fact that the temperature was of greater importance than the disturbance interval indicates that the most important factor in predicting forest species diversity is the amount of available energy on a geographic scale.     

Nitrogen content in feces and the diet of Sika deer on the Boso Peninsula, central Japan

Fecal and rumen nitrogen content of Sika deer (Cervus nippon Temminck 1838) on the Boso Peninsula, central Japan, were analyzed concerning their seasonal fluctuations and local differences at four areas on the Boso Peninsula. The mean fecal nitrogen was high in August and low in February in all the areas. The seasonal variation, however, was relatively small. This may be the result of the following reasons: the Boso deer feed on evergreen broad-leaves containing a high and stable nitrogen content in winter; little snow falls there; and the deer consume foods relatively low in nitrogen in summer. The fecal and rumen nitrogen content in both summer and winter depended on deer density.     

Effect of wave exposure on vegetation abundance, richness and depth distribution of shallow water plants in a New Zealand lake

SUMMARY 1. The effects of physical disturbance in terms of wave exposure, shore slope and substrate mobility on the presence, species richness, cover and depth limits of the low‐growing, shallow water macrophyte community (called the low mixed community) were examined at 41 shore sites in Lake Wanaka, South Island, New Zealand. 2. Wave exposure at a site was described by the previous year's maximum depth of sediment motion and maximum vertical extension of waves, determined from a computer wave model. Shore slope at each site was recorded from ?1 to 0 m depth, and sediment stability was assessed as the cover of small gravel. 3. The low mixed community was only present on sites where the previous year's maximum depth of sediment motion was <8 m, maximum wave run‐up was <0.3 m, shore slope <0.12 m m^?1, and small gravel cover <78%. 4. Species richness, cover and depth limits of the low mixed community decreased with increasing disturbance on the sites. Sixty‐two percentage of the variation in species richness could be explained by physical disturbance variables when all sites were included (N=41). When only sites with a low mixed community were included (N=22), only 18% of the variation was explained. Species richness within sites supporting a low mixed community is thus poorly explained by physical disturbance, whereas presence or absence is better explained. There was no evidence to support the intermediate disturbance hypothesis in this study. 5. Disturbance as a result of waves explained 86% of the variation in cover among sites with a low mixed community and accounted for 68 and 58% of the variation in upper and lower depth limit of the low mixed community, respectively. 6. The models obtained in this study can be used as predictive models for the low mixed community in New Zealand lakes in relation to natural physical disturbance on the shore. By integrating these results with previous studies on the effect of water level fluctuation, we describe a generalised optimum physical habitat for the low mixed community in New Zealand lakes.     

Endogone corticioides sp. nov. from subalpine conifer forests in Japan and China,and its multi-locus phylogeny

Endogone corticioides sp. nov. was described from subalpine conifer forests in Japan and China. This species is similar to E. pisiformis in its yellowish sporocarps and zygosporangia on the equal empty gametangia. Endogone corticioides was clearly distinguished from E. pisiformis based on its resupinate sporocarp, the presence of hyphal peg-like structures on the peridium, and abundant thick-walled hyphae in the gleba. Phylogenetic analysis of the small and large subunit nuclear ribosomal RNA and the elongation factor-1α genes suggested that E. corticioides is closely related to E. pisiformis. Limited vegetative growth from the surface-sterilized sporocarps on agar media was shown.     

Effects of natural disturbance on the regeneration of riparian forests in a Chichibu Mountains,central Japan

Disturbances in the riparian area had a large effect on each stage of regeneration in riparian forests dominated by Fraxinus platypoda in the Chichibu Mountains, central Japan. F. platypoda adapted well to various disturbances in frequency and size such as landslides and canopy gap formation.The spatial distribution and age structure of F. platypoda were studied in relation to the disturbance regime of the riparian zone. The bell-shaped size and age distributions of F. platypoda suggest that F. platypoda trees were established synchronously in a large disturbance site caused by an earthquake landslide about 200 years ago. For the past 200 years, the topography has been stable and the canopy gap has been recovered by advance regenerated saplings.The distribution of F. platypoda saplings was restricted to an abandoned channel and part of a floodplain. These topographical sites formed by gravel provided safe sites for saplings because stream disturbances did not occur for a long time. Channel bars were under low shade stress because of the lack of herbs and a litter layer, which represents a safe site for seedling establishment. However, the seedling bank could have been destroyed by high frequent flooding caused by large typhoons and the establishment of seedlings might have been prevented for a long time.The regeneration process of F. platypoda was explicated based on the gap dynamics theory during the stable period of topography. On the other hand, an even-aged forest was established in a large scale disturbance site.Nomenclature: Ohwi & Kitagawa (1992)     

Topographical pattern of the forest vegetation on a river basin in a warm-temperate hilly region,central Japan

The relationship between tree distribution and topography was examined in a small river basin (3.4 ha) comprising a complex mosaic of topographical units at 10² to 10³ m² order, each of which had a shallow valley bordered by small ridges or breaks of slopes. Twenty-five major woody species were divided into two groups (groups A and B) based on a cluster analysis using the distribution data in the basin. Group A, which mainly consisted of early-successional species, was distributed around the valley sites of the topographical units, while group B, which mainly consisted of late-successional species, was distributed around the ridge sites of the topographical units. This vegetation pattern coincided with erosional condition in the basin. That is, the valley sites were eroded more actively than the ridge sites, as soil depth tended to be thin in the valley sites and thick in the ridge sites, and because large (canopy) trees were restricted in the ridge sites. There was no tendency that group B was replacing group A, and hence it was suggested that repeated disturbance by slope failures or small-scale shallow landslides have prevented compositional change from the early-successional (group A) to the late-successional (group B) species by preventing the invasion of the latter into valley sites.     

The effect of colonization and competition processes on the relation between disturbance and diversity in plant communities

Many theoretical and field studies have emphasized the impact of disturbance in the dynamics and diversity of sessile organism communities. This view is best reflected by the Intermediate Disturbance Hypothesis (IDH), which states that a maximum of diversity is found in ecosystems or communities experiencing intermediate disturbance regimes or at an intermediate stage of development since the last major disturbance event. Although theoretical models based on competitive interactions tend to validate this hypothesis, a recent meta-analysis of field experiments revealed that the mono-modal relationship between disturbance and diversity might not be a general pattern. In this article, we investigate the relationship between disturbance and diversity through the study of patch models, combining two types of competitive interactions: with or without competitive hierarchy, with two mechanisms influencing colonization: negative frequency dependence in colonization rates and immigration. These combinations led to various disturbance-diversity patterns. In the model without competitive hierarchy (founder effect model), a decreasing relationship appeared to be the rule as mentioned in previous studies. In the model with competitive hierarchy, the IDH pattern was obtained for low frequency dependence and low immigration. Nevertheless, high negative frequency dependence in colonization rates led to a decreasing relationship between disturbance and diversity. In contrast, high immigration led to an increasing relationship. The coexistence window (the range of disturbance intensity allowing coexistence) was the widest for intermediate immigration rates. For random species assemblages, patterns with multiple peaks were also possible. These results highlight the fact that the mono-modal IDH pattern should not be considered a rule. Competition and colonization mechanisms have a profound impact on the relationship between disturbance and diversity.     

Structure and regeneration ofFraxinus spaethiana-Pterocarya rhoifolia forests in unstable valleys in the Chichibu Mountains,central Japan

The structure of temperate deciduous forests dominated byFraxinus spaethiana andPterocarya rhoifolia in V-shaped valleys in the Chichibu Mountains, central Japan was studied to clarify their dynamics, with special reference to disturbance. Forest structure and annual ring analysis revealed that the forests have been subjected to mass movement (mud flow) in the past. The size and age structure, and diameter growth curves, indicated thatF. spaethiana is relatively shade tolerant, long living and slow-growing, whileP. rhoifolia is less shade tolerant, short living, but fast growing.Pterocarya rhoifolia appears to depend on large-scale but rare disturbances.Fraxinus spaethiana establishes its seedlings on relatively unstable substrate in V-shaped valleys, but once established, they survive a long time with high shade tolerance. The dominance and coexistence of the two species may primarily depend on the intensity and frequency of geomorphological processes.     

Species richness of woody plants in the landscapes of Central Spain: the role of management disturbances,environment and non‐stationarity

Questions: How important is management disturbance on gamma species richness of woody plants at intermediate landscape scales? How is species richness related to other climatic and biotic factors in the study area? How does the assumption of spatial stationarity affect assessment of relationships among species richness and explanatory variables (e.g. management, biotic and climatic factors) across extensive study areas? Location: Central Spain (regions of Castilla y León, Madrid and Castilla‐La Mancha). Scale: Extent: 150 000 km². Grain: 25 km² (5 × 5‐km cells). Methods: Information from 21 064 plots from the 3SNFI was used to evaluate richness of tree and shrub species at intermediate landscape scales. In addition to variables well known to explain biodiversity, e.g. environmental and biotic factors, effect of management treatments was evaluated by assessing clearcutting, selection cutting, stand improvement treatments and agrosilvopastoral systems (dehesas). Results from GWR techniques were compared with those from OLS regression. Results: Patterns of gamma species richness, although strongly affected by both environmental and biotic variables, were also significantly modified by management factors. Species richness increased with percentage of selection cutting stands and improvement treatments but decreased with percentage of clearcutting stands. Reduced species richness of woody plants was associated with agrosilvopastoral practices. Species richness for trees was closely related to basal area, annual precipitation and topographic complexity; species richness for shrubs was closely related to topographic complexity and agrosilvopastoral systems. Most relationships between species richness and environmental or biotic factors were non‐stationary. Relationships between species richness and management effects tended to be stationary, with a few exceptions. Conclusions: Landscape models of biodiversity in Central Spain were more informative when they accounted for effects of management practices, at least at intermediate scales. In the context of current rural abandonment, silvicultural disturbances of intermediate intensity increased gamma species richness of woody plants. Exclusion of factors such as agrosilvopastoral systems from models could have led to spurious relationships with other spatially co‐varying factors (e.g. summer precipitation). Patterns of spatial variation in relationships, provided by GWR models, allowed formulating hypotheses about potential ecological processes underlying them, beyond generalizations resulting from global (OLS) models.     

Temperature affects male and female potential reproductive rates differently in the sex-role reversed pipefish, Syngnathus typhle

The differences in potential reproductive rate between the sexescan be used to predict the operational sex ratio and the patternsand intensity of mating competition and hence sexual selectionin a population. This article describes how one environmentalcomponent, temperature, affects potential reproductive ratesof the two sexes in the paternally brooding, sex-role reversedpipefish (Syngnathus typhle). Males brooded embryos much longer(on average 58 days) in cold water (about 10°C) than inwarmer water (35 days at about 15°C). As a consequence,the potential reproductive rate (number of eggs brooded perday) of males was significantly higher in warm water. In females,however, potential reproductive rate, i.e., number of eggs producedper day given an unlimited access to mates, was not significantlydifferent between temperatures. In both sexes, potential reproductiverate was positively related to body size. At both temperatures,females had the potential to reproduce faster than males. Asa result, the operational sex ratio will become female biasedand sex-roles reversed, as is the case in this species. Sincetemperature differently influenced the potential reproductiverates of males and females, with the sexual difference largerat lower temperatures, more intense female-female competitionis predicted at low temperatures.     

Secondary succession and effects of clear-logging on diversity in the subtropical forests on Okinawa Island,southern Japan

In order to clarify the recovery process of the subtropical forest on Okinawa Island, southern Japan, biomass accumulation and the successional trend of species diversity with time were investigated by comparing plots of old-growth and clear logged secondary forests. Self-thinning was an important factor in the development of young secondary forests, and the small variance in tree size within a stand was related to the stand not being fully stratified after clear-cutting. A large variance of size structure in old secondary and old-growth forests implies re-initiation of the understorey. Additionally, the trajectory of stand development indicated that the subtropical forest quickly recovered aboveground biomass, which reached its upper limit at about 50 years after disturbance. However, there was a large distinction in species diversity between the secondary forests and old-growth forests. The diversity of forest floor plants did not recover well after being clear-cut. This indicates that management of the subtropical forest should not only take timber-oriented tree species into account, but also the biodiversity in ground flora. The secondary forests were characterized by Castanopsis sieboldii and Schima wallichii, and the monopolization of C. sieboldii through secondary succession had a negative influence on species diversity. Distylium racemosum dominated at the late development stage and was considered a long-lived competing species that reduced the dominance of C. sieboldii and facilitated the co-occurrence of understorey species. Light-demanding pioneer tree species such as S. wallichii that regenerated after logging should be replaced by competitive effects of climax species, and thus relayed floristic change might increase species diversity along secondary succession.Nomenclature: Hatushima and Amano (1994).     

Species composition and distribution of Coleosporium species on the needles of Pinus densiflora at a semi-natural vegetation succession site in central Japan

Coleosporium species cause pine needle rust. Most species have heteromacrocyclic life cycles, and 12 species use Pinus densiflora as aecial hosts. To understand the biology of rust fungi and develop better methods for controlling rust diseases, it is necessary to clarify that which Coleosporium species affect pine trees. However, Coleosporium on pine trees have rarely been identified at the species level because of their morphological similarities. We used polymerase chain reaction - restriction fragment length polymorphism (PCR-RFLP) to clarify the species composition, abundance, and distribution of Coleosporium in a P. densiflora forest. We surveyed a site where several Coleosporium species might complete their life cycles. PCR-RFLP revealed four species on the pines: C. asterum, C. clematidis-apiifoliae, C. lycopodis, and C. phellodendri. Coleosporium phellodendri was distributed throughout the forest and was the most abundant. Aecia of C. phellodendri formed mainly on 2-y-old needles. The abundance and distribution of C. phellodendri appeared to be affected by the longer effective dispersal range of basidiospores and the existence of abundant inoculum sources. The age of leaves where C. phellodendri form aecia mainly was thought to be influenced by the characteristic life cycle, with aecial formation requiring 2?y after basidiospore infection.     

The stable carbon and nitrogen isotopic composition of vegetation in tropical forests of the Amazon Basin, Brazil

Here we present the within-site, seasonal, and interannual variations of the carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope ratios of leaves, wood, bark and litter from four sites in the Amazon region, Brazil. Samples were collected in Manaus (3° 06′07′′ S; 60°01′30′′ W), Ji-Paraná (10°53′07′′ S; 61°57′06′′ W), and Santarém (2°26′35′′ S; 54°42′30′′ W) with mean annual precipitation of 2207, 2040 and 1909 mm respectively. The overall average for all leaf samples was for δ¹³C and for δ¹⁵N (n=756). The leaf δ values at these sites were often but not always statistically distinct from each other. The δ¹³C values varied from to . Pronounced differences in δ¹³C values occurred with height associated with differences in forest structure. The δ¹³C of leaf dry matter showed seasonal variations associated with the length of the dry season, despite the fact that total annual precipitation was similar among the studied sites. Leaf δ¹⁵N values ranged from to a maximum value of , and the Santarém sites showed more enriched values than Manaus and Ji-Paraná sites. No seasonal variation was detected in the δ¹⁵N of leaves, but significant differences were observed among sites and with changes in canopy height. The isotope ratio data are consistent with our current understanding of the roles of light, water availability, and recycling of soil-respired CO₂ influences on δ¹³C and consistent with our understanding that an open nitrogen cycle can lead to high δ¹⁵N values despite a significant number of legumes in the vegetation.     

Phylogeography of the component species of broad-leaved evergreen forests in Japan,based on chloroplast DNA variation

In order to elucidate the past distribution and colonization routes of broad-leaved evergreen (lucidophyllous) forests, we investigated the intraspecific phylogeographic patterns of lucidophyllous forests in Japan and surrounding areas. We selected 6 component species with a similar geographic distributions growing in Castanopsis-dominant forests. We defined possible important refugia during the glacial periods as the regions rich in rare haplotypes (with a frequency of 5% or less), or as regions rich in the number of common haplotypes (with a frequency of more than 5%). We then located the sites of refuge by comparing the intraspecific phylogeographic patterns among 6 component species of lucidophyllous forests with respect to these two parameters (i.e., haplotype uniqueness and the number of haplotypes). The following results were obtained during the course of this study: (1) rare haplotypes were distributed among islands around the main islands of Japan; (2) rare subtypes and the greatest numbers of common haplotypes were observed in Kyushu, a finding which agreed with fossilized pollen data demonstrative of the past existence of refugia in southern Kyushu; and (3) rare haplotypes were found on the Muroto Peninsula, and the second greatest numbers of common haplotypes were observed on the Kii Peninsula, a finding which suggested the existence of additional important refugia along the Pacific coast of Japan during the glacial ages.     

Fire regimes and forest changes in mid and upper montane forests of the southern Cascades, Lassen Volcanic National Park, California, U.S.A.

Aim Spatial and temporal variation in fire regime parameters and forest structure were assessed. Location A 2630‐ha area of mid‐ and upper montane forest in Lassen Volcanic National Park (LVNP). Methods Two hypotheses were tested concerned with fire‐vegetation relationships in southern Cascades forests: (1) fire regime parameters (return interval, season of burn, fire size, rotation period) vary by forest dominant, elevation and slope aspect; and (2) fire exclusion since 1905 has caused forest structural and compositional changes in both mid‐ and upper montane forests. The implications of the study for national park management are also discussed. Results Fire regime parameters varied by forest compositional group and elevation in LVNP. Median composite and point fire return intervals were shorter in low elevation Jeffrey pine (Pinus jeffreyi) (JP) (4–6 years, 16 years) and Jeffrey pine–white fir (Abies concolor) (JP‐WF) (5–10 years, 22 years) and longer in high elevation red fir (Abies magnifica)— western white pine (Pinus monticola) (RF‐WWP) forests (9–27 years, 70 years). Median fire return intervals were also shorter on east‐facing (6–9 years, 16.3 years) and longer on south‐ (11 years, 32.5 years) and west‐facing slopes (22–28 years, 54‐years) in all forests and in each forest composition group. Spatial patterns in fire rotation length were the same as those for fire return intervals. More growing season fires also occurred in JP (33.1%) and JP‐WF (17.5%) than in RF‐WWP (1.1%) forests. A dramatic decline in fire frequency occurred in all forests after 1905. Conclusions Changes in forest structure and composition occurred in both mid‐ and upper montane forests due to twentieth‐century fire exclusion. Forest density increased in JP and JP‐WF forests and white fir increased in JP‐WF forests and is now replacing Jeffrey pine. Forest density only increased in some RF‐WWP stands, but not others. Resource managers restoring fire to these now denser forests need to burn larger areas if fire is going to play its pre‐settlement role in montane forest dynamics.     

Effect of typhoon disturbance on fine litterfall and related nutrient input in a subtropical forest on Okinawa Island, Japan

Typhoons are frequent on Okinawa Island, southwestern Japan. The effects of typhoon disturbance on the patterns of fine litterfall and related nutrient inputs in a subtropical evergreen broad-leaved forest were studied over 5 years from May 1996 to April 2001. Annual fine litterfall averaged 7558 kg ha⁻¹ (range from 6188 to 9439 kg ha⁻¹) for six sampling plots over 5 years, which differed significantly among years (p<0.001) but not among plots (p=0.122). A seasonal maximum was most evident for leaf litter component. Woody litter fell more irregularly through the year, and peak fall varied with typhoon and windstorm. The mean ratio of annual litterfall mass of sexual organs to leaves was 0.06, much lower than that in other tropical and subtropical rain forests. Nutrient concentrations varied in litterfall components, but were not significantly different among plots. The lowest concentrations of N and P in leaf litter were observed in March, which is also the month with the greatest leaf fall. However, the highest concentrations were recorded in typhoon season. Nitrogen and P concentrations were 34% and 106% greater in the green leaves that fell during typhoons than in senescent leaves. Mean nutrient inputs by litterfall were: N 83, P 3.2, K 25, Ca 71, Mg 19, Al 12, Na 10, Fe 0.86 and Mn 3.9 kg ha⁻¹ yr⁻¹, and differed significantly among years for all elements (p<0.0005) and among plots only for K (p<0.05) and Mn (p<0.0001). Typhoon disturbance strongly affected annual fine litterfall and related nutrient inputs, which contributed an average of 30% of the annual litterfall mass, and from 30% to 39% (for different nutrient elements) of annual total nutrient inputs. The results from this study suggest that typhoon-driven maintenance of rapid cycling of P and N and their high availability in soil appears to be an important mechanism to maintain productivity in the subtropical forest on Okinawa Island.     

Species richness and composition of Curculionidae (Coleoptera) in a conifer plantation, secondary forest, and old-growth forest in the central mountainous region of Japan

Weevil (Curculionidae, Coleoptera) species richness and composition were investigated and compared among larch [Larix kaempferi (Lamb.) Carriére] plantations, secondary forests, and old-growth forests in the central mountainous region of Japan. In addition, to assess the effects of forest-management practices, namely thinning and long-rotation logging schedules (long rotation), the weevil assemblages of recently thinned middle-aged and long-rotated larch plantations were compared with those of middle-aged larch plantations. Malaise traps were set in 44 stands of these forest types, and weevils were separated and identified. Several environmental factors other than forest type were also examined. Weevil species richness and diversity indices [Shannon-Wiener diversity index (H) and Simpsons index of diversity (D)] were higher in the secondary forest than in the larch plantation. Because of its wide distribution and higher weevil species richness, the secondary forest contributed to maintaining weevil diversity in this region. Old-growth forest had higher diversity indices (H and D) than did the larch plantation. The secondary forest had the highest number of species in total. Though the number of individuals was the highest in larch plantation, species richness, H and D of the plantation were generally low. Weevil community structure and species composition differed among the three forest types, but the difference in weevil composition between the larch plantation and the other two forest types was the largest. Forest type is probably the most important factor for determining the differences in weevil assemblage, and further, both dominant tree type (coniferous trees versus broad-leaved trees) and the number of mature tree species seem to be important factors for weevil species composition. Among forest management practices, long rotation caused diversity indices (H and D) to increase while thinning appeared to cause only minor changes in the weevil assemblages. Because species richness and species composition of Curculionidae well reflected the differences in forest types and some other environmental factors investigated, this family seems suitable for diversity research in forests. Further research on biodiversity with the use of this family should, therefore, be expected.