Topography-specific emergence of fungal fruiting bodies in warm temperate evergreen broad-leaved forests on Yakushima Island,Japan

We conducted line route censuses of fungal fruiting bodies from August to September in 2005 and 2006 along ridges and valleys and compared the differences in the encounter rates of fungal fruiting bodies (= fruiting bodies seen per census kilometer) between types of topography and between fungal functional groups (i.e., ectomycorrhizal and saprobic fungi) in warm temperate evergreen broad-leaved forests on Yakushima Island, Japan. We found 251 fungal fruiting bodies (26 families, 50 genera, and 65 species) in total, including 51 bodies from Tricholomataceae, 41 from Russulaceae, 25 from Boletaceae, and 19 from Amanitaceae. The encounter rate of ectomycorrhizal fungi was greater at the ridge route (26.7 unit/km) than at the valley route (8.7 unit/km) and that of saprobic fungi was greater at the valley route (25.0 unit/km) than at the ridge route (12.5 unit/km). In addition, we conducted 7-year intermittent sampling and identified 40 families, 96 genera, and 142 species. The topography-specific emergence pattern of the intermittent sampling method was similar to that of the line census method. The fungal species composition in this study was possibly affected by a topographic gradient for both fungal functional groups through soil moisture, nutrient availability, and host tree distribution.     

Nutrient-uptake and nutrient-use efficiency ofPinus thunbergii Parl. along a topographical gradient of soil nutrient availability

To examine responses of a plant species to nutrient availability, we investigated changes in soil nutrient availability, litterfall production and nutrient content in litterfall along a topographic gradient in aPinus thunbergii Parl. plantation. Responses were evaluated in terms of three efficiency indices: (i) nutrient-uptake efficiency (the ratio of nutrient return in litterfall to soil nutrient availability); (ii) nutrient-use efficiency (the ratio of litterfall mass to nutrient return in litterfall); and (iii) nutrient-response efficiency (the ratio of litterfall mass to soil nutrient availability). These indices can distinguish the ability of a species to acquire nutrients and its ability to use them in litterfall production. Nitrogen and phosphorus availabilities in soil were lower in upper slope positions. The three efficiencies were higher in upper slope positions and negatively correlated with soil nutrient availability for both nitrogen and phosphorus. An increase in nutrient-response efficiency was achieved by both increases in nutrient-uptake and nutrient-use efficiencies.     

Pattern of wood litter fall in five forests located along an altitudinal gradient in Central Himalaya

Wood and total litter fall were measured in five forests of the Kumaun Himalaya situated along an altitudinal gradient from 329 m to 1850 m. Total annual wood litter fall, which ranged from 93 to 197 g m², was inversely related with altitude and positively related with forest basal cover. Total litter fall ranged from 405 to 839 g m² yr¹ and was positively related with forest basal cover. Wood litter fall was highest in the rainy season and that of total litter in the summer season. Abscissed litter accounted for a maximum proportion of seasonal wood fall in summer. Monthly non-abscissed wood fall was positively related to the monthly rainfall on each site. The fractions in lower diameter classes (0–5 to 5–10 mm) dominated the annual wood fall in all forests and accounted for 59 to 78% of the total. Fractions in >10 mm size class fall maximally during the rainy months.Nomenclature follows Kanjilal & Gupta (1969) unless otherwise stated.The authors are thankful to Dr Uma Pandey for helpful suggestions and to the Department of Science and Technology, New Delhi for financial support.     

Effects of roads on alpine and subalpine plant species distribution along an altitudinal gradient on Mount Norikura, central Japan

We investigated the effects of roads on alpine and subalpine plant species distribution along an altitudinal gradient on Mount Norikura (3026 m a.s.l.), Japan. We examined the vegetation of herb and tree species shorter than 1.3 m along roadsides and adjacent natural vegetation at 200 m intervals between 1600 and 3000 m a.s.l. The timberline was at 2500 m a.s.l. Although the canopy opening was greater at the roadsides than in the natural vegetation, it was similar above the timberline. Soil cover and litter depth of the soil surface were less at roadsides than the natural vegetation, and gravel and rock cover were greater at roadsides. Species composition changed in similar directions from natural vegetation to roadsides along the altitudinal gradient. This direction was related to canopy opening and litter depth. Liliaceae, Ericaceae and Pinaceae were dominant families in the natural vegetation, and Asteraceae and Poaceae were greatest at the roadsides. Roadside plants were mostly herb species, while tree species increased in natural vegetation. Five exotic species were also observed at the roadsides. Sunny plant species gradually increased with altitude in the natural vegetation, indicated by the increase in canopy opening. By contrast, roadside plants were mostly sunny plant species irrespective of altitude. The number of lowland and montane species increased at the roadsides in the subalpine zone. Thus, roads strongly altered species composition of the natural vegetation along the altitudinal gradient probably because of changes in light and soil-surface conditions for growth and seedling establishment.     

Forest growth along a rainfall gradient in Hawaii: Acacia koa stand structure,productivity, foliar nutrients,and water- and nutrient-use efficiencies

We tested whether variation in growth of native koa (Acacia koa) forest along a rainfall gradient was attributable to differences in leaf area index (LAI) or to differences in physiological performance per unit of leaf area. Koa stands were studied on western Kauai prior to Hurricane Iniki, and ranged from 500 to 1130 m elevation and from 850 to 1800 mm annual precipitation. Koa stands along the gradient had basal area ranging from 8 to 42 m²/ha, LAI ranging from 1.4 to 5.4, and wood increment ranging from 0.7 to 7.1 tonnes/ha/year. N, P, and K contents by weight of sun leaves (phyllodes) were negatively correlated with specific leaf mass (SLM, g m^-2) across sites; on a leaf area basis, N increased whereas P and K decreased with SLM. LAI, aboveground woody biomass increment, and production per unit leaf area (E) increased as phyllode ¹³C became more negative. The ¹³C data suggested that intrinsic water-use efficiency (ratio of assimilation to conductance) increased as water availability decreased. In five of the six sites, phyllode P contents increased as LAI increased, but biomass increment and E were not correlated with phyllode nutrient contents, suggesting that productivity was limited more by water than by nutrient availability. Because vapor pressure deficits increased with decreasing elevation, actual water-use efficiency (ratio of assimilation to transpiration) was lower at drier, low-elevation sites. There was a trade-off between intrinsic water-use efficiency and production per unit of canopy N or P across the gradient. In summary, koa responds to water limitation both by reducing stand LAI and by adjusting gas exchange, which results in increased intrinsic water-use efficiency but decreased E.     

Summer dietary compositions of sika deer on Yakushima Island,southern Japan

Summer food habits of Sika deer (Cervus nippon yakushimae) on Yakushima Island, southern Japan, were studied focusing in particular onPseudosasa owatarii, a dwarf bamboo native to the island. Fecal analyses showed that the food habit of the Sika deer on Yakushima island was plastic: the dietary composition was dominated by graminoids (ca. 90%), particularlyP. owatarii (50%–60%) in the bamboo grassland of theP. owatarii zone above the timber line, whereas leaves and non-leafy parts of woody plants were important in the crytomeria forest and evergreen broad-leaved forest below the timber line. The maximum culm age ofP. owatarii was 8.5 yr, but the majority of leaves (97.8%) were younger than 2.5 yr. Since the weight contribution of the leaves is great (27.7%) and the turnover rate is high, theP. owatarii grassland affords a good foraging place for the Sika deer on Yakushima Island.     

Age structure and regeneration of old growthCryptomeria japonica forests on Yakushima Island

The regeneration process ofCryptomeria japonica forests was studied from surveys of age and initial growth of the coniferous stumps in plots of 2.66 ha in total on Yakushima Island, south Japan. The conifers germinated during particular regeneration periods each of which was shorter than 100 years. The periods repeated themselves several times in each plot, and conifers of each period formed patches. Both tree cutting and natural gap formation of the canopies initiated the regenerations. The saplings that were thought to have germinated before the initiation of the regeneration grew slowly. After the initiation, the saplings that germinated earlier grew more quickly. For the three codominant conifers:Cryptomeria japonica, Tsuga sieboldii, andAbies firma, the shade tolerance was inversely correlated with maximum age. Even the most shade-tolerantA. firma was a gap-dependent species in regeneration periods, and there were no species differences in the colonizing times during the regeneration period. Shade-intolerant long-lived species and shade-tolerant short-lived species coexisted as climax species, not as alternate species in the sere of succession.     

Butterfly diversity and historical land cover change along an altitudinal gradient

Land cover and climate change are both major threats for biodiversity. In mountain ecosystems species have to adapt to fragmented habitats and harsh environmental conditions but so far, altitudinal effects in combination with land cover change have been rarely studied. The objective of this study was to determine the effects of altitude and historical land cover change on butterfly diversity. We studied species richness patterns of butterflies occuring in wetlands and other open habitats along an altitudinal gradient in a low mountain region (340–750 m a.s.l., Bavaria, Germany) with drastic loss of open habitats within the last 40–60 years. We recorded in 27 sites a total of 4,523 individuals of 49 butterfly species and five species of burnet moths. Species richness peaked at mid elevation and increased with patch size. Land cover change was most pronounced at high altitudes, but neither current open habitats, nor the historical loss of open habitats affected the species richness of butterflies. Neither open land specialized butterflies nor generalist and forest species were significantly affected by the loss of open habitats. However, increasing forest area in high altitudes reduces possible refuge open habitats for butterflies at their thermal distribution limits. This could lead to extinction of such butterfly species when temperatures further rise due to global warming.     

Species composition of termites of the Nyika plateau forests, northern Malawi, over an altitudinal gradient

Termites were surveyed at three altitudes (Brachystegia woodland at 1676 m and 1905 m, and Juniper woodland at 2210 m) in forests within the Nyika Plateau, northern Malawi. Sampling was by a standardized 100 m transect protocol. Termite diversity was highest in the mid‐altitude site and lowest in the Juniper forest. The assemblages were dominated by soil‐feeding termites in the Termitidae subfamilies Apicotermitinae and Termitinae, and included one new soldierless Apicotermitinae genus. The structure of the assemblages was clearly due to a mixture of altitudinal and site history factors. This was especially true of the lowest altitude forest where burning and other anthropogenic disturbance factors appear to have reduced termite diversity relative to the mid‐altitude site. The Nyika plateau shows a much higher diversity at mid‐altitudes than similar SE Asian sites, probably due to the larger area of highland in Africa than in SE Asia. In addition, the clade composition of the Nyika assemblages differs completely from that found at similar altitudes in SE Asia. This preliminary study supports the hypothesis that mid‐ to high‐ altitude assemblages in both SE Asia and Africa appear to be derived from depauperated random subsets of the lowland fauna rather than from clades specifically adapted to higher altitudes.     

Regional and local patterns of ectomycorrhizal fungal diversity and community structure along an altitudinal gradient in the Hyrcanian forests of northern Iran

? Altitudinal gradients strongly affect the diversity of plants and animals, yet little is known about the altitudinal effects on the distribution of microorganisms, including ectomycorrhizal fungi. ? By combining morphological and molecular identification methods, we addressed the relative effects of altitude, temperature, precipitation, host community and soil nutrient concentrations on species richness and community composition of ectomycorrhizal fungi in one of the last remaining temperate old-growth forests in Eurasia. ? Molecular analyses revealed 367 species of ectomycorrhizal fungi along three altitudinal transects. Species richness declined monotonically with increasing altitude. Host species and altitude were the main drivers of the ectomycorrhizal fungal community composition at both the local and regional scales. The mean annual temperature and precipitation were strongly correlated with altitude and accounted for the observed patterns of richness and community. ? The decline of ectomycorrhizal fungal richness with increasing altitude is consistent with the general altitudinal richness patterns of macroorganisms. Low environmental energy reduces the competitive ability of rare species and thus has a negative effect on the richness of ectomycorrhizal fungi. Because of multicollinearity with altitude, the direct effects of climatic variables and their seasonality warrant further investigation at the regional and continental scales.     

Inter-male associations in a wild Japanese macaque troop on Yakushima Island,Japan

Adult male association and its annual change were studied in a wild population of Japanese macaques (Macaca fuscata yakui) on Yakushima Island, Japan. Unlike many other Japanese macaque troops, adult troop males frequently maintained proximity and exchanged grooming with one another in both the mating and non-mating seasons, and the dominance relationship rarely appeared in such inter-male associations. The few cases of agonistic interactions occurred mostly when estrous females or food resources were immediately concerned. Although troop males were very intolerant to newly appeared solitary males (new males) during the mating season, close associations were formed between troop males and new males as soon as the mating season terminated. The consort of new males and lower-ranking troop males with estrous females was frequently disturbed, but these males could copulate no less frequently than higher-ranking males. A comparison among macaque species suggests the existence of two forms of inter-male association: (1) the frequent association based on the symmetrical exchange of social behaviors; and (2) the infrequent and asymmetrical association related to the dominance relationship. The form of inter-male association seems to be influenced by whether or not males can keep close associations with females throughout the year.     

Freezing tolerance in grasses along an altitudinal gradient in the Venezuelan Andes

The tropical high Andes experience greater daily temperature oscillations compared to seasonal ones as well as a high frequency of night frost occurrence year round. Survival of organisms, under such environmental conditions, has been determined by selective forces which have evolved into adaptations including avoidance or tolerance to freezing. These adaptations have been studied in different species of trees, shrubs and perennial herbs in páramo ecosystems, while they have not been considered in grasses, an important family of the páramo. In order to understand survival of Poaceae, resistance mechanisms were determined. The study was performed along an altitudinal gradient (2,500–4,200 m a.s.l.) in the páramo. Supercooling capacity and frost injury temperature were determined in nine species in order to establish cold resistance mechanisms. Grasses registered a very low supercooling capacity along the altitudinal gradient, with ice formation between −6 and −3°C. On the other hand, frost injury temperature oscillated between −18 and −7°C. Our results suggest that grasses exhibit freezing tolerance as their main cold resistance mechanism. Since grasses grow at ground level, where greatest heat loss takes place, tolerance may be related to this life form as reported for other small life forms.     

Classification and ordination of plant communities along an altitudinal gradient on the Presidential Range,New Hampshire,USA

An analysis of vegetation along an altitudinal gradient on the Presidential Range, New Hampshire, USA, using the Braun–Blanquet approach followed by multivariate data analysis is presented. Twelve main plant communities have been distinguished. Floristic information is presented in twelve tables and one appendix. The relationships of the communities to complex environmental gradients are analyzed using Correspondence Analysis. Floristic composition and community structure are controlled primarily by the altitudinal gradient (temperature, precipitation), and by mesotopographic conditions (snow accumulation, exposure and cryoturbation, slope position, and soil moisture).     

Structure,composition and regeneration of riparian forest along an altitudinal gradient in northern Iran

In order to protect and understand the regeneration of riparian forests, it is important to understand the environmental conditions that lead to their vegetation differentiation. We evaluated the structure, composition, density and regeneration of woody species in forests along the river Safaroud in Ramsar forest in northern Iran in relation to elevation, soil properties and distance from the river. Using 60 transects located perpendicularly to river and 300 quadrats, we examined forests 0, 50, 100, 150 and 200 m from the stream along an elevation gradient spanning from 350 to 2,400 m a.s.l. We found that total density, regeneration, diameter and basal area of trees were significantly higher in the interior of the forest whereas shrub density was higher close to the river. The uneven-aged forest structure showed no significant differences among gradient from the river to the forest interior. Hydro-geomorphic processes, flooding, the elevation gradient, distance from the river and soil properties were the most important factors that influenced plant community distribution in relation to the river.     

Stable isotopes of lakes and precipitation along an altitudinal gradient in the Eastern Alps

Altitude encompasses broad environmental gradients that influence the isotopic composition of lake water. We selected 55 lakes in the Eastern Alps along an altitudinal gradient [214–2,532 m above sea level (a.s.l.)] to model the isotopic signal of surface water dependent on intrinsic (lake geomorphometry) and extrinsic (air temperature, precipitation) factors. Ordinary and generalised least squared regression were used for statistical analysis. The isotope signal of lake water was lower in spring than in summer and decreased with altitude (?0.21 δ¹⁸O ‰/100 m; ?1.5 δ²H ‰/100 m). This pattern largely depended on temperature and a pseudo-latitude effect. The isotopic signal in monthly precipitation (12 stations; altitudinal gradient 90–2,730 m a.s.l.) generally showed the expected pattern of less enriched values with altitude; however, unusual values were related to weather anomalies. The local meteoric water line was similar to the global meteoric water line as shown by overlapping confidence intervals. By discriminating different elevational bands, we could show that high elevation lakes (>1,500 m a.s.l.) experience different patterns of evaporation with respect to low elevation lakes (<1,500 m a.s.l.). Our study showed that lakes have a unique isotopic fingerprint along an altitudinal gradient, potentially useful for tracing ecological processes and for paleoclimatic studies.     

Short-term signals of climate change along an altitudinal gradient in the South Alps

Short-term changes in plant species number, frequency and composition were studied along an altitudinal gradient crossing four summits from the treeline ecotone to the subnival zone in the South Alps (Dolomites, Italy). Large-scale (summit areas) and small-scale patterns (16 plots of 1 m2/summit) were monitored. After 5 years, a re-visitation of the summit areas revealed a considerable increase of species richness at the upper alpine and subnival zone (10% and 9%, respectively) and relatively modest increases at the lower alpine zone and the treeline ecotone (3% and 1%, respectively). At the small scale, the results were partly different, with species richness decreasing at the lower summits and increasing at the higher summits. The changes can most likely be attributed to climate warming effects and to competitive interactions. The main newcomers at the lower three summits were species from the treeline and the lower altitudinal zones. Only at the highest summit, the newcomers came from the alpine species pool. At the treeline ecotone, the abundance of Pinus cembra, of dwarf shrubs and clonal graminoid species increased. Here, displacements of alpine species may be predicted for the near future. At the higher summits, expansions of the established alpine species and further invasions of species from lower altitudes are forecasted.     

Tree dispersion in oak-dominated forests along an environmental gradient

Summary Spatial pattern was analyzed in seventeen stands of oak-dominated forest to address the hypothesis that species tended to be aggregated under favorable conditions and widely spaced in xeric, nutrient poor conditions. Trees were sampled at 80–100 points in each stand with the distance-to-nearest neighbor method. Soil samples were collected in each stand for analysis of total nitrogen, total phosphorus, total potassium, soil pH, soil texture, and soil organic matter. Growing season precipitation was also recorded from climate stations near each stand. Quercus stellata (Wang.) dominated 10 stands, Q. marilandica (Muenchh.) dominated three stands and these species were codominant in four stands. Principal components analysis identified a soil texture/fertility gradient across the study area. Quercus stellata and all species combined were aggregated in most stands, whereas Q. marilandica was mostly randomly distributed within a stand. Small trees of all species combined tended to be aggregated and large trees were randomly dispersed in all but two stands, suggesting competition. Mean distance between large-large pairs was always greater than mean distance between small-small pairs in all stands, but this difference was only significant in one stand. Correlations between nearest neighbor distance and combined size of nearest neighbors were significant and positive in 12 of 17 stands. In all cases, however, slopes were shallow suggesting that competition is weak in these communities and has a limited effect on spacing of neighboring trees. Contrary to our hypothesis, trees were more aggregated on coarse-textured soils with low organic matter content. For all species combined, degree of aggregation was unrelated to growing season precipitation. Aggregation appears to be common in these forests because environmental stress in many stands reduces growth rates. Trees have not yet reached a size at which competition or other interactions can greatly increase interplant distances and reduce the degree of aggregation. A simple graphical model is developed to describe the relationship between patterns, stress and competition in plant communities.     

Mitochondrial DNA variation along an altitudinal gradient in the greater white-toothed shrew, Crocidura russula

The distribution of mitochondrial control region-sequence polymorphism was investigated in 15 populations of Crocidura russula along an altitudinal gradient in western Switzerland. High-altitude populations are smaller, sparser and appear to undergo frequent bottlenecks. Accordingly, they showed a loss of rare haplotypes, but unexpectedly, were less differentiated than lowland populations. Furthermore, the major haplotypes segregated significantly with altitude. The results were inconsistent with a simple model of drift and dispersal. They suggested instead a role for historical patterns of colonization, or, alternatively, present-day selective forces acting on one of the mitochondrial genes involved in metabolic pathways.     

Changes in structure and composition of evergreen forests on an altitudinal gradient in the Venezuelan Guayana shield

There have been several ecological studies in forests of the Guayana Shield, but so far none had examined the changes in structure and composition of evergreen forests with altitude. This study describes and analyzes the structure, species composition and soil characteristics of forest stands at different altitudinal zones in Southeastern Venezuelan Guayana, in order to explain the patterns and the main factors that determine the structure and composition of evergreen forests along the altitudinal gradient. Inventories of 3 948 big (>10cm DBH) and 1 328 small (5-10cm DBH) woody stems were carried out in eleven plots, ranging from 0.1 to 1.0ha, along a 188km long transect with elevations between 290 and 1 395masl. It has been found that 1) hemiepihytes become more dominant and lianas reduce their dominance with increasing altitude and 2) the forest structure in the study area is size-dependent. Five families and 12 genera represented only 9% of the total number of families and genera, respectively, recorded troughout the gradient, but the two groups of taxa comprised more than 50% of the Importance Value (the sum of the relative density and the relative dominance) of all measured stems. Moreover, the results suggest that low species richness seems to be associated with the dominance of one or few species. Stand-level wood density (WD) of trees decreased significantly with increasing elevation. WD is an indicator of trees'life history strategy. Its decline suggests a change in the functional composition of the forest with increasing altitude. The Canonical Correspondence Analysis (CCA) indicated a distinction of the studied forests on the basis of their altitudinal levels and geographic location, and revealed different ecological responses by the forests, to environmental variables along the altitudinal gradient. The variation in species composition, in terms of basal area among stands, was controlled primarily by elevation and secondarily by rainfall and soil conditions. There are other interacting factors not considered in this study like disturbance regime, biological interactions, productivity, and dispersal history, which could affect the structure and composition of the forests in the altitudinal gradient. In conclusion, it appears that the structural and floristic variability observed in the studied transect is produced by a combination of different climates and randomly expressed local processes interacting across a complex physical landscape.