Variation in species composition and vegetation structure of succulent scrub on Tenerife in relation to environmental variation

Abstract. On Tenerife, the occurrence of environmental gradients over short distances provides a unique opportunity to investigate the relationship between vegetation and environmental factors. In the semi‐arid coastal region of Tenerife, floristic composition, species richness and vegetation structure of perennial plants have been studied in 67 locations covering the existing precipitation gradient. On the island as a whole, variation in species composition could be best explained by mean annual precipitation; at coastal sites, substrate age and soil characteristics also played a significant role. On the other hand, substrate chemistry and the type of eruptive material explained little of the floristic variation. Stand biomass was strongly correlated with mean annual precipitation and was, on the youngest lava flows studied, also affected by substrate age. The native stem succulent species made up the bulk of total biomass along the whole precipitation gradient. Disturbed and undisturbed sites differed significantly in stand biomass and cover. Species richness was correlated with precipitation and substrate age. Distribution of plant functional types was also related to the precipitation gradient. The relative abundance of hemicryptophytes and shrubs with non‐hairy leaves increased with increasing precipitation whereas the ratio of shrubs with hairy/non‐hairy leaves and succulent plants decreased. Some alien plants were quite frequent at disturbed sites but, on the whole, they contributed little to the species spectrum and to the stand biomass. Undisturbed sites remained almost free of introduced species not considering annuals.     

Primary succession of Hawaiian montane rain forest on a chronosequence of eight lava flows

Abstract. The primary-successional sere of a Hawaiian montane rain forest was inferred from an age sequence of eight closely located ‘a’ ā flows (clinker type lava); 8, 50, 140, ca. 300, ca. 400, ca. 1400, ca. 3000 and ca. 9000 yr, on a windward slope of Mauna Loa, Hawaii. All study sites (0.2 ha each) were at 1120 — 1250 m a.s.l. with 4000 mm mean annual rainfall. The 400-yr, 1400-yr, and 9000-yr flows had younger volcanic ash deposits, while the others were pure lava. Comparisons of tree size and foliar nutrients suggested that ash increased the availability of nitrogen, and subsequently standing biomass. An Unweighted Pair Group Cluster Analysis on the samples (flows) using quantitative vascular species composition revealed that clusters were correlated with age regardless of the substrate types (pure lava vs. ash), and an indirect ordination on the samples suggested that the sequence of sample scores along axis 1 was perfectly correlated with the age sequence. Although ash deposits increased biomass, they did not affect the sequence of the successional sere. Both pubescent and glabrous varieties of Metrosideros polymorpha (Myrtaceae) dominated upper canopy layers on all flows ≥ 50 yr and ≤ 1400 yr, but the pubescent variety was replaced by the glabrous on the flows ≥ 3000 yr. Lower layers were dominated initially by a matted fern, Dicranopteris linearis, up to 300 yr, and subsequently by tree ferns, Cibotium spp., to 9000 yr. The cover of Cibotium declined slightly after 3000 yr, while other native herb and shrub species increased. A ‘climax’ stage in the conventional sense was apparently not reached on the observed age gradient, because the sere changed continuously in biomass and species; this divergent successional phenomenon may be unique to Hawaii where the flora is naturally impoverished and disharmonic due to its geographic isolation in contrast to more diverse and harmonic floras in continents.     

Variation in foliar δ13C in Hawaiian Metrosideros polymorpha: a case of internal resistance?

Summary Sun leaves of Metrosideros polymorpha were collected in 51 sites on 9 lava flows that represented gradients of elevation, precipitation, substrate age, and substrate texture on Mauna Loa volcano, Hawai'i. Leaf mass per unit leaf area increased with increasing elevation on all flows, while foliar nitrogen concentration decreased with increasing elevation and increased with increasing substrate age. Foliar ¹³C became less negative with increasing elevation on the wet east-side lava flows, but not the dry northwest-side flows; it did not reflect patterns of precipitation or presumed water availability. ¹³C was very strongly correlated with leaf mass per area across all of the sites. Limited gas-exchange information suggested that calculated c_i/c_a did not decrease with elevation in association with less-negative ¹³C, and photosynthesis per unit of nitrogen was significantly reduced in high-elevation plants. These results are consistent with a substantial internal resistance to CO₂ diffusion in the thick Metrosideros polymorpha leaves in high elevation sites.Published as a contribution to the International Union for Biological Sciences Tropical Mountain Ecosystems Program, and as Carnegie Institution of Washington Department of Plant Biology Publication Number 1065     

Large scale stoat control to protect mohua (

Vegetation succession on 4 recent (1852-1942) montane lava Rows on Mauna Loa, Hawaii, was remeasured 22 years after the first measurement in 1967. Colonisation patterns of vascular plant species were observed on a new lava flow (1984) which overwhelmed part of the earlier studied 1852 flow. An influx of adventive species, positively correlated with flow age, was noted at the remeasured sites; most were herbs and grasses that do not appear to interfere with the succession to Metrosideros- dominated forest. Some indigenous species important in older forest, e.g., Cibotium glaucum, had apparently colonised all four remeasured flows regardless of flow age. Densities and total basal area of Metrosideros polymorpha increased on all flows, but a closed-canopy forest had not yet developed. Vascular plant aggregations comprising a mixture of adventive and indigenous species were found on the 1984 flow associated with soil or logs of the overwhelmed forest. This phenomenon may allow rare individual Metrosideros trees to be in place on a new flow within 10 years of its formation. A closed-canopy, self-thinned Metrosideros forest can develop within 400 years but dieback of colonising Metrosideros individuals and/or invasion of adventive species capable of altering ecosystem processes can delay this process.     

Patterns of Clonal Diversity in Dicranopteris linearis on Mauna Loa,Hawaii1

The clonal mat-forming fern, Dicranopteris linearis (N. L. Burm.) Underw., dominates vast areas of rainforests on the windward slopes of Mauna Loa Volcano on the island of Hawai'i. Because clone size has important ecological and evolutionary consequences in such a dominant species, we used isozyme analysis to investigate clone size and other aspects of genetic diversity and reproduction over a broad range of environmental conditions on primary successional sites (pahoehoe lava substrates). Isozyme analysis provided a measure of the upper limit of clonal size in this interdigitating clonal species. Each 0.5-ha primary successional site on Mauna Loa was comprised of a minimum of two to four clones. Genetic diversity in Dicranopteris was low; of 32 putative loci investigated, only 4 were polymorphic, with 2 or 3 alleles/locus. Over the 17 study locations on Mauna Loa and Kilauea Volcanoes, we identified nine multilocus genotypes based on unique combinations of allozymes. Seven of the nine genotypes were heterozygous for at least one locus, evidence of an intergametophytic mating system. Highly dispersible spores, coupled with intergametophytic mating should promote higher genetic diversity. We propose that the following factors contributed to low genetic diversity: founder effects; extreme isolation from mainland gene pools; high potential for mating among different gametophytes produced from the same sporophyte; relatively low numbers of safe sites for gametophyte establishment over space and time; and long-term reliance on vegetative growth. Leaf phenotypes were associated with genotype, but also with environmental conditions. Enough variability within a genotype existed to support the current treatment of Hawaiian Dicranopteris as one species. Vegetative growth was the primary means by which Dicranopteris covered the landscape. Nevertheless, spore production, gametophyte establishment, and sexual reproduction were absolutely essential for colonization of the few favorable microsites available on pahoehoe lava substrates of Mauna Loa following lava eruptions, dieback, and similar landscape-level disturbances.     

The Mauna Loa environmental matrix: foliar and soil nutrients

Summary The accumulation of total carbon, nitrogen, and phosphorus in soils, available soil nutrients, and foliar nutrients in the native dominant Metrosideros polymorpha were determined across a wide elevational range on 9 lava flows on Mauna Loa, Hawai'i. The flows included a young (<140 y) and an old (>2800 y) áá (rough surface texture) and phoehoe (smooth) flow on the wet east and dry northwest side of the mountain. Soil element pools and nutrient availability increased with flow age independent of climate. The dry sites accumulated organic matter and nutrients more slowly than comparable wet sites, but relative nutrient availability to plants (as indicated by soil assays and foliar nutrients) was greater in the dry sites. Accumulation of soil organic matter and nutrients occurred most rapidly in lowerelevation sites on the young flows, but the largest accumulations occurred at higher elevations on old flows. The range of sites sampled represents a complete and largely independent matrix of major factors governing ecosystem structure and function.     

Rainfall seasonality determines annual/perennial grass balance in vegetation of Mediterranean Iberian

Much recent attention has been focused on the invasion and dominance of annual grass species in areas thought to have been historically dominated by perennial life forms. Explanations of this phenomenon in the literature have focused on two mechanisms favoring annuals: ruderal strategy associated with disturbance, and stress escaping associated with dry sites or deserts. Here I present evidence from vegetation surveys at 50 sites across a 1,200 km band of the Iberian Peninsula—a source region for many invasive annuals—showing that relative annual versus perennial grass composition is not well correlated with degree of disturbance or average annual precipitation. However, annual dominance is strongly and significantly linked to the seasonality of precipitation, in particular the relative intensity of summer drought. Disturbance was significantly associated with annual grass dominance in Iberia, but with much less explanatory power than summer drought intensity. Slope, aspect, and soil parent material were not significantly correlated with annual versus perennial grass dominance. These results suggest that subtle differences in rainfall seasonality largely drive grass composition in herbaceous Mediterranean vegetation. Furthermore, the patterns of annual grass invasion observed in the world’s other Mediterranean climate regions may be associated with similar climatic drivers.     

A landscape perspective of the Hawaiian rain forest dieback

Abstract. Throughout the 1960s and 1970s there was a rapid decline and canopy dieback in the Metrosideros polymorpha dominated rain forest of Hawai'i. An analysis of air photo sets from 1954, 1965, and 1972, covering the windward slopes of Mauna Kea and Mauna Loa, gave support for an alien disease hypothesis. A total demise of the native forest was predicted for the early 1990s. This prediction as well as the disease hypothesis proved to be wrong. Various searches for a single climatic cause also failed to explain the dieback. The spatial dynamics of the dieback phenomenon were newly analyzed with an additional air photo set from 1977 and by using GIS with spatial statistics. Two juxtaposed and climatically similar landscape matrix samples of ca. 200 km², one each on Mauna Loa and Mauna Kea, were subjected to an analysis of landform heterogeneity and superimposed dieback patterns. The Mauna Loa matrix displays up to 15 000 yr old lava flows, while the Mauna Kea matrix displays up to 250 000 yr old substrates. Initiation of dieback occurred simultaneously on both mountains and was highly correlated with poorly-drained sites. The progression of dieback, however, followed a gradient of decreasing soil moisture, which often terminated at clearly recognizable substrate boundaries in the Mauna Loa matrix and moved over well-drained hill sites in the Mauna Kea matrix. Metrosideros dieback spread across the entire spectrum of volcanic substrates and habitat moisture regimes and developed from a smaller into a larger patch mosaic. By 1977, ca. 50 % of the forest area in both sample matrices had gone into dieback. Thereafter, the dieback came to a halt. The domino-type collapse, which frequently came to a halt at volcanic substrate boundaries, indicates that stands in better drained sites were also predisposed to die. Stands on adjoining substrates often survived. Substrates with dieback stands displayed no other obvious vigor-reducing stresses. The canopy trees on such substrates may have a common history, such as a major disturbance (including dieback) that synchronized stand development in the past. Subsequent weather disturbances and other abiotic/endogenous stresses associated with stand maturation, such as nutrient limitations and stand-level senescence, may reinforce a rhythmic synchrony over several generations of canopy cohorts.     

Spatial patterns of microsite colonisation on two young lava flows on Mount Hekla,Iceland

Questions: How does vegetation first establish on newly‐formed lava substrates? Do very small (cm) and meso‐scale (m) variations in the physical environment influence this process and subsequent vegetation development? Location: Mount Hekla, southern Iceland (64°00’ N, 19°40’ W). Methods: Data on vegetation structure and the incidence of ‘safe sites’ suitable for colonisation were collected from high and low points on the surfaces of lava flows emplaced during the 1991 and 2000 A.D. eruptions of Mount Hekla. Effects of flow age and meso‐topographic position on vegetation structure (moss cover, patch density, stem length) were assessed by two‐way analyses of variance. The distributions of colonisation events and available safe sites were analysed using point pattern techniques. Results: Rapid colonisation of the lava surface was observed, despite stressful environmental conditions. The 1991 and 2000 flows differed significantly in vegetation structure, but there were no significant differences in moss cover, patch density and stem length between ‘high’ and ‘low’ sites. Conclusions: Colonisation events are invariably associated with small‐scale irregularities on the surface of the lava. The colonisation process appears to be spatially random. Development of the moss ‘carpet’ proceeds by vertical thickening and lateral growth and coalescence of moss patches that establish in ‘safe sites’. This process is rapid, with close to 100% of available safe sites exploited within 20 years. Topographic position makes no difference to the very early stages of vegetation development and cannot be used to ‘forecast’ the later stages of development.     

Elevational and age gradients in hawaiian montane rainforest: foliar and soil nutrients

Summary Soils and plants were sampled along an elevational gradient from 265–1675 m on a 133-and a 3100-year-old lava flow on Mauna Loa, Hawai'i. Soil organic matter and nutrients accumulated more rapidly at low elevation on the young flow, but reached higher levels at higher elevation on the old flow. Foliar nitrogen and phosphorus concentrations were less and specific leaf weight greater for Metrosideros polymorpha leaves collected at high versus low elevations and on the young versus the old flow. Foliar ¹³C was strongly correlated with specific leaf weight across the range of sites sampled.Published as a contribution to the Tropical Mountain Ecosystem Program of the IUBS Decade of the Tropics     

Ecological and Genetic Implications of Foliar Polymorphism inMetrosideros polymorphaGaud. (Myrtaceae) in a Habitat Matrix on Mauna Loa, Hawaii

Glabrous and pubescent foliar phenotypes ofMetrosideros polymorphaoccursympatrically on Mauna Loa, Hawaii. This coexistence has beensuggested, by earlier workers, to represent distinct genotypes,each of which produces progenies with a homogenous foliar phenotype.We re-examined the question of to what extent siblings fromthe same parents varied in phenotype when grown under the samecommon-garden conditions. Seeds were collected from parentaltrees in a matrix of two substrate ages (114–140 yrvs.approx.1200–3200 yr old) at five elevations from lowland (100m) to treeline (2470 m) on Mauna Loa. Seeds were sown and grownunder identical environmental conditions at 1200 m. In fieldparental populations, two forms converged into pubescent formswith decreasing rainfall (<2000 mm) on both substrates, andinto glabrous forms with increasing soil age where rainfallwas ample. The 4-yr old seedlings from pubescent parents invariablyexpressed both glabrous and pubescent phenotypes for all altitudesand for both substrate ages except for the young 1980-m site.The seedlings from glabrous parents tended to express only theglabrous phenotypes except for the young 1280-m site. Overall,all populations on the young lava flow (glabrous and pubescentindividuals combined if coexisting) could produce progeniesof both phenotypes except at 1980 m. Cuticle thickness on theadaxial lamina surface was positively correlated with the magnitudeof pubescence (P=0.05) among siblings from the same parents,ranging from a mean cuticle thickness of 6.0 µm for glabrousto 15.7 µm for the most pubescent individuals. Greatermagnitudes of pubescence were associated with more negativeosmotic potentials at turgor loss point, and with greater maximumCO₂assimilation rates at the point of light saturation. Thisphysiological and anatomical polymorphism, together with spatiallyand temporally varying selective forces, appears to result inelevationally and successionally patternedM. polymorphapopulationson the wet slope of Mauna Loa. Common-garden experiment; cuticle; foliar pubescence; Hawaii; Metrosideros polymorpha,natural selection; photosynthesis; polymorphism; primary succession; water potential     

Larval habitat for the avian malaria vector Culex quinquefasciatus (Diptera: Culicidae) in altered mid‐elevation mesic‐dry forests in Hawai'i

Effective management of avian malaria (Plasmodium relictum) in Hawai'i's endemic honeycreepers (Drepanidinae) requires the identification and subsequent reduction or treatment of larval habitat for the mosquito vector, Culex quinquefasciatus (Diptera: Culicidae). We conducted ground surveys, treehole surveys, and helicopter aerial surveys from 2001–2003 to identify all potential larval mosquito habitat within two 100+ ha mesic‐dry forest study sites in Hawai'i Volcanoes National Park, Hawai'i; ‘Ainahou Ranch and Mauna Loa Strip Road. At ‘Ainahou Ranch, anthropogenic sites (43%) were more likely to contain mosquitoes than naturally occurring (8%) sites. Larvae of Cx. quinquefasciatus were predominately found in anthropogenic sites while Aedes albopictus larvae occurred less frequently in both anthropogenic sites and naturally‐occurring sites. Additionally, moderate‐size (≈20–22,000 liters) anthropogenic potential larval habitat had >50% probability of mosquito presence compared to larger‐ and smaller‐volume habitat (<50%). Less than 20% of trees surveyed at ‘Ainahou Ranch had treeholes and few mosquito larvae were detected. Aerial surveys at ‘Ainahou Ranch detected 56% (95% CI: 42–68%) of the potential larval habitat identified in ground surveys. At Mauna Loa Strip Road, Cx. quinquefasciatus larvae were only found in the rock holes of small intermittent stream drainages that made up 20% (5 of 25) of the total potential larval habitat. The volume of the potential larval habitat did not influence the probability of mosquito occurrence at Mauna Loa Strip Road. Our results suggest that Cx. quinquefasciatus abundance, and subsequently avian malaria, may be controlled by larval habitat reduction in the mesic‐dry landscapes of Hawai'i where anthropogenic sources predominate.     

Primary succession of the warm-temperate broad-leaved forest on a volcanic island, Miyake-jima, Japan

The primary successional sere of a warm-temperate forest was inferred from an age sequence of four basaltic volcanic substrates (16-, 37- and 125-yr-old lava flows, and volcanic ejecta older than 800 years) at several altitudes (ranging from 30 to 550 m) on Miyake-jima Island. The DCA analysis of species composition indicated that the vegetation is patterned by substrate age and altitude. The vegetation successionally changed fromAlnus sieboldiana shrub, toMachilus thunbergii andPrunus speciosa forest, toCastanopsis sieboldii forest at all altitudes. The rate of succession seems to be slower at higher altitudes. The above-ground biomass was 20 kg/m² (160 m a.s.l.) and 12 kg/m² (300 m a.s.l.) on the 125-yr-old lava flow, and 54 kg/m² (100 m a.s.l.) and 16 kg/m² (350 m a.s.l.) on the old ejecta. However, the large difference of above-ground biomass between low versus high altitudes on the old ejecta might not only reflect climatic differences but also past human disturbances at the higher altitude. In spite of the colder climate of Miyake-jima the rate of above-ground biomass accumulation here was considerably faster than that of a Hawaiian montane tropical evergreen forest on the same basaltic lava substrate. We suggest that the faster above-ground biomass development is due to the facilitation effects of N-fixation byAlnus. The inorganic N soil concentration was extremely high on the 37- and 125-yr-old lava flows whereAlnus was dominant. The foliar N concentration inAlnus was 2% irrespective of substrate age. The deposition of N fromAlnus via litterfall would decrease the soil C/N ratio, which in turn facilitates net soil N mineralization and consequently provides an ample supply of inorganic N to plants. N limitation on vegetation development, which is prevalent during the early stage of succession on volcanic lava flows or similar substrates elsewhere, is thus alleviated.     

Persistent versus transient tree encroachment of temperate peat bogs: effects of climate warming and drought events

Peatlands store approximately 30% of global soil carbon, most in moss‐dominated bogs. Future climatic changes, such as changes in precipitation patterns and warming, are expected to affect peat bog vegetation composition and thereby its long‐term carbon sequestration capacity. Theoretical work suggests that an episode of rapid environmental change is more likely to trigger transitions to alternative ecosystem states than a gradual, but equally large, change in conditions. We used a dynamic vegetation model to explore the impacts of drought events and increased temperature on vegetation composition of temperate peat bogs. We analyzed the consequences of six patterns of summer drought events combined with five temperature scenarios to test whether an open peat bog dominated by moss (Sphagnum) could shift to a tree‐dominated state. Unexpectedly, neither a gradual decrease in the amount of summer precipitation nor the occurrence of a number of extremely dry summers in a row could shift the moss‐dominated peat bog permanently into a tree‐dominated peat bog. The increase in tree biomass during drought events was unable to trigger positive feedbacks that keep the ecosystem in a tree‐dominated state after a return to previous ‘normal’ rainfall conditions. In contrast, temperature increases from 1 °C onward already shifted peat bogs into tree‐dominated ecosystems. In our simulations, drought events facilitated tree establishment, but temperature determined how much tree biomass could develop. Our results suggest that under current climatic conditions, peat bog vegetation is rather resilient to drought events, but very sensitive to temperature increases, indicating that future warming is likely to trigger persistent vegetation shifts.     

Effects of additional N on herbaceous species of desertified steppe in arid regions of China: a four-year field study

Human-induced nitrogen (N) enrichment is impacting the structure and function of grassland ecosystems, although few studies have been conducted in arid and semi-arid ecosystems under field conditions. Here, a 4-year multi-level N addition experiment under ambient precipitation in a typical temperate desert steppe in arid regions of China was designed to evaluate the impact of N enrichment on the herbaceous vegetation community in desert ecosystems. The results showed that species richness declined in response to N addition during all 4 years, while aboveground biomass increased in the relatively wet year (2007) and decreased in the relatively dry years (2008–2010) according to N addition level, as compared to the control. Plant community composition differed among plant functional groups (PFGs) based on added N. Perennial grasses (PG) benefited more compared with annuals and perennial forbs (PF) from N enrichment in terms of biomass production under conditions of high precipitation in 2007. In relatively dry years, biomass production of all PFGs was strictly dependent on inter- and intra-annual precipitation, which led PG to dominate in 2008 (with a dry spring) and PF to dominate in 2009 and 2010. Our results demonstrate that the impact of N enrichment on the herbaceous vegetation community in desert steppes may be strongly dependent on natural precipitation patterns characterized by shifts in plant community composition, particularly in terms of biomass production.     

Relationships of Biomass with Environmental Factors in the Grassland Area of Hulunbuir,China

Many studies have focused on the relationship between vegetation biomass and environmental factors in grassland. However, several questions remain to be answered, especially with regards to the spatial pattern of vegetation biomass. Thus, the distributed mechanism will be explored in the present study. Here, plant biomass was measured at 23 sites along a transect survey during the peak growing season in 2006. The data were analyzed with a classification and regression tree (CART) model. The structural equation modeling (SEM) was conducted to explicitly evaluate the both direct and indirect effects of these critical environmental elements on vegetation biomass. The results demonstrated that mean annual temperature (MAT) affected aboveground biomass (AGB) scored at −0.811 (P<0.05). The direct effect of MAT on belowground biomass (BGB) was −0.490 (P<0.05). The results were determined by SEM. Our results indicate that AGB and BGB in semi-arid ecosystems is strongly affected by precipitation and temperature. Future work shall attempt to take into account the integrated effects of precipitation and temperature. Meanwhile, partitioning the influences of environmental variations and vegetation types are helpful in illuminating the internal mechanism of biomass distribution.     

九龙江西溪漳州段浮游植物组成与环境因子的关系

分别在2010年的丰、平、枯3个时期,从浮游植物种类组成、生物量(叶绿素a含量)及其粒级结构等指标对九龙江漳州段浮游植物组成进行了调查,同时分析了其与环境因子间的关系,评估了该河段的水质质量。结果表明:研究区域微微型浮游植物占总生物量比例与温度存在显著的正相关性(P<0.05),小型浮游植物占总生物量比例与总氮成显著的负相关(P<0.05)。共鉴定出浮游植物7门59种,且在各时期均有差异;丰水期与枯水期相比,蓝藻比例上升(P<0.05),甲藻比例下降(P<0.05);研究水域Shannon-Wiener指数在1.0～3.0,为中度污染。典型对应分析表明,温度和营养盐是影响九龙江西溪漳州段浮游植物群落变化的主要环境因子。     

Analysis of factors regulating ecosystemdevelopment on Mauna Loa using the Century model

We used the Century model to evaluateenvironmental controls over ecosystem developmentduring the first 3500 y of primary succession onpahoehoe (i.e., relatively smooth, solid) lava flowsof wet, windward Mauna Loa, Hawaii. The Century modelis a generalized ecosystem model that simulatescarbon, nitrogen and phosphorus dynamics forplant-soil systems. Preliminary results indicated theneed to modify the model to include the effects ofsoil C accumulation on soil water storage anddrainage. The modified model was parameterized tosimulate observed values of aboveground productivity,biomass and soil element pools on a 3400-y-old site at700 m elevation. Testing the model parameters at 1660m elevation indicated that N inputs were lower andsoil water drainage rates were slower at the higherelevation. We applied the modified and fullyparameterized model to simulate ecosystem attributesduring primary succession at five elevations, andconducted single-factor experiments with the model toidentify the specific influences of variations intemperature, nutrient inputs, and rainfall on modeledecosystem characteristics.Simulated aboveground productivity (ANPP), net N andP mineralization, and biomass element pools allincreased through time at each elevation, and alldeclined with increasing elevation at each point intime. After 3500 y of succession none of theseattributes had reached a stable asymptote, butasymptotes were approached more quickly, andsuccession was therefore faster, at lower than athigher elevations. Simulated soil organic matter(SOM) pools increased with elevation, despite thatplant productivity declined. These results, andsimilar comparisons among rainfall regimes, suggestthat SOM pools were more sensitive to factorscontrolling decay than production rates.Within elevations and temperature regimes, nutrientavailability was the most important factor controllingsimulated rates of plant productivity, biomass, anddetritus accumulation during ecosystem development. Through time, SOM accumulations alleviated nutrientlimitations to plants, but simulated productivityremained highly dependent upon externally suppliednutrients even after 20,000 y. Rainfall had two maineffects on nutrient availability within the model: (1)it increased rates of leaching, and thus depletednutrient supplies; and (2) it exacerbated soil floodingand thereby decreased nutrient turnover rates. Highrainfall on windward Mauna Loa maintains oligotrophicconditions through time despite continuous N and Pinputs.     

Carabid beetles as indicators for shrub encroachment in dry grasslands

Semi-natural grasslands are key habitats for biodiversity conservation in Central Europe. Shrub encroachment is one of the most threatening drivers of grassland degradation and affects soil properties, microclimate, and vegetation with possible impacts on higher trophic levels. We aimed to analyse the impact of shrub encroachment with broom (Cytisus scoparius) on carabid beetle diversity, species composition, and functional traits. In a field study on dry grasslands on the island of Hiddensee (Germany) we studied 15 sites along a gradient of increasing broom encroachment and classified them into three dry grassland types with low, medium, and high shrub cover. Our results provide evidence that shrub encroachment initially has positive effects on species richness and activity densities of dry grassland carabids. Carabid species composition differed among differently shrub-covered dry grassland types, and sites with low and high shrub cover were each characterised by unique carabid assemblages. The species composition of sites with a medium shrub biomass had a transitional character and contained species which are typical for open dry grassland, but also shared species with sites with a high shrub cover. Among functional trait parameters investigated, especially the body size of carabid beetles was related to environmental parameters associated with shrub encroachment. Body size was positively correlated to shrub biomass and soil humidity, but negatively to temperature. Eurytopy values of carabids were related to high litter cover, i.e. habitat generalist (eurytopic) species mainly occurred in densely shrub-encroached sites. In order to preserve unique carabid assemblages of open dry grasslands with stenotopic and smaller species, it is most important to prevent a shrub encroachment higher than about 60% cover. For management we suggest extensive grazing (by cattle, sheep or horses) to prevent shrub encroachment on dry grasslands. In areas with high shrub cover additionally the use of goats or mechanical removal of shrubs might be necessary.     

Cryogenic Minerals in Hawaiian Lava Tubes: A Geochemical and Microbiological Exploration

The Mauna Loa volcano, on the Island of Hawaii, has numerous young lava tubes. Among them, two at high altitudes are known to contain ice year-round: Mauna Loa Icecave (MLIC) and the Arsia Cave. These unusual caves harbor cold, humid, dark, and biologically restricted environments. Secondary minerals and ice were sampled from both caves to explore their geochemical and microbiological characteristics. The minerals sampled from the deep parts of the caves, where near freezing temperatures prevail, are all multi-phase and consist mainly of secondary amorphous silica SiO₂, cryptocrystalline calcite CaCO₃, and gypsum CaSO₄·2H₂O. Based on carbon and oxygen stable isotope ratios, all sampled calcite is cryogenic. The isotopic composition of falls on the global meteoric line, indicating that little evaporation has occurred. The microbial diversity of a silica and calcite deposit in the MLIC and from ice pond water in the Arsia Cave was explored by analysis of ～50,000 small subunit ribosomal RNA gene fragments via amplicon sequencing. Analyses reveal that the Hawaiian ice caves harbor unique microbial diversity distinct from other environments, including cave environments, in Hawaii and worldwide. Actinobacteria and Proteobacteria were the most abundant microbial phyla detected, which is largely consistent with studies of other oligotrophic cave environments. The cold, isolated, oligotrophic basaltic lava cave environment in Hawaii provides a unique opportunity to understand microbial biogeography not only on Earth but also on other planets.