Diplycosia rigidifolia sp. nov. (Ericaceae) from Borneo,Sabah, Malaysia

Diplycosia rigidifolia sp. nov. from Mount Kinabalu, Borneo, Sabah, Malaysia, is described and illustrated. This species is similar to D. urceolata, but differs by its shorter petiole, thicker, strictly elliptic leaf blades, longer pedicels, calyx lobes with sharply acuminate apex, and larger purplish black fruiting calyx. The species is known only from Mt Kinabalu in northern Sabah state, Malaysia.     

The mid‐domain effect matters: simulation analyses of range‐size distribution data from Mount Kinabalu,Borneo

Aim In simulation exercises, mid‐domain peaks in species richness arise as a result of the random placement of modelled species ranges within simulated geometric constraints. This has been called the mid‐domain effect (MDE). Where close correspondence is found between such simulations and empirical data, it is not possible to reject the hypothesis that empirical species richness patterns result from the MDE rather than being the outcome (wholly or largely) of other factors. To separate the influence of the MDE from other factors we therefore need to evaluate variables other than species richness. The distribution of range sizes gives different predictions between models including the MDE or not. Here, we produce predictions for species richness and distribution of range sizes from one model without the MDE and from two MDE models: a classical MDE model encompassing only species with their entire range within the domain (range‐restricted MDE), and a model encompassing all species with the theoretical midpoint within the domain (midpoint‐restricted MDE). These predictions are compared with observations from the elevational pattern of range‐size distributions and species richness of vascular plants. Location Mount Kinabalu, Borneo. Methods The data set analysed comprises more than 28,000 plant specimens with information on elevation. Species ranges are simulated with various assumptions for the three models, and the species simulated are subsequently subjected to a sampling that simulates the actual collection of species on Mount Kinabalu. The resulting pattern of species richness and species range‐size distributions are compared with the observed pattern. Results The comparison of simulated and observed patterns indicates that an underlying monotonically decreasing trend in species richness with elevation is essential to explain fully the observed pattern of richness and range size. When the underlying trend is accounted for, the MDE model that restricts the distributions of theoretical midpoints performs better than both the classical MDE model and the model that does not incorporate geometric constraints. Main conclusions Of the three models evaluated here, the midpoint‐restricted MDE model is found to be the best for explaining species richness and species range‐size distributions on Mount Kinabalu.     

A redescription of Macrostylophora borneensis (Jordan, 1926) (Siphonaptera-Ceratophyllidae-Ceratophyllinae), a flea of the Malaysian sub-region and description of an allied species

This note redescribes M. borneensis and describes M. traubi n. sp. based on the known specimens from the two sampling localities (holotype from Mount Murud and further recorded from Mount Kinabalu by Traub). The clearly allied, but clearly distinct species for the two localities, were recorded.     

Macrostylophora kinabaluae n. sp. (Insecta: Siphonaptera: Ceratophyllidae), a new flea from Sabah (Borneo, Eastern Malaysia)

This species, known only by a single male, is described from Mount Kinabalu and thus is recorded from the same area as Macrostylophora borneensis (Jordan, 1926), teste Traub (1972). It is distinguished from its congeneric and characterized by the absence of eriged setae on the thorax and first abdominal tergits, as well as by the shape of terminal segments and the phallosom. Macrostylophora kinabaluae was found to parasitize the Sciurid rodent Callosciurus prevostii that is widespread throughout most areas in the Malaysian subregion.     

Canopy structure of tropical and sub-tropical rain forests in relation to conifer dominance analysed with a portable LIDAR system

Background and Aims

Globally, conifer dominance is restricted to nutient-poor habitats in colder, drier or waterlogged environments, probably due to competition with angiosperms. Analysis of canopy structure is important for understanding the mechanism of plant coexistence in relation to competition for light. Most conifers are shade intolerant, and often have narrow, deep, conical crowns. In this study it is predicted that conifer-admixed forests have less distinct upper canopies and more undulating canopy surfaces than angiosperm-dominated forests.

Methods

By using a ground-based, portable light detection and ranging (LIDAR) system, canopy structure was quantified for old-growth evergreen rainforests with varying dominance of conifers along altitudinal gradients (200–3100 m a.s.l.) on tropical and sub-tropical mountains (Mount Kinabalu, Malaysian Borneo and Yakushima Island, Japan) that have different conifer floras.

Key Results

Conifers dominated at higher elevations on both mountains (Podocarpaceae and Araucariaceae on Kinabalu and Cupressaceae and Pinaceae on Yakushima), but conifer dominance also varied with soil/substrate conditions on Kinabalu. Conifer dominance was associated with the existence of large-diameter conifers. Forests with higher conifer dominance showed a canopy height profile (CHP) more skewed towards the understorey on both Kinabalu and Yakushima. In contrast, angiosperm-dominated forests had a CHP skewed towards upper canopy, except for lowland dipterocarp forests and a sub-alpine scrub dominated by small-leaved Leptospermum recurvum (Myrtaceae) on Kinabalu. Forests with a less dense upper canopy had more undulating outer canopy surfaces. Mixed conifer–angiosperm forests on Yakushima and dipterocarp forests on Kinabalu showed similar canopy structures.

Conclusions

The results generally supported the prediction, suggesting that lower growth of angiosperm trees (except L. recurvum on Kinabalu) in cold and nutrient-poor environments results in a sparser upper canopy, which allows shade-intolerant conifers to co-occur with angiosperm trees either as emergents or as codominants in the open canopy.     

Comparative study of additive basal area of conifers in forest ecosystems along elevational gradients

We examined the basal area of two life forms (conifers vs. broadleaf trees) along elevational gradients on Yakushima Island, Japan and on two series of geological substrate on Mount Kinabalu, Borneo. On Yakushima, total stand basal area abruptly increased from 700 to 1,050 m in accordance with the high dominance of conifers, indicating the presence of additive basal area of conifers in conifer–broadleaf mixed forests at higher elevations (1,050–1,300 m). Along two substrate series on Kinabalu, some forests at higher elevations (1,860–3,080 m) showed relatively high dominance of conifers, but conifer basal area did not appear to be additive. Conifers were emergents above the canopy of broadleaf trees in mixed forests on Yakushima, but two life forms usually coexisted in the single-story canopy in mixed forests on Kinabalu. Litterfall rate as a surrogate of productivity decreased with decreasing temperature along elevation on both the sites, but the rate of decrease was slower on Yakushima, where mixed forests at higher elevations showed relatively high rates. Thus, we suggest that additive basal area of conifers was linked to their emergent status, and that it enhanced productivity by complementary use of light by two life forms that occupy different stories. On Yakushima, typhoons are a major disturbance, but do not severely limit the height growth of conifers, allowing the development of two-story mixed forests. On Kinabalu, a major disturbance is El Niño-driven drought, and hydraulic limitation to tree height may explain the non-additive and non-emergent nature of conifers.     

The determinants of land snail diversity along a tropical elevational gradient: insularity,geometry and niches

Aim We investigated the patterns of species richness in land snails and slugs along a tropical elevational gradient and whether these patterns correlate with area, elevation, geographic constraints, and productivity. We did so both at the scale at which land snail population processes take place and at the coarser scale of elevational zones. Location Mount Kinabalu (4096 m) and the adjacent Mount Tambuyukon (2588 m) in Kinabalu Park, Sabah, Malaysian Borneo. Methods We used an effort‐controlled sampling protocol to determine land snail and slug species richness in 142 plots of 0.04 ha at elevations ranging from 570 to 4096 m. Extents of elevational ranges were determined by interpolation, extended where appropriate at the lower end with data from lowlands outside the study area. We used regression analysis to study the relationships between species density and richness on the one hand and elevation and area on the other. This was done for point data as well as for data combined into 300‐m elevational intervals. Results Species density (based on the individual samples) showed a decline with elevation. Elevational range length profiles revealed that range lengths are reduced at greater elevations and that a Rapoport effect is absent. Diversity showed a mild mid‐domain effect on Kinabalu, but not on Tambuyukon. When the data were combined into 300‐m elevational intervals, richness correlated more strongly with elevation than with area. Ecomorphospace was seen to shrink with increasing elevation. Main conclusions The elevational species richness patterns show the combined effects of (1) reduced niche diversity at elevations with lower productivity and (2) historical events in which the upward migration of lowland species as well as the speciation of highland endemics took place.     

On the relationships between leaf-litter lignin and net primary productivity in tropical rain forests

We investigated if tropical rainforest trees produced more-lignified leaves in less productive environments using forests on Mount Kinabalu, Borneo. Our investigation was based on two earlier suggestions that slower litter decomposition occurs under less productive forests and that trees under resource limitation invest a large amount of carbon as lignin as a defense substance to minimize the loss from herbivores. When nine forests at different altitudes (700–3100 m) and soil conditions (derived from sedimentary or ultrabasic rocks) but with the same gentle relief position were compared, the concentrations of leaf-litter lignin were positively correlated with litterfall rates and leaf-litter nitrogen concentrations. These patterns would be reinforced in intact leaves if the effects of resorption at the time of leaf shedding were taken into account, because greater magnitude of resorption of mobile elements but not of lignin would occur in less productive environments (i.e. dilution of lignin in intact leaves). These results did not support earlier suggestions to explain the variation of leaf-litter lignin. Instead, we suggest that lower lignin contents are adaptive to recycle minerals without retarding decomposition in less productive environments.     

NaCl‐induced accumulation of glycinebetaine in four subtropical halophytes from Pakistan

Variation in both the total content and the concentration of glycinebetaine in response to increasing soil salinity was studied in the salt‐secreting Atriplex griffithii Moq., the leaf succulent Suaeda fruticosa (L.) Forssk., the stem succulent Haloxylon recurvum Bunge ex Boiss. and the osmotically adjusting desert grass Halopyrum mucronatum (L.) Stapf. collected from a subtropical maritime desert in Pakistan. Glycinebetaine content (mmol kg⁻¹ dry weight) increased with increasing NaCl in Atriplex griffithii, Haloxylon recurvum and Halopyrum mucronatum , but peaked at 600 m M NaCl for Suaeda fruticosa and declined thereafter. Glycinebetaine concentration (mmol l⁻¹ tissue water) increased with increasing salinity in all four halophyte species and was sufficiently high to serve as an osmoticum in all cases.     

Productive phenology of tropical montane forests: Fertilization experiments along a moisture gradient

Phenological responses of leaves and roots were studied in the tropical montane forests of Mount Kinabalu, Borneo. Soil nutrient supply, in addition to the supply of light and water, is a potential abiotic factor influencing plant phenology in the tropics. The main objective of this study was to evaluate the contribution of soil nutrient supply to plant productive phenology. Fertilization experiments, including controls, nitrogen fertilized and nitrogen and phosphorus fertilized treatments, were conducted on three vegetation types in different moisture environments. Responses of leaves and roots were compared among treatments and among vegetation types. Leaf flushing was induced by nitrogen fertilization in the upper montane forest, where extremely wet moisture conditions are associated with cloud cover. This induction of leaf flushing by fertilization was not observed in the other forests. Root growth was suppressed by fertilization when leaf flushing was not induced by fertilization. These results indicate that soil pulsed nutrient release could be a cue for leaf flushing in a tropical wet environment, and that leaf phenology could be regulated by external abiotic factors and root phenology could be regulated by internal plant demands.     

Patterns of species diversity on an oceanic versus a continental island mountain: a hypothesis on species diversification1

Abstract. The vegetation on a wet mountain slope on Haleakala (an oceanic island) is compared with that on Kinabalu (a continental island) to examine relationships between regional floristic richness and α- and β-diversities. The two mountains are similar in their constant tropical climate, generic and family-level floristic elements and geological age of the summit regions, but different in regional floristic richness (rich on Kinabalu vs. poor on Haleakala). α-diversity of canopy and subcanopy tree species was much higher on Kinabalu than in comparable zones on Haleakala. Average turnover rate of species (as logarithmic community similarity) on the slope was one order of magnitude greater on Kinabalu than on Haleakala (0.127 vs. 0.017 per 100 m alt.). While there were genera with wide altitudinal ranges on both mountains, a large proportion of the genera was differentiated into parapatric altitudinal congeners on Kinabalu. By contrast, most genera are altitudinally monotypic on Haleakala. The number of sympatric congeners per genus, and the frequency of multi-specific genera per plot were high on lower slopes but decreased with increasing altitude on Kinabalu, whereas the values were low across all altitudes on Haleakala. These patterns suggest that sympatric and parapatric species radiation was less on Haleakala than on Kinabalu. This may be related to Haleakala's initially poor and disharmonic flora.     

Soil phosphorus fractionation and phosphorus-use efficiencies of tropical rainforests along altitudinal gradients of Mount Kinabalu, Borneo

We studied soil phosphorus (P) fractionation and P-use efficiencies (PUEs) of rainforests along altitudinal gradients (700–3100 m) on two types of parental rocks (sedimentary versus ultrabasic) on Mount Kinabalu, Borneo. Sedimentary rocks were known to contain more quartz (which does not adsorb P) than ultrabasic rocks. The pool (top 30 cm) of total P was always greater on sedimentary (ranging from 34.9 to 72.6 g m^–2) than on ultrabasic (9.0–29.2 g m^–2) rocks at comparable altitudes. Accordingly, the pools of organic P and labile inorganic P were always greater on sedimentary than on ultrabasic rocks. The pool of primary mineral, calcium P increased upslope from 1.7 to 4.3 g m^–2 on sedimentary rock, suggesting that the altitudinal sequence of the sites reflected a decreasing magnitude of soil weathering upslope. The pool of calcium P on ultrabasic rock did not vary consistently with altitude (1.2–2.8 g m^–2), probably reflecting the greater between-site variability of primary mineral P in parent rocks. When all sites were compared, the pool of most labile, bicarbonate-extracted inorganic P increased (ranging from 0.02 to 1.85 g m^–2) with increasing calcium P. Calcium P was therefore considered to be an important P source to the biota on Kinabalu. Gross patterns in the variation of PUE (indexed as the reciprocal of the P concentration in litter) were best explained by the pool size of actively cycling P (total P minus occluded inorganic P). PUE, however, demonstrated distinct altitudinal patterns to generate an intricate conrol of P use pattern by soil P pools and altitude. Received: 2 August 1998 / Accepted: 28 November 1999     

Dynamics,productivity and species richness of tropical rainforests along elevational and edaphic gradients on Mount Kinabalu,Borneo

We studied the dynamics of nine tropical rainforests on Mount Kinabalu, Borneo, at four elevations (700, 1,700, 2,700 and 3,100 m) on various edaphic conditions for four 2-year periods over 8 years (1995–2003), and examined the relationships with above-ground productivity. Mean growth rate of stem diameter, basal area turnover rate and estimated recruitment rate (using growth rate and size distribution) correlated with productivity among the nine forests in all periods. These rates based on growth rates of surviving stems appeared to be good measures of stand turnover. However, observed recruitment rate and mortality (and turnover rate as mean of these rates) based on direct observation of recruits and deaths did not correlate with productivity in some periods. These rates may not be useful as measures of stand turnover given small sample size and short census interval because they were highly influenced by stochastic fluctuation. A severe drought associated with the 1997–1998 El Niño event inflated mortality and depressed mean growth rate, recruitment rate and basal area turnover rate, but had little effect on the correlations between these rates (except mortality) and productivity. Across broad elevational and edaphic gradients on Mount Kinabalu, forest turnover, productivity and species richness correlated with each other, but the causal interpretation is difficult given the different histories and species pools among forests at different elevations.     

Genetic variation of Trigonobalanus verticillata,a primitive species of Fagaceae,in Malaysia revealed by chloroplast sequences and AFLP markers

The genetic variation of Trigonobalanus verticillata, the most recently described genus of Fagaceae, was studied using chloroplast DNA sequences and AFLP fingerprinting. This species has a restricted distribution that is known to include seven localities in tropical lower montane forests in Malaysia and Indonesia. A total of 75 individuals were collected from Bario, Kinabalu, and Fraser's Hill in Malaysia. The sequences of rbcL, matK, and three non-coding regions (atpB-rbcL spacer, trnL intron, and trnL-trnF spacer) were determined for 19 individuals from these populations. We found a total of 30 nucleotide substitutions and four length variations, which allowed identification of three haplotypes characterizing each population. No substitutions were detected within populations, while the tandem repeats in the trnL -trnF spacer had a variable repeat number of a 20-bp motif only in Kinabalu. The differentiation of the populations inferred from the cpDNA molecular clock calibrated with paleontological data was estimated to be 8.3 MYA between Bario and Kinabalu, and 16.7 MYA between Fraser's Hill and the other populations. In AFLP analysis, four selective primer pairs yielded a total of 431 loci, of which 340 (78.9%) were polymorphic. The results showed relatively high gene diversity (H(S) = 0.153 and H(T) = 0.198) and nucleotide diversity (pi(S) = 0.0132 and pi(T) = 0.0168) both within and among the populations. Although the cpDNA data suggest that little or no gene flow occurred between the populations via seeds, the fixation index estimated from AFLP data (F(ST) = 0.153 and N(ST) = 0.214) implies that some gene flow occurs between populations, possibly through pollen transfer.     

Evolutionary dynamics of the leaf phenological cycle in an oak metapopulation along an elevation gradient

It has been predicted that environmental changes will radically alter the selective pressures on phenological traits. Long‐lived species, such as trees, will be particularly affected, as they may need to undergo major adaptive change over only one or a few generations. The traits describing the annual life cycle of trees are generally highly evolvable, but nothing is known about the strength of their genetic correlations. Tight correlations can impose strong evolutionary constraints, potentially hampering the adaptation of multivariate phenological phenotypes. In this study, we investigated the evolutionary, genetic and environmental components of the timing of leaf unfolding and senescence within an oak metapopulation along an elevation gradient. Population divergence, estimated from in situ and common‐garden data, was compared to expectations under neutral evolution, based on microsatellite markers. This approach made it possible (1) to evaluate the influence of genetic correlation on multivariate local adaptation to elevation and (2) to identify traits probably exposed to past selective pressures due to the colder climate at high elevation. The genetic correlation was positive but very weak, indicating that genetic constraints did not shape the local adaptation pattern for leaf phenology. Both spring and fall (leaf unfolding and senescence, respectively) phenology timings were involved in local adaptation, but leaf unfolding was probably the trait most exposed to climate change‐induced selection. Our data indicated that genetic variation makes a much smaller contribution to adaptation than the considerable plastic variation displayed by a tree during its lifetime. The evolutionary potential of leaf phenology is, therefore, probably not the most critical aspect for short‐term population survival in a changing climate.     

942. CYMBIDIUM DAYANUM VAR. ANGUSTIFOLIUM

The widespread and variable Cymbidium dayanum Rchb. f. is discussed. A variety from Mount Kinabalu, var. angustifolium is illustrated, and the new combination is made, based on C. angustifolium Ames & C. Schweinf.     

Asymmetric boundary shifts of tropical montane Lepidoptera over four decades of climate warming

Aim To estimate whether species have shifted at equal rates at their leading edges (cool boundaries) and trailing edges (warm boundaries) in response to climate change. We provide the first such evidence for tropical insects, here examining elevation shifts for the upper and lower boundaries shifts of montane moths. Threats to species on tropical mountains are considered. Location Mount Kinabalu, Sabah, Malaysia. Methods We surveyed Lepidoptera (Geometridae) on Mount Kinabalu in 2007, 42 years after the previous surveys in 1965. Changes in species upper and lower boundaries, elevational extents and range areas were assessed. We randomly subsampled the data to ensure comparable datasets between years. Estimated shifts were compared for endemic versus more widespread species, and for species that reached their range limits at different elevations. Results Species that reached their upper limits at 2500–2700 m (n= 28 species, 20% of those considered) retreated at both their lower and upper boundaries, and hence showed substantial average range contractions (?300 m in elevational extent and ?45 km² in estimated range area). These declines may be associated with changes in cloud cover and the presence of ecological barriers (geological and vegetation transitions) which impede uphill movement. Other than this group, most species (n= 109, 80% of the species considered) expanded their upper boundaries upwards (by an average of 152 m) more than they retreated at their lower boundaries (77 m). Main conclusions Without constraints, leading margins shifted uphill faster than trailing margins retreated, such that many species increased their elevational extents. However, this did not result in increases in range area because the area of land available declines with increasing elevation. Species close to a major ecological/geological transition zone on the mountain flank declined in their range areas. Extinction risk may increase long before species reach the summit, even when undisturbed habitats are available.     

Altitudinal Variation in Leaf Nitrogen Concentration on the Eastern Slope of Mount Gongga on the Tibetan Plateau,China

Mount Gongga spans 6500 m in elevation and has intact and continuous vertical vegetation belts, ranging from subtropical evergreen broad-leaved vegetation to an alpine frigid sparse grass and desert zone. Investigating the altitudinal trends in leaf nitrogen (N) on Mount Gongga can increase our understanding of the global biogeography of foliar N. In this study, 460 leaf samples from mosses, ferns, and seed plants were collected along an altitudinal gradient on the eastern slope of Mount Gongga, and the variation in leaf N concentration (mass basis) with elevation was analyzed. There are considerable differences in leaf N between mosses and ferns, mosses and seed plants, C₄ and C₃ plants, and evergreen and deciduous woody plants. The general altitudial pattern of leaf N in Mount Gongga plants was that leaf N kept increasing until an elevation of about 2200 m above sea level, with a corresponding mean annual temperature (MAT) of 8.5°C, and then decreased with increasing elevation. However, the evergreen woody plants displayed a decline trend in leaf N across the altitude gradient. Our findings provide an insight into the altitudinal variation in leaf N.     

An experimental investigation of enhanced harpacticoid (Copepoda) abundances around isolated seagrass shoots

Summary Premature leaf abscission in Populus was observed in central Washington (USA) about two weeks after the 18 May 1980 ashfall from Mount St. Helens. Leaf abscission was probably a wounding response caused by sandsized ash carried in winds that gusted to 15 m s^-1 7 days after the ashfall. Under glasshouse conditions simulating this environmental regime the leaf epidermis and often part of the mesophyll in Populus nigra L. var. italica Duroi. were abraded, resulting in permanent loss of leaf turgor and decreased stomatal conductance. Substantial abscission of still-green leaves ensued. Leaf abrasion by ash-laden winds, although rare, may result in conspicuous plant damage.     

Soil Phosphorus Fractionation and Phosphorus-Use Efficiency of a Bornean Tropical Montane Rain Forest During Soil Aging With Podozolization

We compared phosphorus (P) dynamics and plant productivity in two montane tropical rain forests (Mount Kinabalu, Borneo) that derived from similar parent materials (largely sedimentary rocks) and had similar climates but differed in terms of soil age. The younger site originated from Quaternary colluvial deposits, whereas the older site had Tertiary-age material. The older site had a distinctive spodic horizon, reduced levels of labile inorganic soil P, higher concentrations of recalcitrant organic soil P, and lower rates of net soil N mineralization. P fertilization led to soil nitrogen (N) immobilization in the P-deficient soil, indicating that soil N mineralization was limited by P at the P-deficient older site. Mean foliar nutrient concentration (on both a weight and an area basis) was similar at the two sites for all elements except P, which was lower at the older site. Aboveground net primary production (ANPP) was lower at the older site than at the younger one; this difference could be explained by the reduced availability of P and N (as down-regulated by P) at the older site. The relatively ample allocation of P and N to leaves, despite the reduced availability at the P-deficient old site, was attributable to its high resorption efficiency. High resorption resulted in lower concentrations of elements in leaf litter—that is, less decomposable low-quality litter. On the other hand, the concentration of leaf litter lignin was considerably lower at the older site; this appeared to be a de facto adaptive mechanism to avoid retarding litter decomposition.