David McClintock Newbery

EVIDENCE OF SPECIES-SPECIFIC NEIGHBORHOOD EFFECTS IN THE DIPTEROCARPACEAE OF A BORNEAN RAIN FOREST

Although accumulating evidence indicates that local intraspecific density-dependent effects are not as rare in species-rich communities as previously suspected, there are still very few detailed and systematic neighborhood analyses of species-rich communities. Here, we provide such an analysis with the overall goal of quantifying the relative importance of inter- and intraspecific interaction strength in a primary, lowland dipterocarp forest located at Danum, Sabah, Malaysia. Using data on 10 abundant overstory dipterocarp species from two 4-ha permanent plots, we evaluated the effects of neighbors on the absolute growth rate of focal trees (from 1986 to 1996) over increasing neighborhood radii (from 1 to 20 m) with multiple regressions. Only trees 10 cm to > 100 cm girth at breast height in 1986 were considered as focal trees. Among neighborhood models with one neighbor term, models including only conspecific larger trees performed best in five out of 10 species. Negative effects of conspecific larger neighbors were most apparent in large overstory species such as those of the genus Shorea. However, neighborhood models with separate terms and radii for heterospecific and conspecific neighbors accounted for more variability in absolute growth rates than did neighborhood models with one neighbor term. The conspecific term was significant for nine out of 10 species. Moreover, in five out of 10 species, trees without conspecific neighbors had significantly higher absolute growth rates than trees with conspecific neighbors. Averaged over the 10 species, trees without conspecific neighbors grew 32.4 cm(2) in basal area from 1986 to 1996, whereas trees with conspecific neighbors only grew 14.7 cm(2) in basal area, although there was no difference in initial basal area between trees in the two groups. Averaged across the six species of the genus Shorea, negative effects of conspecific larger trees were significantly stronger than for heterospecific larger neighbors.

Recruitment limitation constrains local species richness and productivity in dry grassland.

Species coexistence and local-scale species richness are limited by the availability of seeds and microsites for germination and establishment. We conducted a seed addition experiment in seminatural grassland at three sites in southern Switzerland and repeated the experiment in two successive years to evaluate various circumstances under which seed limitation and establishment success affect community functioning. A collection of 144,000 seeds of 22 meadow species including grasses and forbs of local provenance was gathered, and seeds were individually sown in a density that resembled natural seed rain. The three communities were seed limited. Three years after sowing, single species varied in emergence (0-50%), survival (0-69%), and establishment rates (0-27%). One annual and 13 perennial species reached reproductive stage. Low establishment at one site and reduced growth at another site indicated stronger microsite limitation compared to the third site. Recruitment was influenced by differences in abiotic environmental conditions between sites (water availability, soil minerals) and by within-site differences in biotic interaction (competition). At the least water-limited site, sowing resulted in an increase in phytomass due to establishment of short-lived perennials in the second and third years after sowing. This increase persisted over the following two years due to establishment of longer-lived perennials. After sowing in a wetter year with higher phytomass, however, productivity did not increase, because higher intensity of competition in an early phase of establishment resulted in less vigorous plants later on. Due to the generally favorable weather conditions during this study, sowing year had a small effect on numbers of established individuals over all species. Recruitment limitation can thus constrain local-scale species richness and productivity, either by a lack of seeds or by reduced seedling growth, likely due to competition from the established vegetation.

Structure and inferred dynamics of a large grove of Microberlinia bisulcata trees in central African rain forest: the possible role of periods of multiple disturbance events

A 272-ha grove of dominant Microberlinia bisulcata (Caesalpinioideae) adult trees greater than or equal to 50 cm stem diameter was mapped in its entirety in the southern part of Korup National Park, Cameroon. The approach used an earlier-established 82.5-ha permanent plot with a new surrounding 50-m grid of transect lines. Tree diameters were available from the plot but trees on the grid were recorded as being greater than or equal to 50 cm. The grove consisted of 1028 trees in 2000. Other species occurred within the grove. including the associated subdominants Tetraberlinia bifoliolata and T. korupensis. Microberlinia bisulcata becomes adult at a stein diameter of c. 50 cm and at an estimated age of 50 y. Three oval-shaped subgroves with dimensions c. 8 50 in x 13 50 in (90 ha) were defined. For two of them (within the plot) tree diameters were available. Subgroves differed in their scales and intensities of spatial tree patterns, and in their size frequency distributions, these suggesting differing past dynamics. The modal scale of clumping was 40-50 m. Seed dispersal by pod ejection (to c. 50 in) was evident from the semi-circles of trees at the groves edge and from the many internal circles (100-200 m diameter). The grove has the capacity. therefore, to increase at c. 100 m per century. To form its present extent and structure. it is inferred that it expanded and infilled from a possibly smaller area of lower adult-tree density. This possibly happened in three waves of recruitment, each one determined by a period of several intense disturbances. Climate records for Africa show that 1740-50 and 1820-30 were periods of drought, and that 1870-1895 was also regionally very dry. Canopy openings allow the light-demanding and fast-growing ectomycorrhizal M. bisulcata to establish, but successive releases are thought to be required to achieve effective recruitment. Nevertheless, in the last 50 y there were no major events and recruitment in the grove was very poor. This present study leads to a new hypothesis of the role of periods of multiple extreme events being the driving factor for the population dynamics of many large African tree species such as M. bisulcata.

A structural analysis of rain forest at Korup and Douala–Edea, Cameroon

Diameter frequency distributions (dfds) of 197 tree species in Korup National Park and 118 tree species in Douala–Edea Forest Reserve (based on c. 41 000 and 25 000 trees ≥ 30 cm gbh in 135 and 104 plots along four transects at each site, respectively) were individually classified into four groups of species according to their maximum diameters (≥99% trees 1% ≥90cm dbh), and separated into two abundance classes (n≥50 and n = 20–49 (or 15–49). A fifth group, from those species in 3 and 4, was defined by a relative lack of small stems and given detailed attention. Dfds were evaluated for all species together on a transect and site basis, and compared for individual species across transects, and between sites for those group 5 species in common. Analysis focussed on three large ectomycorrhizal species in groups 3 and 4, Microberlinia bisulcata, Tetraberlinia bifoliolata and T. moreliana (Caesalpiniaceae), which were restricted to, and dominated, the forest on the nutrient-poor soils of transect P at Korup. M. bisulcata , the most abundant, was markedly group 5 in character, with very few small trees, whilst the Tetraberlinia species had strong replacement potential. Although there were 27 species (42% of groups 3 & 4) at Korup and 19 (48%) at Douala–Edea in group 5, M. bisulcata was unique in its combination of large size, dfd, mycorrhizal habit, spatial distribution and soil association. The potential replacement of this species was contrasted also with a non-ectomycorrhizal caesalp with similar dfd that occurred on nutrient-richer soils, Erythrophleum ivorense . Knowledge of the Caesalpiniaceae-rich la foret biafrieene of Letouzey (which includes Korup and Douala–Edea) is reviewed, and the distribution, ecology and regeneration of the caesalps discussed. Structure and inferred dynamics of the forest at Korup are discussed with special regard to the ectomycorrhizal species.

Dispersal and microsite limitation in an abandoned calcareous grassland of the southern prealps

Dispersal limitation is often involved when the species composition of a dry abandoned grassland shows a slow response to resumed regular mowing. A seed-addition experiment, using 32 species which do not belong to the local species pool, was performed on Monte San Giorgio (southern Switzerland) to test whether the low recruitment success was due to dispersal limitation or due to unfavourable microsite conditions. In October 1997, 20 species were individually sown in six 3 × 4 m blocks of a 2 × 2 factorial “partial” split-plot design with treatments of abandonment vs. mowing and undisturbed vs. root-removed soil, this last being applied in small naturally-degradable pots. Moreover, 12 species were sown only in the treatments on undisturbed soil. Seedlings of sown and spontaneously germinating seeds were observed on 16 occasions over one 12-month period.Seeds of 31 out of the 32 species germinated. Twenty-four species showed germination rates higher than 5% and different seasonal germination patterns. Established vegetation, especially the tussocks ofMolinia arundinacea, reduced the quality of microsites for germination. Whereas a few species germinated better under the litter ofMolinia arundinacea, many more germinated better under the more variable microsite conditions of a mown grassland.Only a few seedlings of 25 species out of the 31 germinated species survived until October 1998. Seedling survival was negatively affected by litter, unfavourable weather conditions (frost and dry periods followed by heavy rains) and herbivory (slugs and grasshoppers). Tussocks ofMolinia arundinacea, however, tended to protect seedlings.The poor establishment success of “new” species observed in abandoned meadows on Monte San Giorgio after resumed mowing is due to dispersal and microsite limitations.

Reproductive investment and seedling survival of the mast-fruiting rain forest tree, Microberlinia bisulcata A. chev

In the southern part of Korup National Park, Cameroon, the mast fruitingtree Microberlinia bisulcata occurs as a codominant ingroves of ectomycorrhizal Caesalpiniaceae within a mosaic of otherwisespecies-rich lowland rain forest. To estimate the amount of carbon andnutrientsinvested in reproduction during a mast fruiting event, and the consequentialseed and seedling survival, three related field studies were made in 1995.Theseprovided a complete seed and seedling budget for the cohort. Seed productionwasestimated by counting woody pods on the forest floor. Trees produced on average26,000 (range 0–92,000) seeds/tree, with a dry mass of 16.6kg/tree. Seeds were contained in woody pods of mass 307kg/tree. Dry mass production of pods and seeds was 1034 kgha−1, equivalent to over half (55%) of annual leaflitterfall for this species, and contained 13% of the nitrogen and 21% of thephosphorus in annual leaf litterfall. Seed and young-seedling mortality wasinvestigated with open quadratsand cages to exclude vertebrate predators, at two distances from the parenttree. The proportion of seeds on the forest floor which disappeared in thefirst6 wk after dispersal was 84%, of which 26.5% was due to likelyvertebrate removal, 36% to rotting, and 21.5% to other causes. Vertebratepredation was greater close to the stem than 5 m beyond the crown(41 vs 12% of seeds disappearing) where the seed shadow was less dense.Previousstudies have demonstrated an association between mast years at Korup and highdry-season radiation before flowering, and have shown lower leaf-litterfallphosphorus concentrations following mast fruiting. The emerging hypothesis isthat mast fruiting is primarily imposed by energy limitation for fruitproduction, but phosphorus supply and vertebrate predation are regulatingfactors. Recording the survival of naturally-regeneratingM. bisulcata seedlings (6-wk stage) showed that 21% ofseedlings survived to 31 mo. A simple three-stage recruitment model wasconstructed. Mortality rates were initially high and peaked again in each ofthenext two dry seasons, with smaller peaks in the two intervening wet seasons,these latter coinciding with annual troughs in radiation. The very poorrecruitment of M. bisulcata trees in Korup, demonstratedinprevious investigations, appears not to be due to a limitation in seed oryoung-seedling supply, but rather by factors operating at theestablished-seedling stage.

Litter breakdown and mineralization in a central African rain forest dominated by ectomycorrhizal trees

Based on litter mass and litterfall data, decomposition rates for leaveswere found to be fast (k = 3.3) and the turnover times short (3.6mo) on the low-nutrient sandy soils of Korup. Leaf litteroffour ectomycorrhizal tree species (Berlinia bracteosa, Didelotiaafricana, Microberlinia bisulcata and Tetraberliniabifoliolata) and of three non-ectomycorrhizal species(Cola verticillata, Oubanguia alata andStrephonema pseudocola) from Korup were left to decomposein 2-mm mesh bags on the forest floor in three plots ofeachof two forest types forest of low (LEM) and high (HEM) abundance ofectomycorrhizal (caesalp) trees. The litter of the ectomycorrhizal speciesdecayed at a significantly slower rate than that of thenon-ectomycorrhizal species, although the former were richer in P and Nconcentrations of the start. Disappearance rates of the litter layer showed asimilar trend. Ectomycorrhizal species immobilized less N, but mineralized moreP, than non-ectomycorrhizal species. Differences between species groupsin K, Mg and Ca mineralization were negligible. Effect of forest type was clearonly for Mg: mineralization of Mg was faster in the HEM than LEM plots, apattern repeated across all species. This difference was attributed to a muchmore prolific fine root mat in the HEM than LEM forest. The relatively fastrelease of P from the litter of the ectomycorrhizal species suggests that thematmust allow an efficient uptake to maintain P in the forest ecosystem.

Buttress form of the central African rain forest tree Microberlinia bisulcata, and its possible role in nutrient acquisition

Buttressing is a trait special to tropical trees but explanations for its occurrence remain inconclusive. The two main hypotheses are that they provide structural support and/or promote nutrient acquisition. Studies of the first are common but the second has received much less attention. Architectural measurements were made on adult and juvenile trees of the ectomycorrhizal species Microberlinia bisulcata, in Korup (Cameroon). Buttressing on this species is highly distinctive with strong lateral extension of surface roots of the juveniles leading to a mature buttress system of a shallow spreading form on adults. This contrasts with more vertical buttresses, closer to the stem, found on many other tropical tree species. No clear relationship between main buttress and large branch distribution was found. Whilst this does not argue against the essential structural role of buttresses for these very large tropical trees, the form on M. bisulcata does suggest a likely second role, that of aiding nutrient acquisition. At the Korup site, with its deep sandy soils of very low phosphorus status, and where most nutrient cycling takes place in a thin surface layer of fine roots and mycorrhizas, it appears that buttress form could develop from soil-surface root exploration for nutrients by juvenile trees. It may accordingly allow M. bisulcata to attain the higher greater competitive ability, faster growth rate, and maximum tree size that it does compared with other co-occurring tree species. For sites across the tropics in general, the degree of shallowness and spatial extension of buttresses of the dominant species is hypothesized to increase with decreasing nutrient availability.

The role of tree size in the leafing phenology of a seasonally dry tropical forest in Belize, Central America

Leafing phenology of two dry-forest sites on soils of different depth (S = shallow, D = deep) at Shipstern Reserve, Belize, were compared at the start of the rainy season (April-June 2000). Trees 2.5 cm dbh were recorded weekly for 8 wk in three 0.04-ha plots per site. Ten species were analysed individually for their phenological patterns, of which the three most common were Bursera simaruba, Metopium brownei and Jatropha gaumeri. Trees were divided into those in the canopy (> 10 cm dbh) and the subcanopy ( 10 cm dbh). Site S had larger trees on average than site D. The proportion of trees flushing leaves at any one time was generally higher in site S than in site D, for both canopy and subcanopy trees. Leaf flush started 2 wk earlier in site S than site Dfor subcanopy trees, but only 0.5 wk earlier for the canopy trees. Leaf flush duration was 1.5 wk longer in site S than site D. Large trees in the subcanopy flushed leaves earlier than small ones at both sites but in the canopy just at site D. Large trees flushed leaves earlier than small ones in three species and small trees flushed leaves more rapidly in two species. Bursera and Jatropha followed the general trends but Metopium, with larger trees in site Dthan site S, showed the converse with onset of flushing 1 wk earlier in site Dthan site S. D ifferences in response of the canopy and subcanopy trees on each site can be accounted for by the predominance of spring-flushing or stem-succulent species in site S and a tendency for evergreen species to occur in site D. Early flushing of relatively larger trees in site D most likely requires access to deeper soil water reserves but small and large trees utilize stored tree water in site S.

Relaxation of species‐specific neighborhood effects in Bornean rain forest under climatic perturbation

Evidence of negative conspecific density dependence (NDD) operating on seedling survival and sapling recruitment has accumulated recently. In contrast, evidence of NDD operating on growth of trees has been circumstantial at best. Whether or not local NDD at the level of individual trees leads to NDD at the level of the community is still an open question. Moreover, whether and how perturbations interfere with these processes have rarely been investigated. We applied neighborhood models to permanent plot data from a Bornean dipterocarp forest censused over two 10-11 year periods. Although the first period was only lightly perturbed, a moderately strong El Niño event causing severe drought occurred in the first half of the second period. Such events are an important component of the environmental stochasticity affecting the region. We show that local NDD on growth of small-to-medium-sized trees may indeed translate to NDD at the level of the community. This interpretation is based on increasingly negative effects of bigger conspecific neighbors on absolute growth rates of individual trees with increasing basal area across the 18 most abundant overstory species in the first period. However, this relationship was much weaker in the second period. We interpreted this relaxation of local and community-level NDD as a consequence of increased light levels at the forest floor due to temporary leaf and twig loss of large trees in response to the drought event. Mitigation of NDD under climatic perturbation acts to decrease species richness, especially in forest overstory and therefore has an important role in determining species relative abundances at the site.