El Niño droughts and their effects on tree species composition and diversity in tropical rain forests

In this study I investigated the effects of the extreme, 1997/98 El Niño related drought on tree mortality and understorey light conditions of logged and unlogged tropical rain forest in the Indonesian province of East Kalimantan (Borneo). My objectives were to test (1) whether drought had a significant effect on tree mortality and understorey light conditions, (2) whether this effect was greater in logged than in undisturbed forest, (3) if the expected change in tree mortality and light conditions had an effect on Macaranga pioneer seedling and sapling densities, and (4) which (a)biotic factors influenced tree mortality during the drought. The 1997/1998 drought led to an additional tree mortality of 11.2, 18.1, and 22.7% in undisturbed, old logged and recently logged forest, respectively. Mortality was highest in logged forests, due to extremely high mortality of pioneer Macaranga trees (65.4%). Canopy openness was significantly higher during the drought than during the non-drought year (6.0, 8.6 and 10.4 vs 3.7, 3.8 and 3.7 in undisturbed, old logged and recently logged forest, respectively) and was positively correlated with the number of dead standing trees. The increase in light in the understorey was accompanied by a 30 to 300-fold increase in pioneer Macaranga seedling densities. Factors affecting tree mortality during drought were (1) tree species successional status, (2) tree size, and (3) tree location with respect to soil moisture. Tree density and basal area per surface unit had no influence on tree mortality during drought. The results of this study show that extreme droughts, such as those associated with El Niño events, can affect the tree species composition and diversity of tropical forests in two ways: (1) by disproportionate mortality of certain tree species groups and tree size classes, and (2) by changing the light environment in the forest understorey, thereby affecting the recruitment and growth conditions of small and immature trees.     

Colonization of subtidal macroalgae in a fucalean-dominated algal assemblage,southwestern Australia

The influence of fucalean canopy species and dominant understory macroalgae on algal colonization was investigated to evaluate whether layering contributes to patterns in algal diversity. Patterns in recruitment were compared among total-clearing, understory-removal, canopy-removal, and undisturbed plots (plot area = 0.25 m²), using a randomized block design in depths <10 m and 10–20 m at Woody Island, Western Australia. To evaluate if propagules were available in the water column above the canopy layer, settlement plates (plate area = 0.04 m²) were deployed in depths <10 m, 10–20 m, and >20 m. A total of 198 macroalgal species was recorded. Biomass of the understory species Osmundaria prolifera Lamouroux and Botryocladia sonderi Silva was similar between canopy-removal and undisturbed plots. Diversity of macroalgae was similar in the presence and absence of a canopy layer. Taxa found in the canopy showed different patterns in recruitment. Cystoseiraceae recruited predominantly in total-clearings in both depth strata. Sargassaceae recruited most abundantly in depths <10 m. Density of canopy taxa on settlement plates was similar with depth (20–30 juveniles per plate), and juveniles were mainly Cystoseiraceae. In contrast to kelp beds or forests, patterns in algal colonization appeared to be maintained by environmental factors or processes other than the direct effects of layering in the subtidal fucoid-dominated assemblages at Woody Island. Handling editor: K. Martens     

Short-term changes in tree populations in a tropical evergreen forest at Varagalaiar, Western Ghats, India

Within a 30-ha permanent plot of a tropical evergreen forest in the Anamalais, Indian Western Ghats, all the 13415 trees 30cmgbh (belonging to 153 species) were identified and tagged during 1997–1998. This communication reports the results of tree population changes based on two annual censuses (1999 and 2000) of the plot after the initial census, and compares the results with other tropical forest sites. One species, Ficus beddomei, disappeared from the plot and there was no new addition of species. Of the 96 species that showed changes in their population density, for 12 species recruitment and mortality matched. Tree recruitment (5 treesha^–1year^–1) exceeded mortality (4 treesha^–1year^–1) during the two-year period. Four modes of tree deaths were recorded in our site. The per cent mortality of trees differed between tree size classes during each recensus. The plot tree density increased progressively in the two-year period, indicating that shifts in species composition and density in natural forests, without major catastrophic disturbance, occur slowly.     

Quantifying and testing coexistence mechanisms arising from recruitment fluctuations

Temporal fluctuations in recruitment are involved in two distinct coexistence mechanisms, the storage effect and relative nonlinearity of competition, which may act simultaneously to stabilize species coexistence. It is shown that comparisons of recruitment variation between species at high versus low densities can test whether these mechanisms are responsible for stable coexistence. Moreover, under certain circumstances, these comparisons can measure the total coexistence stabilizing effect of the mechanism. These comparisons are clearest for the situation of an invader (a species perturbed to low density) in the presence of its competitors, termed residents. Then average invader-resident differences in the variances of log recruitment, potentially weighted by adult survival rates and species' sensitivities to competition, are proportional to the overall stabilizing effect of the storage effect and relative nonlinearity of competition. Less effective comparisons are available for species naturally at high and low densities or with substantial mean differences in average fitness. These developments lead also to a technique of partitioning the long-term low-density growth rate of a species into community average measures of stabilizing mechanisms, deviations from these measures, and other factors. The community average measure is argued as most appropriate for understanding the ability of a coexistence mechanism to stabilize coexistence. Individual species' deviations from the community average indicate the ways in a which a coexistence mechanism may affect average fitness differences between species either enhancing or diminishing the ability of a given set of species to coexist, depending on other factors. This approach provides a general new tool for analyzing species coexistence.     

Habitat structure,conspecific presence and spatial variation in the recruitment of a temperate reef fish

Pronounced spatial variation in recruitment occurs in many marine invertebrate and fish populations and is thought to be critical to the demography of these species. In this study I examined the importance of habitat structure and the presence of conspecific residents to spatial variation in larval settlement and recruitment in a temperate fish Tautogolabrus adspersus. I define settlement as the movement of individuals from the water column to the benthic habitat, while I refer to recruitment as numbers of individuals surviving some arbitrary period of time after settlement. Experiments in which standard habitats were stocked with conspecifics showed that resident conspecifics were not an important factor contributing to small-scale variability in recruitment. Further correlative analyses demonstrated that large-scale variation in recruitment could not be explained by variability in older age classes. By contrast, manipulations of macroalgal structure within a kelp bed demonstrated that recruitment was significantly higher in habitats with a dense understory of foliose and filamentous algae than in habitats with only crustose algae. Understory algae varied in their pattern of disperison among sites, and the dispersion of fish matched that of the plants. In order to determine the effects of differences in patterns of algal dispersion on the demography of associated T. adspersus populations, I used experimental habitat units to manipulate patterns of dispersion. Settlement was significantly greater to randomly placed versus clumped habitats; however, no differences in recruitment between random and clumped habitats were detected. Because recruitment is a function of the numbers of settlers minus the subsequent loss of settlers, rates of mortality or migration must have been higher in the randomly placed habitats. These results are counter to the current paradigm for reef fishes which suggests that larval settlement is the crucial demographic process producing variability in population abundance. In this experiment patterns of settlement were modified by varying the patch structure of the habitat.Contribution number 278 from the Center for Marine Biology, University of New Hampshire     

Nuthatches (Sitta europaea: Aves) in forest fragments: demography of a patchy population

Breeding density, local survival and summer recruitment of nuthatches were evaluated in a population scattered over many small (1–30 ha) forest fragments, and compared with study plots inside larger forests. Since most young birds settle outside the fragment in which they were born this population corresponds to the “patchy population” concept implying that patterns in abundance may be better explained by processes at the population level than by metapopulation processes. Mean breeding density was c. 50% lower in fragments and decreased with regional isolation (distance from larger forests) but not with local isolation (distance to nearby fragments). Local survival of adults and established (i.e. territorial) 1st-year birds was not related to forest size or isolation. However, fewer young birds settled in summer in the fragments compared with a large forest. This difference probably reflects high mortality during the sensitive dispersal phase. Moreover, the observed number of recruits and their estimated survival was insufficient to maintain the breeding population, suggesting significant net immigration from larger forests. This “rescue effect” explains why densities are affected by regional, but not local isolation. Received: 14 December 1998 / Accepted: 1 March 1999     

Competing associations in bacterial warfare with two toxins

Simple combinations of common competitive mechanisms can easily result in cyclic competitive dominance relationships between species. The topological features of such competitive networks allow for complex spatial coexistence patterns. We investigate self-organization and coexistence in a lattice model, describing the spatial population dynamics of competing bacterial strains. With increasing diffusion rate the community of the nine possible toxicity/resistance types undergoes two phase transitions. Below a critical level of diffusion, the system exhibits expanding domains of three different defensive alliances, each consisting of three cyclically dominant species. Due to the neutral relationship between these alliances and the finite system size effect, ultimately only one of them remains. At large diffusion rates the system admits three coexisting domains, each containing mutually neutral species. Because of the cyclical dominance between these domains, a long term stable coexistence of all species is ensured. In the third phase at intermediate diffusion the spatial structure becomes even more complicated with domains of mutually neutral species persisting along the borders of defensive alliances. The study reveals that cyclic competitive relationships may produce a large variety of complex coexistence patterns, exhibiting common features of natural ecosystems, like hierarchical organization, phase transitions and sudden, large-scale fluctuations.     

Mortality of pregnant females in Arikara villages: osteological evidence

High infant mortality and high mortality for late adolescent and young adult females suggest that obstetrical hazards may be one explanation for differences in male and female mortality curves. This possibility is investigated in Arikara skeletal series by determining the frequency of females who died with fetal remains in utero. Two females (0.9%) were so identified. Examination of the females and the fetal remains do not provide evidence that stress of childbearing was the cause of death in these cases.     

The natural distribution of survival

Great similarities in survival patterns permit the Gompertz and other established equations to describe parts of mortality curves in various species. These patterns appear non-random and invite inference of biological meaning, though no unifying explanation is agreed.Under the theory described here, linear decline of an initial quantity of species or strain-specific redundant reserve interacts with extrinsic hazards via a ‘nested binomial’ model, which is presented both in a simple, four parameter form, and a more complex form that incorporates inter-individual and inter-functional biological variation.This approach demonstrates exponential rises in mortality, late-life deceleration and Strehler-Mildvan correlation. Biological variation within the complex model, specifically in the redundancy decay rate parameter, is shown to generate mortality plateaux, while outlier phenotypes produce mortality decelerations, supporting inter-individual heterogeneity as the cause of these phenomena. The model is robust to large variations in organism complexity, and to broad intra-population hazard variation. Specific parameters appear analogous to observed elements of ageing, and a central role for redundancy depletion provides a context for longevity genes and rapid evolution of increased lifespan.This approach offers a unifying model for a great variety of ageing phenomena across a wide range of species.     

Temporal and spatial patterns in emergence and early survival of perennial plants in the Sonoran Desert

Seedling emergence and survival of 15 perennial species were studied for six years in a 557-m² permanent plot at Tumamoc Hill, Arizona, USA, an ungrazed site in the northern Sonoran Desert. The minimum rain required for germination and emergence ranged from 17.5 to 35.6 mm. Few species emerged in every year of the study. First-year survival averaged across all 15 species was 3.7%; only 0.1% of seedlings lived as long as four years. The odds of survival in the first year improved with increased rain. About three times as many seedlings died from predation as desiccation. In 2-m² subplots, mortality of three woody species in the first 30 days after emergence appeared to be independent of seedling density. Short-, moderate-, and long-lived species displayed distinct survival strategies. Long-lived species compensated for generally poor seedling survival by frequent germination and emergence. Moderate-lived species exhibited highly episodic germination and emergence, a potentially risky behavior that might have been offset to some extent by relatively good long-term survival. Short-lived species had the highest seedling survival. Because these species can bloom in their first year, good early survival meant that some individuals were able to reproduce before they died.     

Behavior and physiology of coexistence in two species of Dascyllus (Pomacentridae,Teleostei)

Synopsis In the Gulf of Aqaba Dascyllus aruanus and D. marginatus, form distinct populations organised into harems of single males and groups of females, living permanently in Stylophora and Acropora corals. Each harem consists of 4 sexual casts: dominating males, adult females, sex-changing fish and juveniles. The sexual performance of each adult fish in such social units is controlled by the presence or absence of a dominating male. In corals occupied by mixed groups of both species, only those females spawn that belong to the same species as the dominating male. Direct observation in the sea shows that such situations are produced by indiscriminative settling mechanisms of groups of juvenile fish. Evidence is also presented, demonstrating that dispersion of coral colonies suitable for occupancy and predation by piscivorous fish can modulate such complicated reproduction.     

Demographic analyses of the dusky shark, <Emphasis Type="Italic">Carcharhinus obscurus,</Emphasis> in the Northwest Atlantic incorporating hooking mortality estimates and revised reproductive parameters

Demographic analyses were conducted for the dusky shark, Carcharhinus obscurus, in the Northwest Atlantic. Fishery-independent, fishery-dependent, and previously published data were used to estimate age specific by-catch mortality and reproductive cycle for incorporation into the analyses. Correspondence analyses were conducted for viability (dead or alive), shark length, month, soaktime, and region. Dead sharks were associated with small size, long soak times, and the spring season, while live sharks were associated with large size and shorter soak times. Data on reproduction suggested a 3-year reproductive cycle consisting of a 2-year gestation period and a 1-year resting period. Litter sizes ranged from 3–12 embryos. Decreasing age-specific fishing mortality and a 3-year reproductive cycle were used in age structured life tables which incorporated stochasticity in life history parameters through Monte Carlo simulation to estimate annual population growth. Demographic analyses showed that the dusky shark population would decline even at low levels of fishing mortality despite low natural mortality for neonates. The demographics of this species make it among the most vulnerable to excessive fishing mortality, and suggest stringent regulatory measures may be required to recover collapsed populations.     

Demography of the Juang tribal population of Orissa

Demographic analysis of genealogical data collected in 1954 for 23 Juang villages was undertaken employing indirect estimation techniques and computer projection methodology. Results indicated that this group did not feature the historically high fertility levels associated with Indian tribal groups, although fertility was higher than previously reported for the Juang. The population did feature a mortality differential, with worse mortality conditions than the Indian national population at this time. Reversed sexual mortality differentials, common in South Asian populations, were also present for the Juang. Computer projection investigation revealed a steadily growing population, in contrast to some Indian tribal groups faced with extinction.