Seed fate and seedling dynamics after masting in two African rain forest trees

How the effects of biotic factors are moderated by abiotic factors, and their consequences for species interactions, is generally understudied in ecology. A key abiotic feature of forests is regular canopy disturbances that create temporary patches, or “gaps,” of above-average light availability. Co-occurring in lowland primary forest of Korup National Park (Cameroon), Microberlinia bisulcata and Tetraberlinia bifoliolata are locally dominant, ectomycorrhizal trees whose seeds share predator guilds in masting years. Here, we experimentally tested the impact of small mammal predators upon seedling abundance, growth, and survivorship. In 2007, we added a fixed density of seeds of each species to exclosures at 48 gap–understory locations across 82.5 ha within a large Microberlinia grove, and at 15 locations outside it. For both species, small mammals removed more seeds in gaps than in understory, whereas this was reversed for seeds killed by invertebrates. Nonetheless, Microberlinia lost twice as many seeds to small mammals, and more to invertebrates in exclosures, than Tetraberlinia, which was more prone to a pathogenic white fungus. After six weeks, both species had greater seedling establishment in gaps than understory, and in exclosures outside compared to exclosures inside the grove. In the subsequent two-year period, seedling growth and survivorship peaked in exclosures in gaps, but Microberlinia had more seedlings' stems clipped by animals than Tetraberlinia, and more than twice the percentage of leaf area damaged. Whereas Microberlinia seedling performance in gaps was inferior to Tetraberlinia inside the grove, outside it Microberlinia had reduced leaf damage, grew taller, and had many more leaves than Tetraberlinia. No evidence was found for “apparent mutualism” in the understory as seedling establishment of both species increased away from (>25 m) large stems of either species, pointing to “apparent competition” instead. In gaps, Microberlinia seedling establishment was lower near Tetraberlinia than conspecific adults because of context-dependent small mammal satiation. Stage-matrix analysis suggested that protecting Microberlinia from small mammals could increase its population growth rate by 0.06. In the light of prior research we conclude that small mammals and canopy gaps play an important role in promoting species coexistence in this forest, and that their strong interaction contributes to Microberlinia's currently very poor regeneration.

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 fruiting tree Microberlinia bisulcata occurs as a codominant in groves of ectomycorrhizal Caesalpiniaceae within a mosaic of otherwise species-rich lowland rain forest. To estimate the amount of carbon and nutrients invested in reproduction during a mast fruiting event, and the consequential seed and seedling survival, three related field studies were made in 1995. These provided a complete seed and seedling budget for the cohort. Seed production was estimated by counting woody pods on the forest floor. Trees produced on average 26,000 (range 0-92,000) seeds/tree, with a dry mass of 16.6 kg/tree. Seeds were contained in woody pods of mass 307 kg/tree. Dry mass production of pods and seeds was 1034 kg ha(-1), equivalent to over half (55%) of annual leaf litterfall for this species, and contained 13% of the nitrogen and 21% of the phosphorus in annual leaf litterfall. Seed and young-seedling mortality was investigated with open quadrats and cages to exclude vertebrate predators, at two distances from the parent tree. The proportion of seeds on the forest floor which disappeared in the first 6 wk after dispersal was 84%, of which 26.5% was due to likely vertebrate removal, 36% to rotting, and 21.5% to other causes. Vertebrate predation 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. Previous studies have demonstrated an association between mast years at Korup and high dry-season radiation before flowering, and have shown lower leaf-litterfall phosphorus concentrations following mast fruiting. The emerging hypothesis is that mast fruiting is primarily imposed by energy limitation for fruit production, but phosphorus supply and vertebrate predation are regulating factors. Recording the survival of naturally-regenerating M. bisulcata seedlings (6-wk stage) showed that 21% of seedlings survived to 31 mo. A simple three-stage recruitment model was constructed. Mortality rates were initially high and peaked again in each of the next two dry seasons, with smaller peaks in the two intervening wet seasons, these latter coinciding with annual troughs in radiation. The very poor recruitment of M. bisulcata trees in Korup, demonstrated in previous investigations, appears not to be due to a limitation in seed or young-seedling supply, but rather by factors operating at the established-seedling stage.

Tree size and fecundity influence ballistic seed dispersal of two dominant mast-fruiting species in a tropical rain forest

Seed production, seed dispersal, and seedling recruitment are integral to forest dynamics, especially in masting species. Often these are studied separately, yet scarcely ever for species with ballistic dispersal even though this mode of dispersal is common in legume trees of tropical African rain forests. Here, we studied two dominant main-canopy tree species, Microberlinia bisulcata and Tetraberlinia bifoliolata (Caesalpinioideae), in 25 ha of primary rain forest at Korup, Cameroon, during two successive masting events (2007/2010). In the vicinity of c. 100 and 130 trees of each species, 476/580 traps caught dispersed seeds and beneath their crowns c. 57,000 pod valves per species were inspected to estimate tree-level fecundity. Seed production of trees increased non-linearly and asymptotically with increasing stem diameters. It was unequal within the two species’ populations, and differed strongly between years to foster both spatial and temporal patchiness in seed rain. The M. bisulcata trees could begin seeding at 42–44 cm diameter: at a much larger size than could T. bifoliolata (25 cm). Nevertheless, per capita life-time reproductive capacity was c. five times greater in M. bisulcata than T. bifoliolata owing to former’s larger adult stature, lower mortality rate (despite a shorter life-time) and smaller seed mass. The two species displayed strong differences in their dispersal capabilities. Inverse modelling (IM) revealed that dispersal of M. bisulcata was best described by a lognormal kernel. Most seeds landed at 10–15 m from stems, with 1% of them going beyond 80 m (<100 m). The direct estimates of fecundity significantly improved the models fitted. The lognormal also described well the seedling recruitment distribution of this species in 121 ground plots. By contrast, the lower intensity of masting and more limited dispersal of the heavier-seeded T. bifoliolata prevented reliable IM. For this species, seed density as function of distance to traps suggested a maximum dispersal distance of 40–50 m, and a correspondingly more aggregated seedling recruitment pattern ensued than for M. bisulcata. From this integrated field study, we conclude that the reproductive traits of M. bisulcata give it a considerable advantage over T. bifoliolata by better dispersing more seeds per capita to reach more suitable establishment sites, and combined with other key traits they explain its local dominance in the forest. Understanding the linkages between size at onset of maturity, individual fecundity, and dispersal capability can better inform the life-history strategies, and hence management, of co-occurring tree species in tropical forests.

Herbivores limit the population size of big-leaf mahogany trees in an Amazonian forest

The Janzen–Connell hypothesis proposes that specialized herbivores maintain high numbers of tree species in tropical forests by restricting adult recruitment so that host populations remain at low densities. We tested this prediction for the large timber tree species, Swietenia macrophylla, whose seeds and seedlings are preyed upon by small mammals and a host-specific moth caterpillar Steniscadia poliophaea, respectively. At a primary forest site, experimental seed additions to gaps – canopy-disturbed areas that enhance seedling growth into saplings – over three years revealed lower survival and seedling recruitment closer to conspecific trees and in higher basal area neighborhoods, as well as reduced subsequent seedling survival and height growth. When we included these Janzen–Connell effects in a spatially explicit individual-based population model, the caterpillar's impact was critical to limiting Swietenia's adult tree density, with a > 10-fold reduction estimated at 300 years. Our research demonstrates the crucial but oft-ignored linkage between Janzen–Connell effects on offspring and population-level consequences for a long-lived, potentially dominant tree species.

Light and seed size affect establishment of grove-forming ectomycorrhizal rain forest tree species

• Regeneration of the dominant ectomycorrhizal tree Microberlinia bisulcata in groves in Korup, Central Africa, is very poor. The hypothesis was tested that this species is more shade intolerant than other co-occurring species. • In two 1-yr trials, each with M. bisulcata and four other species at a nursery close to Korup, growth was measured under five PAR levels, with ± added P and ± watering in the dry season. In parallel experiments the effects of PAR with two R : FR ratios were investigated. • Increasing PAR had a consistent effect on the rates of increase in plant mass and on changes in the other variables. Doubling soil P, watering and halving the R : FR ratio had almost no effect. However, across species, mass at low PAR and relative growth rate related positively and negatively, respectively, to seed mass. • One contributing factor for the poor recruitment of M. bisulcata is therefore its low survival and slow growth at low PAR, due to its small seed size. The two codominant ectomycorrhizal grove species of Tetraberlinia, with larger seeds, were less affected by low PAR.