Effects of Treefall Gap Disturbances on Ant Assemblages in a Tropical Montane Cloud Forest

The study of gap dynamics and the effects of gaps on diversity has been at the center of tropical ecology for decades. While most studies have focused on the responses of plant species and communities to gap formation, in this study, we consider the effects of treefall gap disturbances on leaf litter ant assemblages in a Neotropical montane cloud forest. We sampled leaf litter ant assemblages and estimated a suite of abiotic parameters in 12 large (>80‐m²) treefall gaps across a chronosequence and in 12 paired adjacent intact forest sites in the Monteverde Cloud Forest Preserve in Costa Rica. No species were more common in gaps than in intact forests, and in fact, species that were common in gaps were also among the most common in forests. The Chao2 estimate of species richness, however, was higher in gap sites than in intact forest sites. In addition, ant assemblages in gap sites did not become more similar to those in adjacent intact sites as gaps aged. In contrast to other studies, our work demonstrates that ant assemblages in the Monteverde Cloud Forest Preserve are weakly affected by the formation of treefall gaps. Together, these results indicate that treefall gap dynamics probably play little role in promoting ant diversity at more regional scales, or coexistence among species at more local scales.     

Planting Seedlings in Tree Islands Versus Plantations as a Large‐Scale Tropical Forest Restoration Strategy

Planting tree seedlings in small patches (islands) has been proposed as a method to facilitate forest recovery that is less expensive than planting large areas and better simulates the nucleation process of recovery. We planted seedlings of four tree species at 12 formerly agricultural sites in southern Costa Rica in two designs: plantation (entire 50 × 50 m area planted) and island (six patches of three sizes). We monitored seedling survival, height, and canopy area over 3 years. To elucidate mechanisms influencing survival and growth, we measured soil and foliar nutrients, soil compaction, and photosynthesis. Survival of all species was similar in the two planting designs. Seedling height and canopy area were greater in plantations than islands at most sites, and more seedlings in islands decreased in height due to damage incurred during plot maintenance. Survival, height, and canopy area were both site‐ and species‐specific with the two N‐fixing species (Inga edulis and Erythrina poeppigiana) greater than the other species (Terminalia amazonia and Vochysia guatemalensis). Foliar N was higher in Terminalia and Vochysia in sites where Inga growth was greater. Soil nutrients, however, explained a small amount of the large differences in growth across sites. Leaf mass per area was higher in islands, and P use efficiency was higher in plantations. Our results show advantages (good seedling survival, cheaper) and disadvantages (more seedling damage, slightly lower growth) to the island planting design. Our study highlights the importance of replicating restoration strategies at several sites to make widespread management recommendations.     

Age and Long-term Growth of Trees in an Old-growth Tropical Rain Forest, Based on Analyses of Tree Rings and 14C1

In an old‐growth tropical wet forest at La Selva, Costa Rica, we combined radiocarbon (¹⁴C) dating and tree‐ring analysis to estimate the ages of large trees of canopy and emergent species spanning a broad range of wood densities and growth rates. We collected samples from the trunks of 29 fallen, dead individuals. We found that all eight sampled species formed visible growth rings, which varied considerably in distinctiveness. For five of the six species for which we combined wood anatomical studies with ¹⁴C‐dates (ring ages), the analyses demonstrated that growth rings were of annual formation. The oldest tree we found by direct ring counting was a Hymenolobium mesoamericanum Lima (Papilionaceae) specimen, with an age of ca. 530 years at the time of death. All other sampled individuals, including very large trees of slow‐growing species, had died at ages between 200 and 300 years. These results show that, even in an everwet tropical rain forest, tree growth of many species can be rhythmic, with an annual periodicity. This study thus raises the possibility of extending tree‐ring analyses throughout the tropical forest types lacking a strong dry season or annual flooding. Our findings and similar measurements from other tropical forests indicate that the maximum ages of tropical emergent trees are unlikely to be much greater than 600 years, and that these trees often die earlier from various natural causes.     

Seedling Mortality in Hawaiian Rain Forest: The Role of Small‐Scale Physical Disturbance1

Most montane rain forests on the island of Hawaii consist of a closed canopy formed by Cibotium spp. tree ferns beneath an open canopy of emergent Metrosideros polymorpha trees. We used artificial seedlings to assess the extent to which physical disturbance caused by the senescing fronds of tree ferns and the activities of feral pigs might limit tree regeneration. Artificial seedlings were established terrestrially (N= 300) or epiphytically (N = 300) on tree fern stems. Half of the seedlings on each substrate were in an exclosure lacking feral pigs and half were in forest with pigs present. After one year, the percentage of seedlings damaged was significantly greater among terrestrial seedlings (25.7%) than epiphytic seedlings (11.3%). Significantly more terrestrial seedlings were damaged in the presence of pigs (31.3%) than in the absence of pigs (20.0%). Senescing fronds of tree ferns were responsible for 60.3 percent of the damaged seedlings. Physical disturbance is potentially a major cause of seedling mortality and may reduce the expected half‐life of a seedling cohort to less than two years.     

Impacts of Vegetation and Precipitation on Throughfall Heterogeneity in a Tropical Pre‐Montane Transitional Cloud Forest

Precipitation throughfall (TF) plays an important role in the water balance of tropical forests. This study used 164 gauges to quantify precipitation and TF variability in a tropical pre‐montane transitional cloud forest on the Caribbean slope of the Cordillera Tilarán, Costa Rica, to identify the ecological and meteorological drivers of this variability. Daily TF measurements were taken from 28 June to 17 July 2012 and 12 June to 16 July 2013, for a total of 39 precipitation events. The total mean TF was 87.9 percent and TF at individual gauges ranged from 22.7 percent to 245.7 percent. Leaf area index (LAI) was calculated above each gauge using hemispheric photography for a mean study‐site LAI of 7.7. There was no statistically significant relationship between LAI and TF. However, the amount of TF was positively correlated with precipitation intensity, while the variability of TF was negatively correlated with precipitation intensity. Our calculations indicate that at least 61 gauges are required to obtain mean TF estimates with less than 5 percent error. This study demonstrates that TF is highly spatially heterogeneous due to multiple compounding effects.     

Modification of Vegetative Phenology in a Tropical Semi‐deciduous Forest by Abnormal Drought and Rain1

The control of vegetative phenology in tropical trees is not well understood. In dry forest trees, leaf abscission may be enhanced by advanced leaf age, increasing water stress, or declining photoperiod. Normally, it is impossible to dissect the effects of each of these variables because most leaves are shed during the early dry season when day length is near its minimum and leaves are relatively old. The 1997 El‐Niño Southern Oscillation caused a ten‐week long, severe abnormal drought from June to August in the semi‐deciduous forests of Guanacaste, Costa Rica. We monitored the effect of this drought on phenology and water status of trees with young leaves and compared modifications of phenology in trees of different functional types with the pattern observed during the regular dry season. Although deciduous trees at dry sites were severely water stressed (Ψstem < ‐7MPa) and their mesic leaves remained wilted for more than two months, these and all other trees retained all leaves during the abnormal drought. Many trees exchanged leaves three to four months earlier than normal during the wet period after the abnormal drought and shed leaves again during the regular dry season. Irrigation and an exceptional 70 mm rainfall during the mid‐dry season 1998/1999 caused bud break and flushing in all leafless trees except dormant stem succulents. The complex interactions between leaf age and water stress, the principal determinants of leaf abscission, were found to vary widely among trees of different functional types.     

Reproductive Ecology of Understory Species in a Tropical Montane Forest in Costa Rica1

The reproductive ecology of nine hermaphroditic understory species in a tropical montane Quercus forest was studied at two sites (2300 and 2600 m elev.) in the Cordillera de Talamanca, Costa Rica. Flower life span, studied in six species, averaged 4.4 d. This is longer than flower life spans found in the Monteverde cloud forest (2.7 d) and comparable to flower life spans found for arctic and alpine species. We studied the breeding system in five species and found no self-incompatible species. Four species proved self-compatible, and three of these showed autogamy. The main diurnal insect pollinator was the bumblebee Bombus ephippiatus. Natural fruit set was low (8-32%) in six species with few seeds per fruit, while two many-seeded species showed a high rate of fruit set (90 and 96%). The incidence of pre–dispersal seed predation was high; the percentage of seeds infested in four species ranged from 8 to 56 percent.     

Inner‐crown Microenvironments of Two Emergent Tree Species in a Lowland Wet Forest1

Vascular epiphyte communities, comprising up to 25 percent of tropical forest flora, contribute to plant diversity and thus ecosystem‐level processes; however, one of the proximal determinants of those communities, microclimate, is little studied. Here we present the first comprehensive study of microclimates in the inner crowns of two emergent tree species, Hyeronima alchorneoides and Lecythis ampla, at La Selva Biological Station, Costa Rica. We examined photon flux density, temperature, vapor pressure, and humidity in inner‐crown branches during the wet and dry seasons and during the wet‐season leafless phase of Lecythis. In both seasons, the percentage daily PFD in foliated Lecythis crowns (9%, wet season; 11%, dry season) was significantly higher than in Hyeronima crowns (5%, both seasons), with the leafless wet‐season PFD of Lecythis reaching 23 percent of full sun. Temperature and vapor pressure varied less in Hyeronima than in Lecythis crowns during the dry season. Microenvironmental conditions for epiphytes within Hyeronima crowns were more spatially and temporally homogeneous and were more buffered from ambient conditions than within Lecythis crowns. Growing conditions within the crowns of the same trees and among different trees were measurably different and are likely to affect the structure and composition of the resident epiphyte communities.     

Ants Induce Domatia in a Rain Forest Tree (Vochysia vismiaefolia)1

In Amazonian rain forest trees of Vochysia vismiaefolia (Vochysiaceae), ants were found to induce twig structures that resembled classical ant domatia. This phenomenon is novel for ant‐plants, which commonly develop domatia without the activity of ants. Eight species of ants were recorded inside the domatia of six individual trees, but domatia were most numerous and characteristic when induced and inhabited by an undescribed species of Pseudomyrmex on two trees. To investigate the mechanism of domatium growth, we drilled holes into young twigs: the expansion of the twig diameter surrounding the holes was significantly accelerated, comparable to domatia formation. Domatia induction is discussed as a putative step in the evolution of ant‐plants.     

Increasing Liana Abundance and Basal Area in a Tropical Forest: The Contribution of Long‐distance Clonal Colonization

Recent evidence suggests that liana abundance and biomass are increasing in Neotropical forests, representing a major structural change to tropical ecosystems. Explanations for these increases, however, remain largely untested. Over an 8‐yr period (1999–2007), we censused lianas in nine, 24 × 36 m permanent plots in old‐growth and selectively logged forest at La Selva Biological Station, Costa Rica to test whether: (1) liana abundance and basal area are increasing in this forest; (2) the increase is being driven by increased recruitment, decreased mortality, or both; and (3) long‐distance clonal colonization explains the increase in liana abundance and basal area. We defined long‐distance clonal colonization as lianas that entered and rooted in the plots as vegetative propagules of stems that originated from outside or above the plot, and were present in 2007, but not in 1999 or 2002. Our hypotheses were supported in the old‐growth forest: mean liana abundance and BA (≥1 cm diameter) increased 15 and 20 percent, respectively, and clonal colonization from outside of the plots contributed 19 and 60 percent (respectively) to these increases. Lianas colonized clonally by falling vertically from the forest canopy above or growing horizontally along the forest floor and re‐rooting—common forms of colonization for many liana species. In the selectively logged forest, liana abundance and BA did not change, and thus the pattern of increasing lianas may be restricted to old‐growth forests. In summary, our data support the hypothesis that lianas are increasing in old‐growth forests, and that long‐distance clonal colonization is a major contributor.     

Heterogeneity of Light Availability and its Effects on Simulated Carbon Gain of Tree Leaves in a Small Gap and the Understory in a Tropical Rain Forest1

To clarify the small-scale heterogeneity of light regimes in a rain forest, photosynthetic photon flux density (PFD) was measured at 1-min intervals during six days at 12 microsites in each of two plots, a small gap and an understory in Pasoh Forest Reserve, Peninsular Malaysia. Frequency distribution of microsite PFD was unimodal with the peak value between 16 and 32 μmol/m²/sec in the small gap, but between 8 and 16 μmol/m²/sec in the understory. In the small gap, PFD was more variable among microsites; total daily PFD and daily sunfleck PFD exceeding 10 μmol/ m²/sec tended to be higher (P <0.05; t-test) compared to those in the understory. Sunfleck PFD exceeding 50 μmol/ m²/sec, however, showed no difference between the two plots. Diffuse PFD transmittance, defined as the ratio of PFD in the forest to that measured at 43 m above ground during the periods 0800-0810 and 1750-1800 h, was significantly higher in the small gap than in the understory plot. Diffuse PFD transmittance was also positively correlated with microsite total daily PFD. To examine the effects of the subtle heterogeneity of light regimes on leaf carbon gain, we simulated carbon gain by sun and shade leaves in a typical shade-tolerant species, Brosimum aticastrum Sw. (Moraceae). Despite the similarity in total daily PFD, total daily carbon gain was considerably higher in the gap than in the understory for both sun and shade leaves. This study suggests that frequency distribution of PFD is critical in describing microsite PFD regimes and determining leaf carbon gain in the tropical forest floor.     

The Role of Arboreal Seed Dispersal Groups on the Seed Rain of a Lowland Tropical Forest1

Most tropical plants produce fleshy fruits that are dispersed primarily by vertebrate frugivores. Behavioral disparities among vertebrate seed dispersers could influence patterns of seed distribution and thus forest structure. This study investigated the relative importance of arboreal seed dispersers and seed predators on the initial stage of forest organization–seed deposition. We asked the following questions: (1) To what degree do arboreal seed dispersers influence the species richness and abundance of the seed rain? and (2) Based on the plant species and strata of the forest for which they provide dispersal services, do arboreal seed dispersers represent similar or distinct functional groups? To answer these questions, seed rain was sampled for 12 months in the Dja Reserve, Cameroon. Seed traps representing five percent of the crown area were erected below the canopies of 90 trees belonging to nine focal tree species: 3 dispersed by monkeys, 3 dispersed by large frugivorous birds, and 3 wind‐dispersed species. Seeds disseminated by arboreal seed dispersers accounted for ca 12 percent of the seeds and 68 percent of the seed species identified in seed traps. Monkeys dispersed more than twice the number of seed species than large frugivorous birds, but birds dispersed more individual seeds. We identified two distinct functional dispersal groups, one composed of large frugivorous birds and one composed of monkeys, drop dispersers, and seed predators. These groups dispersed plants found in different canopy strata and exhibited low overlap in the seed species they disseminated. We conclude it is unlikely that seed dispersal services provided by monkeys could be compensated for by frugivorous birds in the event of their extirpation from Afrotropical forests.     

The Role of Dissolved Organic Carbon, Dissolved Organic Nitrogen, and Dissolved Inorganic Nitrogen in a Tropical Wet Forest Ecosystem

Although tropical wet forests play an important role in the global carbon (C) and nitrogen (N) cycles, little is known about the origin, composition, and fate of dissolved organic C (DOC) and N (DON) in these ecosystems. We quantified and characterized fluxes of DOC, DON, and dissolved inorganic N (DIN) in throughfall, litter leachate, and soil solution of an old-growth tropical wet forest to assess their contribution to C stabilization (DOC) and to N export (DON and DIN) from this ecosystem. We found that the forest canopy was a major source of DOC (232 kg C ha^–1 y^–1). Dissolved organic C fluxes decreased with soil depth from 277 kg C ha^–1 y^–1 below the litter layer to around 50 kg C kg C ha^–1 y^–1 between 0.75 and 3.5m depth. Laboratory experiments to quantify biodegradable DOC and DON and to estimate the DOC sorption capacity of the soil, combined with chemical analyses of DOC, revealed that sorption was the dominant process controlling the observed DOC profiles in the soil. This sorption of DOC by the soil matrix has probably led to large soil organic C stores, especially below the rooting zone. Dissolved N fluxes in all strata were dominated by mineral N (mainly NO₃⁻). The dominance of NO₃^– relative to the total amount nitrate of N leaching from the soil shows that NO₃^– is dominant not only in forest ecosystems receiving large anthropogenic nitrogen inputs but also in this old-growth forest ecosystem, which is not N-limited.