Shiels A.B.

Early successional woody plants facilitate and ferns inhibit forest development on Puerto Rican landslides

Walker, L.R., Landau, F.H., Velázquez, E., Shiels,
A.B. and Sparrow, A.D. (2010). Early successional
woody plants facilitate and ferns inhibit forest
development on Puerto Rican landslides. Journal
of Ecology 98, 625-35.

1. The experimental removal of early successional species can explain how plant communities change over time. 2. During a 7.3-year period, early successional woody species, scrambling ferns and tree ferns were removed from a total of 10 landslides in the Luquillo Experimental Forest in north-eastern Puerto Rico. 3. Early successional woody plants in combination with tree ferns decreased species richness and cover of forbs and increased richness of late-successional woody plants compared to removals, facilitating long-term forest development. 4. Dense stands of scrambling ferns decreased both forb and woody plant richness compared to removals, inhibiting forest development. 5. Stands of monospecific tree ferns initially increased woody plant richness compared to removals, but overall decreased woody plant richness and cover, inhibiting forest development. 6. Synthesis. Early successional species both facilitate and inhibit succession on tropical landslides, but detailed predictions of successional trajectories remain elusive and are influenced by stochastic processes including arrival order, the life-form of colonizing species and their competitive interactions.

Soil factors predict initial plant colonization on Puerto Rican landslides

Shiels, A.B., West, C.A., Weiss, L., Klawinski, P.D. &
Walker, L.R. 2008. Soil factors predict initial plant
colonization on Puerto Rican landslides. Plant
Ecology 195: 165–178.

Tropical storms are the principal cause of landslides in montane rainforests, such as the Luquillo Experimental Forest (LEF) of Puerto Rico. A storm in 2003 caused 30 new landslides in the LEF that we used to examine prior hypotheses that slope stability and organically enriched soils are prerequisites for plant colonization. We measured slope stability and litterfall 8–13 months following landslide formation. At 13 months we also measured microtopography, soil characteristics (organic matter, particle size, total nitrogen, and water-holding capacity), elevation, distance to forest edge, and canopy cover. When all landslides were analyzed together, plant biomass and cover at 13 months were not correlated with slope stability or organic matter, but instead with soil nitrogen, clay content, waterholding capacity, and elevation. When landslides were analyzed after separating by soil type, the distance from the forest edge and slope stability combined with soil factors (excluding organic matter) predicted initial plant colonization on volcaniclastic landslides, whereas on diorite landslides none of the measured characteristics affected initial plant colonization. The life forms of the colonizing plants reflected these differences in landslide soils, as trees, shrubs, and vines colonized high clay, high nitrogen, and low elevation volcaniclastic soils, whereas herbs were the dominant colonists on high sand, low nitrogen, and high elevation diorite soils. Therefore, the predictability of the initial stage of plant succession on LEF landslides is primarily determined by soil characteristics that are related to soil type.

Plant responses to simulated hurricane impacts in a subtropical wet forest, Puerto Rico

Shiels, Aaron B.; Zimmerman, Jess K.; García-Montiel, Diana C.; Jonckheere, Inge; Holm, Jennifer; Horton, David; Brokaw, Nicholas. 2010. Plant responses to simulated hurricane impacts in a subtropical wet forest, Puerto Rico. Journal of Ecology. doi: 10.1111/j.1365-2745.2010.01646.x.

1. We simulated two key components of severe hurricane disturbance, canopy openness and detritus deposition, to determine the independent and interactive effects of these components on woody plant recruitment and forest structure. 2. We increased canopy openness by trimming branches and added or subtracted canopy detritus in a factorial design. Plant responses were measured during the 4-year study, which followed at least 1 year of pre-manipulation monitoring. 3. The physical conditions of canopy openness and detritus deposition in our experiment resembled the responses to Hurricane Hugo, a severe category 4 hurricane that struck this forest in 1989. 4. Canopy detritus deposition killed existing woody seedlings and provided a mechanical barrier that suppressed seedling recruitment. The increase in understorey light caused by canopy trimming stimulated germination from the seed bank and increased seedling recruitment and density of pioneer species several hundred-fold when hurricane debris was absent. Many significant interactions between trimming and detritus deposition were evident from the manner in which seedling density, recruitment and mortality changed over time, and subsequently influenced the composition of woody stems (individuals ‡ 1 cmd.b.h.). 5. When the canopy was trimmed, stem densities increased> 2-fold and rates of recruitment into the stem size class increased> 25-fold. Trimming had no significant effect on stem mortality. The two dominant species that flourished following canopy trimming were the pioneer species Cecropia schreberiana and Psychotria berteriana. Deposition of canopy detritus had little effect on stems, although basal area increased slightly when detritus was added. There were no evident effects of the interactions between canopy trimming and detritus deposition on stems. 6. Synthesis. The separate and interactive effects of canopy openness and detritus deposition result in variable short-term trajectories of forest recovery. However, the short interval of increased canopy openness due to hurricane impacts and its influence on the recruitment of pioneer trees is the dominant factor that drives short-termrecovery and may alter long-term structure and composition of the forest.

A Canopy Trimming Experiment in Puerto Rico: The Response of Litter Invertebrate Communities to Canopy Loss and Debris Deposition in a Tropical Forest Subject to Hurricanes

Richardson, Barbara A.; Richardson, Michael J.; Gonzalez, Grizelle; Shiels, Aaron B.; Srivastava, Diane S. 2010. A canopy trimming experiment in Puerto Rico: the response of litter invertebrate communities to canopy loss and debris deposition in a tropical forest subject to hurricanes. Ecosystems. 13: 286-301.

Hurricanes cause canopy removal and deposition of pulses of litter to the forest floor. A Canopy Trimming Experiment (CTE) was designed to decouple these two factors, and to investigate the separate abiotic and biotic consequences of hurricane-type damage and monitor recovery processes. As part of this experiment, effects on forest floor invertebrate communities were studied using litterbags. Canopy opening resulted in increased throughfall, soil moisture and light levels, but decreased litter moisture. Of these, only throughfall and soil moisture had returned to control levels 9 months after trimming. Canopy opening was the major determinant of adverse changes in forest floor invertebrate litter communities, by reducing diversity and biomass, irrespective of debris deposition, which played a secondary role. Plots subjected to the most disturbance, with canopy removed and debris added, had the lowest diversity and biomass. These two parameters were higher than control levels when debris was added to plots with an intact canopy, demonstrating that increased nutrient potential or habitat complexity can have a beneficial effect, but only if the abiotic conditions are suitable. Animal abundance remained similar over all treatments, because individual taxa responded differentlyto canopy trimming. Mites, Collembola, and Psocoptera, all microbiovores feeding mainly on fungal hyphae and spores, responded positively, with higher abundance in trimmed plots, whereas all other taxa, particularly predators and larger detritivores, declined in relative abundance. Litterbag mesh size and litter type had only minor effects on communities, and canopy trimming and debris deposition explained most variation between sites. Effects of trimming on diversity, biomass, and abundance of some invertebrate taxa were still seen when observations finished and canopy closure was complete at 19 months. This suggests that disturbance has a long-lasting effect on litter communities and may, therefore, delay detrital processing, depending on the severity of canopy damage and rate of regrowth.

Landsliding and Its Multiscale Influence on Mountainscapes

Restrepo, Carla; Walker, Lawrence R.; Shiels, Aaron B.; Bussmann, Rainer; Claessens, Lieven; Fisch, Simey; Lozano, Pablo; Negi, Girish; Paolini, Leonardo; Poveda, Germán; Ramos-Sharrón, Carlos; Ritcher, Michael; Velázquez, Eduardo. 2009. Landsliding and its multiscale influence on mountainscapes. Bioscience. 59(8): 685-698.

Landsliding is a complex process that modifies mountainscapes worldwide. Its severe and sometimes long-lasting negative effects contrast with the less-documented positive effects on ecosystems, raising numerous questions about the dual role of landsliding, the feedbacks between biotic and geomorphic processes, and, ultimately, the ecological and evolutionary responses of organisms. We present a conceptual model in which feedbacks between biotic and geomorphic processes, landslides, and ecosystem attributes are hypothesized to drive the dynamics of mountain ecosystems at multiple scales. This model is used to integrate and synthesize a rich, but fragmented, body of literature generated in different disciplines, and to highlight the need for profitable collaborations between biologists and geoscientists. Such efforts should help identify attributes that contribute to the resilience of mountain ecosystems, and also should help in conservation, restoration, and hazard assessment. Given the sensitivity of mountains to land-use and global climate change, these endeavors are both relevant and timely.
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