Ecology

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.

Abstract: 
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.

Invertebrate communities in a tropical rain forest canopy in Puerto Rico following Hurricane Hugo

Schowalter, T. W., and L. M. Ganio. 1999. Invertebrate communities
in a tropical rain forest canopy in Puerto Rico
following Hurricane Hugo. Ecological Entomology 24:
191–201.

Abstract: 
1. Canopy invertebrate responses to Hurricane Hugo, tree species, and recovery time were examined at the Luquillo Experimental Forest in Puerto Rico during 1991–92 and 1994–95. Six tree species representing early and late successional stages were examined in paired plots representing severe hurricane disturbance (most trees toppled) and light hurricane disturbance (all trees standing and most branches intact). 2. Hurricane disturbance affected invertebrate abundances significantly. Sap-suckers and molluscs were more abundant, and defoliators, detritivores, and emergent aquatic insects were less abundant in recovering tree-fall gaps than in intact forest during this 5-year period. These changes in functional organisation are consistent with comparable studies of arthropod responses to canopy removal during harvest in temperate forests. 3. Tree species also affected invertebrate abundances significantly, but invertebrate communities did not differ significantly between the three early successional and three later successional tree species. 4. Most taxa showed significant annual variation in abundances, but only two Homoptera species showed a significant linear decline in abundance through time, perhaps reflecting long-term trends during recovery. 5. Leaf area missing, an indicator of herbivore effect on canopy processes, showed significant seasonal and annual trends, as well as differences among tree species and hurricane treatments. Generally, leaf area missing peaked during the wet season each year, but reached its highest levels during an extended drought in 1994. Leaf area missing also tended to be higher on the more abundant tree species in each disturbance treatment. 6. Herbivore abundances and leaf area missing were not related to concentrations of nitrogen, phosphorous, potassium, or calcium in the foliage. 7. This study demonstrated that invertebrate community structure and herbivory are dynamic processes that reflect the influences of host species and variable environmental conditions.

Separating the effects of forest type and elevation on the diversity of litter invertebrate communities in a humid tropical forest in Puerto Rico

RICHARDSON, BARBARA A.; RICHARDSON, MICHAEL J.; SOTO-ADAMES, FELIPE N. 2005. Separating the effects of forest type and elevation on the diversity of litter invertebrate communities in a humid tropical forest in Puerto Rico.. Journal of Animal Ecology 74, :926-936.

Abstract: 
1. The primary effects of climatic conditions on invertebrate litter communities, and the secondary effects of different forest types, were distinguished by using the sierra palm as a control in a natural experiment along an elevational gradient in the Luquillo Mountains. These mountains have three well-defined forest types along the gradient, with the palm occurring as stands within each forest. 2. Palm litter samples were richer in nutrients, particularly phosphorus, than nonpalm litter, significantly so at higher elevations where leaching would have been expected. In nonpalm litter, mineral concentrations were significantly lower at higher elevations. 3. Animal abundance mirrored the pattern of mineral amounts and declined significantly in mid- and high-altitude forests, but did not decline with increasing elevation in palm stands. A pulse of post-hurricane litterfall was reflected in the high abundance of Coleoptera and Isoptera the following year. 4. The species richness of communities (Margalef’s index) declined with increasing elevation in nonpalm forest litter, but was remarkably similar in palm litter at all elevations. 5. Palm litter communities were more similar to each other (Sørensen’s index) than nonpalm communities, which became less similar with increasing elevation. 6. The differences observed from the lower slopes to the summits, in animal abundance, species richness and the uniformity of communities, are better explained by the contribution of forest composition to the chemical and physical nature of litter and forest heterogeneity, rather than to direct effects of temperature and rainfall differences.

The Bromeliad Microcosm and the Assessment of Faunal Diversity in a Neotropical Forest

The Bromeliad Microcosm and the Assessment of Faunal Diversity in a Neotropical Forest
Barbara A. Richardson
Biotropica
Vol. 31, No. 2 (Jun., 1999), pp. 321-336

Abstract: 
The faunas of tank bromeliads were sampled over two years in three forest types at different elevations in the Luquillo Experimental Forest, Puerto Rico, and the diversity of their animal communities compared. Bromeliad plants behaved as islands in that, within forests, the species richness and abundance of their animal communities were significantly and positively correlated with increase in plant size. The amount of canopy debris they accumulated was similarly correlated with increase in plant size. Overall diversity was lowest in the dwarf forest, where plants were uniformly small. Animal communities were stable from year to year, and could be characterised for each forest type and for compartments within the plant. They showed a pattern of high dominance, which increased with elevation (McNaughton index 37, 54, and 73, respectively, for the tabonuco, palo colorado, and dwarf forest). Alpha-diversity for sites sampled in each year reflected net primary productivity (NPP) of the forest, declining with increasing elevation when animal abundance measures were used (jackknife estimates of Simpson's diversity index 6.54 & 11.04 [tabonuco], 3.53 & 6.22 [palo colorado], and 2.75 & 2.17 [dwarf forest]). Species richness over the two years, however, was highest in the intermediate palo colorado forest (187 species), compared to 146 and 88 in the tabonuco and dwarf forests, respectively. These figures were close to jackknife estimates of maximum species richness. The difference in species richness between tabonuco and palo colorado forests was significant in one year only. In addition to NPP, other factors, such as litter quality and the structural complexity of the habitat in the palo colorado forest, may have influenced species richness. The most abundant species in individual plants were also the most widely occurring, confirming known patterns of abundance and distribution in other functional groups. Diversity within bromeliad microcosms at different elevations supported known relationships between diversity, productivity, and habitat complexity along gradients and was not related to differences in the total bromeliad habitat available for colonization.

On the relative importance of pool morphology and woody debris to distributions of shrimp in a Puerto Rican headwater stream

Pyron, M., Covich, A.P. and Black, R.W. 1999. On the relative importance of pool morphology and
woody debris to distributions of shrimp in a Puerto Rican headwater stream. Hydrobiologia, 402:
207–215.

Abstract: 
In this paper, we report the sizes and distributional orientation of woody debris in a headwater rainforest stream in the Luquillo Experimental Forest (LEF), Puerto Rico. We also provide results of a 4-month study of a wood addition experiment designed to increase cover for benthic macroinvertebrates (freshwater shrimp). We added branch-sized woody debris to 20 pools in three streams.We trapped four species of freshwater shrimp (two species of benthic detritivores and two predatory shrimp species) during each of the 4 months following wood additions. An analysis of pool morphology (maximum depth, surface area and volume) provided a useful predictor of shrimp abundances. In general, numbers of shrimps increased with sizes of stream pools. A repeated measures ANOVA demonstrated no effect of woody debris additions on total numbers of shrimp per pool area. Two detritivore species (Atya lanipes, a filter feeder and Xiphocaris elongata, a shredder) decreased in abundance with increased woody debris and there was no statistical relationship between woody debris additions and predators (Macrobrachium carcinus and M. crenulatum). Small woody debris additions may have altered flow velocities that were important to filter-feeding Atya at the microhabitat scale, although the overall velocities within pools were not altered by wood additions. Lower numbers of Atya and Xiphocaris in two of the three streams may result from the occurrence of two predaceous fishes (American eel and mountainmullet) and more predatoryMacrobrachium in these streams. One likely interpretation of the results of this study is that the stream pools in these study reaches had sufficient habitat structure provided by numerous rock crevices (among large rocks and boulders) to provide refuge from predators. Addition of woody debris did not add significantly to the existing structure. These results may not apply to stream channels with sand and gravel substrata where crevices and undercut banks are lacking and where woody debris often plays a major role by providing structure and refuge.

Linking Species and Ecosystems: Different Biotic Assemblages Cause Interstream Differences in Organic Matter

Linking Species and Ecosystems: Different Biotic Assemblages Cause Interstream Differences in Organic Matter
C. M. Pringle, Nina Hemphill, W. H. McDowell, Angela Bednarek and James G. March
Ecology
Vol. 80, No. 6 (Sep., 1999), pp. 1860-1872

Abstract: 
Here we test the hypothesis that differences in macrobiotic assemblages can lead to differences in the quantity and quality of organic matter in benthic depositional environments among streams in montane Puerto Rico. We experimentally manipulated biota over a 30–40 d period in two streams with distinctly different macrobiotic assemblages: one characterized by high densities of omnivorous shrimps (Decapoda: Atyidae and Xiphocarididae) and no predaceous fishes, and one characterized by low densities of shrimps and the presence of predaceous fishes. To incorporate the natural hydrologic regime and to avoid confounding artifacts associated with cage enclosures/exclosures (e.g., high sedimentation), we used electricity as a mechanism for experimental exclusion, in situ. In each stream, shrimps and/or fishes were excluded from specific areas of rock substrata in four pools using electric “fences” attached to solar-powered fence chargers. In the stream lacking predaceous fishes (Sonadora), the unelectrified control treatment was almost exclusively dominated by high densities of omnivorous shrimps that constantly ingested fine particulate material from rock surfaces. Consequently, the control had significantly lower levels of inorganic sediments, organic material, carbon, and nitrogen than the exclusion treatment, as well as less variability in these parameters. Tenfold more organic material (as ash-free dry mass, AFDM) and fivefold more nitrogen accrued in shrimp exclosures (10.6 g AFDM/m2, 0.2 g N/m2) than in controls (1.1 g AFDM/m2, 0.04 g N/m2). By reducing the quantity of fine particulate organic material and associated nitrogen in benthic environments, omnivorous shrimps potentially affect the supply of this important resource to other trophic levels. The small amount of fine particulate organic matter (FPOM) that remained in control treatments (composed of sparse algal cells) was of higher quality than that in shrimp exclosures. This is evidenced by the significantly lower carbon-to-nitrogen (C/N) ratio (an indicator of food quality, with relatively low C/N indicating higher food quality) in the control relative to the shrimp exclosure treatment. In contrast, the stream with predaceous fishes (Bisley) was characterized by very low numbers of shrimps, and macrobiota had no significant effect on benthic sediments, organic matter, C, N, and C/N. All parameters were highly variable through time, with levels and ranges in variability similar to the shrimp exclusion treatment in the Sonadora. Our experimental results are consistent with findings of an independent survey of six streams in four different drainages. Four streams that had an abundance of omnivorous shrimps, but lacked predaceous fishes, had extremely low levels of fine benthic organic and inorganic material. In contrast, two streams that had low densities of shrimps and contained predaceous fishes had significantly higher levels. Results show a strong linkage between species and ecosystem characteristics: interstream differences in the quantity and quality of fine benthic organic matter resources were determined by the nature of the macrobiotic assemblage. Furthermore, patterns in the distribution of shrimp assemblages reflected landscape patterns in the benthic depositional environment among streams.

Marine dispersal determines the genetic population structure of migratory stream fauna of Puerto Rico: evidence for island-scale population recovery processes

Cook, Benjamin D.; Bernays, Sofie; Pringle, Catherine M.; Hughes, Jane M. 2009. Marine dispersal determines the genetic population structure of migratory stream fauna of Puerto Rico: evidence for island-scale population recovery processes. Journal of the North American Benthological Society. 28(3): 709-718.

Abstract: 
Various components of island stream faunas, including caridean shrimps, fish, and gastropods, undertake obligate amphidromous migration, whereby larvae are released in upstream freshwater reaches, drift downstream to estuaries or marine waters, then migrate upstream as postlarvae to freshwater adult habitats. Longitudinal migration from estuaries to headwaters is well documented for many amphidromous species, but the degree of among-river marine dispersal is poorly known for most species. We need better understanding of the potential for marine dispersal in population processes of amphidromous species, particularly recolonization and population recovery in impacted lotic systems, such as those on Puerto Rico, because some theories of dispersal for species with marine larvae predict high rates of self-recruitment. We tested population genetic predictions for widespread marine larval dispersal and self-recruitment to the natal river for 11 amphidromous species, including shrimps, fish, and a gastropod, in Puerto Rico. Population genetic analysis of mitochondrial DNA data showed high rates of gene flow among rivers and indicated that marine dispersal determines the population genetic structure of all 11 species. Difficulty in recruiting to oceanic currents promotes closed population structures in some marine species, but larvae of amphidromous species entrained in downstream river flow might be delivered more readily to ocean currents. Population recovery processes occurred at the island scale rather than at the river scale, but further studies are needed to identify whether population recovery processes are likely at larger spatial scales (e.g., among islands). River management strategies should maintain environmental flows that allow larval export, maintain longitudinal dispersal pathways over dam spillways and via subterranean passages, and maintain open and healthy estuaries.

Litterfall and Decomposition in Relation to Soil Carbon Pools Along a Secondary Forest Chronosequence in Puerto Rico

Ostertag, R.; Marín-Spiotta, E.; Silver, W.L.; Schulten, J. 2008. Litterfall and decomposition in relation to soil carbon pools along a secondary forest chronosequence in Puerto Rico. Ecosystems. 11:701-714.

Abstract: 
Secondary forests are becoming increasingly widespread in the tropics, but our understanding of how secondary succession affects carbon (C) cycling and C sequestration in these ecosystems is limited. We used a well-replicated 80-year pasture to forest successional chronosequence and primary forest in Puerto Rico to explore the relationships among litterfall, litter quality, decomposition, and soil C pools. Litterfall rates recovered rapidly during early secondary succession and averaged 10.5 (± 0.1 SE) Mg/ha/y among all sites over a 2-year period. Although forest plant community composition and plant life form dominance changed during succession, litter chemistry as evaluated by sequential C fractions and by 13C-nuclear magnetic resonance spectroscopy did not change significantly with forest age, nor did leaf decomposition rates. Root decomposition was slower than leaves and was fastest in the 60-year-old sites and slowest in the 10- and 30-year-old sites. Common litter and common site experiments suggested that site conditions were more important controls than litter quality in this chronosequence. Bulk soil C content was positively correlated with hydrophobic leaf compounds, suggesting that there is greater soil C accumulation if leaf litter contains more tannins and waxy compounds relative to more labile compounds. Our results suggest that most key C fluxes associated with litter production and decomposition re-establish rapidly—within a decade or two—during tropical secondary succession. Therefore, recovery of leaf litter C cycling processes after pasture use are faster than aboveground woody biomass and species accumulation, indicating that these young secondary forests have the potential to recover litter cycling functions and provide some of the same ecosystem services of primary forests.

the role of omnivory in a neotropical stream: separating diurnal and nocturnal effects

The Role of Omnivory in a Neotropical Stream: Separating Diurnal and Nocturnal Effects
Catherine M. Pringle and Toshihide Hamazaki
Ecology
Vol. 79, No. 1 (Jan., 1998), pp. 269-280

Abstract: 
The role of omnivory in structuring communities is potentially great in lowland neotropical streams that are characterized by an abundance of macroconsumers that consume both insects and algae. Here, we separate effects of natural densities of diurnal fishes and nocturnal shrimps in structuring the benthic community of a stream draining Costa Rica's Atlantic slope. We experimentally manipulated the spatial and temporal access of fishes and shrimps to benthic resources, in situ, using electric "fences" powered by solar-powered fence chargers. Both fishes and shrimps significantly reduced inorganic sediment mass, organic ashfree dry mass (AFDM), densities of larval Chironomidae, and total insects: their combined effects were greater than effects of either group alone, and there was no significant interaction. Fishes shifted algal community composition from diatoms to green and blue-green algae and benthic insect communities towards chironomids, while shrimps had no significant effect on community composition. Effects of fishes were generally greater than those of shrimps, and this is due, in part, to higher natural densities and foraging pressures of fishes. Furthermore, shrimps foraged for significantly longer periods of time in the treatment where fishes were excluded than in the combined fish and shrimp access treatment, suggesting that diurnally feeding fishes are strong "interactors," mediating resource availability to nocturnally feeding shrimps. Natural erosion and sediment-mediated effects of macroconsumers (both direct and indirect) also affected algal communities: a manual sediment removal experiment resulted in significant reductions of diatom biovolume and increases in the filament length of green and blue-green algae. Our results show the importance of: (1) assessing macroconsumer effects in a relatively natural depositional environment subject to background erosion and sloughing (i.e., in this case by using electric exclosures); (2) evaluating effects of natural densities of both diurnal and nocturnal macroconsumers through time in the context of these abiotic effects; and (3) distinguishing between the response of different types of algal resources (e.g., diatoms vs. green and blue-green algae), which are differentially affected by sedimentation and erosion. Cage experiments, short-term observations, or one-time sampling of undifferentiated "algae" may artificially overestimate trophic effects and underestimate abiotic effects. We found no evidence of a trophic cascade. Our findings are in agreement with the theoretical prediction that large-sized omnivores have strong direct trophic (feeding) effects, both on smaller primary consumers (insects) and basal resources (algae).

Effects of drought and hurricane disturbances on headwater distributions of palaemonid river shrimp (Macrobrachium spp.) in the Luquillo Mountains, Puerto Rico

Covich, Alan P.; Crowl, Todd A.; Heartsill-Scalley, Tamara 2006. Effects of drought and hurricane disturbances on headwater distributions of palaemonid river shrimp (Macrobrachium spp.) in the Luquillo Mountains, Puerto Rico.. J. N. Am. Benthol. Soc., 25(1):99-107.

Abstract: 
Extreme events (hurricanes, floods, and droughts) can influence upstream migration of macroinvertebrates and wash out benthic communities, thereby locally altering food webs and species interactions. We sampled palaemonid river shrimp (Macrobrachium spp.), dominant consumers in headwaters of the Luquillo Mountains of northeastern Puerto Rico, to determine their distributions along an elevational gradient (274–456 m asl) during a series of disturbances (Hurricane Hugo in 1989, a drought in 1994, and Hurricane Georges in 1998) that occurred over a 15-y period (19882002).We measured shrimp abundance 3 to 6 times/y in Quebrada Prieta in the Espiritu Santo drainage as part of the Luquillo Long-Term Ecological Research Program. In general, Macrobrachium abundance declined with elevation during most years. The lowest mean abundance of Macrobrachium occurred during the 1994 drought, the driest year in 28 y of record in the Espiritu Santo drainage. Macrobrachium increased in abundance for 6 y following the 1994 drought. In contrast, hurricanes and storm flows had relatively little effect on Macrobrachium abundance.
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