Invasive rodents are among the most ubiquitous and problematic species introduced to islands; more than 80% of the world's island groups have been invaded. Introduced rats (black rat, Rattus rattus; Norway rat, R. norvegicus; Pacific rat, R. exulans) are well known as seed predators but are often overlooked as potential seed dispersers despite their common habit of transporting fruits and seeds prior to consumption. The relative likelihood of seed predation and dispersal by the black rat, which is the most common rat in Hawaiian forest, was tested with field and laboratory experiments. In the field, fruits of eight native and four non-native common woody plant species were arranged individually on the forest floor in four treatments that excluded vertebrates of different sizes. Eleven species had a portion (3-100%) of their fruits removed from vertebrate-accessible treatments, and automated cameras photographed only black rats removing fruit. In the laboratory, black rats were offered fruits of all 12 species to assess consumption and seed fate. Seeds of two species (non-native Clidemia hirta and native Kadua affinis) passed intact through the digestive tracts of rats. Most of the remaining larger-seeded species had their seeds chewed and destroyed, but for several of these, some partly damaged or undamaged seeds survived rat exposure. The combined field and laboratory findings indicate that many interactions between black rats and seeds of native and non-native plants may result in dispersal. Rats are likely to be affecting plant communities through both seed predation and dispersal.
and 4 718 Inglestone Court, Manchester, MO 63021, USA Summary 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 cm d.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-term recovery and may alter long-term structure and composition of the forest.
The diets of sympatric rodents partially define their realized niches. Identifying items in stomachs of introduced rodents helps determine rodents' trophic positions and species most at risk of consumption. In the Hawaiian Islands, which lacked rodents prior to human arrival, three rodents (Rattus rattus or black rat, R. exulans or Pacific rat, Mus musculus or house mouse) commonly coexist in native habitats where they consume a wide range of plants and animals. These three rodent species were trapped in montane forest for 2.5 years; their stomach contents were analyzed to determine short-term diets (n = 12-95 indiv. per species), and isotopic fractions of d 15 N and d 13C in their bone collagen were analyzed to further estimate their trophic positions (n = 11-20 indiv. per species). For all three species, [75 % of individuals had plants and [90 % had arthropods in their stomachs, and significant differences in mean relative abundances were found for food items in stomachs among all three rodents. Rodents may be dispersing some native and non-native seeds, including the highly invasive Clidemia hirta. Most identifiable arthropods in rodent stomachs were non-native, and no stomachs contained birds, snails, or lizards. The d 15 N and d 13 C signatures were consistent with trophic feeding differences revealed from stomach contents. Dietary niche differentiation by coexisting rodent species is evident in this forest, with Pacific rats being intermediate between the mostly carnivorous house mouse and the mostly herbivorous black rat; such findings can help forecast rodent impacts and direct management efforts in ecosystems where these invasive animals coexist.
Landslides result in the loss of vertical vegetative structure, soil nutrients, and the soil seed bank. These losses impede timely recovery of tropical forest communities. In this study we added bird perches to six Puerto Rican landslides with three types of surfaces (bare, climbing fern, grass) in an effort to facilitate inputs of forest seeds through bird dispersal and to accelerate plant succession. Numbers of bird‐dispersed forest seeds were significantly higher in plots beneath introduced perches than in control plots. Perches did not increase forest seedling densities compared with control plots. Seven different species of birds were observed on introduced perches. Because 99% of the seed inputs to controls and perch plots in the six landslides were wind‐dispersed seeds (mostly graminoids), perches can improve landslide restoration if woody plants establish and shade out the dominant graminoid and climbing fern ground cover. Although increasing seed inputs from forest species is a critical step in accelerating revegetation of landslides, we suggest that supplemental restoration techniques be applied in addition to bird perches to promote forest recovery.
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.
a b s t r a c tTo date, it is not clear which are the factors that most influence tropical forest recovery from hurricanes. Increased canopy openness and increased detritus (debris) deposition are two of the most likely factors, but due to their simultaneous occurrence during a hurricane, their relative effects cannot be separated without a manipulative experiment. Hence, in the Luquillo Experimental Forest (LEF) of Puerto Rico, the Luquillo Long-Term Ecological Research Program (LTER) has undertaken experimental manipulations in replicated 30 Â 30 m plots to simulate the major effects of hurricane disturbance-increased canopy openness and debris addition to the forest floor. Using a factorial experiment enabled investigation of the separate and combined effects of canopy opening and debris on this wet tropical forest; the experimental outcomes may help direct forest management decisions in similar disturbance-prone environments. In this first article of the special issue, we (1) provide details of the design and methodology for this manipulative experiment (the Canopy Trimming Experiment, CTE), (2) report some principal abiotic responses to treatments, and (3) introduce the subject areas of the 12 additional CTE manuscripts in this special issue. The physical conditions created by canopy and understory treatment and the amounts of debris added to CTE plots were similar to the LEF's conditions following Hurricane Hugo (a category 4 storm) in 1989; although more wood and a 37% (1.5 cm) deeper litter layer was present in the CTE. Our selective cutting and removal of the forest canopy above 3 m, which included trimming 234 palm trees and 342 non-palm trees, greatly altered the understory micro-environment by increasing light levels and decreasing litter moisture for 18 months; throughfall and soil moisture were elevated in trim plots for 3 months. In plots where the canopy was trimmed and the debris (6 kg m À2) was added to the forest floor, the canopy debris persisted on the forest floor for at least 4 years; debris decomposed more quickly in plots with intact canopies. The diverse collection of papers in this special issue provide mechanistic understandings of response patterns of tropical forest biota (microbes, plants, animals) and processes (decomposition, herbivory, nutrient cycling, primary production) to canopy and understory disturbance that resembles a major (Pcategory 3) hurricane. Although measurements for this experiment are ongoing to further identify the mechanisms of long-term forest change resulting from hurricanes, we include findings up to the first seven years post-treatment at this time.Published by Elsevier B.V.
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