Effects of Hurricanes on Rare Plant Demography in Fire‐Controlled Ecosystems

Hurricanes have dramatic effects on forest vegetation, but their effects on shrublands have rarely been studied. We analyzed the effects of three 2004 hurricanes—among the strongest on record in Florida—on vital rates of 12 rare plant species of pyrogenic interior Florida scrub and sandhill. Tree damage varied by vegetation type (being highest in areas with Pinus clausa) and was associated with debris deposition. Most rare species were minimally impacted by hurricanes. The two most frequently damaged species were the shrubs Prunus geniculata (11% of individuals) and Asimina obovata (7%); both were resilient to damage. Prunus geniculata had little mortality during the hurricane year but damaged plants had a temporary (1‐yr) reduction in relative growth rate. Prunus geniculata flowering was unaffected by hurricane damage. Hurricane damage had no effects on vital rates of A. obovata, Eriogonum longifolium var. gnaphalifolium, or Chrysopsis highlandsensis. Other species suffered little or no observable hurricane damage. Of 12 species analyzed, nine had similar annual survival in hurricane and nonhurricane years. Relatively low survival in the hurricane year (compared with other years) was linked to prehurricane drought or prescribed fire in two of three species. Thus, the 2004 hurricanes did not have important effects on populations of interior Florida scrub and sandhill plants, especially herbaceous species. This is in marked contrast to dramatic demographic responses to fire in central Florida and strong effects of hurricanes in coastal Florida, highlighting that these different disturbances may have divergent effects on vegetation and populations over short distances.     

Species-specific Seedling Responses to Hurricane Disturbance in a Puerto Rican Rain Forest1

Seedling dynamics were followed in a Puerto Rican forest for 20 months following a severe hurricane to study the interactive effects of hurricane debris, nutrients, and light on seedling diversity, density, growth, and mortality. Three treatments (debris removal, an unaltered control with hurricane debris, and chemical fertilization added to hurricane debris) altered levels of forest debris and soil nutrients. Canopy openness was measured twice using hemispherical photographs of the canopy. We examined the demographic responses of six common species to treatments over time. Seedling densities increased for all six species but the only significant treatment effects were increased densities of the pioneer tree Cecropia and the shrub Palicourea in the debris removal treatment. Seedling growth declined with declining light levels for four species but not for the pioneer tree Alchornea or the non‐pioneer tree Dacryodes. Only Cecropia and the non‐pioneer tree Chionanthus had treatment effects on growth. Mortality also differed among species and tended to be highest in the fertilized plots for all but Cecropia and Dacryodes. We found only some of the expected differences between pioneer and non‐pioneer plants, as each species had a unique response to the patchy distributions of organic debris, nutrients, and light following the hurricane. High local species diversity was maintained through the individualistic responses of seedlings after a disturbance.     

Linking responses of native and invasive plants to hurricane disturbances: implications for coastal plant community structure

Hurricane disturbances produce significant changes in forest microclimates, creating opportunities for seedling regeneration of native and invasive plant species alike. However, there is limited information on how changes in microclimates and pre-existing forest conditions affect native and invasive plants responses to hurricane disturbances. In this manipulative study, we examined the responses of three common shrub/small stature tree species, two of which are native to the coastal region of the southeastern USA (Baccharis halimifolia and Morella cerifera) and one that is invasive (Triadica sebifera), to two key components of hurricane disturbance (canopy damage and saline storm surge). In a greenhouse, we grew seedlings of these species under a range of shade levels that mimicked pre-and post-hurricane canopy conditions for wet pine forest and mixed hardwood forest, two forest communities common in coastal areas of the southeastern USA. Seedlings were subjected to saline storm surges equivalent to full strength sea water for 3 days. Seedling responses (mortality and growth) to the treatments were monitored for 16 months. All species benefitted from higher canopy openness. Storm surge effects were short-lived and seedlings readily recovered under high light conditions. The storm surge had stronger negative effects on survival and growth of all species when coupled with high shade, suggesting storm surge has greater negative impacts on seedlings where hurricane winds cause minimal or no canopy damage. The invasive T. sebifera was by far more shade tolerant than the natives. Survival of T. sebifera seedlings under highly shaded conditions may provide it a competitive edge over native species during community reassembly following tropical storms. Differential responses of native and invasive species to hurricane disturbances will have profound consequences on community structure across coastal forest stands, and may be regulated by legacies of prior disturbances, community structure, extent of canopy damage, and species’ tolerance to specific microclimates.

     

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.     

Early post-hurricane stand development in Fringe mangrove forests of contrasting productivity

We examined the immediate effects of a hurricane (Hurricane Andrew, August 1992) in a coastal landscape in sub-tropical Florida, and then monitored stand recovery in Fringe mangrove sites of different productive capacity for 9 years after the disturbance. Structural impacts of the hurricane were confined almost entirely to forests within 200–300 m of the coast. Mortality and damage were concentrated on canopy individuals. Following the hurricane, rapid canopy recovery and the early onset of competition among Fringe forest stems, as evidenced by relatively high mortality of smaller individuals, magnified the initial dominance of hurricane survivors and early-established seedlings over later cohorts, and limited recruitment to the brief period prior to canopy closure. Changes in the relative abundance of the two dominant mangrove species following disturbance varied strongly along the productivity gradient. The shade-tolerant Rhizophora mangle L. generally became the overwhelming canopy dominant in the competitive environment of the recovering Coastal Fringe forest following hurricane, but the shade-intolerant Laguncularia racemosa (L.) C.F. Gaertn was better represented in less productive Interior Fringe sites, where canopy closure was delayed. Site productivity is an important determinant of the success of mangrove species during post-hurricane stand development, and consequently of the zonation of communities in the coastal landscape.     

Seedling and Sapling Dynamics of Treefall Pits in Puerto Rico1

Seedling and sapling dynamics in a Puerto Rican rain forest were compared between forest understory and soil pits created by the uprooting of 27 trees during Hurricane Hugo. Soil N and P, organic matter, and soil moisture were lower and bulk densities were higher in the disturbed mineral soils of the pits than in undisturbed forest soils ten months after the hurricane. No differences in N and P levels were found in pit or forest soils under two trees with N–fixing symbionts (Inga laurina and Ormosia krugii) compared to soils under a tree species without N–fixing sym–bionts (Casearia arborea), but other soil variables (Al, Fe, K) did vary by tree species. Forest plots had greater species richness of seedlings (<10 cm tall) and saplings (10–100 cm tall) than plots in the soil pits (and greater sapling densities), but seedling densities were similar between plot types. Species richness and seedling densities did not vary among plots associated with the three tree species, but some saplings were more abundant under trees of the same species. Pit size did not affect species richness or seedling and sapling densities. Recruitment of young Cecropia schreberiana trees (>5 m tall) 45 months after the hurricane was entirely from the soil pits, with no tree recruitment from forest plots. Larger soil pits had more tree recruitment than smaller pits. Defoliation of the forest by the hurricane created a large but temporary increase in light availability. Recruitment of C. schreberiana to the canopy occurred in gaps created by the treefall pits that had lower soil nutrients but provided a longer–term increase in light availability. Treefall pits also significantly altered the recruitment and mortality of many understory species in the Puerto Rican rain forest but did not alter species richness.     

Ecosystem recovery in terra firme forests after cutting and burning: a comparison on species richness, floristic composition and forest structure in the Jaú National Park, Amazonia

Six hectares, three in a primary forest and three in a 40 year old secondary forest were inventoried for all trees with Diameter at Breast Height (DNH) of 10 cm or greater in a terra firme forest 200 km north-east of Manaus, central Amazonia in order to compare the difference between structure, species richness and floristic composition. Both species richness and tree density were significantly higher in the upland forest than in the secondary forest. The forest structure pattern analysed (DBH, basal area and estimated dry biomass) did not differ significantly between the two forest types. Similarity indices at species level were only 14%. In the 3 ha of primary forest the number of species varied from 137 to 159, the number of individuals from 639 to 713, total basal area from 32.8 to 40.2 m² and estimate total of above-ground dry biomass (AGBM) from 405 to 560 tons per ha. In the 3 ha of secondary forest, the number of species varied from 86 to 90, the number of individuals from 611 to 653, total basal area from 28.8 to 39.9 m² and estimated total AGBM from 340 to 586 tons per ha. Family Importance Value (FIV) is the sum of relative density, dominance and richness of a family. The most important families in relation to FIV were Burseraceae, Chrysobalanaceae, Lecythidaceae, Myristicaceae, Bombacaceae, Fabaceae and Mimosaceae in the 3 ha of primary forest, while Burseraceae, Lecythidaceae, Sapotaceae, Arecaceae and Cecropiaceae were the most important families in the 3 ha of secondary forest. Importance Value Index (IVI) is the sum of relative density, dominance and frequency of a species. Alexa grandiflora (Caesalpiniaceae), Sckronema micranthum (Bombacaceae) and Pourouma guianensis (Cecropiaceae) were the most important species in relation IVI, in the primary forest, while Eschweilera grandiflora (Lecythidaceae), Protium apiculatum (Burseraceae) and Bertholletia excelsa (Lecythidaceae) were the most important species in the secondary forest. We conclude that species richness was significandy different between the two forests, but that forest structure patterns analysed in this study (DBH, basal area and dry biomass) were similar. This demonstrates that 40 years was sufficient time for the secondary forest to recover the original structure of the primary forest, but not the original species richness. The low species similarity between the two forests indicates that the floristic composition was quite distinct and that the mixture of primary forest and disturbed forest has led to an increase in total species diversity.     

Growth and mortality patterns in a thinning canopy of post-hurricane regenerating rain forest in eastern Nicaragua (1990-2005)

One of the strongest hypothesis about the maintenance of tree species diversity in tropical areas is disturbance. In order to assess this, the effect of intensive natural disturbances on forest growth and mortality in a thinning canopy was studied after the landfall of hurricane Joan in 1988. We evaluated the growth and mortality rates of the 26 most common tree species of that forest in eastern Nicaragua. Permanent plots were established at two study sites within the damaged area. Growth and mortality rates of all individual trees > or = 3.18cm diameter at breast height were assessed annually from 1990 to 2005. During this period the forest underwent two phases: the building phase (marked by increased number of individuals of tree species present after the hurricane) and the canopy thinning phase (marked by increased competition and mortality). Our results from the thinning phase show that tree survival was independent of species identity and was positively related to the increase in growth rates. The analysis of mortality presented here aims to test the null hypothesis that individual trees die independently of their species identity. These findings were influenced by the mortality observed during the late thinning phase (2003-2005) and provide evidence in favor of a non-niche hypothesis at the thinning phase of forest regeneration.     

Simulated hurricane‐induced changes in light and nutrient regimes change seedling performance in Everglades forest‐dominant species

Wind damage from cyclones can devastate the forest canopy, altering environmental conditions in the understory that affect seedling growth and plant community regeneration. To investigate the impact of hurricane‐induced increases in light and soil nutrients as a result of canopy defoliation, we conducted a two‐way factorial light and nutrient manipulation in a shadehouse experiment. We measured seedling growth of the dominant canopy species in the four Everglades forest communities: pine rocklands (Pinus elliottii var densa), cypress domes (Taxodium distichum), hardwood hammocks, and tree islands (Quercus virginiana and Bursera simaruba). Light levels were full sun and 50% shade, and nutrient levels coupled with an additional set of individuals that were subjected to a treatment mimicking the sudden effects of canopy opening from hurricane‐induced defoliation and the corresponding nutrient pulse. Seedlings were measured weekly for height growth and photosynthesis, with seedlings being harvested after 16 weeks for biomass, leaf area, and leaf tissue N and ¹³C isotope ratio. Growth rates and biomass accumulation responded more to differences in soil nutrients than differences in light availability, with largest individuals being in the high nutrient treatments. For B. simaruba and P. elliottii, the highest photosynthetic rates occurred in the high light, high nutrient treatment, while T. distichum and Q. virginiana photosynthetic rates were highest in low light, high nutrient treatment. Tissue biomass allocation patterns remained similar across treatments, except for Q. virginiana, which altered above‐ and belowground biomass allocation to increase capture of limiting soil and light resources. In response to the hurricane simulation treatment, height growth increased rapidly for Q. virginiana and B. simaruba, with nonsignificant increases for the other two species. We show here that ultimately, hurricane‐adapted, tropical species may be more likely to recolonize the forest canopy following a large‐scale hurricane disturbance.     

Dendrochronology of lianas of the Leguminosae family from the Atlantic Forest, Brazil

Information about plant growth, development and age forms the basis for understanding complex forest ecological processes. Although lianas play an important role in tropical forests, little is known about their growth and development from either climatic or ecological perspectives. Therefore, we studied the growth rings in Legume liana species collected in a mountainous Atlantic Forest in southeastern Brazil. Four of the eight studied species did not show cambial variants, three had a lobed stem, and one had a furrowed xylem. Distinct growth rings were observed in all species. Semi-ring porosity, marginal parenchyma, fibrous zone and radially flattened latewood cells were the main characteristic features of these growth rings. Species without cambial variants, including Dalbergia frutescens, Piptadenia adiantoides, P. micracantha and Senegalia tenuifolia, showed very distinct growth rings visible in macroscopic and microscopic analysis. Ring-width time series and cambial wound assessment were performed to analyze periodicity and dendrochronology. The species with cambial variants, S. grandistipula, S. lacerans, S. martiusiana and S. pedicellata, also showed distinct growth rings, however, sometimes barely detectable or not detected at all. Cambial wounding, cross-dating and climate-growth relationships indicated the annual nature of growth rings in species without cambial variant. Cross-dating between radii within one individual and between individuals was successful, and the synchronized series enabled us to build species chronologies and a mean chronology. Climate-growth analysis revealed significant correlations between chronologies and precipitation, indicating that available moisture is the main factor determining growth rates of lianas in the Atlantic forest.     

Responses of the mangrove fern <Emphasis Type="Italic">Acrostichum danaeifolium</Emphasis> Langsd. &; Fisch. (Pteridaceae,Pteridophyta) to disturbances resulting from increased soil salinity and Hurricane Georges at the Jobos Bay National Estuarine Research Reserve,Puerto Rico

Mortality of plants in a mangrove forest dominated by the tree Avicennia germinans began to appear at the southern coastal edge of the forest in the central watershed of the Jobos Bay National Estuarine Research Reserve in Puerto Rico in 1977. The dieback was attributed to salinities in excess of 70 ppt and gradually continued north until some tree mortality was observed in 1995 at the forest’s landward interface with agricultural land. Mortality of the common understory mangrove fern Acrostichum danaeifolium occurred at a much slower rate than tree mortality. Remediation at the forest’s edge was recommended in 1997 and a study to monitor A. danaeifolium at 4-month intervals was begun in January 1998 to determine how the ferns would respond to hydrologic change. Hurricane Georges passed over the reserve in September 1998, so the effects of this disturbance were assessed as well. In May 1998, 4 months after tagging individual plants, the mean number of leaves plant⁻¹ at the impact site was only half that for plants in a control area in the nearby undisturbed eastern watershed. Fertile (spore-bearing) leaf production in both watersheds was very low accounting for plant fertility of 7% or less. Short leaf lengths in the impact site combined with smaller plant leaf counts and much lower leaf production rates in one area of the impact site had resulted in biomass production at the control site that was 18 times greater than in the impact site. After Hurricane Georges plant mortality in the impact area increased with individuals of A. danaeifolium found dead on every monitoring date except September 1999. However, when means for all variables measured in both sections of the forest were compared within 8 months after the hurricane there was an increase in all growth characteristics except sterile leaf length. Relative plant fertility increased from 10% before the hurricane to over 50% by May 2000. The fertile leaf production rate of these plants increased nine-fold by May 1999, and though it declined slowly thereafter it was still almost three time pre-hurricane levels at the end of the study almost 2 years later. Biomass production rates, though very low in the 4 months after the hurricane, had increased to five times pre-hurricane levels by the end of the study in January 2001. Efforts at remediation of the hypersaline conditions that were completed in May 1999 at the landward edge of the impact area appeared to have little effect on the ferns. For A. danaeifolium, plant mortality and decreases in plant size characteristics were the most notable responses to high salinity levels. In contrast, increases in fertile leaf and biomass production rates were the most significant response to hurricane disturbance.     

Demography of the long‐lived conifer Agathis ovata in maquis and rainforest,New Caledonia

Abstract. The endemic New Caledonian conifer Agathis ovata occurs as an emergent tree in fire‐prone shrublands (maquis), and fire‐sensitive rainforest. Growth, survivorship and recruitment over 5 yr were compared for populations from forest and maquis on ultramafic substrates in New Caledonia to investigate whether demographic behaviour varied in response to the strongly contrasting forest and shrubland environments. Growth of seedlings and of small (30–100 cm height) and large (100 cm height; 5 cm DBH) saplings was slow, but varied significantly among stages, site types and years. The greatest difference in growth rates was among stages, seedlings growing 0.34 cm.yr^?1, small saplings 1.06 cm.yr^?1 and large saplings 2.13 cm.yr^?1. Tree DBH increased by only 0.05 cm.yr^?1 and, based on these rates, individuals with DBH of 30 cm are estimated to be more than 700 yr old. Few trees (3.5%) produced cones in any year and seedling recruitment was low, but some recruitment was recorded each year in both maquis and forest. Rates of recruitment per parent were highest in forest (1.28.yr^?1, cf 0.78.yr^?1), but the higher density of trees in maquis meant that overall recruitment was greater there (92 ha^?1.yr^?1, cf 56 ha^?1.yr^?1). Seedling mortality ranged from 0.9 to 2.9% among years with no significant difference between maquis and forest. No sapling mortality was recorded, but annual tree mortality ranged from 0 to 1.4%. Evidence from a recently burned site indicated that while trees may survive fire, seedlings and saplings do not. Post‐fire seedling recruitment per ha from surviving trees was four times lower than in unburned sites, but growth rates were four times higher. Similar demographic attributes, including high survivorship, low growth rate and low rates of recruitment over a long reproductive life, characterize Agathis ovata populations in both maquis and rainforest in New Caledonia and are indicative of a broad tolerance of light environments that is unusual among tree species. These demographic attributes help to explain the long‐term persistence of the species in these strongly contrasting habitats.     

Intra- and inter-specific variation in canopy photosynthesis in a mixed deciduous forest

 Within the same forest, photosynthesis can vary greatly among species and within an individual tree. Quantifying the magnitude of variation in leaf-level photosynthesis in a forest canopy will improve our understanding of and ability to model forest carbon cycling. This information requires extensive sampling of photosynthesis in the canopy. We used a 22-m-tall, four-wheel-drive aerial lift to reach five to ten leaves from the tops of numerous individuals of several species of temperate deciduous trees in central Massachusetts. The goals of this study were to measure light-saturated photosynthesis in co-occurring canopy tree species under field conditions, and to identify sampling schemes appropriate for canopy tree studies with challenging logistics. Photosynthesis differed significantly among species. Even though all leaves measured were canopy-top, sun-acclimated foliage, the more shade-tolerant species tended to have lower light-saturated photosynthetic rates (P _max) than the shade-intolerant species. Likewise, leaf mass per area (LMA) and nitrogen content (N) varied significantly between species. With only one exception, the shade-tolerant species tended to have lower nitrogen content on an area basis than the intolerant species, although the LMA did not differ systematically between these ecological types. Light-saturated P _max rates and nitrogen content, both calculated on either an area or a mass basis, and the leaf mass to area ratio, significantly differed not only among species, but also among individuals within species (P<0.0001 for both). Differences among species accounted for a greater proportion of variance in the P _max rates and the nitrogen content than the differences among individuals within a species (58.5–78.8% of the total variance for the measured parameters was attributed to species-level differences versus 5.5–17.4% of the variance was attributed to differences between individual trees of a given species). Furthermore, more variation is accounted for by differences among leaves in a single individual tree, than by differences among individual trees of a given species (10.7–30.4% versus 5.5–17.4%). This result allows us to compare species-level photosynthesis, even if the sample size of the number of trees is low. This is important because studies of canopy-level photosynthesis are often limited by the difficulty of canopy access. As an alternative to direct canopy access measurements of photosynthesis, it would be useful to find an ”easy-to-measure” proxy for light-saturated photosynthetic rates to facilitate modeling forest carbon cycling. Across all species in this study, the strongest correlation was between nitrogen content expressed on an area basis (mmol m^–2, N _area) and light-saturated P _max rate (μmol m^–2 s^–1, P _maxarea) (r ²=0.511). However, within a given species, leaf nitrogen was not tightly correlated with photosynthesis. Our sampling design minimized intra-specific leaf-level variation (i.e., leaves were taken only from the top of the canopy and at only one point in the season). This implies that easy-to-measure trends in nitrogen content of leaves may be used to predict the species-specific light-saturated P _max rates. Received: 16 March 1996 / Accepted: 16 August 1996     

Long-Term Growth and Succession in Restored and Natural Mangrove Forests in Southwestern Florida

We compared colonization, growth and succession from 1989 to 2000 in a restored mangrove site and in gap and closed canopy sites in a natural mangrove forest. The restored site was created in 1982 and planted with Rhizophora mangle (≈2 m⁻²) propagules. By 1989, Laguncularia racemosa, with densities up to 12.9 tree m⁻², was a dominant in all plots, although densities were greater at edge plots relative to inner plots, and near open water (west plots) relative to further inland (east plots), and in tall mangrove plots relative to scrub plots. Rhizophora mangle (1989 tree densities about 2 m⁻²) was a codominant in inner and scrub plots, while Avicennia germinans had the lowest densities (<1 tree m⁻²) in all plots. From 1989 to 2000 L. racemosa experienced reduced recruitment and apparent density-dependent mortality of canopy individuals in plots with high initial densities. Scrub plots experienced high rates of colonization by R. mangle and L. racemosa, rapid growth in height of all species (1989–1996), followed by a dieoff of L. racemosa in later years (1997–2000) as the canopy came to resemble that of tall mangrove plots. Colonization and growth rates were lower in gap and closed canopy regions of the natural forest relative to rates in the restored site. After 11 years, densities of L. racemosa were 10–20× lower and R. mangle slightly less in the gap relative to densities in tall mangrove plots in the restored site at the same age. Although the restored stand had converged with the natural forest by 2000 in terms of some factors such as species richness, vegetation cover, litterfall, and light penetration, trees were still much smaller and stem densities much higher. Full development of mature structure and ecological function will likely require decades more development.     

High Mortality for Rare Species Following Hurricane Disturbance in the Southern Yucatán

Hurricanes are an important part of the natural disturbance regime of the Yucatán Peninsula with the potential to alter forest structure and composition, yet investigations of species‐level responses to severe winds are limited in this region. The effect of a category 5 hurricane (Hurricane Dean, 21 August 2007) on dry tropical forests across the southern Yucatán was examined with respect to tree damage, mortality, and sprouting. Damage was assessed 9–11 mo following the hurricane in 92 (500 m²) plots stratified by wind speed and normalized difference vegetation index (NDVI) change classes over a 25,000 km² study area. We investigated the relative importance of biotic (i.e., species, size, and wood density) and abiotic (i.e., wind speed) factors to better explain patterns of damage. Overall mortality was low (3.9%), however, mortality of less common species (8.5%) was elevated more than fourfold above that of 28 common species (1.8%), indicating immediate selective consequences for community composition. Species varied in the degree and type of damage experienced, with susceptibility increasing with tree diameter and height. Wood density influenced damage patterns only in areas where a critical threshold in storm intensity was exceeded (wind speeds ≥210 km/h). Although overall, damage severity increased with wind speed, common coastal species were more resistant to damage than species distributed farther inland. Our findings suggest that selective pressure exerted by frequent hurricane disturbance has, and will, continue to impact the floristic composition of forests on the Yucatán Peninsula, favoring certain wind‐resistant species. Abstract in Spanish is available at http://www.blackwell‐synergy.com/loi/btp .     

Changes in forest bird abundance,community structure and composition following a hurricane in Sweden

The effect that extreme natural events have on biological diversity is relatively poorly known. We used a before–after control‐impact (BACI) design to analyze changes in bird abundances and communities following Hurricane Gudrun, which struck southern Sweden in January 2005, felling 75 million m³ of forest and causing damage to 5% of forested areas (half a million hectares) in a few hours. We used recent measures of impact in combination with classical BACI contrasts to analyze bird count data from a monitoring program in Sweden. We investigated changes in the abundance of 17 species commonly found in forests, along with changes in species composition and functional structure of the bird community. In total, we considered 34 response variables and examined whether responses were immediate or long‐term. There was no evidence of a strong effect of the hurricane on the abundances of six species. Estimates of the effects on five species were too uncertain to draw inferences. We detected positive and negative effects of the hurricane on the abundances of the remaining six species, but the magnitude of effects often was small. Generally, the effects were in the expected direction: negative on birds associated with mature forest and positive on birds associated with open land or young forest. We found evidence of an increase in the proportion of species that nest on the ground and a decrease in the proportion of species that nest in cavities and trees. In contrast, the hurricane had no discernible effect on functional measures of diversity (richness, evenness or divergence), or on communities’ reproductive or morphological characteristics. Our results suggest that the hurricane affected bird populations and communities, but the magnitude of effects was generally small.     

Influence of Forest Type and Tree Species on Canopy‐Dwelling Beetles in Budongo Forest,Uganda1

Beetles were collected on 64 trees of four species (Cynometra alexandri C. H. Wright, Rinorea beniemis (Welwitsch ex Olivier) Kuntze, Teclea nobilis Delile, and Trichilia rubescens Olivier) in Budongo Forest, Uganda, using an insecticidal fogging technique. Selected tree species were abundant, taxonomically not closely related, and different in the shape of leaves, growth form, and size, with heights between 7 and 35 m. Trees were fogged in an old primary forest stand, in an area of secondary forest where selective logging was performed, and in a swamp forest. Eight conspecific trees per forest type were fogged. A total of 29,736 beetles were collected from all trees that could be assigned to 1433 (morpho)‐species; 41.6 percent were singletons and 89.6 percent of species were found with less than ten individuals. Abundant beetle taxa included Latridiidae (N= 4093), Chrysomelidae (3952), Staphylinidae (2931), Apioninae (2621), and Curculionidae (2457). Most species‐rich groups were Staphylinidae (N= 196 spp.), Curculionidae (189), and Chrysomelidae (148). Abundance increased in the order: primary < secondary < swamp forest. Due to the relatively high dominance of some species in the secondary forest, species richness increased in the order: secondary < primary < swamp forest. Beta diversity measures and factor analysis showed distinct differences among forest types but higher similarity of beetle communities on different tree species within one forest type. The taxonomic distribution of beetles in the secondary forest was more heterogeneous than in the primary forest. Analyses of the data revealed low host specificity even for phytophagous beetles, underlining the importance of habitat structure and chance effects on the spatial distribution of beetles in the canopy of Budongo Forest.     

Land use history, hurricane disturbance, and the fate of introduced species in a subtropical wet forest in Puerto Rico

Tropical forests are suffering from increasing intensities and frequency of disturbances. As a result, non-native species accidentally introduced or intentionally planted for farming, plantations, and ornamental purposes may spread and potentially invade undisturbed native forest. It is not known if these introduced species will become invasive, as a result of recurrent natural disturbances such as hurricanes. Using data from three censuses (spanning 15 years) of a 16-ha subtropical wet forest plot, we investigated the impact of two hurricanes on populations of plant species that were planted in farms and plantations that were then abandoned and from the natural spread of species introduced into Puerto Rico in the past. The populations of four species (Citrus paradis, Mangifera indica, Musa sp., and Simarouba glauca) changed little over time. Six species (Artocarpus altilis, Calophyllum calaba, Genipa americana, Hibiscus pernambucensis, Syzygium jambos, and Swietenia macrophylla) declined between the first two censuses after Hurricane Hugo, then increased again in Census 3 after Hurricane Georges. Spathodea campanulata gradually increased from census to census, while Coffea arabica declined. These introduced species represent only a small part of the forest basal area and few show signs of increasing over time. The number of stems per plant, new recruits, and the growth rates of these introduced species were within the ranges of those for native plant species. The mortality rates over both census intervals were significantly lower for introduced species (<5% year⁻¹) than for native ones (15% year⁻¹). Many new recruits established after Hurricane Hugo (prior to this study) had opened the forest canopy and these high mortality rates reflect their death as the canopy recovered. Only Swietenia macrophylla and Syzygium jambos showed an increase in stem numbers in the closed canopy area of forest that had suffered limited human disturbance in the past. A future increase in frequency of disturbance may enable greater stem numbers of introduced species to establish, while lower-mortality rates compared to native species, may allow them to persist during inter-hurricane intervals. An increase in the population of introduced species, especially for those that grow into large trees, may have an impact on this tropical forest in the future.     

Regeneration in fringe mangrove forests damaged by Hurricane Andrew

Mangrove forests along many tropical coastlines are frequently andseverely damaged by hurricanes. The ability of mangrove forests to regeneratefollowing hurricanes has been noted, but changes that occur in vegetationfollowing disturbance by hurricane winds and storm tides have not been studied.We measured changes in plant community structure and environmental variables intwo fringe mangrove forests in south Florida, USA that experienced high windvelocities and storm tides associated with Hurricane Andrew (August1992). Loss of the forest canopy stimulated regeneration via seedlinggrowth and recruitment, as well as resprouting of some trees that survived thehurricane. Initial regeneration differed among species in both forests:Rhizophora mangle L. regenerated primarily via growth ofseedlings present at the time of the hurricane (i.e., release of advancerecruits), but many trees of Avicennia germinans(L.) Stearn and Laguncularia racemosa Gaertn.f.resprouted profusely from dormant epicormic buds. In one forest, which wasformerly dominated by Laguncularia, high densities ofRhizophora seedlings survived the hurricane and grew toform dense stands of saplings and small trees ofRhizophora. In the other forest, there were lowerdensitiesof surviving Rhizophora seedlings (possibly due tohigher storm tide), and extensive bare areas that were colonized byAvicennia, Laguncularia, andherbaceous species. This forest, predominantly Rhizophoraat the time of the hurricane, now contains stands of saplings and small treesofall three species, interspersed with patches dominated by herbaceous plants.These findings indicate that moderately damaged fringe forests may regenerateprimarily via release of Rhizophora advance recruits,leading to single-species stands. In severely damaged forests, seedlingrecruitment may be more important and lead to mixed-species stands.Regeneration of mangrove forests following hurricanes can involve differentpathways produced by complex interactions between resprouting capability,seedling survival, post-hurricane seedling recruitment, and colonizationby herbaceous vegetation. These differences in relative importance ofregeneration pathways, which may result in post-hurricane forestsdifferent from their pre-hurricane structure, suggest that models forregeneration of mangrove forests will be more complex than directregeneration models proposed for other tropical forests whereregeneration after hurricanes is dominated by resprouting.     

Differences in endemic insect assemblages among vegetation types on a small island of the oceanic Ogasawara Islands

Natural vegetation is often replaced by invasive alien plants on isolated oceanic islands. To determine how invasive alien plants affect insect diversity, we compared flying insects captured using Malaise traps among different vegetation types on a small island (Nishijima; 0.49 km²) in the oceanic Ogasawara (Bonin) Islands in the north‐western Pacific. The numbers of individuals and species, and the species composition of pollinators (bees), predators (wasps) and wood borers (cerambycid, mordellid and elaterid beetles) were compared among three vegetation types: Casuarina equisetifolia (an invasive alien tree) forest, natural forest and natural grassland (forest edge), during two seasons (June and October–November 2005). In traps, 80.0, 66.7, 87.5, 85.7 and 100.0% of bee, wasp, cerambycid, mordellid and elaterid beetle species, respectively, were endemic to the Ogasawara Islands. Grassland had the highest wasp and bee species richness, whereas natural forest had the highest species richness of wood‐boring beetles. The C. equisetifolia forest had the poorest species richness for most insect groups (except mordellid beetles). More individuals of most insect groups (except bees) were captured in June than in October–November. More individual bees and wasps were captured in grassland than in forests, whereas more individual mordellid and elaterid beetles were captured in forests than in grassland. The number of cerambycid individuals did not differ among vegetation types. Redundancy analysis suggested that most insect species preferred natural forest or grassland to alien forest. Therefore, further invasion of natural grassland and forest by the alien tree C. equisetifolia may negatively affect the endemic insect fauna of Nishijima.