Small Gap Creation in Belizean Mangrove Forests by a Wood–Boring Insect1

We investigated the role of wood-boring insects in the creation of light gaps within mangrove forests. We compared the frequency of gaps caused by wood borers to other gap-forming processes and characterized the physical attributes of light gaps in mangrove forests on small islands in Belize. Methods of quantifying light gaps included aerial photography, ground surveys, and experimental plots. Small light gaps (≤12 m²) were very common in Rhizophom mangle fringe, comprising almost 22 percent of these forests. Rhizophora mangle gaps were smaller than gaps in Avicennia germinans forests. In R. mangle forests, gaps were caused by branch death, and in A. germinans forests, gaps were caused primarily by downed trees. More than 91 percent of the gap-forming branches and boles in the R. mangle fringe were killed by a wood-boring cerambycid beetle, Elaphidion mimeticum, indicating that it is the major cause of small-scale disturbances in these forests. No trees or branches in the A. germinans forest were attacked by this beetle. In R. mangle forests, small gaps had significantly higher light levels and soil temperatures than areas under the closed canopy; however, soil conditions for sulfide concentrations, porewater salinity, and redox potentials were similar in small gaps and under the closed canopy. Survival of R. mangle, A. germinans, and Laguncularia racemosa seedlings was also higher inside these small gaps, indicating their importance in regeneration of mangrove forests. Feeding by wood borers is thus an important type of indirect herbivory in mangrove forests, with a critical role in ecological processes such as gap dynamics.     

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.     

Seedling survivorship,growth, and response to disturbance in Belizean mangal

Species zonation patterns across tidal gradients in mangrove forests are formed by successful seedling establishment and maintained by replacement of adults by conspecific seedlings. These two processes rarely have been examined experimentally in neotropical mangal. We studied survivorship and growth of seedlings of two species of mangrove, Rhizophora mangle L. and Avicennia germinans (L.) Steam, across a tidal gradient in Belize, Central America. Propagules of each species were planted in common gardens at tidal elevations corresponding to lowest low water (LLW), mean water (MW), and highest high water (HHW). Sixty-nine percent of Rhizophora seedlings planted at MW and 56% of those planted at LLW survived 1 year. Forty-seven percent of MW Avicennia seedlings also survived 1 year. No individuals of either species survived at HHW, and neither did any LLW Avicennia seedlings. Among the surviving Rhizophora seedlings, LLW seedlings grew more rapidly in terms of height, diameter, leaf production, and biomass than did MW seedlings. Insect herbivory was twice as high on MW seedlings as on LLW Rhizophora seedlings. We also examined the response of established Rhizophora seedlings to experimental removal of the adult Rhizophora canopy. Seedlings in canopy removal areas had higher survivorship, grew twice as fast, produced more leaves, and had less than half the herbivory of seedlings growing beneath an intact canopy. These results provide insights into underlying causes and maintenance of zonation in Caribbean mangrove forests.     

Differential predation on mangrove propagules in open and closed canopy forest habitats

Amounts of seed predation by grapsid crabs (Brachyura: Grapsidae) on two species of mangroves (Aegiceras corniculatum and Avicennia marina) were compared among different habitats in an Australian mangrove forest. For Avicennia, comparisons were between canopy gaps and the adjacent forest understory for six, mid intertidal, gaps of different sizes. For Aegiceras the comparisons were among canopy gaps in the high intertidal; open, accreting mud/sand banks where mangroves were colonizing in the low intertidal; and in the forest understory in both the high and low intertidal zones. These were repeated in the high salinity (35\%) downstream portion and the low salinity (0–5\%) upstream portion of a tidal river.Predation on Avicennia was significantly higher in the understory than in adjacent canopy gaps. Within a canopy opening, predation was greatest in the smallest gaps and lowest in the largest gaps. Predation on Aegiceras was greater in the high intertidal compared to the low intertidal, but no differences were found between river mouth and upstream locations. In the high intertidal zone of the forest, there were no differences in predation between canopy gap or forest understory sites for Aegiceras. In the low intertidal zone, however, significant differences in amount of predation were found between habitats. More Aegiceras propagules were consumed in the understory than on adjacent accreting sandbanks.Frequency of tidal inundation, which in turn affects the amount of time available to forage, is hypothesized to account for differences in predation between low and high intertidal forests and between small and large canopy gaps. Our results also suggest that shade intolerance in these two species may actually reflect an escape from predators, successful when the seeds are dispersed into open areas such as canopy gaps or mud banks.     

Gap formation, microsite variation and the conifer seedling occurrence in a subalpine old-growth forest, central Japan

To evaluate the effects of canopy gaps and forest floor microsites (soil, fallen logs, root-mounds, buttresses and stumps) on regeneration of subalpine forests, the gap regeneration and seedling occurrence of conifers (Abies mariesii, Abies veitchii, Picea jezoensis var. hondoensis and Tsuga diversifolia) were studied in two stands of a subalpine old-growth forest, central Japan. The percentage of gap area to total surveyed area was 11.2–11.3% in the stands. Gap regeneration was not common for P. jezoensis var. hondoensis and T. diversifolia. In contrast, gap regeneration by advanced regeneration was relatively common for Abies. Seedling occurrence of P. jezoensis var. hondoensis and T. diversifolia was restricted on elevated surfaces such as stumps and root-mounds, while Abies seedlings could occur on soil as well as on elevated surfaces. Rotten stumps were the most favorable microsites for conifer seedling occurrence, which covered small area in the forest floor. Although canopy gaps were not always favorable for seedling occurrence, all conifer seedlings were larger under canopy gaps than under closed canopy. Canopy gaps and forest floor microsites clearly affected seedling occurrence and growth of conifers. This suggests that regeneration of conifers is related to the difference of growth advantage under canopy gaps and favorable microsites for seedling occurrence.     

Plant-animal interactions and the structure and function of mangrove forest ecosystems*

Trophic interactions involving plants and animals in tropical mangrove forests have important controlling influences on several population, community and ecosystem-level processes. Insect herbivores remove up to 35% of leaf area from some mangrove tree species and can cause the death of seedlings. Leaf chemistry and toughness and soil nutrient status all appear to be important in explaining the between- and among-species variance in leaf damage. Insects also attack and damage, mainly by boring, a large proportion of mangrove seeds. Shadehouse experiments have shown that such post-dispersal predation can have a significant effect on seedling survival, growth and biomass allocation to leaves, stems and roots. Sesarmid crabs are also responsible for severe post-dispersal seed predation. In field trials, crabs consumed more than 70% of the seeds of five tree species. For four of these five species there was an inverse relationship between seed predation rate and the dominance of conspecific adult trees, while the within-site distribution pattern of one tree species appears to be partially controlled by crabs. The same crab species also consume 30–80% (depending on forest type and intertidal elevation), of the annual litter fall in mangrove forests and, thus, have an important role in controlling the rate of remineralization of detritus within forests and the export of particulate matter from the forests to other nearshore habitats. The other major component of litter in the forests is wood, which is broken down relatively rapidly by teredinid molluscs (shipworms). More than 90% of the weight loss from decomposing trunks of Rhizophora species during the first four years of decay is through ingestion by teredinids. The annual turnover of dead wood mass in Rhizophora forests is equivalent to that of the processing of leaf detritus by crabs. Because of the relatively low species richness of trees and consumers in tropical mangrove forests, they are likely to serve as productive sites for further investigations of the influence of plant-animal interactions on the dynamics of tropical forests.     

ECOPHYSIOLOGY OF TROPICAL RHODOPHYTES. I. MICROSCALE ACCLIMATION IN PIGMENTATION1,2

Microscale pigment adjustments to a tropical photosynthetically active radiation and ultraviolet (UV) environment by the intertidal turf algae Ahnfeltiopsis concinna (J. Ag.) Silva et DeCew and Laurencia mcdermidiae (J. Ag) Abbott were promoted by thalli densities that self-shade the under story portions of the same diminutive axes. Tissues of A. concinna from canopy microsites had significantly reduced levels of phycoerythrin, phycocyanin, and allophycocyanin compared to tissues from understory microsites of the same axes. Tissues of L. mcdermidiae from canopy microsites had reduced levels of only phycoerythrin compared to tissues from understory microsites. These alterations coupled with enhanced levels of carotenoid and UV-absorbing compounds in tissues from canopy compared to tissues from understory microsites indicated a pattern of remarkably sensitive photoacclimation over the ≤10-cm axes of these turf-forming rhodophytes. Microscale variation in the in vivo UV absorbance capabilities for turfs of A. concinna and L. mcdermidiae was directly related to the amount of extractable UV-absorbing compounds. An in vivo absorbance signature at ～345 nm appears to provide a method to quickly and accurately gauge the potential UV-shielding capacity of primary producers even at remarkably fine ecological scales. The capacity for highly responsive biochemical adjustments that result in marked canopy–understory distinctions coupled with a turf morphology may be crucial for macroalgal tolerance of physiological stresses associated with tropical intertidal zones. This responsive capacity allows for enhanced photoprotective mechanisms in tissues from canopy microsites while optimizing irradiance capture in deeply shaded tissues from understory microsites < 10 cm away.     

ECOPHYSIOLOGY OF TROPICAL RHODOPHYTES. II. MICROSCALE ACCLIMATION IN PHOTOSYNTHESIS1

Parameters of photosynthesis vs. irradiance curves varied markedly between tissues from microsites along the < 10-cm axes of the tropical intertidal red algae Ahnfeltiopsis concinna (J. Ag.) Silva et DeCew and Laurencia mcdermidiae (J. Ag.) Abbott. Differences in photosynthetic performance between tissues from canopy and understory microsites indicates that L. mcdermidiae exhibited an expected sun-to-shade acclimation but over the space of < 10 cm. Respiration, I_c, I_k, and P_max values were significantly lower in tissues from the understory relative to tissues from the canopy of L. mcdermidiae, while photosystem I (PS I) sizes (PSU I) were significantly higher in tissues from understory microsites. Quantum efficiency was unchanged. Ahnfeltiopsis concinna, in contrast, exhibited higher α in tissues from understory rather than canopy microsites. The values of P_max for tissues from the canopy of A. concinna were not higher than the understory, while PSU O₂(PS II size) of tissues from canopy microsites were unusually higher than those of understory microsites. These characteristics suggest a high degree of irradiance stress in tissues from the canopy of A. concinna, the highest tidal alga in Hawaiian coastal zones. Acclimation to high photosynthetically active radiation and ultraviolet irradiance levels especially in tropical regions appears to be an essential mechanism(s) for stress resistance and persistence of intertidal algae. Algal turfs acclimate at microscales in part fostered by their dense stands that create sharp irradiance gradients as well as adjust physiologically to canopy irradiance levels as mechanisms for optimal photosynthetic performance and stress tolerance.     

Isolation of endophytes from leaves of <Emphasis Type="Italic">Neolitsea sericea</Emphasis> in broadleaf and conifer stands

Endophytic fungi were isolated from leaves of Neolitsea sericea, a major lauraceous tree in the laurel forests of southern Kyushu, collected from the understory layer of broadleaf and conifer stands. Cytosphaera sp. and a species of Ascomycetes in leaf blade segments, plus a xylariaceous species and Phomopsis spp. in petiole segments, were isolated at relatively high frequency. In general, isolation frequencies of endophytes were higher in petiole than blade segments. In blade segments, patterns of endophyte isolation were quite different among stands, while relatively similar in petiole segments. Significant effects of sampling sites or canopy vegetation were rarely detected. the understory layer of laurel forests. Neolitsea sericea is an evergreen broadleaf lauraceous tree, widely distributed in areas of eastern Asia. In southern Kyushu, it is one of the most common trees, growing both as canopy and understory species. In this study, endophytes were isolated from the leaves of N. sericea growing in the understory layer of conifer and broadleaf forest stands to survey the endophytes of N. sericea leaves and to examine the effect of the canopy layer on endophytic mycobiota in understory plants.     

Stand structure influences nekton community composition and provides protection from natural disturbance in Micronesian mangroves

Structurally complex mangrove roots are thought to provide foraging habitat, predation refugia, and typhoon protection for resident fish, shrimp, and crabs. The spatially compact nature of Micronesian mangroves results in model ecosystems to test these ideas. Tidal creek nekton assemblages were compared among mangrove forests impacted by Typhoon Sudal and differing in stand structure. Structurally complex Rhizophora spp. stands were predicted to support higher densities and different communities of nekton and to provide greater protection from typhoons compared to less complex Sonneratia alba/Bruguiera gymnorrhiza stands. Lift net data revealed that structural complexity did not support greater nekton densities, but did support significantly different nekton assemblages. The cardinalfish Apogon ceramensis and goby Oxyurichthys lonchotus had significantly higher densities in S. alba/B. gymnorrhiza mangrove creeks, whereas the silverside Atherinomorus lacunosus and diogenid crabs had significantly higher densities in Rhizophora spp. creeks. Similar nekton densities 17 and 4 months after the typhoon in Rhizophora spp. creeks provided indirect evidence that structural complexity increased protection for resident nekton from disturbances. Findings indicate that studies of structural complexity and nekton densities may be better served when individual species are compared and that diverse mangrove tree assemblages will support diverse nekton assemblages that may be more resilient to disturbance.     

The population biology of Bromus tectorum in forests: effect of disturbance,grazing, and litter on seedling establishment and reproduction

Summary The effect of tree canopy, understory, herbivores, and litter depth on seedling establishment, survival, and reproduction of the alien grass, Bromus tectorum (cheatgrass), was examined in a series of experiments in four forest habitat types in western North America. Higher recruitment, survival, and reproduction on clearcuts, which would be expected if the overstory alone is limiting the distribution of cheatgrass in forests, were not observed. Removing the understory in an otherwise undisturbed Pinus ponderosa forest did, however, increase the emergence of B. tectorum, but plants in these experimentally-created openings were more vulnerable to grazing by small mammals. In contrast, removing the sparse understory in an Abies forest neither enhanced recruitment nor increased the incidence of grazing of B. tectorum seedlings. Regardless of the forest habitat, most grazed plants died before maturity; even fewer grazed plants produced seeds. Litter depth influenced both recruitment and biomass production: both the rate of germination and the size of resultant seedlings were lower on thick litter (6 cm) compared to results on thin litter (1.5 cm). In the more open Pinus ponderosa and Pseudotsuga menziesii forests, cheatgrass colonization may often occur in openings in the understory alone. Colonization in the more shady A. grandis and Thuja plicata forests is unlikely, however, unless the opening extends through both the understory and the overstory. As a result, cheatgrass is unlikely to increase in any of these forests unless the scale and incidence of disturbance increases substantially.     

Cultivation of Non‐timber Forest Products Alters Understory Light Availability in a Humid Tropical Forest in Mexico1

The planting of non‐timber forest products (NTFPs) in the understory of tropical forests is promoted in many regions as a strategy to conserve forested lands and meet the economic needs of rural communities. While the forest canopy is left intact in most understory plantations, much of the midstory and understory vegetation is removed in order to increase light availability for cultivated species. We assessed the extent to which the removal of vegetation in understory plantations of Chamaedorea hooperiana Hodel (Arecaceae) alters understory light conditions. We also examined how any changes in light availability may be reflected by changes in the composition of canopy tree seedlings regenerating in understory plantations. We employed a blocked design consisting of four C. hooperiana plantation sites; each site was paired with an adjacent, unmanaged forest site. Hemispherical canopy photographs were taken and canopy tree seedlings were identified and measured within 12 3 × 2 m randomly placed plots in each site for a total of 96 plots (4 blocks × 2 sites × 12 plots). Plantation management did not affect canopy openness or direct light availability but understory plantations had a higher frequency of plots with greater total and diffuse light availability than unmanaged forest. Comparisons of canopy tree seedling composition between understory plantations and unmanaged forest sites were less conclusive but suggest that management practices have the potential to increase the proportion of shade‐intolerant species of tree seedlings establishing in plantations. Given the importance of advanced regeneration in gap‐phase forest dynamics, these changes may have implications for future patterns of succession in the areas of forest where NTFPs are cultivated.     

Environmental Effects of Canopy Gap Formation in High-Rainfall Mangrove Forests1

This study investigated the importance of gap formation in mangrove swamps on the island of Kosrae, Federated States of Micronesia, in order to understand better both natural processes of forest development and the effects of harvesting trees for firewood in these wetlands. Measurements were concentrated in seven plots located near four rivers: three in fringe zones and four in basin zones. Each plot was a cluster of five points and covered an area of ca 1.3 ha. From every point in each of the seven plots, the nearest canopy gap ≥10 m² was located; 25 of the 35 gaps were formed by harvesting. Porewater salinity was significantly higher under the canopy in fringe mangrove forests than in basin mangrove forests. Although gaps were small (mean gap size = 158 m²; median gap size = 92 m²), soil temperatures were significantly higher in gaps of both zones. Soil redox potential was significantly lower and porewater salinity significantly higher in the gaps than under the canopy in the basin zone only. Higher porewater salinity may be attributed to high evaporation rates from the soil and high transpiration rates from trees surrounding gaps. There were significantly more seedlings in gaps than under the canopy only in the fringe zones. Although gap formation alters the soil environment of Kosraean mangrove swamps, high freshwater input may buffer these effects in basin mangrove swamps by reducing porewater salinity. Current harvesting rates do not appear to be changing canopy species composition, but large gaps, especially in mangrove forests in more arid areas, may lead to major changes.     

Light limitation creates patchy distribution of an invasive grass in eastern deciduous forests

Species interactions and their indirect effects on the availability and distribution of resources have been considered strong determinants of community structure in many different ecological systems. In deciduous forests, the presence of overstory trees and shrubs creates a shifting mosaic of resources for understory plants, with implications for their distribution and abundance. Determination of the ultimate resource constraints on understory vegetation may aid management of these systems that have become increasingly susceptible to invasions by non-native plants. Microstegium vimineum (Japanese grass) is an invasive annual grass that has spread rapidly throughout the understory of forests across the eastern United States since it was first observed in Tennessee in 1919. M. vimineum occurs as extensive, dense patches in the understory of eastern deciduous forests, yet these patches often exhibit sharp boundaries and distinct gaps in cover. One example of this distributional pattern was observed relative to the native midstory tree Asimina triloba (pawpaw), whereby dense M. vimineum cover stopped abruptly at the drip line of the A. triloba patch and was absent beneath the A. triloba canopy. We conducted field and greenhouse experiments to test several hypotheses regarding the causes of this observed pattern of M. vimineum distribution, including allelopathy, seed dispersal, light limitations, and soil moisture, texture, and nutrient content. We concluded that light reduction by the A. triloba canopy was the environmental constraint that prevented establishment of M. vimineum beneath this tree. Whereas overstory tree canopy apparently facilitates the establishment of this shade-tolerant grass, the interaction of overstory canopy with midstory canopy interferes with M. vimineum by reducing the availability of sunflecks at the ground layer. It is likely that other midstory species influence the distribution and abundance of other herb-layer species, with implications for management of understory invasive plant species.     

The abundance and growth of Avicennia marina and Rhizophora stylosa in the low shore zone of Darwin Harbour,Northern Territory

Abstract The growth and distribution of the two mangrove seedling species, Avicennia marina (Forsk.) Vierh. and Rhizophora stylosa Griff., were studied in the low-shore zone at three locations along two estuaries of Darwin Harbour to determine how growth varied along the estuary and within the habitats of the low shore zone. The low shore environment was heterogenous for factors such as salinity and light, with substantial variation within and between locations. Seedlings of Avicennia were in higher densities in light gaps than in the adjacent forest, and density in both habitats decreased towards the mouth of the estuary. Growth of Avicennia seedlings, as measured through in situ growth trials and plant harvests, increased towards the mouth of the estuary. Results for Rhizophora seedlings were more complex although some similar trends were evident. This may have been due to the greater early survival of Rhizophora seedlings compared to Avicennia seedlings. The importance of propagule origin was investigated, for Rhizophora, by transplanting propagules between locations along one estuary. Increased seedling survival was observed at the propagule source site; however, this effect was subtle and probably not the most important factor affecting seedling survival at a site.     

Holocene mangrove dynamics and sea-level changes in northern Brazil, inferences from the Taperebal core in northeastern Pará State

A 450 cm sediment core from Taperebal, in the mangrove region of northeastern Pará State in northern Brazil has been studied through pollen analysis in order to reconstruct mangrove development and dynamics and to infer relative sea-level (RSL) changes during the Holocene. Six AMS radiocarbon dates, which provide a somewhat limited age control with some uncertainties, suggest early and late Holocene deposits interrupted by a hiatus between them. A patchy vegetation of coastal Amazon rain forest, restinga, salt marsh and some mangrove, which was dominated by Avicennia, covered the study area during the early Holocene period. The occurrence of an early Avicennia dominated mangrove phase has not been reported so far from other sites in northern Brazil. During the mid Holocene mangroves mostly replaced the former coastal Amazon rain forest, restinga and some salt marsh vegetation, reflecting the rise in the RSL. Rhizophora trees expanded markedly and Avicennia became rare. In the sediment core there is apparently a gap between the depths of 115 and 85 cm (possibly starting between 5900 and 5750 b.p.). The deposits above 85 cm are of modern age and were probably deposited during the last decades. This gap can be explained by the lowering of the RSL as is shown for other northern Brazilian coastal sites. The deposition of sediments during the last decades suggests that the modern RSL is high compared to other periods in the Holocene. Pollen data from these deposits show that Rhizophora trees dominate the mangrove forests, also indicating a high RSL.     

Combined Effects of Host Plant Quality and Predation on a Tropical Lepidopteran: A Comparison between Treefall Gaps and the Understory in Panama

In tropical forests, light‐gaps created from treefalls are a frequent source of habitat heterogeneity. The increase in productivity, through gap formation, can alter food quality, predation and their impact on insect herbivores. We hypothesized that in gaps, herbivores would be less resource‐limited and more predator limited, whereas in the understory, we predicted the reverse. In this study, we investigate the combined effects of food quality and predation on the lepidopteran larva Zunacetha annulata feeding on its host plant Hybanthus prunifolius in two habitats; sunny treefall gaps and the shaded understory in Panama. In bioassays, Z. annulata feeding on sun leaves ate 22 percent less leaf area, grew 25 percent faster, and had higher pupal weights than larvae feeding on shade leaves. However, shade leaves had higher nitrogen content and specific leaf area. In gaps, predation was 26.4 percent compared to 13.8 percent in the understory. Larvae on understory plants traveled greater distances and spent more time searching and traveling than larvae on gap plants. These differences in behavior are consistent with lower predation risk and lower quality food in the understory. Using data from bioassays and field experiments we calculated 0.22 percent and 1.02 percent survival to adulthood for larvae in gaps and the understory, respectively. In conclusion, although these habitats were in close proximity, we found that larvae in the understory are more resource‐limited and larvae in gaps are more predator limited.     

Conservation value of timber quality versus associated non-timber quality stands for understory diversity in Nothofagus forests

Conservation strategies of forested landscapes must consider biodiversity of the included site types, i.e. timber-quality forests and associated non-timber-quality stands. The objectives were to characterize forest overstory structure in timber-quality versus associated non-timber-quality stands; and to compare their understory communities. Six forest types were sampled in Nothofagus forests of Tierra del Fuego (Argentina): two timber-quality N. pumilio forests, and four associated non-timber-quality stands (edge, N. antarctica, wetlands and streamside forests). Overstory structure and understory vegetation (species richness, frequencies, cover and biomass) were characterized during spring and summer seasons. Analysis of variance and multivariates were carried out. Overstory structure differed across the site types, with higher tree size, canopy closure and tree volume in timber-quality stands. Fifty-one understory plant species were observed, but understory variables varied with site types, especially wetlands (highest native and exotic richness, cover and biomass, and 25% of exclusive species). Site types were grouped in three: N. antarctica stands, streamside stands and the other N. pumilio forests according to multivariate analysis. Forty three percent of plants were distributed in all site types, and all timber-quality forest understory species were present in some associated non-timber-quality stands. Timber-quality N. pumilio forests have a marginal value for understory conservation compared to associated non-timber-quality stands, because these last include all the plants observed in timber-quality forests and also possess many exclusive species. Therefore, protection of associated non-timber-quality stands during forest management planning could increase understory conservation at landscape level, and these could be better reserves of understory diversity than retentions of timber-quality stands.     

Loss of oak dominance in dry-mesic deciduous forests predicted by gap capture methods

Gap capture methods predict future forest canopy species composition from the tallest trees growing in canopy gaps rather than from random samples of shaded understory trees. We used gap capture methods and a simulation approach to forecast canopy composition in three old oak forests (Quercus spp.) on dry-mesic sites in southern Wisconsin, USA. In the simulation, a gap sapling is considered successful if it exceeds a threshold height of 13–17 m (height of maximum crown width of canopy trees) before its crown center can be overtopped by lateral crown growth of mature trees. The composition of both the tallest gap trees and simulated gap captures suggests that 68–90% of the next generation of canopy trees in the stands will consist of non-Quercus species, particularly Ulmus rubra, Carya ovata and Prunus serotina. Quercus species will probably remain as a lesser stand component, with Quercus alba and Quercus rubra predicted to comprise about 19% of successful gap trees across the three stands. Several methods of predicting future canopy composition gave similar results, probably because no gap opportunist species were present in these stands and there was an even distribution of species among height strata in gaps. Gap trees of competing species already average 11–13 m tall, and mean expected time for these trees to reach full canopy height is only 19 years. For these reasons, we suggest that dominance will shift from oaks to other species, even though late successional species (e.g., Acer and Tilia) are not presently common in the understories of these stands.     

When sex is not enough: ecological correlates of resprouting capacity in congeneric tropical forest shrubs

In moist tropical forests resprouting may be an important component of life history, contributing to asexual reproduction through the clonal spread of individuals derived from shoot fragments. However, in contrast to other ecosystems where resprouting is common, the ecological correlates of resprouting capacity in tropical forests remain largely unexplored. In this study we characterized shade tolerance, resprouting capacity and sexual reproductive success of eight co-occurring Piper species from lowland forests of Panama. In field experiments we found that shade-tolerant Piper species had a higher capacity to regenerate from excised or pinned stem fragments than light-demanding species in both gap and understory light conditions. In contrast, shade-tolerant species had lower recruitment probabilities from seeds, as a consequence of lower initial seed viability, and lower seedling emergence rates. All Piper species needed gap conditions for successful seedling establishment. Of 8,000 seeds sown in the understory only 0.2% emerged. In gaps, seed germination of light-demanding species was between 10 and 50%, whereas for shade-tolerant species it was 0.5–9.8%. We propose that the capacity to reproduce asexually from resprouts could be adaptive for shade-tolerant species that are constantly exposed to damage from falling litter in the understory. Resprouting may allow Piper populations to persist and spread despite the high rate of pre-dispersal seed predation and low seed emergence rates. Across Piper species, we detected a trade-off between resprouting capacity and the annual viable seed production per plant but not with annual seed mass produced per plant. This suggests that species differences in sexual reproductive success may not necessarily result from differential resource allocation. Instead we suggest that low sexual reproductive success in the understory may in part reflect reduced genetic diversity in populations undergoing clonal growth, resulting in self-fertilization and in-breeding depression.