Development of Tree Regeneration in Fern-dominated Forest Understories After Reduction of Deer Browsing

Dennstaedtia punctilobula (hay‐scented fern) can act as a native invasive species in forests in eastern North America where prolonged deer browsing occurs in stands with partially open overstory canopies. Ferns dominate the understory with a 60‐cm tall canopy, with little regeneration of native tree species. It has been hypothesized that, once established, ferns may continue to inhibit tree regeneration after deer browsing has been reduced. To test this hypothesis, we documented the pattern of recovery of the tree seedling understory in plantations of Pinus strobus (white pine) and Pinus resinosa (red pine) on the Quabbin Reservation watershed protection forest in central Massachusetts, where after 40 years of intensive deer browsing the deer herd was rapidly reduced through controlled hunting. Dense fern understories occur on nearly 4,000 ha of the predominantly oak–pine forest. Three years after deer herd reduction, stands with the highest density fern cover (77% of plots with>90% cover) had significantly fewer seedlings at least 30 cm in height, compared with stands with lower fern density, and those seedlings consisted almost entirely of Betula lenta (black birch) and white pine. Height growth analysis showed that black birch and white pine grew above the height of the fern canopy in 3 and 6 years, respectively. In contrast, two common species, Fraxinus americana (white ash) and Quercus rubra (red oak), grew beneath the dense fern cover for 5 years with height growth less than 5 cm/yr after the first year. A study of spring phenology indicated that the ability of black birch to grow through the fern canopy might have been due to its early leaf development in spring before the fern canopy was formed, in contrast to oak and ash with delayed leaf development. Thus, the ferns showed differential interference among species with seedling development after reduction of deer browse.     

Long‐term tree fern dynamics linked to disturbance and shade tolerance

Question: Do New Zealand tree ferns have recognizable shade tolerance niches? Location: Lowland temperate rain forest of New Zealand (41°20′S, 174°58′E). Methods: Growth, death and recruitment of five tree fern species were estimated from a 38‐year record of stem heights, collected within a 2.25‐ha block of forest, and electron transport rates (ETR) of photosystem II of fronds were measured. Results: Two species of Cyathea were comparatively common (603 and 351 stems in total) and two were comparatively rare (155 and 17 stems in total) on the site. The common species had lower rates of growth, recruitment and mortality than the rare species, had skewed age distributions typical of shade‐tolerant species and were probably recruited soon after a catastrophic earthquake in 1855. The two rare species were failing to recruit under closed forests; their age distributions indicated that all had regenerated long after the earthquake. ETR were higher for faster‐growing than for the shade‐tolerant species. A tree fern that regenerates vegetatively from aerial buds, Dicksonia squarrosa, was common on the site (361 stems in total). Its age distribution suggested it was relatively shade tolerant, but its mortality and recruitment rates were much higher than those of the two shade‐tolerating Cyathea species, suggesting that this multi‐stemmed species functions differently from the monopodial Cyathea species. Conclusions: New Zealand Cyathea tree ferns occupy distinct niches along a shade tolerance spectrum and their relative abundances are strongly influenced by disturbance history. The study provides evidence that tree fern species differ strongly in their responses to canopy disturbance and are not ecologically equivalent.     

Soil conservation in Polylepis mountain forests of Central Argentina: Is livestock reducing our natural capital?

Mountain forests and their soils provide ecological services such as maintenance of biodiversity, provision of clean water, carbon capture and forage for livestock rearing, which is one of the principal economic activities in mountain areas. However, surprisingly little is known about livestock impact in South American mountain forest soils. With the aim of understanding how livestock and topography influence patterns of forest cover, soil compaction, soil loss and soil chemical properties, we analysed these parameters in 100 Polylepis australis woodland plots situated in the humid subtropical mountains of Central Argentina. We used distance from the nearest ranch as an objective index of historical livestock impact and measured standard topographic variables. Our main results reveal that distance from ranch in all cases partly explains tree canopy cover, soil loss, soil compaction and soil chemical properties; suggesting a strong negative effect of livestock. Intermediate altitudes had more tree canopy cover, while landscape roughness – a measure of the variability in slope inclination and aspect – was negatively associated to soil impedance and acidity, and positively associated to soil organic matter content. Finally, flatter areas were more acid. We conclude that livestock has had a substantial influence on forest soil degradation in the Mountains of Central Argentina and possibly other similar South American mountains. Soil degradation should be incorporated into decision making when considering long‐term forest sustainability, or when taking into account retaining livestock for biodiversity conservation reasons. Where soil loss and degradation are ongoing, we recommend drastic reductions in livestock density.     

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.     

Post‐fire succession and 20th century reduction in fire frequency on xeric southern Appalachian sites

Abstract. We document post‐fire succession on xeric sites in the southern Appalachian Mountains, USA and assess effects of 20th century reduction in fire frequency on vegetation structure and composition. Successional studies over 18 yr on permanent plots that had burned in 1976–1977 indicate that tree mortality and vegetation response varied with fuel load and fire season. In the first three years after fire, hardwood sprouts dominated tree regeneration. On sites where summer and autumn fires reduced litter depth to less than 1 cm, densities of shade‐intolerant Pinus seedlings increased steadily over this period. 4 to 8 yr after fire, large numbers of newly established seedlings and sprouts had grown to 1 – 10 cm DBH. By year 18 growth of these saplings led to canopy closure on most sites. Herbaceous cover and richness peaked in the first decade after fire, then declined. On similar sites that had not burned in more than 50 yr, regeneration of shade‐intolerant Pinus spp. and mean cover and richness of herbs were considerably lower than those observed on recently burned plots. Reconstructions of landscape conditions based on observed post‐fire succession and 20th century changes in fire regime suggest that reductions in fire frequency circa 1940 led to substantial changes in forest structure and decreases in cover and richness of herbaceous species.     

Interactive effects of introduced herbivores and post‐flowering die‐off of bamboos in Patagonian Nothofagus forests

Question: In November 2000, Chusquea culeou, a bamboo species dominating Andean forest understories in southern Argentina and Chile, massively flowered and died over a north‐south distance of ca. 120 km. Because bamboo is the major forage for large herbivores in these forests, we examined the interactive influences of the bamboo die‐off and herbivory by introduced cattle on understory and tree regeneration. Location: Lanín National Park, Argentina. Methods: Permanent plots, in and outside livestock exclosures, were installed in a Nothofagus dombeyi forest in patches of flowered and non‐flowered C. culeou. Plots were monitored over four years for changes in understory composition and tree seedling densities and heights. Results: After the C. culeou die‐off, new establishment of N. dombeyi was low, both with and without herbivory. Livestock alone directly increased N. dombeyi seedling mortality through physical damage. However, tree seedling browse ratings and height growth were interactively affected by bamboo flowering and herbivory; unfenced plots in flowered bamboo patches had the shortest seedlings, highest browse ratings, and lowest tree seedling annual growth rates. Understory cover was higher where livestock were excluded, and this effect was intensified in the patches of flowered bamboo. Neither herbivory nor bamboo flowering resulted in major changes in species composition, with the exception of Alstroemeria aurea. Conclusion: Effects of livestock on N. dombeyi regeneration were contingent on flowering of C. culeou. Prior to introduction of livestock, N. dombeyi regeneration was probably successful beneath canopy gaps during windows of opportunity following bamboo die‐off, but now livestock impede tree regeneration. Herbivory during bamboo withering periods also produces more open understories, particularly affecting palatable heliophyllous herb species such as Alstroemeria aurea. The results underscore the importance of assessing herbivore impacts on tree regeneration during relatively short periods of potential tree regeneration immediately following rare bamboo flowering and die‐off.     

Gap-phase dynamics and coexistence of a long-lived pioneer and shade-tolerant tree species in the canopy of an old-growth coastal temperate rain forest of Chiloé Island, Chile

Aim A major question with regard to the ecology of temperate rain forests in south‐central Chile is how pioneer and shade‐tolerant tree species coexist in old‐growth forests. We explored the correspondence between tree regeneration dynamics and life‐history traits to explain the coexistence of these two functional types in stands apparently representing a non‐equilibrium mixture. Location This study was conducted in northern Chiloé Island, Chile (41.6° S, 73.9° W) in a temperate coastal rain forest with no evidence of stand disruption by human impact. Methods We assessed stand structure by sampling all stems within two 50 × 20 m and four 5 × 100 m plots. A 600‐m long transect, with 20 uniformly spaced sampling points, was used to quantify seedling and sapling densities, obtain increment cores, and randomly select 10 tree‐fall gaps. We used tree‐ring analysis to assess establishment periods and to relate the influences of disturbances to the regeneration dynamics of the main canopy species. Results Canopy emergent tree species were the long‐lived pioneer Eucryphia cordifolia and the shade‐tolerant Aextoxicon punctatum. Shade‐tolerant species such as Laureliopsis philippiana and several species of Myrtaceae occupied the main canopy. The stem diameter distribution for E. cordifolia was distinctly unimodal, while for A. punctatum it was multi‐modal, with all age classes represented. Myrtaceae accounted for most of the small trees. Most tree seedlings and saplings occurred beneath canopy gaps. Based on tree‐ring counts, the largest individuals of A. punctatum and E. cordifolia had minimum ages estimated to be > 350 years and > 286 years, respectively. Shade‐tolerant Myrtaceae species and L. philippiana had shorter life spans (< 200 years). Most growth releases, regardless of tree species, were moderate and have occurred continuously since 1750. Main conclusions We suggest that this coastal forest has remained largely free of stand‐disrupting disturbances for at least 450 years, without substantial changes in canopy composition. Release patterns are consistent with this hypothesis and suggest that the disturbance regime is dominated by individual tree‐fall gaps, with sporadic multiple tree falls. Long life spans, maximum height and differences in shade tolerance provide a basis for understanding the long‐term coexistence of pioneer and shade‐tolerant tree species in this coastal, old‐growth rain forest, despite the rarity of major disturbances.     

Birds,Cattle, and Bracken Ferns: Bird Community Responses to a Neotropical Landscape Shaped by Cattle Grazing Activities

Large areas of tropical moist forests have been converted to cattle pastures, generating complex landscapes where different habitats are represented by small patches with an uneven spatial distribution. Here, we describe how bird communities respond to the different elements present in a livestock landscape that was originally dominated by tropical moist forest. We surveyed six habitats: open pastures, pastures with shrubs, early‐ and middle‐secondary forests, mature forest, and pastures invaded by bracken ferns (Pteridium aquilinum). Bird diversity was high in secondary and mature forests, and low in fern‐invaded sites and open pastures. Fern‐dominated sites had the lowest bird species richness, and trophic guild diversity of all habitats. Habitat structure affected both bird species richness and densities in similar ways. Tree species richness was the habitat attribute that had a bigger positive effect on bird species richness. Bird community structure varied among sampled habitats, separating habitats in two major groups (forests and pastures). Our data indicate that bracken fern‐invaded pastures were the worst habitat condition for avian communities. To increase bird diversity, we recommend to eliminate or manage bracken fern and to increase shrub and tree cover in open pastures to provide food resources and shelter for birds. Finally, we encourage the maintenance of secondary and mature forest remnants as a strategy to conserve resident birds within a landscape dominated by livestock activities.     

Cover and growth habit of Polylepis woodlands and shrublands in the mountains of central Argentina: human or environmental influence?

Aim To determine whether the cover and growth habit of the main forest forming species (Polylepis australis BITT.) in a mountain range with low human population density is mainly affected by anthropogenic activities or by environmental influences. Location Central Argentina. Methods Using GIS and field surveys we established 146 plots of 30 × 30 m located in five river basins differing in human impact. We measured P. australis cover, growth habit of each individual (number of basal ramifications), index of long term human impact (percentage of rock exposed by soil erosion due to livestock and fires), evidence of logging, fire scars, local relief, percentage of rock outcrops and altitude above sea level. We analysed the influence of independent variables on P. australis cover and growth habit (average number of basal ramifications per plot) using correlations and General Linear Models. Results Polylepis australis cover was greater at intermediate altitudes above sea level and in areas with reduced long term human impact. Contrastingly local relief, percentage of rock outcrops and logging in the recent past did not have a major influence on P. australis abundance. Growth habit varied in complex patterns. Individuals with fewer ramifications were found in valley bottoms and more disturbed basins, while more ramifications were found at mid‐ and upper slopes and well preserved basins. In valley bottoms, ramifications decreased with increasing altitude whereas the opposite trend was observed for mid‐slopes. Ramifications were positively related to fires in two river basins and in mid‐ and upper slopes but not in valley bottoms. Fire impact was always less in valley bottoms than at mid‐ and upper slopes. Main conclusions Human impact had a major role on P. australis cover, while growth habit was determined by complex combinations of potentially cumulative natural and anthropogenic factors. Even in sparsely populated mountains, both human impact and their interaction with natural environmental gradients influence plant communities and need to be understood for effective management.     

The effects of foraging by the superb lyrebird (Menura novae-hollandiae) in Eucalyptus regnans forests at Beenak,Victoria

Abstract In an early spar-stage stand of Eucalyptus regnans at Beenak, Victoria, foraging by lyrebirds in bare floor areas on steep slopes results in a complex microtopography of excavations, accumulations and terracettes. About 200 t ha^?1 of litter and top soil may be displaced an average of 70 cm downhill per year. Magnetic ferruginous pisolite was used as a marker to monitor progressive soil movement over 3 years. Very little disturbance occurred in areas of dense ground fern, but in bare areas the whole forest floor may be turned over every 20 months. In the site studied, foraging activity by lyrebirds varied seasonally and topographically. Disturbance by other biotic agents was minimal. The mean depth of soil cultivation was about 10 cm and litter was frequently buried or mixed intimately with soil. Since buried leaf litter decays more quickly than that on the surface, lyrebird foraging is likely to increase the rate of nutrient cycling. The small, steep clifflets left at the uphill limits of each scratch microsite provide litter-free niches for the establishment of tree fern prothalli and shade-tolerant herbs. All stages in the growth of the rough tree fern, Cyathea australis, were present in bare floor areas, but in dense ground fern patches, young stages were confined to rotten logs and upturned root balls. Since dense tree fern development tends to diminish the cover of dense ground fern, lyrebird foraging activity may maintain an accessible food resource which would otherwise diminish with increased ground fern cover in these forests in the course of secondary succession after fire.     

Do we need livestock grazing to promote <Emphasis Type="Italic">Polylepis australis</Emphasis> tree recruitment in the Central Argentinean Mountains?

South American Polylepis mountain forests are endangered due to centuries of logging, fire clearance, and intensive livestock grazing. Polylepis australis (Rosaceae) dominates the tree canopy in the Central Argentinean Mountains, where livestock have been excluded or reduced to very low densities over wide areas. Lowered plant and bird diversity in these abandoned grazing areas suggest that some livestock grazing may be beneficial for conservation purposes. Our study aimed to determine whether this may also be true for forest regeneration. In 300 plots of 1 m² distributed in sub-regions of high to moderate grazing pressure as well as grazing exclusion, we recorded the presence of P. australis seedlings and grazing indicators. Topographical parameters as well as vegetation and soil characteristics were also assessed. Seedling frequencies were highest in the region with moderate grazing, intermediate with grazing exclusion, and the lowest with high grazing pressures. Logistic regression models revealed that seedling occurrence was highest close to seeder trees, with intermediate litter coverage and at intermediate altitudes above sea level. Since grazing ultimately affects seeder tree occurrence through browsing and reduces litter cover and predominates in higher areas, we conclude that grazing is an important driving factor for P. australis regeneration. We confirm previous studies determining high grazing pressure to be detrimental for P. australis regeneration, but also provide data suggesting that its complete exclusion may not be recommendable either. Management should aim to alternate between a moderate grazing intensity to promote seedling recruitment, with subsequent exclusion to accelerate the growth of the established seedlings.     

Leaf structure of Syzygium spp. (Myrtaceae) in relation to site affinity within a tropical rain forest

This study examined four species of Syzygium (S. firmum, S. makul, S. operculatum, S. rubicundum) Myrtaceae, a tree genus that dominates the canopy of rain forests of south‐west Sri Lanka. Syzygium spp. occupy differing habitats with relation to succession and forest topography. We examined differences in leaf morphology and physiology in response to amount of shade, an important environmental variable affecting Syzygium distribution within the forest. To study change in leaf structure and physiology, environmental shelters were constructed simulating forest shade that differed in quality, quantity and duration. Seedlings were exposed to: (i) 0% shade (full sun, FS), red : far red (R : FR) ratio 1.27; (ii) 65% shade (large opening, LO) with direct sunlight similar to the centre of a large canopy opening, R : FR ratio 1.27; (iii) 82% shade (small opening, SO) with direct sunlight similar to the centre of a small canopy opening, R : FR ratio 1.27; (iv) 58% uniform light shade (LS) with a quality similar to the outside edge of a large canopy opening, R : FR ratio 1.05; (v) 85% uniform medium shade (MS) with a quality similar to the inside forest edge of a large canopy opening, R : FR ratio 0.97; (vi) 99% uniform deep shade (DS) similar to that of the forest understorey, R : FR ratio 0.23. The shelters were constructed in a large open area at the field station of the Sinharaja World Heritage site, Sri Lanka. Seedlings of each species were grown for two years in their respective shade treatments before physiological, morphological and anatomical measurements were made on leaves. Variation in leaf structure and physiology between the species was associated with differences in shade‐tolerance and water‐use. All species increased in photosynthesis rates and dimensions in leaf structure (leaf blade and cuticle thickness, stomatal density, thickness of upper and lower epidermis, and thickness of palisade mesophyll) with decrease in shade. In contrast, stomatal conductivity was highest in the DS (99% shade) treatment. Leaves of Syzygium firmum were thickest and largest in area. S. firmum also had highest photosynthesis in the SO (82% shade) treatment. S. firmum was the most shade‐tolerant of all species: it grows well in low shade and its leaf structure suggests it to be the most conservative in water‐use of the Syzygium spp. In the forest S. firmum can persist in the forest shade as established seedlings, but grows best within canopy openings of late‐seral rain forest. Leaves of S. operculatum were thinnest but had highest stomatal densities of the four species. S. operculatum is considered shade‐intolerant, with a leaf structure suggesting it to be prone to desiccation, and by implication susceptible to drought. S. operculatum is found along streams within early seral rain forest habitat, often originating on stream banks after land clearance for cultivation. In the FS (0% shade) treatment, S. rubicundun had highest photosynthesis rates and greatest number of leaves but smallest leaf area of the Syzygium species. S. rubicundum is more shade‐intolerant but more efficient in water‐use than S. operculatum. S. rubicundum is a mid‐seral canopy tree of the midslope stands that are thought to have originated after catastrophic windthrows or swidden cultivation. The leaf physiology and structure of S. makul suggests it to be both moderately shade‐tolerant and conservative in water‐use. It is the most widely distributed Syzygium species across the topography of late‐seral rain forest. We suggest forest disturbance and hydrology are important environmental factors that influence distribution of Syzygium species across the topography. Results from this study contribute to a body of knowledge suggesting that canopy tree species of rain forests in south‐west Sri Lanka have discrete affinities to topography and differences in successional status, and that adaptations in leaf structure and physiology are indicative of such phenomena. © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141 , 365–377.     

Mushroom diversity in sustainable shade tea forest and the effect of fire damage

A survey of the biodiversity of wild macrofungi, including edible species yields, was carried out from 1 May to 30 September 2007 at four different forest types (in mainly Miang tea forest). The plots 100 m², comprised a tea garden with a few planted canopy tree species (37.2% canopy cover), a cultivated sustainable tea forest (80.2% canopy cover), an abandoned sustainable tea forest (89.8% canopy cover), and an abandoned sustainable tea forest that had suffered fire damage (72.9% canopy cover). All visible mushrooms were collected during weekly forays. Macro-characters of the fungi were annotated and photographed by digital camera and the fungi were identified to genera and morphospecies (e.g. Agaricus sp.1, Agaricus sp.2). The fresh weight of wild edible mushrooms produced in the plots was also recorded during this period. The biodiversity of macrofungi in abandoned sustainable tea forest was highest with 115 species in 47 genera, followed by cultivated sustainable tea forest with 85 species in 42 genera and fire damaged abandoned sustainable tea forest with 48 species in 25 genera, while only 19 species belonging to 15 genera were found in the tea garden. Twenty-one species belonging to nine genera with a total of 60.9 kg of edible fungi were collected throughout this study. The fresh weight of edible mushrooms was recorded with the highest fresh weight (25.3 kg) collected from the abandoned sustainable tea forest, followed by 18.2 kg in the cultivated sustainable tea forest, while in the fire damaged sustainable tea forest 16.4 kg were collected and only 1 kg was collected in the tea garden. Abandoned sustainable tea forest contained a greater shade tree biodiversity and higher canopy cover than other plots. This area has a generally higher humidity, a greater ground litter cover, a greater number of microhabitats, a greater number of trees that can form mycorrhizal associations and this probably accounts for the higher diversity of macrofungi and production of edible mushrooms. The fire damaged sustainable tea forest had lower mushroom diversity and edible mushroom production possibly due to the loss of litter and lower tree diversity. The tea garden supported a poor diversity of mushrooms and almost no edible mushrooms. Shade tea forest (Miang tea forest) is a good method to produce tea in a sustainable way as it maintains diversity of mushrooms and other organisms and could be developed as an alternative to shade coffee.     

Differential survival among life stages contributes to co‐dominance of Abies mariesii and Abies veitchii in a sub‐alpine old‐growth forest

Questions: Are there interspecific differences in mortality and recruitment rates across life stages between two shade‐tolerant dominant trees in a sub‐alpine old‐growth forest? Do such differences in demography contribute to the coexistence and co‐dominance of the two species? Location: Sub‐alpine, old‐growth forest on Mt. Ontake, central Honshu, Japan. Methods: From 1980 to 2005, we recorded DBH and status (alive or dead) of all Abies mariesii and A. veitchii individuals (DBH ≥ 5 cm) in a 0.44‐ha plot. Based on this 25 year census, we quantified mortality and recruitment rates of the two species in three life stages (small tree, 5 cm ≤ DBH < 10 cm; subcanopy tree, 10 cm ≤ DBH < 20 cm; canopy tree, DBH ≥ 20 cm). Results: Significant interspecific differences in mortality and recruitment rates were observed in both the small tree and sub‐canopy tree stages. In this forest, saplings (< 5 cm DBH) are mostly buried by snow‐pack during winter. As a consequence, saplings of A. mariesii, which is snow and shade tolerant, show higher rates of recruitment into the small tree stage than do those of A. veitchii. Above the snow‐pack, trees must tolerate dry, cold temperatures. A. veitchii, which can more readily endure such climate conditions, showed lower mortality rate at the subcanopy stage and a higher recruitment rate into the canopy tree stage. This differential mortality and recruitment among life‐stages determines relative dominance of the two species in the canopy. Conclusion: Differential growth conditions along a vertical gradient in this old forest determine survival of the two species prior to reaching the canopy, and consequently allow co‐dominance at the canopy stage.     

Fruit biomass availability along a forest cover gradient

Habitat loss is the main driver of the current high rate of species extinction, particularly in tropical forests. Understanding the factors associated with biodiversity loss, such as the extinction of species interactions and ecological functions, is an urgent priority. Here, our aim was to evaluate how landscape‐scale forest cover influences fruit biomass comparing different tree functional groups. We sampled 20 forest fragments located within landscapes with forest cover ranging from 2 to 93 percent in the Atlantic forest of southern Bahia, Brazil. In each fragment, we established five plots of 25 × 4 m and carried out phenological observations on fleshy fruit throughout 1 year on all trees ≥5 cm dbh. We estimated fruit availability by direct counting of all fruits and derived fruit biomass from this count. We used spatial mixed linear models to evaluate the effects of forest cover on species richness, abundance, and fruit biomass. Our results indicated that forest cover was the main explanatory variable and negatively influenced the total richness and abundance of zoochoric and shade‐tolerant but not shade‐intolerant species. A linear model best explained variations in richness and abundance of total and shade‐tolerant species. We also found that forest cover was positively correlated with the fruit biomass produced by all species and by the shade‐tolerant assemblages, with linear models best explaining both relationships. The loss of shade‐tolerant species and the lower fruit production in fragments with lower landscape‐scale forest cover may have implications for the maintenance of frugivore, seed dispersal service, and plant recruitment.     

Influence of within‐patch habitat quality on high‐Andean Polylepis bird abundance

Habitat restoration strategies for fragmented high Andean forest landscapes must consider the influence of within‐patch habitat quality on bird abundance. I examined vegetation and bird abundance at three locations within a highly fragmented Polylepis forest landscape in the Cordillera Vilcanota, southern Peru. Across the landscape, there was significant variation in the vegetation structure of Polylepis forest patches of different size categories, especially in terms of tree girth, tree height, tree density, and canopy vegetation structure. Principal Component Analysis extracted five factors of habitat quality, which together accounted for 74.2% of the variability within 15 habitat variables. Polylepis bird species differed in their responses to habitat quality but, overall, variation in Polylepis bird abundance was not fully captured by the range of habitat quality variables. Tall, dense vegetation cover was clearly important for 11 conservation‐important species, a high density of large trees was important for 10 species and primary forest ground cover was important for eight species. Habitat quality exhibited no significant influence on the abundance of only one species –Asthenes urubambensis. The abundance of seven species was associated with lower elevation forest, but only one species was associated with higher elevation forest. Management of habitat quality in large and medium remnant forest patches throughout the Cordillera Vilcanota, particularly in the 3800–4200 m elevation range, will be a cornerstone in ensuring the persistence of the majority of conservation‐important bird species populations.     

Tall shrub and tree expansion in Siberian tundra ecotones since the 1960s

Circumpolar expansion of tall shrubs and trees into Arctic tundra is widely thought to be occurring as a result of recent climate warming, but little quantitative evidence exists for northern Siberia, which encompasses the world's largest forest‐tundra ecotonal belt. We quantified changes in tall shrub and tree canopy cover in 11, widely distributed Siberian ecotonal landscapes by comparing very high‐resolution photography from the Cold War‐era ‘Gambit’ and ‘Corona’ satellite surveillance systems (1965–1969) with modern imagery. We also analyzed within‐landscape patterns of vegetation change to evaluate the susceptibility of different landscape components to tall shrub and tree increase. The total cover of tall shrubs and trees increased in nine of 11 ecotones. In northwest Siberia, alder (Alnus) shrubland cover increased 5.3–25.9% in five ecotones. In Taymyr and Yakutia, larch (Larix) cover increased 3.0–6.7% within three ecotones, but declined 16.8% at a fourth ecotone due to thaw of ice‐rich permafrost. In Chukotka, the total cover of alder and dwarf pine (Pinus) increased 6.1% within one ecotone and was little changed at a second ecotone. Within most landscapes, shrub and tree increase was linked to specific geomorphic settings, especially those with active disturbance regimes such as permafrost patterned‐ground, floodplains, and colluvial hillslopes. Mean summer temperatures increased at most ecotones since the mid‐1960s, but rates of shrub and tree canopy cover expansion were not strongly correlated with temperature trends and were better correlated with mean annual precipitation. We conclude that shrub and tree cover is increasing in tundra ecotones across most of northern Siberia, but rates of increase vary widely regionally and at the landscape scale. Our results indicate that extensive changes can occur within decades in moist, shrub‐dominated ecotones, as in northwest Siberia, while changes are likely to occur much more slowly in the highly continental, larch‐dominated ecotones of central and eastern Siberia.     

The complexity of forest borders determines the understorey vegetation

Questions

What are the most important drivers of plant species richness (gamma‐diversity) and species turnover (beta‐diversity) in the field layer of a forest edge? Does the tree and shrub species richness structure and complexity affect the richness of forest and grassland specialist species?

Location

Southeast Sweden.

Methods

We sampled 50 forest edges with different levels of structural complexity in agricultural landscapes. In each border we recorded trees, shrubs and herb layer species in a 50‐m transect parallel with the forest. We investigated species composition and species turnover in relation to the proportions of gaps in the border and the diversity of trees and shrubs.

Results

Total plant species richness in the field layer was mainly explained by the proportion of gaps to areas with full canopy cover and tree diversity. Increasing number of gaps promoted higher diversity of grassland specialist species within the field layer, resulting in open forest borders with the highest overall species richness. Gaps did however have a negative impact on forest species richness. Conversely, increasing forest species richness was positively related to tree diversity, but the number of grassland specialist species was negatively affected by tree diversity.

Conclusions

Managing forest borders, and therefore increasing the area of semi‐open habitats in fragmented agricultural landscapes, provides future opportunities to create a network of suitable habitats for both grassland and deciduous forest specialist species. Such measures therefore have the potential to increase functional connectivity and support dispersal of species in homogeneous forest/agricultural landscapes.     

Tree invasion in managed tropical forests facilitates endemic species

Aim To determine whether different abundances of introduced species of Cinchona (Rubiaceae) affect species composition and facilitate species richness in managed tropical forests, to test whether any facilitative effects on understorey species depend on forest type, and to investigate whether facilitative effects can be attributed to the ‘substitutive facilitation model’. Location Makawao Forest Reserve on Maui, Hawai’i, USA. Methods Cinchona species (Cinchona pubescens and Cinchona calisaya) were mapped within various forest types. In three forest types (ageing Eucalyptus and Pinus plantations, and near‐natural Acacia koa forests), we analysed environmental parameters (e.g. canopy cover, litter cover, pH value and soil depth) and the species composition of Cinchona‐invaded and non‐invaded plots; data were compared based on Cinchona cover and forest types. Habitat modelling for several endemic species and tree ferns was carried out to test whether Cinchona cover is an important variable for the probability of occurrence of these endemics. Results Cinchona species have naturalized mainly in Eucalyptus and Pinus plantations and Acacia koa forests and here add an additional shrub layer. In contrast to other studies, we revealed facilitative effects of Cinchona on native species within all forest types. Species richness is about 20% higher in invaded plots than in non‐invaded plots, and these show a nearly 50% higher proportion of endemic species, including tree ferns. The proportion of endemics even increases with increasing Cinchona cover. For several endemics, Cinchona is found to be an important variable for the probability of occurrence, and the removal of Cinchona cover as an explanatory variable lowers the model fit. In addition to Cinchona, variables delineating vegetation structure and light availability have a strong effect on the model fit. Main conclusions In the structurally simplified Hawaiian forests studied, Cinchona facilitated endemic species in accordance with the ‘substitutive facilitation model’. This contrasts with the results of an earlier study in the naturally treeless Galápagos highlands, which revealed a sharp decrease in the abundance of endemics under Cinchona canopy. These results illustrate that, through the same structural change (addition of a vegetation layer), an invasive species may exert divergent effects across different ecosystem types. The facilitation of endemic understorey species by invasive tree species in managed forests leads to a dilemma in conservation but also to new perspectives for ecosystem restoration.     

Water inputs across a tropical montane landscape in Veracruz,Mexico: synergistic effects of land cover,rain and fog seasonality,and interannual precipitation variability

Land‐cover change can alter the spatiotemporal distribution of water inputs to mountain ecosystems, an important control on land‐surface and land‐atmosphere hydrologic fluxes. In eastern Mexico, we examined the influence of three widespread land‐cover types, montane cloud forest, coffee agroforestry, and cleared areas, on total and net water inputs to soil. Stand structural characteristics, as well as rain, fog, stemflow, and throughfall (water that falls through the canopy) water fluxes were measured across 11 sites during wet and dry seasons from 2005 to 2008. Land‐cover type had a significant effect on annual and seasonal net throughfall (NTF <0=canopy water retention plus canopy evaporation; NTF >0=fog water deposition). Forest canopies retained and/or lost to evaporation (i.e. NTF<0) five‐ to 11‐fold more water than coffee agroforests. Moreover, stemflow was fourfold higher under coffee shade than forest trees. Precipitation seasonality and phenological patterns determined the magnitude of these land‐cover differences, as well as their implications for the hydrologic cycle. Significant negative relationships were found between NTF and tree leaf area index (R²=0.38, P<0.002), NTF and stand basal area (R²=0.664, P<0.002), and stemflow and epiphyte loading (R²=0.414, P<0.001). These findings indicate that leaf and epiphyte surface area reductions associated with forest conversion decrease canopy water retention/evaporation, thereby increasing throughfall and stemflow inputs to soil. Interannual precipitation variability also altered patterns of water redistribution across this landscape. Storms and hurricanes resulted in little difference in forest‐coffee wet season NTF, while El Niño Southern Oscillation was associated with a twofold increase in dry season rain and fog throughfall water deposition. In montane headwater regions, changes in water delivery to canopies and soils may affect infiltration, runoff, and evapotranspiration, with implications for provisioning (e.g. water supply) and regulating (e.g. flood mitigation) ecosystem services.