Seedling assemblages and the alternative successional pathways experienced by Atlantic forest fragments

Background: Habitat loss and fragmentation have been argued to drastically alter the composition of tree assemblages inhabiting small forest fragments but the successional trajectory experienced by such edge-affected habitats remains controversial.

Aims: Here we examine whether small fragments (3.4–91.2 ha) support seedling assemblages more similar to those in 10–70-year-old secondary forests than to those in mature forests, in order to infer to what extent fragments move toward early successional systems.

Methods: Using 59 0.1-ha plots distributed in a fragmented landscape of Brazilian Atlantic forest, we evaluated species richness and functional and taxonomic composition of seedling assemblages in 20 small forest fragments, 19 stands of secondary forest and 20 stands of mature forests in the interior of an exceptionally large fragment (ca. 3500 ha).

Results: Small fragments presented the least species-rich seedling assemblages (17.2 ± 5.7 species), followed by secondary (22.5 ± 5.3), and mature forest (28.4 ± 5.3). Small fragments had seedling assemblages with functional and taxonomic composition more similar to those in secondary than in mature forest. Small fragments had a greater relative richness and abundance of pioneer trees (ca. 40% more), vertebrate-dispersed (6–25%), and those bearing medium-sized seeds (30–70%), while large-seeded species and individuals were reduced (>50% decrement) in comparison to seedling assemblages in mature forest.

Conclusions: By comparing seedlings across a wide range of successional habitats we offer evidence that small forest fragments are experiencing an alternative successional pathway towards an early-successional system with reduced plant diversity.     

Effects of habitat transitions on rainforest bird communities across an anthropogenic landscape mosaic

We compared bird community responses to the habitat transitions of rainforest‐to‐pasture conversion, consequent habitat fragmentation, and post‐agricultural regeneration, across a landscape mosaic of about 600 km² in the eastern Australian subtropics. Birds were surveyed in seven habitats: continuous mature rainforest; two size classes of mature rainforest fragment (4–21 ha and 1–3 ha); regrowth forest patches dominated by a non‐native tree (2–20 ha, 30–50 years old); two types of isolated mature trees in pasture; and treeless pasture, with six sites per habitat. We compared the avifauna among habitats and among sites, at the levels of species, functional guilds, and community‐wide. Community‐wide species richness and abundance of birds in pasture sites were about one‐fifth and one‐third, respectively, of their values in mature rainforest (irrespective of patch size). Many measured attributes changed progressively across a gradient of increased habitat simplification. Rainforest specialists became less common and less diverse with decreased habitat patch size and vegetation maturity. However, even rainforest fragments of 1–3 ha supported about half of these species. Forest generalist species were largely insensitive to patch size and successional stage. Few species reached their greatest abundance in either small rainforest fragments or regrowth. All pastures were dominated by bird species whose typical native habitats were grassland, wetland, and open eucalypt forest, while pasture trees modestly enhanced local bird communities. Overall, even small scattered patches of mature and regrowth forest contributed substantial bird diversity to local landscapes. Therefore, maximizing the aggregate rainforest area is a useful regional conservation strategy.     

Arthropod Diversity in Lama Forest Reserve (South Benin), a Mosaic of Natural, Degraded and Plantation Forests

Arthropod assemblages were examined in Lama forest reserve, a protected area situated in the Dahomey gap, southern Benin, composed of plantations, degraded forest and remnants of natural forest. The objectives were to compare assemblages in relation to forest type and use, to elucidate the value of forest plantations for biodiversity conservation and to identify indicator species for specific forest habitats. Arthropods were collected over an 11-month period, using standardized sets of traps (pitfall, emergence, Malaise and flight intercept traps). Nine different habitats were studied, including natural and degraded forest, forest plantations (Tectona grandis and Senna siamea) of different age, and isolated forest fragments. Our analysis focused on detritivorous and xylophagous arthropods but also included ground beetles and heteropterans, totalling 393 species. We found no differences in species richness among natural and degraded forest habitats in the centre of the reserve (Noyau central). Outside of the Noyau central, species richness was highest in old teak plantations and isolated forest fragments and lowest in young teak and fuelwood plantations. Detrended correspondence analysis (DCA) separated three main groups: (1) natural forest, (2) degraded forest and young plantations, and (3) old plantations and isolated forest fragments. Multiple regression of DCA scores of the first two axes on environmental variables identified one natural and three disturbance-related predictors of arthropod assemblages in Lama forest: soil type (texture), canopy height, naturalness (proportion of Guineo-Congolian plant species) and understorey vegetation cover. We identified 15 indicator species for six different forest habitats. The highest numbers were found in abandoned settlements and old teak plantations. β-diversity was similar among the three DCA ordination groups (degraded forest excluded). Values for β-diversity were relatively high, suggesting that all major forest habitats contribute significantly to regional species pools and should therefore be protected. To enhance arthropod diversity, we propose that management practices in Lama forest should aim to encourage the development of species-rich understorey vegetation of the Guineo-Congolian phytogeographical region.     

‘Ecologically complex carbon’– linking biodiversity values,carbon storage and habitat structure in some austral temperate forests

Aim We assessed how avian biodiversity and above‐ground carbon storage were related in different forest age‐classes, including mature stands (> 100 years), in a managed, mixed‐species eucalypt forest. Location Gippsland, south‐eastern Australia. Methods In 50 2‐ha stands ranging in age from ≤ 5 years to mature stands > 100 years, we undertook repeated avian surveys, performed detailed habitat measurements and estimated amounts of above‐ground carbon. Extensive wildfire reduced the number of sites to 28 (seven in each of four age classes) upon which analyses and inferences were made. We also analysed data on carbon storage and some bird responses from previously published studies. Results Mature vegetation (> 100 years) had the greatest richness, abundance and biomass of birds. Key ecological resources, such as tree‐hollows for nesting, generally occurred mostly in stands > 60 years. Avian richness per unit of above‐ground carbon storage was relatively low for stands of 20–60 years. While above‐ground carbon storage appeared to increase in a monotonic fashion as stands age and mature, there were quantum increases in all measures of avian biodiversity in mature stands (> 100 years). Main conclusions Our results suggest that carbon is organized in a different way, with substantially greater biodiversity benefits, in very old stands. Mature vegetation simultaneously maximizes both avian biodiversity and above‐ground carbon storage. These results bolster arguments for allocating highest priorities to the preservation of old‐growth forest stands rather than alternative investments (e.g. reafforestation for carbon sequestration).     

Landscape context of plantation forests in the conservation of tropical mammals

Plantation forests have been expanding in many tropical and subtropical environments. Howerver, even when they replace less wildlife friendly land uses such as pastures and annual crops, the biodiversity levels of pristine natural habitats often have not been recovered. Here we addressed how the landscape context of plantation forests located in South-eastern Brazil affects species richness and community resilience of medium and large size mammals. The area covered by native habitat fragments surrounding plantation forests is positively related to functional richness, including the presence of species more vulnerable to extinction in fragmented landscapes. In addition, the degree of aggregation of plantation forest stands is negatively related to more vulnerable species. No primates were recorded in our seven plantation forest sites (ranging from 272 to 24,921 ha), even when they were seen in native habitat fragments adjacent to commercial tree stands. Two invasive species (Sus scrofa and Lepus capensis) were recorded in four plantation forest sites. The impoverishment of fauna in plantation forests is due to two factors. First, plantation forests generally are structurally simplified habitats when compared to highly diverse tropical forests. Secondly, the isolation from habitat fragments which act as source of individuals in the landscape precludes the establishment of individual in plantation forest. We also highlighted the management practices to improve the complexity of vegetation in commercial tree stands should be taken cautiously, insofar as reduced productivity per area entails a greater demand for land. Thus, an alternative would be intensify the management of the commercial tree stands for wood production together with the restoration of adjacent areas set aside to conservation and native habitat fragments protection.     

Biodiversity values of abandoned teak,Tectona grandis plantations in southern Western Ghats: Is there a need for management intervention?

Abandoned plantations could potentially support a large number of native tree species through succession and restore the original tree community. In order to assess the ability of abandoned teak plantations to recover through regeneration, teak stands from 29 to 80 years old were sampled for seedling and sapling density, species richness and the use by large ungulates in the southern Western Ghats using 10 m circular plots. The influence of the forest-plantation edge was also studied. There was regeneration of a species-rich tree community in the understorey of abandoned teak plantations. However, regeneration was arrested, and thereby the large girth-class tree community remained species-poor. There was no significant change in the tree species richness with distance from natural forest, suggesting that the forest-plantation edge had little influence on the penetration of native tree species inside plantations. Asian elephant and Indian gaur dung densities were significantly lower in the plantations than in the forest. Indian gaur and sambar used the younger plantations intensively, and the density of their dung was negatively correlated with age of the plantation. Abandonment of mature teak stands arrested the succession of native trees. We provide evidence that abandoned teak plantations might not serve as suitable habitats for large herbivores during the dry months of the year in the region. The study highlights the need for active management of mature teak plantations inside wildlife reserves, in order to promote succession and improve the habitat for wild flora and fauna in the Western Ghats.     

Use of Fruiting Trees by Birds in Continuous Forest and Riparian Forest Remnants in Los Tuxtlas,Veracruz, Mexico1

The composition of plant species in fragmented landscapes may be influenced by the pattern of visitation by birds to fruiting trees and by the movement of seeds among and within fragments. We compared bird visitation patterns to two tree species (Dendropanax arboreus, Araliaceae; Bursera simaruba, Burseraceae) in continuous forest and remnants of riparian vegetation in a region dominated by pasture in Los Tuxtlas, Veracruz, Mexico. We quantified frequency of visitation, fruit consumption, consistency of visitation (percentage of total tree observation periods during which a given bird species was recorded), and species composition of birds at individuals of both tree species in continuous forest and riparian remnants. Bird visitation rate, species richness, and fruit consumption rates were similar within both tree species in the two habitats. Species assemblages at D. arboreus were different between continuous forest and remnants. Species assemblages at B. simaruba did not differ by habitat. Our results demonstrate that habitat disturbance may influence avian visitation patterns, which may in turn affect subsequent recruitment patterns in some tree species. Our results, however, were not consistent between the tree species, suggesting that it is difficult to generalize concerning the effects of forest disturbance on avian species assemblages in fruiting trees.     

Human impact diminishes seedling species richness in Kakamega Forest, Kenya

Anthropogenic forest fragmentation and other kinds of human disturbance, such as selective logging, can reduce the diversity of plant and animal species. To evaluate the impact of fragmentation and small-scale disturbance on forest regeneration, we assessed species richness and total abundance of adult trees in comparison with seedlings in the heavily fragmented and disturbed Kakamega Forest, western Kenya. In nine differently disturbed 1-ha study blocks distributed across the main forest and fragments, we mapped all trees >10 cm in diameter at breast height. Additionally, we established ninety 1-m² seedling plots within these 1-ha study blocks which were monitored over 2.5 years. We recorded altogether 74 species of adult trees (30–43 per block) and 64 seedling species (24–41 per block). Neither fragmentation nor small-scale disturbance had an impact on adult tree species richness or total tree abundance. Yet, fragmentation and especially small-scale disturbance significantly reduced seedling species richness, particularly of late-successional species. While human impact did not affect diversity of adults, the impoverished species richness of seedlings suggests a reduced potential for regeneration and a loss of tree diversity in the long-term.     

Bryophyte species richness and composition in young forests regenerated after clear-cut logging versus after wildfire and spruce budworm outbreak

The disturbance regime in mixed-wood forests of eastern Canada is characterized by both natural disturbances including wildfires and insect outbreaks as well as forestry. The understanding of how understorey plant assemblages respond to different disturbances is mostly limited to short-term wildfire-logging comparisons of vascular plants. Here, we compare patterns of species richness and composition of four bryophyte guilds in young forests (approx. 40 years old) regenerating after clear-cut logging, wildfire, and spruce budworm outbreak. In addition, young forests were compared with mature spruce-fir dominated stands (approx. 90 years old). Although similar in overall species richness at the scale of 1,000 m² all young forest types were compositionally distinct with fewer species than mature forests. Stands developed after spruce budworm outbreaks had the highest canopy cover values and the highest surface area of coarse woody debris. These stands had similar numbers of woody debris species as mature forests and were closest to mature forests in species composition. Wildfire-disturbed sites were dominated by deciduous trees and a high number of treebase species. Finally, young managed forest had the highest number of forest floor bryophytes at the scale of 100 m² among the three young forest types, but was compositionally far from mature forests in their woody debris flora. In conclusion, young forests regenerating after natural disturbances are distinctly different from young forests regenerated after clear-cutting and if natural disturbances are eliminated certain species (e.g., epixylic and treebase species) might become more restricted to older stands in the landscape.     

The effect of forest age and habitat structure on the ground‐dwelling ant assemblages of lowland poplar plantations

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Land use,fallow period and the recovery of a Caatinga forest

Caatinga vegetation continues to be converted into mosaics of secondary forest stands, but the affect of this process on biodiversity has not yet been examined. We used 35 regenerating and old‐growth stands of Caatinga to examine the recovery of plant assemblages subsequent to slash‐and‐burn agriculture and cattle ranching/pasture in northeastern Brazil. Plant assemblages were contrasted in terms of community structure (stem density/basal area/species richness/diversity), functional (leaf habit/reproductive traits) and taxonomic composition. Soil attributes were also examined to infer potential drivers for cross‐habitat differences. As expected, plant assemblages clearly differed across a large set of community‐level attributes, including all trait categories relative to leaf habit and reproduction (pollination syndrome/floral color, size, type). Overall, old‐growth forest stands supported distinct and more diverse assemblages at the plot and habitat level; e.g., long‐lived tree species were almost exclusively found in old‐growth forest stands. For most attributes, plant assemblages subsequent to pasture exhibited intermediate values between those exhibited by old‐growth forest and those of agriculture‐related stands. Surprisingly, soils exhibited similar fertility‐related scores across habitats. Our results indicate that: (1) sprouting/resprouting represents an important mechanism of forest regeneration; (2) assemblage‐level attributes suggest recovery at distinct rates; (3) forest regeneration implies community‐level changes in both vegetative and reproductive functional attributes, including directional changes; (4) Caatinga is not able to completely recover in a period of 15‐yr following land abandonment; and (5) historical land use affects recovery rates and successional pathways/taxonomic trajectories. Seasonally dry tropical forests may intrinsically cover a wide range of patterns relative to successional model, recovery rates and successional pathways.     

Structural and Floristic Heterogeneity in a 30-Year-Old Costa Rican Rain Forest Restored on Pasture Through Natural Secondary Succession

Relatively little information exists on neotropical secondary rain forests that have progressed beyond the pioneer stages of succession, or on the potential of natural regeneration to restore forest on large areas. We determined the structural and floristic characteristics (10 cm dbh) of a 30‐year‐old secondary forest developing on a 32.5 ha pasture on hilly terrain, abandoned after use of moderate intensity. Ten 0.24 ha sample plots covered the range of site conditions. The forest was dominated by long‐lived pioneer tree species; overall, the majority of species (70%) was vertebrate dispersed but the majority of individuals (52%) was of wind‐dispersed species. Tree species, including the dominants, were a mixture of those present in old‐growth and adventives colonizing from agricultural land. The forest was very heterogeneous. Vochysia ferruginea‐dominated stands characterized slopes with soils of high exchangeable acidity, while the adventive Cordia alliodora dominated sites with gentler topography and soils of lower acidity. Structural differences between the two forest types were slight, but Cordia forest had significantly greater species diversity and absolute and relative abundances of vertebrate‐dispersed tree species than Vochysia forest, which had significantly greater absolute and relative abundances of wind‐dispersed tree species. These latter differences between forest types, as well as the wide structural variation of the forest as a whole, were probably largely due to spatial and temporal variation in seed rains, some of it linked to the characteristics of the dominant species. Rain forest restoration on large pastures may depend greatly on wind dispersal and adventive tree species, and techniques for silvicultural diagnosis must be developed as a basis for the management of heterogeneous successional stands. Studies of early colonization of pastures should be expanded to focus on the causes of heterogeneity in older forests.     

Diversity of understory birds in old stands of native and Eucalyptus plantations

We compared the vegetation structure between old (>70 year) stands of planted diversified native forests and stands of Eucalyptus tereticornis embedded in a mosaic of Eucalyptus stands. We then tested for differences in the abundance, species richness, species composition, and ecological traits (forest dependence, sensitivity to forest fragmentation, and diet) of the understory bird assemblages inhabiting both kinds of stands. We expected differences in the structure of the bird assemblages because of the different origins and management strategies (contrary to native stands, Eucalyptus stands were selectively logged in the past). Three stands of each habitat (native and Eucalyptus) were sampled with mist nets during 11 months. Eucalyptus stands had a denser understory, whereas native plantations had a more developed vertical structure and a greater density of native trees. The abundance distribution of bird species was more homogeneous in Eucalyptus than in native stands. Eucalyptus had slightly higher species richness (36 species) than native stands (32 species). The composition of species and the occurrence of the diet, forest dependence, and sensitivity to forest fragmentation categories were similar between habitats. Some bird species (e.g. Turdus leucomelas), however, were more abundant in one habitat over the other. Old stands of Eucalyptus and planted native forest can harbor a diverse bird community similar in structure but not exactly equivalent for individual bird species. Planting native diversified forests and keeping set‐aside stands of the exotic tree should be viewed as complementary rather than alternative strategies for maintaining bird diversity within plantations.     

Comparing bird assemblages in large and small fragments of the Atlantic Forest hotspots

The Atlantic Forest (AF) is one of the five most threatened and megadiverse world hotspots. It is arguably the most devastated and highly threatened ecosystem on the planet. The vast scope of habitat loss and extreme fragmentation in the AF hotspots has left intact very few extensive and continuous forested fragments. We compared bird assemblages between small (<100 ha) and large (>6,000 ha) forest remnants, in one of the largest AF remnants in Argentina. We performed 84 point-counts of birds in four large fragments (LF) and 67 points in 25 small fragments (SF). We recorded 4,527 bird individuals belonging to 173 species; 2,632 belonging to 153 species in LF and 1,897 in 124 species in SF. Small fragments suffered a significant loss of bird richness, mainly forest dependent species, but the birds abundance did not decrease, due to an increase in abundance of forest independent and semi-dependent bird species (edge and non forest species) that benefit from forest fragmentation. The bird guilds of frugivores, undestory, terrestrial and midstory insectivores, nectarivores and raptors, and the endemic species of AF were area sensitive, decreasing significantly in richness and abundance in the SF. Terrestrial granivores were the only guild positively affected by forest fragmentation, containing mainly edge species, which forage in open areas or borders including crops. Our first observations on fragmentation effects on bird assemblages in the southernmost Argentinean Atlantic Forests did not validate the hypothesis on pre-adaptation to human disturbances in the bird communities of AF. On the contrary, we observed that forest dependent, endemic and several sensitive bird guilds were strongly affected by fragmentation, putting in evidence the vulnerability to the fragmentation process and the necessity to conserve large remnants to avoid reduction of the high biodiversity of AF birds.     

Differences in seed rain composition in small and large fragments in the northeast Brazilian Atlantic Forest

Tropical forests are seriously threatened by fragmentation and habitat loss. The impact of fragment size and forest configuration on the composition of seed rain is insufficiently studied. For the present study, seed rain composition of small and large forest fragments (8–388 ha) was assessed in order to identify variations in seed abundance, species richness, seed size and dispersal mode. Seed rain was documented during a 1‐year period in three large and four small Atlantic Forest fragments that are isolated by a sugarcane matrix. Total seed rain included 20,518 seeds of 149 species of trees, shrubs, palms, lianas and herbs. Most species and seeds were animal‐dispersed. A significant difference in the proportion of seeds and species within different categories of seed size was found between small and large fragments. Small fragments received significantly more very small‐sized seeds (<0.3 cm) and less large‐seeded species (>1.5 cm) that were generally very rare, with only one species in small and eight in large fragments. We found a negative correlation between the inflow of small‐sized seeds and the percentage of forest cover. Species richness was lower in small than in large fragments, but the difference was not very pronounced. Given our results, we propose changing plant species pools through logging, tree mortality and a high inflow of pioneer species and lianas, especially in small forest fragments and areas with low forest cover. Connecting forest fragments through corridors and reforestation with local large‐seeded tree species may facilitate the maintenance of species diversity.     

Would protecting tropical forest fragments provide carbon and biodiversity cobenefits under REDD+?

Tropical forests store vast amounts of carbon and are the most biodiverse terrestrial habitats, yet they are being converted and degraded at alarming rates. Given global shortfalls in the budgets required to prevent carbon and biodiversity loss, we need to seek solutions that simultaneously address both issues. Of particular interest are carbon‐based payments under the Reducing Emissions from Deforestation and Forest Degradation (REDD+) mechanism to also conserve biodiversity at no additional cost. One potential is for REDD+ to protect forest fragments, especially within biomes where contiguous forest cover has diminished dramatically, but we require empirical tests of the strength of any carbon and biodiversity cobenefits in such fragmented systems. Using the globally threatened Atlantic Forest landscape, we measured above‐ground carbon stocks within forest fragments spanning 13 to 23 442 ha in area and with different degrees of isolation. We related these stocks to tree community structure and to the richness and abundance of endemic and IUCN Red‐listed species. We found that increasing fragment size has a positive relationship with above‐ground carbon stock and with abundance of IUCN Red‐listed species and tree community structure. We also found negative relationships between distance from large forest block and tree community structure, endemic species richness and abundance, and IUCN Red‐listed species abundance. These resulted in positive congruence between carbon stocks and Red‐listed species, and the abundance and richness of endemic species, demonstrating vital cobenefits. As such, protecting forest fragments in hotspots of biodiversity, particularly larger fragments and those closest to sources, offers important carbon and biodiversity cobenefits. More generally, our results suggest that macroscale models of cobenefits under REDD+ have likely overlooked key benefits at small scales, indicating the necessity to apply models that include finer‐grained assessments in fragmented landscapes rather than using averaged coarse‐grained cells.     

Landscape‐level tree cover predicts species richness of large‐bodied frugivorous birds in forest fragments

Large‐bodied frugivorous birds play an important role in dispersing large‐sized seeds in Neotropical rain forests, thereby maintaining tree species richness and diversity. Conversion of contiguous forest land to forest fragments is thought to be driving population declines in large‐bodied frugivores, but the mechanistic drivers of this decline remain poorly understood. To assess the importance of fragment‐level versus local landscape attributes in influencing the species richness of large‐bodied (>100 g) frugivorous birds, we surveyed 15 focal species in 22 forest fragments (2.7 to 33.6 ha, avg. = 16.0 ha) in northwest Ecuador in 2014. Fragment habitat variables included density of large trees, canopy openness and height, and fragment size; landscape variables included elevation and the proportion of tree cover within a 1 km radius of each fragment. At both the individual species level, and across the community of 12 species of avian frugivore we detected, there was higher richness and probability of presence in fragments with more tree cover on surrounding land. This tendency was particularly pronounced among some endangered species. These findings corroborate the idea that partially forested land surrounding fragments may effectively increase the suitable habitat for forest‐dwelling frugivorous birds in fragmented landscapes. These results can help guide conservation priorities within fragmented landscapes, with particular reference to retaining trees and reforesting to attain high levels of tree cover in areas between forest patches.     

Recovery of Amphibian and Reptile Assemblages During Old‐Field Succession of Tropical Rain Forests

Conversion of tropical forests to agriculture affects vertebrate assemblages, but we do not know how fast or to what extent these assemblages recover after field abandonment. We addressed this question by examining amphibians and reptiles in secondary forests in southeastern Mexico. We used chronosequence data (12 secondary forests fallow for 1–23 yr and 3 old‐growth forest sites) to analyze successional trajectories and estimate recovery times of assemblage attributes for amphibians and reptiles. We conducted 6 surveys at each site over 14 mo (1200 person‐hours) and recorded 1552 individuals, including 25 species of amphibians and 36 of reptiles, representing 96 and 74 percent of the expected regional number of species, respectively. Abundance, species richness, and species diversity of amphibians increased rapidly with successional age, approaching old‐growth forest values in < 30 yr. Species richness and species diversity of reptiles reached old‐growth forest values in < 20 yr. By contrast, the abundance of reptiles and the assemblage composition of amphibians and reptiles recovered more slowly. Along the chronosequence, we observed more species replacement in reptile assemblages than in amphibian assemblages. Several species in the old‐growth forest were absent from secondary forests. Dispersal limitation and harsh conditions prevailing in open sites and early successional environments appear to preclude colonization by old‐growth forest species. Furthermore, short fallow periods and isolation of forest remnants lead to the formation of new assemblages dominated by species favored by human disturbances.     

Effect of fragmentation on the community structure of epixylic bryophytes in Atlantic Forest remnants in the Northeast of Brazil

The effects of fragmentation and edge effects on the floristic composition, richness, diversity and abundance of epixylic bryophytes (growing on decaying wood) were investigated in ten fragments of Atlantic Forest remnants in the Northeast of Brazil. In each fragment, four perpendicular 100 m transects were demarcated. Along these transects, samples of bryophytes growing on decaying wood were collected. The forest fragments were grouped in three size classes (small: <100 ha; medium: 100–500 ha; large: >500 ha). Correlation and multivariate analysis were undertaken between bryophyte flora and fragment metrics (size, form, isolation, altitude variation, nuclear area and secondary vegetation percentage and distance from the edge). A total of 99 species of bryophytes, 52 liverworts and 47 mosses were registered. The statistical results confirming fragment size is an important factor in epixylic community structure. Therefore, composition, richness, diversity and abundance can be better explained by a junction of all studied landscape factors. Bryophyte richness, the percentage of samples with the greatest coverage of decaying wood and shade-tolerant species’ distribution, were not correlated to distance from the forest edge. This suggests that edge effects are not linear or can be detected beyond 100 m from the edge, which is very important for inclusion in future studies.     

Understorey vegetation development in North Finnish Picea forests after disturbance: re‐analysis of Sirén's data

Abstract. Sirén (1955) studied understorey species composition, tree stand properties and humus‐layer thickness in 64 unlogged forest stands on topographically and pedologically comparable sites. The stands were of even age (6 – 300 yr), stocked with the first or second tree generation after wildfire. The view of Sirén and several authors after him, that the vegetation of old‐growth boreal Picea forests is homogeneous on a broad scale, was examined by applying, in parallel, the partial variants of two ordination methods (DCA and PCA) to Sirén's vegetation data. Two main vegetation gradients were found: a major gradient running from recently burnt plots with prominence of pioneer species to plots with stand age > 100 yr, a well stocked tree layer and a thick humus layer, dominance of feather‐mosses and ample occurrence of shade‐tolerant as well as light‐preferring vascular plant species, and a second gradient along which first‐ and second‐generation plots segregate. The more prominent element of Betula trees in first‐ than in second‐generation stands < 100 yr contributed to the latter. A minor third gradient related to humus‐layer thickness was recovered by partial DCA only. The main vegetation gradient reappeared in separate ordinations of data from 47 mature forest stands (> 100 yr), but without being correlated with forest age. Variation among mature‐forest stands in the importance of pioneer species is considered mainly to be brought about by fine‐scale disturbance processes such as tree uprooting. Increasing importance of factors operating on within‐stand scales [development of a varied gap structure and stronger gradients in tree influence (radiation at ground level), soil moisture, soil depth and nutrient availability] with time is also reflected in the second and third mature‐forest ordination axes. Possible implications of the results for conservation of biological diversity and monitoring of changes in boreal forests are discussed.