Structure of insect communities inhabiting old‐growth forest specialist bracket fungi

1. This paper describes the structure of the insect communities inhabiting two old‐growth forest specialist bracket fungi, Amylocystis lapponica and Fomitopsis rosea. To study the consequences of old‐growth forest fragmentation on community structure, non‐fragmented and fragmented forest areas in eastern and southern Finland were compared. 2. Both fungal species are inhabited by more than 50 insect species. The most abundant species in the A. lapponica community is a melandryid beetle Hallomenus sp., and in the F. rosea community a tineid moth Agnathosia mendicella. The occurrence of several nationally or globally rare species underscores the importance of specific insect microhabitats, such as the specialist fungi A. lapponica and F. rosea, for the persistence of specialised insects in old‐growth forests. 3. The species composition in both fungal species differs greatly from each other. The food webs are characterised by one numerically dominant host‐specific primary fungivore, which implies that fungivorous species inhabiting bracket fungi may be more specialised on a particular host than is generally thought. 4. The community structure was the same in the highly fragmented and in the non‐fragmented study areas, probably because fragmentation has occurred relatively recently.     

Spore deposition of wood-decaying fungi: importance of landscape composition

The abundance of many species of wood-decaying fungi has decreased dramatically in Swedish boreal forests over the last century. Therefore, we have investigated the relationship between the spore dispersal of wood-decaying fungi and two key features of landscape composition, namely the amount and age of old forest stands at different spatial scales. Spore deposition was monitored in two regions, using sampling methods based on recording the dikaryotisation of monokaryotic mycelia on nutrient agar and wood discs. The studied species, all mainly confined to Norway spruce in the boreal forest zone, were Fomitopsis pinicola, Fomitopsis rosea , Gloeoporus taxicola , Phlebia centrifuga and Trichaptum laricinum . The study of forestry intensity showed for both regions, that the spore deposition for F. pinicola , F. rosea and G. taxicola was higher in circular plots (radius 2-km) with a high proportion of old Norway spruce forest (>80 yr) than in plots with a lower proportion of old forest. Analysis of the variation in spore deposition of F. rosea and P. centrifuga in relation to the proportion of old spruce forest within 1, 2 and 3 km of the spore sampling point showed that the proportion of old forest within a 3-km radius explained more of the variation than the proportions within 1- and 2-km radii. In addition, the proportion of forest older than 140 yr explained more of the variation than the proportion of younger forests. Thus, the results show that the spore deposition of the studied species strongly depends on the landscape composition at both regional and local scales. Further, the spore deposition at the local scale was best explained by the proportion of >140 yr old spruce forest, which exceeds the common harvest rotation period.     

Will forest conservation areas protect functionally important diversity of fungi and lichens over time?

Incorporating functional values in biodiversity monitoring systems could add novel perspectives of the status of biodiversity in conservation areas. Stable frequencies of large foliose nitrogen-fixing cyanolichens likely have positive effects on the nitrogen budget of forests and provide food, material and shelter for invertebrates, gastropods and birds. Stable volumes of deadwood and frequencies of associated fungi provide an important supporting function for ecosystem services such as nutrient cycling, carbon storage and soil formation. Based on regional monitoring data from boreal old-growth forest nature reserves and key habitats, we tested for changes in the frequency of various functionally important substrates and species over time. We detected significant reductions in the frequency of indicator cyanolichens occurring on deciduous trees already after 10 years in key habitats, despite non-significant changes in their host substrates. Frequencies of indicator pendulous lichens Alectoria sarmentosa and Bryoria nadvornikiana had also decreased in key habitats, despite overall stable volumes of large conifer host trees. Lichen reductions were more pronounced in the smaller key habitats compared to the larger formally protected nature reserves, likely due to degrading fragmentation and isolation effects. In contrast to these lichens, the average frequencies of old-growth forest indicator fungi decaying coniferous deadwood and common fungi on deciduous trees (Fomes fomentarius) and coniferous trees (Fomitopsis pinicola) remained unchanged. The studied cyanolichens and fruiting fungi generally had similar extinction rates over 10 years, whilst only cyanolichens had substantially lower colonization rates. Amid a severely fragmented landscape, conservation areas seem to struggle in preserving some of the basic old-growth forest values.     

Population genetics of Fomitopsis rosea– a wood-decay fungus of the old-growth European taiga

The genetic structure of the wood-decay fungus Fomitopsis rosea (Alb. et Schw. Fr.) Karst is presented for populations sampled in Russia, Sweden and Finland. A total of 11 variable arbitrary primed (AP)–PCR markers were found to be segregated in a 1:1 ratio. The genotype for each fruiting body was inferred from the genotypes of the haploid single-spore isolates. Observed heterozygosity was lowest in southern Fennoscandia, the area with the longest history of forestry management. Overall, there was a deficit of heterozygotes compared with frequencies of heterozygotes predicted from Hardy–Weinberg expectations. When we partitioned the deficit of heterozygotes into the hierarchical genetic structure by means of F -statistics we found that the possible causes for it were both limited gene flow between regions and local nonrandom mating within all populations. Future management for the conservation of F. rosea populations and how the results relate to the life cycle of basidomycete fungi are discussed.     

Initial fungal colonizer affects mass loss and fungal community development in Picea abies logs 6 yr after inoculation

Picea abies logs were inoculated with Resinicium bicolor, Fomitopsis pinicola or left un-inoculated and placed in an old-growth boreal forest. Mass loss and fungal community data were collected after 6 yr to test whether simplification of the fungal community via inoculation affects mass loss and fungal community development. Three techniques were used to survey communities: (1) observation of fruiting structures; (2) culturing on media; and (3) cloning and sequencing of ITS rDNA. Fruit body surveys detected the smallest number of species (18, 3.8 per log), DNA-based methods detected the most species (72, 31.7 per log), and culturing detected an intermediate number (23, 7.2 per log). Initial colonizer affected community development and inoculation with F. pinicola led to significantly greater mass loss. Relationships among fungal community composition, community richness and mass loss are complex and further work is needed to determine whether simplification of fungal communities affects carbon sequestration in forests.     

Rapid Recovery of Biomass, Species Richness, and Species Composition in a Forest Chronosequence in Northeastern Costa Rica

Secondary forests are a vital part of the tropical landscape, and their worldwide extent and importance continues to increase. Here, we present the largest chronosequence data set on forest succession in the wet tropics that includes both secondary and old-growth sites. We performed 0.1 ha vegetation inventories in 30 sites in northeastern Costa Rica, including seven old-growth forests and 23 secondary forests on former pastures, ranging from 10 to 42 yr. The secondary forest sites were formerly pasture for intervals of <1–25 yr. Aboveground biomass in secondary forests recovered rapidly, with sites already exhibiting values comparable to old growth after 21–30 yr, and biomass accumulation was not impacted by the length of time that a site was in pasture. Species richness reached old-growth levels in as little as 30 yr, although sites that were in pasture for > 10 yr had significantly lower species richness. Forest cover near the sites at the time of forest establishment did not significantly impact biomass or species richness, and the species composition of older secondary forest sites (>30 yr) converged with that of old growth. These results emphasize the resilience of tropical ecosystems in this region and the high conservation value of secondary forests.
     

Fomitopsis incarnatus sp. nov. based on generic evaluation of Fomitopsis and Rhodofomes

A new polypore in the genus Fomitopsis was discovered in Kangwon Province, Korea. The species was morphologically similar to Fomitopsis rosea and F. cajanderi, but the pinkish white pore surface, the size and shape of the pores and the number of sterigmata were different enough for it to be distinguished from the recorded species of Fomitopsis. Based on the results of morphological and phylogenetic analyses, this new polypore is proposed as Fomitopsis incarnatus sp. nov.     

Forest fragmentation leads to food web contraction

Fragmentation and loss of habitat are critical components of the global change currently threatening biodiversity and ecosystem functioning. We studied the effects of habitat loss through fragmentation on food web structure, by constructing and analyzing plant‐herbivore and host‐parasitoid food webs including more than 400 species and over 120 000 feeding records, in 19 Chaco Serrano remnants of differing areas. Food web structure was altered by habitat fragmentation, with different metrics being affected depending on interaction type, and with all changes being driven by the reduced size of networks in smaller fragments. Only connectance varied in both quantitative and qualitative analyses, being negatively related to area. In addition, the interactions were represented by proper successive subsets, modulated mainly by resource availability (plant–herbivore) or consumer specialization (host–parasitoid), as forest size decreased. The results suggest that habitat loss has led to food web contraction around a central core of highly‐connected species, for plant–herbivore as well as for host–parasitoid systems. The study provides new insights into the effects of human perturbations on complex biological systems.     

林业活动和森林片断化对甲虫多样性的影响及保护对策

森林片断化是造成生物多样性丧失的主要原因之一,而林业活动是导致森林片断化的重要因素,同时也在森林恢复中起重要作用。本文从小尺度、局域尺度以及生物地理尺度（大尺度）3个生态尺度分析林业活动和森林片断化对甲虫多样性的影响。在小尺度下,林业活动能够通过改变森林生境或微生境的类型和特性而影响甲虫物种分布。在局域尺度下,林业活动（尤其是森林砍伐）往往能提高许多甲虫类群（如步甲）的物种丰富度（α多样性）,这主要与来自周围环境物种扩散以及保留了若干耐受新环境能力较强的森林物种有关;然而,对森林生境依赖性很强的特有种受到了森林片断化的负面影响,面临局域种群灭绝的危险。在生物地理尺度下,林业活动（伐木或森林恢复）使森林生境单一化、异质性降低,从而导致对森林生境变化敏感的物种种群数量降低甚至灭绝。基于以上结果,可以归纳出3个基本原则用于指导林业管理,既能保证林业经济收益,又能维持森林生物多样性。首先,保留大面积的原始森林作为特有种的栖息环境基地,为这些物种在将来森林恢复后重新定居提供资源;其次,由于保护区内原始森林面积有限,且所代表的生境类型有限,所以发展依据自然干扰模式的新伐木方法十分必要;最后,依据自然规律（如火灾）进行森林恢复和天然演替,避免森林的单一化,丰富森林生境类型。     

Insights on the functional composition of specialist and generalist birds throughout continuous and fragmented forests

A decline in species number often occurs after forest fragmentation and habitat loss, which usually results in the loss of ecological functions and a reduction in functional diversity in the forest fragments. However, it is uncertain whether these lost ecological functions are consistently maintained throughout continuous forests, and so the importance of these functions in continuous forests remains unknown. Point counts were used to assess both the taxonomic and functional diversity of specialist and generalist birds from sampling in a continuous primary forest compared with forest fragments in order to investigate the responses of these groups to forest fragmentation. We also measured alpha and beta diversity. The responses of specialists and generalists were similar when we assessed all bird species but were different when only passerines were considered. When examining passerines we found lower total taxonomic beta diversity for specialists than for generalists in the continuous forest, while taxonomic beta diversity was higher in the fragmented forest and similar between bird groups. However, total functional beta‐diversity values indicated clearly higher trait regularity in continuous forest for specialists and higher trait regularity in fragments for generalists. Specialists showed significantly higher functional alpha diversity in comparison with generalists in the continuous forest, while both groups showed similar values in fragments. In passerines, species richness and alpha functional diversity of both specialist and generalist were explained by forest connectivity; but, only fragment size explained those parameters for specialist passerines. We suggest that considering subsets of the community with high similarity among species, as passerines, provides a better tool for understanding responses to forest fragmentation. Due to the regularity of specialists in continuous forest, their lost could highly affect functionality in forest fragments.     

The effect of deforestation on the genetic diversity and structure in <Emphasis Type="Italic">Acer saccharum</Emphasis> (Marsh): Evidence for the loss and restructuring of genetic variation in a natural system

The level and distribution of genetic diversity can be influenced by species life history traits and demographic factors, including perturbations that might produce population bottlenecks. Deforestation and forest fragmentation are common sources of population disturbance in contemporary populations of forest ecosystems. Although the genetic effects of forest fragmentation and deforestation have been examined by assessing levels of genetic variation in forest fragments that remain after logging, few considerations have been made of the populations that re-colonize once-cleared areas. Here we examine the effects of human-mediated population bottlenecks on the level and distribution of genetic diversity in natural populations of the long-lived forest tree species, Acer saccharum (sugar maple). We compared genetic variation and structure for populations of sugar maple found within old-growth forested area and in area that has re-colonized since logging. In this study the percent polymorphic loci and allelic richness estimates were reduced in the logged populations compared to old-growth populations. Jackknifed estimates of population genetic differentiation showed significantly higher differentiation among logged populations, with this result being consistently seen when individuals within populations were grouped according to diameter at breast height. The result of decreased genetic variation and higher levels of genetic structure among logged populations suggests that even one extensive bout of logging can alter the level and distribution of genetic variation in this forest tree species.     

Microclimate and tree community linked to differences in lepidopteran larval communities between forest fragments and continuous forest

Aim  We aim to assess the impact of forest fragmentation on lepidopteran larval community and study the associations of microclimate and tree community with lepidopteran assemblage.
Location  Kibale National Park, Uganda.
Methods  We investigated the effects of forest fragmentation on leaf herbivory, density of lepidopteran caterpillars, species richness and diversity as well as the composition of herbivorous lepidopteran larval community. Microclimate, size of the fragment, distance to the continuous forest, and tree diversity were studied as possible explanatory factors. We sampled 10 Neoboutonia macrocalyx Pax. (Euphorbiaceae) trees in each fragment during dry and rainy season, total of four times, in a year to cover the seasonal variation.
Results  The rates of herbivory, total larval density and species richness were significantly lower in the forest fragments than in the continuous forest but species diversity expressed as Fisher's alpha did not differ. The dominance structure and community composition of the larval communities in the fragments was different from that of the continuous forest. None of the differences we observed were related to the fragment area or distance to the continuous forest. Instead, we found an indication of association between the herbivore and the tree communities. The fragments had significantly lower humidity during most of the day and higher temperature during the afternoons (14–17 h), which might partially explain the differences in lepidopteran larval communities.
Main conclusions  Decreased larval density and species richness as well as differences in the community composition and structure all highlight the importance of large continuous forest areas for maintaining larval biodiversity.     

Spatial and temporal changes in bird assemblages in forest fragments in an eastern Amazonian savannah

We investigated the effects of forest fragmentation on bird assemblages in an Amazonian savannah landscape with forest fragments that have been isolated for more than 100 years. The study was conducted in areas surrounding the village of Alter do Chão (2°31′S, 55°00′W), Santarém, Brazil. Bird surveys and measurements of tree density were undertaken in 25 areas, with 19 plots in forest fragments of different sizes and six in an area of continuous forest. Data on forest‐fragment size, perimeter, and isolation were obtained from a georeferenced satellite image. Variation in number of bird species recorded per plot was not related to vegetation structure (tree density). The number of bird species recorded per plot increased significantly only with fragment area, but was not influenced by fragment shape or degree of isolation, even when considering species from the savannah matrix in the analysis. Fragments had fewer rare species. Multivariate ordination analyses (multiple dimensional scaling, [MDS]) indicated that bird species composition changed along a gradient from small to large forest fragments and continuous‐forest areas. In the Amazonian savannah landscapes of Alter do Chão, the organization and composition of bird assemblages in forest fragments are affected by local long‐term forest‐fragmentation processes. Differences in the number of bird species recorded per plot and assemblage composition between forest fragments and continuous forest were not influenced by forest structure, suggesting that the observed patterns in species composition result from the effects of fragmentation per se rather than from preexisting differences in vegetation structure between sites. Nevertheless, despite their long history of isolation, the forest fragments still preserve a large proportion (on average 80%) of the avifauna found in continuous‐forest areas. The fragments at Alter do Chão are surrounded by natural (rather than planted) grassland, with many trees in the savannah matrix and the landscape has vast areas covered by forest, which may have helped to ameliorate the influences of forest fragmentation.     

Deforestation Trends of Tropical Dry Forests in Central Brazil

Tropical dry forests are the most threatened forest type in the world yet a paucity of research about them stymies development of appropriate conservation actions. The Paranã River Basin has the most significant dry forest formations in the Cerrado biome of central Brazil and is threatened by intense land conversion to pastures and agriculture. We examined changes in Paranã River Basin deforestation rates and fragmentation across three time intervals that covered 31 yr using Landsat imagery. Our results indicated a 66.3 percent decrease in forest extent between 1977 and 2008, with an annual rate of forest cover change of 3.5 percent. Landscape metrics further indicated severe forest loss and fragmentation, resulting in an increase in the number of fragments and reduction in patch sizes. Forest fragments in flatlands have virtually disappeared and the only significant forest remnants are mostly found over limestone outcrops in the eastern part of the basin. If current patterns persist, we project that these forests will likely disappear within 25 yr. These patterns may be reversed with creation of protected areas and involvement of local people to preserve small fragments that can be managed for restoration.     

Parasitoids on the loose – experimental lack of support of the parasitoid movement hypothesis

Forest fragmentation can disrupt important ecological processes both within and among species. Of particular interest is the extent to which fragmentation decouples economically important pest species from their parasites. In an influential paper, Roland and Taylor (1997) advanced the hypothesis that forest fragmentation at a relatively fine spatial scale would disrupt the ability of parasitoids to aggregate in response to host density, thereby facilitating the build-up of host populations at isolated sites. Our study provides the first experimental test of this notion. We augmented existing densities of forest tent caterpillar individuals at two types of sites: isolated forest fragments and continuous tracts of forest. Of the four parasitoid species studied by Roland and Taylor (1997), two were encountered in our experiment. For the sarcophagid fly Arachnidomyia aldrichi and the tachinid Carcelia malacosomae, we found no difference in parasitism rates among forest fragments and continuous forest tracts. A third tachinid species unique to this study, Lespesia frenchii , is known to be a broad generalist with respect to host choice. In this species, we observed significantly elevated parasitism in forest fragments, and higher parasitism at the forest edge than within the forest. All of these findings suggest complex effects of forest fragmentation on parasitoid movement, and warn against broad generalizations from observational studies. Forest fragmentation may affect movement patterns in a multitude of ways, significantly obscuring or modifying the simplest mechanisms proposed by Roland and Taylor (1997).     

Diversity of coleoptera of the bracket fungus Fomitopsis pinicola in a Norwegian spruce forest

The beetle fauna of 299 sporocarps of the bracket fungus Fomitopsis pinicola in a 200 ha spruce forest in southeastern Norway was investigated in relation to sporocarp, tree and forest variables. The sporocarps contained 36 species of beetles, of which six species are on the Norwegian Red List. Of 12,373 individual beetles collected, 91% were Cis glabratus. Plots of species accumulation curves suggested that there may be more than 60 beetle species present in F. pinicola in the area, but that probably all the specialist Ciidae were found. The major factor influencing beetle diversity turned out to be the level of dead wood at and in the vicinity of the sampling site, with a higher number of species per unit volume of sporocarps in areas with high levels of dead wood. There were also significantly more red-listed species in those areas. Analysing the species occurrence with stepwise logistic regression, we show preferential habitat selection of the six most abundant species of Ciidae. Conservation of beetles associated with bracket fungi using amount of dead wood as a surrogate measurement is discussed.     

Forest fragmentation effects on palm diversity in central Amazonia

1 The effects of fragmentation on quantitative measures of floristic diversity in a palm community were examined in the Biological Dynamics of Forest Fragments study area in central Amazonia. Three 1-ha, three 10-ha, two 100-ha and three continuous forest reserves, distributed among three sites, were surveyed. In each reserve, 10 20 × 20 m plots were sampled, resulting in a total of 110 plots representing 4.4 sampled hectares.
2 The taxon composition of this palm community was dominated by stemmed, understorey palms. A total of 23 225 individuals from 36 taxa was recorded; five of the taxa were not sampled in continuous forest.
3 Taxa richness did not vary across reserve size or sites unless taxa not sampled in the continuous forest were removed from the analysis. Smaller forest fragments then harboured fewer taxa in the seedling stage than large forest fragments or continuous forest, despite the short time since isolation (10–15 years). There was a significant effect of location on the number of taxa per plot for all life stages, but only seedling and total were significantly affected by reserve size.
4 Reserve size did not affect the Shannon and Evenness indices. Reserves of similar sizes were floristically more similar than reserves of very different sizes.
5 Palms are important for the structure and composition of the forest. Their conservation may require the establishment of a number of large reserves.     

Population genetics of the wood-decay fungus Phlebia centrifuga P. Karst. in fragmented and continuous habitats

The basidiomycete Phlebia centrifuga is a wood-decay fungus characteristic for unmanaged old-growth forests of spruce, a habitat that has become increasingly fragmented due to forest management. The aim of this study was to investigate the genetic population structures of P. centrifuga in both continuous and fragmented habitats, and estimate the potential impact of fragmentation on the genetic diversity of the fungus. Three hundred fifteen single spore isolates (representing 47 spore families and 33 single isolates) from eight populations across northern Europe (Russia, Finland, and Sweden) were screened with seven microsatellite markers and arbitrary primed polymerase chain reaction with the M13 minisatellite. The two molecular methods generally gave the same pattern for the genetic population structure. There were no significant differences between the observed and the expected heterozygosities, and the inbreeding coefficient (FIS) did not indicate any inbreeding. The fixation index (FST) revealed a general pattern with little to moderate genetic differentiation for the majority of populations, while the southernmost Swedish population Norra Kvill was the only one showing high differentiation from about half of the other populations. Swedish population Fiby with the shortest distance to the continuous habitat was moderately differentiated from most of the others and to the largest extent differed from geographically closest population of Norra Kvill. The results indicate that the fragmentation of old-growth forest in Russia and Finland is more recent than the fragmentation in Sweden, and the genetic population structures of P. centrifuga in northern Europe might be related to differences in forest landscape dynamics between the two areas.     

Evaluation of the monophyly of Fomitopsis using parsimony and MCMC methods

To evaluate the monophyly of Fomitopsis and elucidate phylogenetic relationships of its members, partial nuclear large subunit (partial 28S) ribosomal RNA genes were sequenced from 10 species of Fomitopsis and 15 related species. Phylogenetic analyses indicated that Fomitopsis was phylogenetically heterogeneous and its members were divided into three subgroups. The constrained tree excluding F. palustris (the type species of Pilatoporus) from Fomitopsis core group was rejected, thus rejecting the taxonomic concept to segregate Pilatoporus from Fomitopsis. The monophyly of taxa belonging to F. rosea complex was rejected, thus rejecting the complex definition based on morphological similarities. The exclusion of Piptoporus betulinus (the type species of Piptoporus) from Fomitopsis core group was rejected and Piptoporus proved to be heterogeneous in both best MP and MAP trees. The monophyly of F. officinalis with Fomitopsis core group also was rejected. Fomitopsis officinalis was closely related to Antrodia xantha and formed an independent lineage from Fomitopsis core group at the basal position of brown rotting fungi comprising Antrodia, Daedalea, Fomitopsis, Piptoporus and Postia. The MAP tree topologyobtained from MCMC computation of Bayesian inference was similar to the one of the best MP tree based on the parsimony analysis but showed a higher likelihood score in the Kishino-Hasegawa test and reflected better evolutionary patterns for the phylogeny of Fomitopsis.