Early response of understory vegetation to the mass dieback of Norway spruce in the European lowland temperate forest

Spruce-dominated forests are commonly exposed to disturbances associated with mass occurrences of bark beetles.The dieback of trees triggers many physical and chemical processes in the ecosystem resulting in rapid changes in the vegetation of the lower forest layers.We aimed to determine the response of non-tree understory vegetation to the mass dieback of Norway spruce(Picea abies)in the first years after the disturbance caused by the European spruce bark beetle(Ips typographus)outbreak.Our study area was the Białowieża Biosphere Reserve covering the Polish part of the emblematic Białowieża Forest,in total 597km^(2).The main data source comprised 3,900 phytosociological relevés(combined spring and summer campaigns)collected from 1,300 systematically distributed forest sites in 2016–2018–the peak years of the bark beetle outbreak.We found that the understory responded immediately to mass spruce dieback,with the most pronounced changes observed in the year of the disturbance and the subsequent year.Shade-tolerant forest species declined in the initial years following the mass spruce dieback,while hemicryptophytes,therophytes,light-demanding species associated with non-forest seminatural communities,as well as water-demanding forest species,expanded.Oxalis acetosella,the most common understory species in the Białowieża Forest,showed a distinct fluctuation pattern,with strong short-term expansion right after spruce dieback,followed by a gradual decline over the next 3–4 years to a cover level 5 percentage points lower than before the disturbance.Thus,our study revealed that mass spruce dieback selectively affects individual herb species,and their responses can be directional and non-directional(fluctuation).Furthermore,we demonstrated that the mass dieback of spruce temporarily increases plant species diversity(α-diversity).

The effects of understory vegetation on P availability in Pinus radiata forest stands: A review

In many second-rotation Pinus radiata forest planta-tions, there has been a steady trend towards wider tree spacing and an increased rate of application of P fertiliser. Under these regimes, the potential for understory growth is expected to in-crease through increased light and greater nutrient resources. Therefore, understory vegetation could become a more signifi-cant component of P cycling in P. radiata forests than under closely-spaced stands. Studies have shown that growth rates and survival of trees is reduced in the presence of understory vegeta-tion due to the competition of understory vegetation with trees. Other studies have suggested that understory vegetation might have beneficial effects on nutrient cycling and conservation within forest stands. This review discusses the significance of understory vegetation in radiata pine forest stands, especially their role in enhancing or reducing P availability to forest trees.

Investigation and Analysis of Understory Vegetation in Casuarina equisetifolia Forests in Northern Hainan Province

Understory vegetation in the Casuarina equisetifolia forests at different ages was sampled and analyzed fi rstly, and then the structural features, biomass, and distribution characteristics of carbon stock of the understory vegetation, as well as the quantity and thickness of the understory litter were studied. The results show that the understory vegetation in the C. equisetifolia forests is mainly composed of herbaceous plants, and there are nearly no shrubs in the study area. Moreover, the quantity of herbaceous plants rises obviously with the increase of forest age. There is a big difference between the woodlands at different ages in total biomass of herbaceous plants, and the maximum of total biomass can be found in 15-year-old woodland, reaching 1.68 t/hm^2; the biomass of overground parts of herbaceous plants is higher than that of underground parts. There is a big difference between the woodlands at different ages in thickness of the understory litter, and the thickness of understory litter in 10-yearold woodland is the highest, reaching 6.67 cm. The maximum quantity of the understory litter has reached 14.69 t/hm^2, and the thickness of the half decomposed litter is larger than that of the undercomposed litter in most woodlands.

Relationship between topography and the distribution of understory vegetation in a Pinus massoniana forest in Southern China

The poor growth of understory vegetation and the severe losses of soil and water in Pinus massoniana forests have recently become serious concerns in an area in southern China with eroded red soil.The influence of topography on the spatial distribution of vegetation,however,has received little attention.This study combined several multivariate analyses to discern the complicated relationship between understory vegetation and topography.Thirty-six plots(10 m×10 m)were sampled in a field survey of the vegetation and topography in the central red-soil region.The distributions of the understory vegetation differed significantly amongst the topographies.Most plants grew in gullies,and few grew on ridges.The low coverage(25.2%)and number of species(5 per plot)of the vegetation on ridges was due to serious soil erosion.Surface curvature and slope aspect were the first and second most important topographic factors,respectively,affecting the distribution of the vegetation.The relationship between topography and distribution could be described by a linear model.Surface curvature or slope aspect alone,however,could only explain 22.2-59.2%of the variance in distribution.The adaptation of vegetation to specific topographies should be considered for restorations of P.massoniana forests in the study area.The results of this study will be helpful for selecting potential sites for seeding and vegetation restoration to improve the ecology of the study area.Further studies will be needed to identify the mechanism of the distribution of the understory vegetation in these P.massoniana forests.

Effect of abiotic factors on understory community structures in moist deciduous forests of northern India

Understory vegetation controls, in a significant way, the regeneration of overstory trees, carbon sequestration and nutrient retention in tropical forests. Development and organization of understory vegetation depend on climate, edaphic and biotic factors which are not well correlated with plant community structures. This study aimed to explore the relationships between understory vegetation and abiotic factors in natural and planted forest ecosystems. A non-metric multidimensional scaling(NMS) ordination technique was applied to represent forest understory vegetation among five forest communities, i.e., a dry miscellaneous forest(DMF), a sal mixed forest(SMF), a teak plantation(TP), a low-land miscellaneous forest(LMF) and a savanna area(SAV) of the Katerniaghat Wildlife Sanctuary, located in northern India. Microclimatic variables, such as photosynthetically active radiation(PAR), air temperature(AT), soil temperature(ST), ambient atmospheric CO 2 concentration, absolute air humidity(AH), physical and chemical soil properties as well as biological properties were measured. Understory species were assessed via 100 random quadrats(5 m × 5 m) in each of the five forests in which a total of 75 species were recorded encompassing 67 genera from 37 families, consisting of 32 shrubs and 43 plant saplings. DMF was the most dense forest with 34,068 understory individuals per ha of different species, whereas the lowest understory population(13,900 per ha) was observed in the savanna. Ordination and correlation revealed that microclimate factors are most important in their effect compared to edaphic factors, on the development of understory vegetation in the various forest communities in the north of India.

Consistent response of nematode communities to management of coniferous plantations

Background:Crop-tree thinning (CTT) is a forest management practice aiming at enhancing the growth of target trees in plantations by harvesting neighboring trees. Along with the positive effect on tree growth, thinning represents a disturbance, which likely affects belowground organisms and may feed back to stand productivity.However, the impact of CTT on the belowground food web is poorly understood. Since nematodes are species-rich and abundant belowground organisms, occupying a variety of trophic positions in soil food webs and being sensitive to disturbances, they serve as ecological indicators of ecosystem disturbance.Results:We studied the effect of CTT on the soil nematode community structure in pine (Pinus massoniana Lamb.),Chinese fir (Cunninghamia lanceolata (Lamb.) Hook.) and cypress (Cupressus funebris Endl.) plantations in the Sichuan Basin three years after applying CTT. Crop-tree thinning significantly increased the abundance of soil nematodes in each of the plantations. Moreover, CTT significantly increased the relative abundance of herbivorous nematodes in each of the plantations. Furthermore, CTT increased the proportion of stress tolerators (c-p 1)and enrichment opportunists (c-p 2) and reduced the maturity, structure and enrichment indices of nematodes in Chinese fir and cypress plantations, while only subtle changes were observed in pine plantations. Interestingly,across plantations, the effects of CTT on soil nematode communities were mainly due to changes in microbial biomass nitrogen and understory vegetation diversity.Conclusions:Forest management practices resulting in more open canopies uniformly affect soil food webs by changing the quantity and quality of resources associated with increased understory cover and diversity as well as microbial food. These insights increase our understanding of the impacts of forest management on the structure and functioning of belowground communities, which may contribute to management and conservation policies for more sustainable forestry.

Effect of stand density and humus removal on regeneration of Pinus densiflora seedlings in a mixed conifer and broad-leaved forest

We studied the impact of stand density and forest floor humus removal on regeneration ofPinus densiflora in a mixed co- nifer and broad-leaved forest on the Daimonji-Yama Mountain, Kyoto, Japan. Three levels of stand density were set, i.e., a clear-cut site, a heavily thinned site and a lightly thinned site. In each density treatment, comparisons were carried out between humus removal （A0-free） and humus intact （A0-intact） subplots. We counted the emergence of sown pine seeds and measured survival and growth of pine seedlings over the next four years. In addition, light conditions and the recovery of understory were monitored. Results show that thinning intensity and humus removal promoted the establishment and growth of seedlings. One exception, however, was that seedling growth was minimal in the heavily thinned A0-intact subplots. This was due to a strong response of understory vegetation growth on the treatment combination, which hindered the pine seedling growth. Furthermore, we found that the canopy openness measured at 1.3-m height remained at least 35% for the heavily thinned site in 2008. The results suggest that the removal of the A0 layer after heavy thinning is the most effective and labor-saving operation for P. densifiora regeneration.

Experimental evidence on biodiversity impacts of variable retention forestry,prescribed burning,and deadwood manipulation in Fennoscandia

Intensive forest management has been applied in most Fennoscandian forests for a period of almost one felling rotation.This paradigm has produced even-aged and even-structured forests of different successional stages that cover about 90%of forest land.At the same time,wildfires have been nearly eliminated in most of the Fennoscandian nature.Consequently,hundreds of species are red-listed because of forest management.To support these species,forest management requires improvements.Variable retention forestry and habitat restoration have been suggested to mitigate negative effects of forest management on biodiversity,and these have been practiced to some extent during the past few decades.Here,we review experimental results on the effects of variable retention and two restoration measures(prescribed burning and artificial addition of coarse woody debris)on different species groups in Fennoscandia.Our key findings are as follows:(i)Many species respond positively to felling within a few years,apparently due to released and often ephemeral resources,such as fresh residue and stumps.Species associated with shady conditions are negatively impacted,but any retention supports many of these,and their species composition remains almost unaffected with 50-70%retention of the initial tree volume.(ii)These effects remain detectable for at least 10-30 years or,according to some studies,nearly 100 years,e.g.,in polypore fungi.(iii)Initial effects of prescribed burning on most species groups(apart from pyrophiles)are negative,but within 10-15 years post-fire sites begin to support many rare and threatened deadwood-dependent species.Epiphytic lichens,however,remain negatively affected.(iv)Artificial addition of deadwood(mostly high stumps)supports a wide spectrum of deadwood-dependent species,but the species composition differs from that of naturally died trees.(v)Moisture and micro-habitat variation are crucial for forest species at harvested sites,at least in forests dominated by Norway spruce.We conclude that felling method as such is of little importance for threatened forest species,although retention mitigates many negative effects.These species require microclimatic continuity,and maintenance and active increase of legacies,such as deadwood of different qualities(species,downed/standing,snag/log/stump,decay stage),very old trees,and tree species mixtures.