Arctic warming on two continents has consistent negative effects on lichen diversity and mixed effects on bryophyte diversity

Little is known about the impact of changing temperature regimes on composition and diversity of cryptogam communities in the Arctic and Subarctic, despite the well‐known importance of lichens and bryophytes to the functioning and climate feedbacks of northern ecosystems. We investigated changes in diversity and abundance of lichens and bryophytes within long‐term (9–16 years) warming experiments and along natural climatic gradients, ranging from Swedish subarctic birch forest and subarctic/subalpine tundra to Alaskan arctic tussock tundra. In both Sweden and Alaska, lichen diversity responded negatively to experimental warming (with the exception of a birch forest) and to higher temperatures along climatic gradients. Bryophytes were less sensitive to experimental warming than lichens, but depending on the length of the gradient, bryophyte diversity decreased both with increasing temperatures and at extremely low temperatures. Among bryophytes, Sphagnum mosses were particularly resistant to experimental warming in terms of both abundance and diversity. Temperature, on both continents, was the main driver of species composition within experiments and along gradients, with the exception of the Swedish subarctic birch forest where amount of litter constituted the best explanatory variable. In a warming experiment in moist acidic tussock tundra in Alaska, temperature together with soil ammonium availability were the most important factors influencing species composition. Overall, dwarf shrub abundance (deciduous and evergreen) was positively related to warming but so were the bryophytes Sphagnum girgensohnii, Hylocomium splendens and Pleurozium schreberi; the majority of other cryptogams showed a negative relationship to warming. This unique combination of intercontinental comparison, natural gradient studies and experimental studies shows that cryptogam diversity and abundance, especially within lichens, is likely to decrease under arctic climate warming. Given the many ecosystem processes affected by cryptogams in high latitudes (e.g. carbon sequestration, N₂‐fixation, trophic interactions), these changes will have important feedback consequences for ecosystem functions and climate.     

The effects of wood decay type on the growth of bryophyte gametophytes

Deadwood is an important habitat for bryophytes in boreal and subalpine forests. The type of decay in wood (white, brown, and soft rot) caused by fungal colonizers has been revealed to affect bryophyte communities. However, little is known about the effects of decay type on the growth of bryophytes. We tested the effect of wood decay type on gametophyte growth for two common bryophyte species, Scapania bolanderi Austin and Pleurozium schreberi (Brid.) Mitt., which dominate the logs in subalpine coniferous forest on Mt. Ontake, in central Honshu, Japan. We used pot culture experiments in an open-sky nursery field. After eight months of cultivation, the growth of S. bolanderi was larger on brown rot wood than white rot wood, but the growth of P. schreberi was not. Mixed cultures of the two species also showed greater growth on brown rot wood. However, growth of S. bolanderi was significantly smaller than P. schreberi in mixed culture. These results suggest that brown rot wood enhances growth of S. bolanderi, but growth may be reduced under competition from P. schreberi. The results are in agreement with the field observation that brown rot wood has a positive association with S. bolanderi coverage on deadwood.     

Effects of reindeer grazing on understorey vegetation in dry Pinus sylvestris forests

Abstract. Data on floristic composition and environmental variables were collected in floristically homogeneous oligotrophic pine (Pinus sylvestris) forests with heath-like under- storey vegetation in eastern Fennoscandia, and ordinated by non-linear multidimensional scaling (NMDS) in order to study the effect of lichen grazing by reindeer on the understorey vegetation. The study sites included areas with varying grazing pressure, as well as 50-yr old grazing exclosures. Sites rich in respectively bryophytes and lichens were placed at opposite ends of the ordination axes, and heavily grazed sites were placed in between them. Reindeer grazing increased the abundance of bryophytes, especially Dicranum spp. and Pleurozium schreberi. Grazing changed the vegetation to the extent that it resembled more mesotrophic sites, but this did not show any relationship with tree volume or other site productivity indicators. This was observed both in the ordination and, in a more compelling way, when exclosures with adjacent grazed areas were compared. No such signs were evident at ungrazed sites, where especially Cladina spp. spatially replace Cladonia spp. and tiny bryophytes like Barbilophozia spp., Polytrichum spp. and Pohlia nutans during succession. Cladina stellaris had almost disappeared from the most intensively grazed sites. The soil at ungrazed sites was characterized by high Al and Fe concentrations and bryophyte-rich sites by high Mn concentrations. Shannon's diversity index, depth of humus layer and proportion of bare ground also increased in sites getting richer in bryophytes.     

Growth and nutrition of black spruce seedlings in response to disruption of Pleurozium and Sphagnum moss carpets in boreal forested peatlands

In boreal forested peatlands, we disturbed Sphagnum spp. and Pleurozium schreberi carpets to see how disturbance influenced substrate physico-chemistry, and growth and foliar nutrition of planted Picea mariana seedlings. Carpets were hand disturbed using gardening tools to a depth of approximately 25 cm. Carpet disturbance was aimed at disrupting only the organic layer and did not result in the mixing of organic matter with mineral soil. Disturbed carpets, whether P. schreberi or Sphagnum spp., were warmer than undisturbed carpets and had a lower cover of ericaceous shrubs. Pleurozium schreberi carpets had a higher decomposition index than Sphagnum spp. carpets, whereas disturbance had no effect on this variable. Pleurozium schreberi had higher N_tot and dissolved organic N concentrations (DON) than Sphagnum spp., whereas disturbance increased NH ₄ ⁺ availability in both substrates. Moss disruption increased seedling growth rates as well as their foliar N and P concentrations in both substrates and these variables remained higher in P. schreberi than in Sphagnum spp. within a given treatment. Seedling growth was positively correlated to substrate N_tot, NH ₄ ⁺ and DON concentrations, and to foliar N and P concentrations, and negatively to substrate C/N and ericaceous shrub cover. Disruption of the moss carpets without mineral soil mixing improved black spruce seedling growth and nutrition in both moss types but the superiority of P. schreberi compared to Sphagnum spp. as a growing substrate remained present.     

Decomposition rates of bryophytes in managed boreal forests: influence of bryophyte species and forest harvesting

The slow decomposition rate of boreal forest floor bryophytes contributes both to maintaining high soil C reserves as well as affecting conditions for tree growth by maintaining excessively high soil water content, cooling the soil and slowing nutrient cycles. In this study, mass loss of three bryophyte species (Pleurozium schreberi, Sphagnum capillifolium, S. fuscum) was measured in unharvested, partial cut and low-retention cut forest blocks. Mesh decomposition bags containing the three species and wood sticks were placed at two depths in colonies of either P. schreberi or S. capillifolium (environment) in the three harvest treatments and retrieved after two growing seasons. Mass loss was primarily related to substrate type (P. schreberi > S. capillifolium > wood sticks > S. fuscum) and secondarily to depth. Harvest treatment and environment (P. schreberi or S. capillifolium) only weakly affected sphagna mass loss. The weak effect of harvest treatment suggests that conditions created by low retention cuts do not to stimulate decomposition in this system and are not important enough to stimulate carbon loss, or to counteract paludification. On the other hand, the strong effect of bryophyte type indicates that conditions affecting bryophyte colonization and succession are of great importance in driving carbon and nutrient cycles.     

Additional reports for Italian moss flora

As a result of the study of both recent collections and a review of herbarium specimens, the following taxa are reported for the first time from various Italian regions: Hedwigia stellata Hedenäs, Hedwigia ciliata var. leucophaea Bruch & Schimp., Pleurozium schreberi (Willd. ex Brid.) Mitt. and Ptychostomum compactum Hornsch. var. compactum. The last taxon is new for southern Italy, while the report of Pleurozium schreberi confirms the occurrence of this species in southern Italy after more than half a century. Moreover, Crossidium laxefilamentosum Frey & Kürschner represents the second report of this species from Italy.     

On the Distribution of <Emphasis Type="Italic">Pleurozium schreberi</Emphasis> (Bryophyta,Hylocomiaceae) in the East European Plain and Eastern Fennoscandia

Pleurozium schreberi is one of the most common moss species in the forest area. It is dominant in the moss layer of blueberry and wood sorrel forests. It can occur in small quantities in almost all types of forests (even in bogs). It is also a typical component of the moss layer in tundra. The article considers the distribution of Pleurozium schreberi in the East European Plain and Eastern Fennoscandia. On the basis of literature sources on the occurrence of the species in different regions (according to point data), a model map of species distribution using the kriging-method has been created. The overlaying of the model map on the maps of spatial distribution of climatic parameters and vegetation zones in this area has revealed that the biogeographical preferences of the species. P. schreberi is characterized by its highest distribution in the forest zone. It often occurs here and represents a phytocenotically active species. The occurrence of Pleurozium schreberi dramatically decreases in the transition from the forest to the steppe zone, where it is a rather rare species, growing exclusively in pine and birch pegs. This species disappears in the open steppe. From the steppe zone to the south, the occurrence of Pleurozium schreberi gradually decreases with increase in summer temperatures and decrease in precipitation and with forest disappearance. In the north, where the species is highly active, its range abruptly ends on the coast of the Arctic Ocean. This pattern of distribution of Pleurozium schreberi is associated both with cenotic preferences and with climate: it becomes rare in regions with summer temperatures higher than +23°C and annual precipitation of less than 400 mm.     

Epiphytic lichens and bryophytes of forest ecosystems in Tuscany (Central Italy)

The distribution of epiphytic bryophytes and lichens on three species of trees (Quercus ilex, Quercus cerris and Fagus sylvatica, representing the forest ecosystems of central Italy at low, medium and high elevations, respectively) was examined with respect to host specificity and habitat differences. Although most species were host-specific, the results suggested that habitat characteristics are more important than phorophyte properties in determining the distribution of lichens and bryophytes. Factors affecting the diversity of the epiphytic flora were different for lichens and bryophytes. In the case of lichens, climatic parameters and forest monospecificity were important factors, whereas tree cover (i.e. shade) and to a certain extent the number of available trees seemed more important for bryophytes.     

Fungal biomass associated with the phyllosphere of bryophytes and vascular plants

Little is known about the amount of fungal biomass in the phyllosphere of bryophytes compared to higher plants. In this study, fungal biomass associated with the phyllosphere of three bryophytes (Hylocomium splendens, Pleurozium schreberi, Polytrichum commune) and three vascular plants (Avenella flexuosa, Gymnocarpium dryopteris, Vaccinium myrtillus) was investigated using ergosterol content as a proxy for fungal biomass. Phyllosphere fungi accounted for 0.2-4.0 % of the dry mass of moss gametophytes, representing the first estimation of fungal biomass associated with bryophytes. Significantly more fungal biomass was associated with the phyllosphere of bryophytes than co-occurring vascular plants. The ergosterol present in moss gametophytic tissues differed significantly between species, while the ergosterol present in vascular plant leaf tissues did not. The photosynthetic tissues of mosses had less associated fungal biomass than their senescent tissues, and the magnitude of this difference varied in a species-specific manner. The fungal biomass associated with the vascular plants studied varied significantly between localities, while that of mosses did not. The observed differences in phyllosphere community biomass suggest their size could be affected by host anatomical and physiological attributes, including micro-niche availability and chemical host defenses, in addition to abiotic factors like moisture and nutrient availability.     

Pleurozium schreberi as an ecological indicator of polybrominated diphenyl ethers (PBDEs) in a heavily industrialized urban area

Polybrominated diphenyl ethers (PBDEs) are toxic contaminants with a persistent character and adverse effects on humans and wildlife. Therefore, the deposition of these chemicals in vegetation must be carefully controlled. Our objective was to determine PBDE concentrations (BDEs 28, 47, 66, 85, 99, 100, 153, 154, 183 and 209) in Pleurozium schreberi collected in a heavily industrialized urban agglomeration. High PBDE concentrations in the moss confirm the presence of active sources of atmospheric pollution in an area tested. The distribution of these xenobiotics was related to the vegetation cover being lower in sites surrounded by forests which indicates that PBDEs may have a tendency to be trapped from the air by tree leaves and needles. Congener profiles in P. schreberi were dominated by BDE 209 which was for 79% (in case of the coke smelter) to 95% (in case of the chemical plant) part of the total PBDE burden in this moss. The principal component and classification analysis classifying the concentration of PBDEs in P. schreberi allowed us to distinguish the pattern of these compounds characteristic for the origin of pollution. P. schreberi may be used as a bioindicator for PBDEs in areas contaminated with these chemicals.     

Large‐scale changes in abundance of terricolous bryophytes and macrolichens in Finland

Abstract. Major changes in forest floor vegetation were identified on the basis of three nationwide surveys conducted as part of national forest inventories in 1951–1953, 1985–1986 and 1995. These surveys provided objectively selected, statistically representative samples of all forested land in Finland. The 1951–1953 data consist of over 10000 sample plots, while the later surveys were conducted on ca. 3000 permanent plots. Changes in relative abundance of dominant species (i.e. in the proportions of species of the total cover of forest floor vegetation) were analysed across biogeographical provinces. Spatial correlation, systematic sampling, partial re‐measurement and multiple testing were taken into account in assessment of the statistical significance of the observed changes. The most notable changes in forest floor vegetation were a decrease in the relative abundance of Hylocomium splendens and an increase in Dicranum polysetum. In N Finland, where forests are grazed by semi‐domestic reindeer, we observed a decline in the abundance of Cladina lichens and an increase in Dicranum mosses. Peltigera aphthosa declined throughout the country. Polytrichum juniperinum, Pohlia nutans, and Brachythecium species, which occupy disturbed sites or grow on litter, increased in abundance. The relative abundance of Sphagnum species decreased, particularly in W Finland, where Pleurozium schreberi was favoured. A major decline in S. fuscum was also recorded in C and E Finland. Many of the changes detected in this study are apparently related to intensified forest management; but solely on the basis of this study, its effects cannot be distinguished from those of other large‐scale environmental changes.     

Species richness of vascular plants, bryophytes and lichens in dry grasslands: The effects of environment, landscape structure and competition

We studied the relative importance of local habitat conditions and landscape structure for species richness of vascular plants, bryophytes and lichens in dry grasslands on the Baltic island of Öland (Sweden). In addition, we tested whether relationships between species richness and vegetation cover indicate that competition within and between the studied taxonomic groups is important. We recorded species numbers of vascular plants, bryophytes and lichens in 4 m² plots (n=452), distributed over dry grassland patches differing in size and degree of isolation. Structural and environmental data were collected for each plot. We tested effects of local environmental conditions, landscape structure and vegetation cover on species richness using generalized linear mixed models. Different environmental variables explained species richness of vascular plants, bryophytes and lichens. Environmental effects, particularly soil pH, were more important than landscape structure. Interaction effects of soil pH with other environmental variables were significant in vascular plants. Plot heterogeneity enhanced species richness. Size and degree of isolation of dry grassland patches significantly affected bryophyte and lichen species richness, but not that of vascular plants. We observed negative relationships between bryophyte and lichen species richness and the cover of vascular plants. To conclude, effects of single environmental variables on species richness depend both on the taxonomic group and on the combination of environmental factors on a whole. Dispersal limitation in bryophytes and lichens confined to dry grasslands may be more common than is often assumed. Our study further suggests that competition between vascular plants and cryptogams is rather asymmetric.     

The herb and dwarf shrubs colonization of decaying logs in subalpine forest in the Polish Tatra Mountains

This is a study of the colonization pattern of herbs and dwarf shrubs on rotten logs in subalpine spruce forests (Plagiothecio Piceetum) in the Tatra Mountains. On four study plots (total area 1.43 ha.) all dead logs were measured and the decomposition stage was estimated using the 8-degree scale. For each log the cover of all vascular species, bryophytes and lichens was determined according to the methods of classical phytosociology. Constancy and an index of coverage were calculated for all vascular species growing on logs. The total volume of logs was relatively high (93 m³ ha^–1) and constituted 22% of the volume of living trees. Logs and stumps covered 411 m² ha^–1. These values are similar to those known from natural spruce forest from Carpathians and Scandinavia. The 8 stages of decomposition were equally represented, which indicates a constant supply of dead wood to the forest floor over time. The colonization of dead wood starts with lichens, followed by bryophytes and finally herbs and tree saplings. The first vascular plant colonists of dead logs appear at decay stage nr. 3 at least 20 years after tree death. The most suitable condition for most of the herb species corresponds to decay stage nr. 6 ca. 50 years after tree death. The herb cover is distinctively dominated by Vaccinium myrtillus. Simultaneously with herb species, tree seedlings colonize the logs. Constancy and abundance of Norway spruce saplings increases with advanced decomposition. It seems that the herb cover of logs does not hinder the regeneration of spruce.     

Variation in responses to temperature treatments ex situ of the moss Pleurozium schreberi (Willd. ex Brid.) Mitt. originating from eight altitude sites in Hokkaido,Japan

Thermal acclimatisations are important for the survival and growth of individuals and populations but seldom studied for different populations of bryophytes. The aims of this study were to (I) investigate if responses to temperature treatments were independent of the site sampled or if the intra- and inter-population variation in responses were larger than the responses to the temperature treatments (control, press, and pulse), and to (II) examine if experimental responses varied, depending on the sampled sites. We collected samples of the circumpolar bryophyte species, Pleurozium schreberi (Willd. ex Brid.) Mitt., originating from eight altitude sites on Mt. Oakan in Hokkaido, Japan, and exposed them to three different temperature treatments ex situ for four weeks. Thermal acclimatisation was estimated by measuring responses in growth length increase, biomass increase, number of branches, and the maximum quantum yield of PS II (Fv/Fm). We found that responses to temperature treatments were dependent on the site sampled, and that differences were most pronounced in the length increase. Results also shows that the responses to experimental treatments may differ between sites. Our results therefore raise important concerns regarding the general validity of both ex situ and in situ experiments when performed on a single or a limited number of sites.     

Impact of management intensity on non-vascular epiphyte diversity in cacao plantations in western Ecuador

A first study on the biodiversity of non-vascular epiphytes in cacao (Theobroma cacao L.) plantations in western Ecuador yielded 112 species (51 bryophytes, 61 lichens). Epiphyte assemblages of cacao plantations resembled those of tropical rain forests but species richness was usually lower and individual species were found at lower heights on the trunks. The vast majority of the species are widespread neotropical or pantropical species; one species, Spruceanthus theobromae (Spruce) Gradst., is endemic to cacao plantations of western Ecuador. Differences in management intensity, by manual removal of epiphytes from tree trunks (limpia), had a significant impact on epiphyte species diversity. Total species richness was significantly reduced in plantations with high management intensity, due mainly to the decreased diversity of lichens and liverworts; moss diversity was not affected by management regime. Total percentage cover of bryophytes was highest in plantations with low management intensity, while lichen cover was greatest in plantations with high management intensity. Crustose lichens and smooth mats growing closely appressed to the substrate were little affected by the limpia and their growth may have been promoted when larger-sized species were removed, by reducing competition. Cacao plantations with low and moderate management intensity serve as an important substitute habitat for ecological specialists (sun epiphytes, shade epiphytes) of the rain forest and are of considerable significance for their conservation.     

Distribution and floristics of moss- and lichen-dominated soil crusts in a patterned Callitris glaucophylla woodland in eastern Australia

The distribution and abundance of soil crust lichens and bryophytes was examined in a patterned Callitris glaucophylla woodland in eastern Australia. Twenty-one lichen species and 26 bryophyte species were collected within thirty quadrats along a sequence of runoff, interception and runoff zones. Crust cover was significantly greatest in the interception zones (79.0 %), followed by the runoff zones (24.0 %), and lowest in the groved, runon zones (6.6 %). Lichens and bryophytes were distributed across all geomorphic zones, and, although there were significantly more moss species in the interception zones (mean = 9.1) compared with either the runoff (4.2) or runon (3.2) zones, the number of lichen species did not vary between zones. Ordination of a reduced data set of 32 species revealed a separation of taxa into distinct groups corresponding to the three geomorphic zones. Canonical correspondence analysis (CCA) of the 32 species and thirteen environmental variables revealed that the most important factors associated with the distribution of species were sheet and scarp erosion, soil stability and coherence, litter cover and crust cover. Surface cracking, microtopography and plant cover were of intermediate importance. The CCA biplot revealed that the timbered runon zones (groves) were dominated by `shade-tolerant' mosses Fissidens vittatus and Barbula hornschuchiana, whilst the heavily eroded runoff zones supported sparse populations of `erosion tolerant' lichens (Endocarpon rogersii) and mosses (Bryum argenteum and Didymodon torquatus). Interception zones supported a rich suite of `crust forming' mosses and lichens capable of tolerating moderate inundation by overland flow. Two other groups of taxa were identified by this analysis: the `pioneer' group, comprising mainly nitrogen-fixing lichens which occupy the zone of active erosion at the lower edge of the groves, and the `opportunists' dominated by liverworts, occupying the shallow depressions or bays at the margins of the groves and the interception zones. This study confirms that the non-vascular lichens and bryophytes in these arid soil crusts, are, like the vascular plants, strongly patterned according to geomorphic zone, being most strongly associated with soil surface and erosional features.     

The effect on mosses of transplantation to different latitudes

Abstract

Using net assimilation rate as a measure of physiological activity, reciprocal transplants of Hylocomium splendens, Pleurozium schreberi and Racomitrium lanuginosum between Yläne (60°55′N) and Kevo (69°45′N) and between Kevo and Spitzbergen (78°13′N) were used to investigate adaptation to latitude. It was found that physiological stress increased northwards and that adaptation to different latitudes was largely due to acclimation. The importance of day length and temperature to mosses in extreme latitudes is discussed.     

Hidden crown jewels: the role of tree crowns for bryophyte and lichen species richness in sycamore maple wooded pastures

Tree crowns typically cover the vast majority of the surface area of trees, but they are rarely considered in diversity surveys of epiphytic bryophytes and lichens, especially in temperate Europe. Usually only stems are sampled. We assessed the number of bryophyte and lichen species on stems and in crowns of 80 solitary sycamore maple trees (Acer pseudoplatanus) at six sites in wooded pastures in the northern Alps. The total number of species detected per tree ranged from 13 to 60 for bryophytes, from 25 to 67 for lichens, and from 42 to 104 for bryophytes and lichens considered together. At the tree level, 29 % of bryophyte and 61 % of lichen species were recorded only in the crown. Considering all sampled trees together, only 4 % of bryophyte, compared to 34 % of lichen species, were never recorded on the stem. Five out of 10 red-listed bryophyte species and 29 out of 39 red-listed lichen species were more frequent in crowns. The species richness detected per tree was unexpectedly high, whereas the proportion of exclusive crown species was similar to studies from forest trees. For bryophytes, in contrast to lichens, sampling several stems can give a good estimation of the species present at a site. However, frequency estimates may be highly biased for lichens and bryophytes if crowns are not considered. Our study demonstrates that tree crowns need to be considered in research on these taxa, especially in biodiversity surveys and in conservation tasks involving lichens and to a lesser degree also bryophytes.     

Response of feather moss associated N2 fixation and litter decomposition to variations in simulated rainfall intensity and frequency

Feather mosses in boreal forests form a dense ground‐cover that is an important driver of both nutrient and carbon cycling. While moss growth is highly sensitive to moisture availability, little is known about how moss effects on nutrient and carbon cycling are affected by the dynamics of moisture input to the ecosystem. We experimentally investigated how rainfall regimes affected ecosystem processes driven by the dominant boreal feather moss Pleurozium schreberi by manipulating total moisture amount, frequency of moisture addition and moss presence/absence. Moisture treatments represented the range of rainfall conditions that occur in Swedish boreal forests as well as shifts in rainfall expected through climate change. We found that nitrogen (N) fixation by cyanobacteria in feather mosses (the main biological N input to boreal forests) was strongly influenced by both moisture amount and frequency, and their interaction; increased frequency had greater effects when amounts were higher. Within a given moisture amount, N fixation varied up to seven‐fold depending on how that amount was distributed temporally. We also found that mosses promoted vascular litter decomposition rates, concentrations of litter nutrients, and active soil microbial biomass, and reduced N release into soil solution. These effects were usually strongest under low moisture amount and/or frequency, and revealed a buffering effect of mosses on the decomposer subsystem under moisture limitation. These results highlight that both the amount and temporal distribution of rainfall, determine the effect of feather mosses on ecosystem N input and the decomposer subsystem. They also emphasize the role of feather mosses in mediating moisture effects on decomposer processes. Finally, our results suggest that projected shifts in precipitation in the Swedish boreal forest through climate change will result in increased moss growth and N₂ fixation but a reduced dependency of the decomposer subsystem on feather moss cover for moisture retention.     

Nonvascular contribution to ecosystem NPP in a subarctic heath during early and late growing season

Bryophytes and lichens abound in many arctic ecosystems and can contribute substantially to the ecosystem net primary production (NPP). Because of their growth seasonality and their potential for growth out of the growing season peak, bryophyte and lichen contribution to NPP may be particularly significant when vascular plants are less active and ecosystems act as a source of carbon (C). To clarify these dynamics, nonvascular and vascular aboveground NPP was compared for a subarctic heath during two contrasting periods of the growing season, viz. early-mid summer and late summer-early autumn. Nonvascular NPP was determined by assessing shoot biomass increment of three moss species (Hylocomium splendens, Pleurozium schreberi and Dicranum elongatum) and by scaling to ecosystem level using average standing crop. For D. elongatum, these estimates were compared with production estimates obtained from measurements of shoot length increase. Vascular NPP was determined by harvesting shrub and herb apical growth and considering production due to stem secondary growth of shrubs. Hylocomium splendens and Pleurozium schreberi showed highest biomass growth in late summer, whereas for D. elongatum this occurred in early summer. Maximum relative growth rates were ca. 0.003–0.007 g g⁻¹ d⁻¹. For D. elongatum, production estimates from length growth differed from estimations from biomass growth, likely because of an uncoupling between length growth and biomass shoot growth. Nonvascular NPP was 0.37 and 0.46 g dry weight m⁻² d⁻¹, in early and late summer, respectively, whereas in the same periods vascular NPP was 3.6 and 1.1 g dry weight m⁻² d⁻¹. The contribution of nonvascular NPP to total aboveground NPP was therefore minor in early summer but substantial in late summer, when 25% of the C accumulated by the vegetation was incorporated into nonvascular plant tissue. The expected global change-induced reduction of nonvascular plant biomass in subarctic heath is likely therefore to enhance C release during the late part of the growing season.