Tree regeneration and future dynamics of the laurel forest on Tenerife,Canary Islands

Abstract. We studied two sites in the laurel forest of Tenerife to predict future changes in canopy composition. We used two projection methods: Horn's ‘Markovian Projection’, which utilizes information on juveniles in the vicinity of canopy trees, and a 'Stand Projection’, which ignores such information. We performed these projections both including and excluding a-sexual regeneration. Although all of our projections predict a change in species composition, inclusion of a-sexual reproduction decreased the magnitude of successional change. The persistence of Prunus lusitanica and Ilex canariensis appears to be highly dependent on a-sexual regeneration. Both the Markov- and stand projections predict a slight convergence in species composition between the two sites when only sexual regeneration is considered, and also a higher dominance of the shade-tolerant species Laurus azorica. When a-sexual regeneration is also considered, some divergence is shown, with less projected change in the stand projection than in the Markov projection. In spite of some differences between the models, general patterns such as an increase of shade-tolerant species (Laurus azorica and Prunus lusitanica) and a decrease of shade-intolerant species (Erica arborea, Erica scoparia and Myrica faya) are consistent.     

Treefall Gaps and Regeneration Composition in the Laurel Forest of Anaga (Tenerife): a Matter of Size?

The size of treefall gaps is an important determinant of regeneration composition in tropical and temperate forests. Preliminary studies in the laurel forest of Tenerife have shown that small gaps (<100 m²) were the most numerous. However, due to this small size, no significant differences were found between regeneration in gaps and regeneration below the canopy. Because infrequent large gaps (>100 m²) are present in the laurel forest, we analyzed the regeneration in these large uncommon gaps, considering their potentially important role in the dynamics of the system. Our main hypothesis is that large gaps are important disturbance to ensure the regeneration and stablishment of shade intolerant species. Only five gaps larger than 100 m² (ranging from 125–268 m²) were found in the study area. Data from a further 20 small gaps (<100 m²), analysed in a previous study, was also included. Control plots were examined close to the gaps in order to determine regeneration below the closed canopy. We did not find a significant difference between regeneration density in the gaps (<100 m²) and regeneration below the canopy in the control plots. Contrary to our expectations, regeneration was lower in the large gaps than under the canopy. The open canopy in the large gaps increases light intensity, and has a negative effect on the germination and growth of shade-tolerant tree species like Viburnum tinus (although non-statistically significant); however, the increase in light intensity is not sufficient to stimulate the germination of shade-intolerant tree species. The effects of treefall gaps in the dynamics of the laurel forest of Anaga should be not considered as significant in comparison to other factors such as human disturbances or infrequent disturbances (land slides or hurricanes).     

Forest development in canopy gaps in old-growth beech (Nothofagus) forests,New Zealand

Abstract. Tree size and age structures, treefall and canopy gap characteristics, and regeneration responses to treefalls were compared for three stands of old-growth beech (Nothofagus) forest dominated by N. fusca and N. menziesii on the South Island, New Zealand. Treefall gaps (up to 1000 m²) were most often caused by standing trees killed by drought and/or insect attack, or by trees snapped by wind. The causes of gap formation and the size and age distributions of treefall gaps varied between localities because of spatial and temporal differences in the histories of disturbance. At Fergies Bush where drought-related dieback had produced many large gaps with standing dead trees, gaps were generally young. Conversely, at Station Creek, small, old gaps formed by bolesnap dominated the disturbance regime. At Rough Creek, gaps of all ages and sizes were found along with an almost complete fern cover, and abundant shrubs and occasional subcanopy hardwood trees. Although overall patterns of regeneration were unrelated to differences in gap size, the relative abundance of N. fusca and N. menziesii varied between localities according to the seemingly minor differences in forest structure and disturbance history described above. Interpretations of regeneration response to gap parameters, therefore, need to account for differences in disturbance history between sites. Differences in the disturbance history between localities will also influence rates of gap closure, and because closure rates are used to estimate forest turnover times, meaningful comparisons of disturbance regimes for different forest types can only be made if this intersite variability is addressed.     

Gap dynamics and regeneration strategies in <Emphasis Type="Italic">Juniperus-Laurus</Emphasis> forests of the Azores Islands

We asked the following questions regarding gap dynamics and regeneration strategies in Juniperus-Laurus forests: How important are gaps for the maintenance of tree diversity? What are the regeneration strategies of the tree species? Thirty canopy openings were randomly selected in the forest and in each the expanded gap area was delimited. Inside expanded gaps the distinction was made between gap and transition zone. In the 30 expanded gaps a plot, enclosing the gap and transition zone, was placed. In order to evaluate the differences in regeneration and size structure of tree species between forest and expanded gaps, 30 control plots were also delimited in the forest, near each expanded gap. In the 60 plots the number of seedlings, saplings, basal sprouts and adults of tree species were registered. Canopy height and width of adult individuals were also measured. The areas of the 30 gaps and expanded gaps were measured and the gap-maker identified. Juniperus-Laurus forests have a gap dynamic associated with small scale disturbances that cause the death, on average, of two trees, mainly of Juniperus brevifolia. Gap and expanded gap average dimensions are 8 and 25 m², respectively. Gaps are of major importance for the maintenance of tree diversity since they are fundamental for the regeneration of all species, with the exception of Ilex azorica. Three types of regeneration behaviour and five regeneration strategies were identified: (1) Juniperus brevifolia and Erica azorica are pioneer species that regenerate in gaps from seedlings recruited after gap formation. However, Juniperus brevifolia is a pioneer persistent species capable of maintaining it self in the forest due to a high longevity and biomass; (2) Laurus azorica and Frangula azorica are primary species that regenerate in gaps from seedlings or saplings recruited before gap formation but Laurus azorica is able to maintain it self in the forest through asexual regeneration thus being considered a primary persistent species; (3) Ilex azorica is a mature species that regenerates in the forest.     

Laurel forests in Tenerife,Canary Islands

 Stand structure and leaf area distribution of a laurel forest in the Agua García mountains of Tenerife are described. The site is situated at 820 m a.s.l., faces NNE, and has a humid mediterranean climate. Summer droughts are mitigated by relatively high air humidity and clouds. The natural mixed hardwood forest is composed of six major tree species: Laurus azorica (Seub.) Franco, Persea indica (L.) Spreng, Myrica faya Ait., Erica arborea L. and two species of Ilex (I. platyphylla Webb & Berth. and I. canariensis Poivet.). The experimental stand had a density of 1693 trees ha^– 1, a basal area of 33.7 m²ha^– 1, and a cumulated volume of above-ground parts of trees of 231 m³ ha^– 1 with a corresponding dry mass of 204 ton ha^– 1. Diameters at breast height ranged from 6 to 46 cm. Mean concentration of plant dry mass per volume was 1.17 kg m^– 3. The vertical pattern of leaf area distribution in individual trees for all tree species was characterized by a Gaussian-like curve. Stand leaf area index was 7.8. These evergreen, broad-leaved (laurisilva or lucidophyllous) forests represent a relic forest that was widespread in the Mediterranean region some 20 million years ago. Our data illustrate some of the structural characteristics of this historically widespread forest type. Received: 2 December 1994 / Accepted: 6 November 1995     

Study on canopy disturbance regime and mechanism of tree species diversity maintenance in the lower subtropical evergreen broad-leaved forest,South China

ABSTRACT

The canopy disturbance, the gap environment, gap regeneration and maintenance of tree species diversity in the lower subtropical evergreen broad-leaved forest (LSEBF) of South China were studied in this paper. The most common manner of gap formation in the forest was by stem breakage. Most gaps were formed by two gap makers. The sizes of most expanded gaps (EG) and canopy gaps (CG) were in the range of 100 – 300 m² and 50 – 100 m², respectively. The ecological factors in gaps were analysed on the basis of contrasting measurement of the microclimatic regimes in gaps of different sizes and in non-gap stands. Tree species in the LSEBF were classified into 5 ecological species groups on the bases of their changes in order of importance values in gaps and in non-gap stands. Most of the species reached their peak of regeneration density around the gap sizes of 100 m² and 500 m². The curves of regeneration density vs. gap age for major species revealed two types. Regeneration densities of most species, and species diversity indices in gaps were greater than those in non-gap stands.     

Community dynamics of evergreen broadleaf forests in southwestern Japan. I. Wind damaged trees and canopy gaps in an evergreen oak forest

The rates of treefall and canopy opening in the evergreen oak forest in southwestern Japan were determined by studying the number and size distribution of overstory trees, wind damaged trees, and canopy gaps in a belt transect in the Kasugayama Forest Reserve in Nara City. Thirty three percent of the overstory trees wereCastanopsis cuspidata. The total area of canopy gaps was about 20% of the total land area in the study area. The ages of the gaps were determined by counting the annual rings of various kinds of trees growing in gaps. By comparing gap ages with meteorological data, it became evident that gap formation was mainly caused by strong typhoons. The mean time interval between strong typhoons visiting the forest reserve, 6.57 years, was determined by applying the MNY method to the meteorological data. The treefall rate and the mean area of canopy openings per year were 0.84 overstory trees/ha·year and 55.6 m²/ha·year, respectively. The mean residence time of the forest canopy was about 180 years.     

Tree-ring distinctness,dating potential and climatic sensitivity of laurel forest tree species in Tenerife Island

Macaronesian laurel forests are the only remnants of a subtropical palaeoecosystem dominant during the Tertiary in Europe and northern Africa. These biodiverse ecosystems are restricted to cloudy and temperate insular environments in the North Atlantic Ocean. Due to their reduced distribution area, these forests are particularly vulnerable to anthropogenic disturbances and changes in climatic conditions. The assessment of laurel forest trees’ response to climate variation by dendrochronological methods is limited because it was assumed that the lack of marked seasonality would prevent the formation of distinct annual tree rings. The aims of this study were to identify the presence of annual growth rings and to assess the dendrochronological potential of the most representative tree species from laurel forests in Tenerife, Canary Islands. We sampled increment cores from 498 trees of 12 species in two well-preserved forests in Tenerife Island. We evaluated tree-ring boundary distinctness, dating potential, and sensitivity of tree-ring growth to climate and, particularly, to drought occurrence. Eight species showed clear tree-ring boundaries, but synchronic annual tree rings and robust tree-ring chronologies were only obtained for Laurus novocanariensis, Ilex perado subsp. platyphylla, Persea indica and Picconia excelsa, a third of the studied species. Tree-ring width depended on water balance and drought occurrence, showing sharp reductions in growth in the face of decreased water availability, a response that was consistent among species and sites. Inter-annual tree-ring width variation was directly dependent on rainfall input in the humid period, from previous October to current April. The four negative pointer years 1995, 1999, 2008 and 2012 corresponded to severe drought events in the study area. This study gives the first assessment of dendrochronological potential and tree-ring climate sensitivity of tree species from the Tenerife laurel forest, which opens new research avenues for dendroecological studies in Macaronesian laurel forests.     

Ilex Canariensis Poir. (Aquifoliaceae) Post-dispersal Seed Predation in the Canary Islands

Alien species have many negative effects on insular ecosystems worldwide. We investigated Ilex canariensis post-dispersal seed predation by introduced rats (Rattus spp.) in relict forests of the Canary Islands at different spatial scales: among microhabitats within the same forest, among forest types within the same island, and among different islands of the archipelago. Seed predation intensity was very high (>70%) in all cases considered, irrespective of the spatial scale. We did not find significant differences between forest interior, edges or gaps, as well as between different forest types in four islands of the archipelago. Comparatively low predation intensity was found in El Hierro island, where more than 50% of the seeds survived at the end of the experiment, while highest seed predation was observed in Tenerife island. It is concluded that post-dispersal seed predation by rats, due to its extent and intensity, could have an important effect on Ilex canariensis recruitment, especially in successional areas where this light tolerant tree can naturally establish.     

Exploring the effect of host tree identity on epiphyte bryophyte communities in different Canarian subtropical cloud forests

This study explores the host tree preferences of epiphyte bryophyte communities in two key types of evergreen cloud forests on La Gomera (Canary Islands, Spain): ericaceous versus broadleaved laurel forest. By comparing two pairs of tree species (Erica arborea and Myrica faya vs. Ilex canariensis and Laurus novocanariensis) that occur in both forest types, we quantitatively examined whether epiphyte-host relationships change with the type of forest. In 51 ancient forest stands, the low-trunk bryophyte composition on the two host tree groups was analysed with both non-parametric procedures and ordination analyses. Our results highlight that the influence of local environmental conditions on the structuring of epiphyte communities may be different depending on the host tree identity. Whilst the epiphyte composition of the host tree group formed by I. canariensis and L. novocanariensis differed significantly between forest types depending on solar exposure, we failed to find variations for the other host tree group, excepting in those localities wherein mist precipitation was extremely high. Our findings highlight the importance of the natural diversity of tree species along environmental gradients, as well as of water availability at different spatial scales for epiphytic bryophytes in montane cloud forests.     

Spatial dynamics of regeneration in a conifer/broad‐leaved forest in northern Japan

Abstract. This study deals with stand dynamics over a 6‐yr period in a conifer/broad‐leaved mixed forest in Hokkaido, northern Japan. The annual rates of gap formation and recovery were 81.3 m²/ha and 66.7 m²/ha, respectively and turnover time of the canopy was 125 yr. The recruitment processes of the component species in this cool‐temperate forest were governed by different canopy types: gap, canopy edge and closed canopy. Magnolia obovata regenerated in canopy edges, and Acer mono and Prunus ssiori regenerated in canopy edges and gaps. The results suggested that the mosaic structure made up of closed canopy, canopy edge and gap was related to various regeneration niches. Abies sachalinensis had high mortality rates, initiating gap expansion. The transition probabilities from closed canopy or canopy edge to gap for deciduous broad‐leaved trees were lower than for A. sachalinensis, which implies that the difference in degeneration patterns of conifer and broad‐leaved canopies contributes to the heterogeneity of spatial structure in the mixed forests. Spatial dynamics were determined by a combination of gap expansion by A. sachalinensis (neighbour‐dependent disturbance) and gap formation by deciduous broad‐leaved trees (random disturbance).     

Forest gap formation and closure along an altitudinal gradient in Tongariro National Park,New Zealand

24 treefall gaps accumulated over a 10 year period along an altitudinal transectcovering 4.6ha on Mt. Hauhungatahi, Tongariro National Park, New Zealand were described quantitatively in terms of the area of damage (‘expanded gap’), the canopy opening (‘Tight-gap’) and the size of the root mound. Tree mortality and branch loss following cyclone Bola, 1988, were recorded. In each gap saplings were ranked by species according to their vigour. Pre-gap and post-gap vertical and horizontal branch growth rates were calculated. Effects in the subalpine forest (> 1050 m) were compared with those in the montane zone. Tree mortality was highly episodic, associated with major storms, and patchy. Falling canopy trees destroyed, on average, 1.3 additional trees (> 10 cm diameter at 1 m). About half the trees were uprooted and the remainder broken off. Uprooted angiosperm (canopy) trees frequently resprouted from their bases, gymnosperms rarely. Expanded gap area averaged 56 m² in the sub-alpine forest and 88 m² in the montane zone. Median expanded gap areas were about twice those of light gaps. Gap size frequency distribution was highly skewed. The largest gap was formed by a single Dacrydium cupressinum which destroyed six other trees creating a gap of ca. 0.03 ha. Expanded gaps, light gaps, and root mounds comprised 4.5, 2.8 and 0.1 % of the forest area in the sub-alpine zone, and 3.8, 2.5 and 0.06 % in the montane forest. These values represent 10 years of accumulation, and imply light gap ‘return times’ of 360 years for the sub-alpine and 400 years for the montane forest. These periods are in agreement with the known longevities of the canopy and emergent trees. Vertical shoot growth rate was about twice that in the horizontal plane, and both increased following gap formation. The relative increase was greatest in the subalpine forest. Using the measured growth rates it is estimated that gaps of median dimensions are filled by lateral extension growth in 31–44 yr. Saplings require longer to reach the mean canopy height and consequently require large (multiple tree) gaps or sequential gap events.     

Gap dynamics in climaxFagus crenata forests

Gap characteristics and gap regeneration were studied in several climaxFagus crenata forests in Japan. 278 gaps were observed. Gaps covered 12% of the total land area of 20.05 ha. Gap density was 13.9 gaps per ha and, mean gap size was 92.0 m². Smaller gaps were much more frequent than larger ones. Gaps larger than 400 m² were rare. Most gaps were created by the death of single trees. Canopy trees died more often standing or with broken trunks than by uprooting, although uprooted trees were relatively abundant in the site with poor soil drainage and in the site on upper slope. Differences of gap regeneration behaviour were recognized among tree species.F. crenata regenerates in gaps from saplings recruited before gap creation and can replace not only its own gaps but also gaps of other species. Most species other thanF. crenata andMagnolia obovata could not regenerate in their own gaps. More successful regeneration ofF. crenata may occur in gaps smaller than 200 m², althought it regenerated in a wide range of gap size. However, increased relative density ofF. crenata in the canopy layer seems to prevent its successful regeneration. Gap regeneration of other species did not clearly depend on a species-specific gap size.     

Gap characteristics and gap regeneration in primary evergreen broad-leaved forests of western Japan

Gap characteristics and regeneration in gaps were studied in some primary evergreen broad-leaved forests of the warm temperate zone in western Japan. Total observed 161 gaps covered 15.7% of the total land area of 8.23 ha. Gap density was 19. 6 gaps ha⁻¹ and mean gap size was 80.3 m². Smaller gaps (<80 m²) were much more frequent than larger ones, and gaps larger than 400 m² were rare. Gaps created by the death or the injury of single trees were 79.5%. Canopy trees died most often with broken trunks and not so often by uprooting or leaving standing-dead. Different types of gap regeneration behaviour were recognized among the major canopy tree species, though gap regeneration of the common evergreen broad-leaved tree species did not clearly depend on a species-specific gap size.Distylium racemosum, which occurred in equal importance (about 25%) in the canopy layer of each study stand, regenerates in gaps from saplings recruited before gap creation and can replace not only its own gaps but also gaps of other species. Therefore, it can be considered a typical climax species in this forest type of western Japan.Persea thunbergii, which can reproduce vegetatively, showed a similar type of gap regeneration behaviour.Castanopsis cuspidata can replace itself with relatively higher frequency by means of vegetative reproduction (stump sprouting) after gap creation.Quercus acuta andQuercus salicina did not regenerate under the current gap-disturbance regime. Though the frequency of uprooting is low, soil disturbance by uprooting seems to be important for the perpetuation of the pioneer tree species,Fagara ailanthoides, which recruits from buried seeds in the soil     

Gap characteristics and gap regeneration in a subalpine coniferous forest on Mt Ontake,central Honshu,Japan

Gap characteristics and gap regeneration were studied in three mature stands belonging to different community types in a subalpine coniferous forest on Mt Ontake, central Honshu, Japan. Gap disturbance regimes were remarkably similar among stands studied; percentage gap area to surveyed area, gap density and mean gap size were 7.3–8.5%, 17.8–20.0 ha⁻¹ and 40.8–42.5 m², respectively. The gap size class distributions were also similar and showed a strong positive skewness with a few large and many small gaps; gaps <40m² were most frequent and gaps >200m² were rare. Forty-five to 66% of gaps were due to the death of single canopy trees. Canopy trees more often died leaving standing dead wood (40–5.7%) or broken trunks (43–49%). Shade tolerantAbies mariesii andAbies veitchii, frequently, andTsuga diversifolia, less frequently, regenerated in gaps, from advance regenerations recruited before gap formation.Picea jezoensis var.hondoensis may regenerate in gaps, from new individuals recruited after gap formation. The breakage of denseSasa coverage and the mineral soils exposed by the uprooted plants that form gaps might provide regeneration opportunities for shade intolerantBetula. Of the important species limited to the ridge site,Chamaecyparis obtusa, frequently, andThuja standishii, infrequently, regenerated from plants alreadyin situ. Regeneration ofPinus parviflora was not seen. Based on the gap characteristics and gap regeneration behaviour of each species described, stand dynamics in each stand are discussed.     

Effects of Treefall Gap Disturbances on Ant Assemblages in a Tropical Montane Cloud Forest

The study of gap dynamics and the effects of gaps on diversity has been at the center of tropical ecology for decades. While most studies have focused on the responses of plant species and communities to gap formation, in this study, we consider the effects of treefall gap disturbances on leaf litter ant assemblages in a Neotropical montane cloud forest. We sampled leaf litter ant assemblages and estimated a suite of abiotic parameters in 12 large (>80‐m²) treefall gaps across a chronosequence and in 12 paired adjacent intact forest sites in the Monteverde Cloud Forest Preserve in Costa Rica. No species were more common in gaps than in intact forests, and in fact, species that were common in gaps were also among the most common in forests. The Chao2 estimate of species richness, however, was higher in gap sites than in intact forest sites. In addition, ant assemblages in gap sites did not become more similar to those in adjacent intact sites as gaps aged. In contrast to other studies, our work demonstrates that ant assemblages in the Monteverde Cloud Forest Preserve are weakly affected by the formation of treefall gaps. Together, these results indicate that treefall gap dynamics probably play little role in promoting ant diversity at more regional scales, or coexistence among species at more local scales.     

Gap characteristics and gap regeneration in subalpine old-growth coniferous forests,central Japan

Gap characteristics and gap regeneration were studied in three old-growth stands of subalpine coniferous forests in the northern Yatsugatake and the northern Akaishi mountains, central Japan. With the results of the present study and those of a previous study conducted in another locality, general features of gap characteristics and gap regeneration behavior of major tree species in subalpine coniferous forests of central Japan were summarized and discussed. Of the total 237 gaps investigated in the 14.48 ha of forested area, the percentage gap area to surveyed area, gap density and mean gap size were 7.3%, 17.2 ha⁻¹, and 43.3 m², respectively. The gap size distributions were similar among stands and showed a strong positive skewness with a few large and many small gaps; gaps <40m² were most frequent and those >200 m² were rare. Gaps due to the death of multiple canopy trees comprised 44.7% of the total ones. Canopy trees died in various states; standing dead (42.6%) or trunk broken (43.7%) were common and uprooted (12.2%) was an uncommon type of death of canopy trees. These figures indicate that general features of gap characteristics in this forest type are the low proportion of gap area and the high proportions of small gap size and multiple-tree gap formation. In general, shade-tolerantAbies frequently, andTsuga, infrequently, regenerate in gaps from advance regenerations recruited before gap formation, whilePicea and shade-intolerantBetula possibly regenerate in gaps from new individuals recruited after gap formation. Gap successors of conifers occurred in a wide range of gap size and did not show the clear preference to species specific gap size. In old-growth stands without large-scale disturbance (≥0.1 ha in area) of subalpine coniferous forests of central Japan, major tree species may coexist with their different gap-regeneration behaviors and, probably, different life history traits.     

Spatial patterns of trees and juveniles in a laurel forest of Tenerife, Canary Islands

Spatial patterns are important characteristics of the forest and theycan reveal such things as successional status and ecological characteristics ofthe species. We tested the hypothesis that spatial distribution will bedifferent, depending on whether the species is intolerant or tolerant to shade.We assessed the spatial distribution of trees (> 4 cm dbh) andjuveniles in eight laurel forest plots. A univariate spatial analysis(performed with Ripley's K₁) showed that all tree species havesignificantaggregation at short distances (2 m). Nevertheless, two groups ofspecies could be differentiated: Erica scoparia,Myrica faya and Ilex canariensisshowed a tendency for aggregation at large distances (larger than 6m)while L. azorica and Prunuslusitanicashowed aggregation only at shorter distances. Ripley's BivariateK_1,2 analyses showed no significant differences in the spatialdistribution ofanalyzed species pairs from a null model. Only Laurusazoricahad a sufficient sample size for analysis of juvenile distribution. Aunivariateanalysis revealed thatL. azorica seedlings (stems < 50 cm high)were clumped in some plots up to 5 m, but this was not consistent.Saplings (stems > 50 cm high and < 4 cm dbh)didnot show strong clumping even at short distances. L.azoricasaplings had no significant aggregation with, nor repulsionfrom, adults of the same or different species. Spatial patterns of the speciesshould be considered in the development of restoration plans of the laurelforest 90%of which has disappeared or been intensively disturbed on Tenerife Island.     

Understanding causes of tree growth response to gap formation: ∆13C-values in tree rings reveal a predominant effect of light

Key message

Carbon isotope ratios in growth rings of a tropical tree species show that treefall gaps stimulate diameter growth mainly through changes in the availability of light and not water. The formation of treefall gaps in closed canopy forests usually entails considerable increases in light and nutrient availability for remaining trees, as well as altered plant water availability, and is considered to play a key role in tree demography. The effects of gaps on tree growth are highly variable and while usually stimulatory they may also include growth reductions. In most studies, the causes of changes in tree growth rates after gap formation remain unknown. We used changes in carbon isotope ¹³C discrimination (Δ¹³C) in annual growth rings to understand growth responses after gap formation of Peltogyne cf. heterophylla, in a moist forest of Northern Bolivia. We compared growth and Δ¹³C of the 7 years before and after gap formation. Forty-two trees of different sizes were studied, half of which grew close (<10 m) to single treefall gaps (gap trees), the other half more than 40 m away from gaps (controls). We found variable responses among gap trees in growth and Δ¹³C. Increased growth was mainly associated with decreased Δ¹³C, suggesting that the growth response was driven by increased light availability, possibly in combination with improved nutrient availability. Most trees showing zero or negative growth change after gap formation had increased Δ¹³C, suggesting that increased water stress did not play a role, but rather that light conditions had not changed much or nutrient availability was insufficient to support increased growth. Combining growth rates with Δ¹³C proved to be a valuable tool to identify the causes of temporal variation in tree growth.     

Burrowing seabirds affect forest regeneration, Rangatira Island, Chatham Islands, New Zealand

The forests of Rangatira Island (218 ha) in the Chatham Islands are a critical breeding site for a number of rare and threatened forest bird species, but are also home to more than three million seabirds, which could significantly affect forest regeneration processes. We surveyed the forests of Rangatira Island by establishing 40 permanent forest plots, estimated seabird density through burrow counts, and analysed soil properties. To determine if seabirds were impacting on forest regeneration, we established exclosures (0.25 m²) in 30 of the forest plots, and examined the role of canopy gaps in forest regeneration. The tallest current forest (c. 15 m), dominated by Plagianthus chathamicus, has mostly regenerated since stock were removed in 1959. Mean burrow density was estimated to be 1.19 per square metre, all soils were highly acidic (pH 3.36–5.18), and burrow density was positively correlated with soil phosphorus. Seedling density of woody species in seabird exclosures measured after 9, 24 and 33 months was significantly higher than in the adjacent non-gap plots, and seedling density was positively associated with reduced canopy cover. Seedling densities were also significantly higher in canopy gaps than in adjacent non-gap plots, but seabird burrow density was significantly lower in gaps. These results suggest that canopy gaps allow forest regeneration despite the negative impacts of seabird burrowing. However, the gap makers, largely senescing Olearia traversii, are slowly disappearing from the forests. The cohort of Plagianthus that has regenerated following farm abandonment may progressively collapse, allowing regeneration to continue in small openings, but there is also the potential for a catastrophic blowdown. This might have serious implications for forest-dwelling birds, invertebrates, and plants.