Age and Long-term Growth of Trees in an Old-growth Tropical Rain Forest, Based on Analyses of Tree Rings and 14C1

In an old‐growth tropical wet forest at La Selva, Costa Rica, we combined radiocarbon (¹⁴C) dating and tree‐ring analysis to estimate the ages of large trees of canopy and emergent species spanning a broad range of wood densities and growth rates. We collected samples from the trunks of 29 fallen, dead individuals. We found that all eight sampled species formed visible growth rings, which varied considerably in distinctiveness. For five of the six species for which we combined wood anatomical studies with ¹⁴C‐dates (ring ages), the analyses demonstrated that growth rings were of annual formation. The oldest tree we found by direct ring counting was a Hymenolobium mesoamericanum Lima (Papilionaceae) specimen, with an age of ca. 530 years at the time of death. All other sampled individuals, including very large trees of slow‐growing species, had died at ages between 200 and 300 years. These results show that, even in an everwet tropical rain forest, tree growth of many species can be rhythmic, with an annual periodicity. This study thus raises the possibility of extending tree‐ring analyses throughout the tropical forest types lacking a strong dry season or annual flooding. Our findings and similar measurements from other tropical forests indicate that the maximum ages of tropical emergent trees are unlikely to be much greater than 600 years, and that these trees often die earlier from various natural causes.     

Annual growth rings, rainfall-dependent growth and long-term growth patterns of tropical trees from the Caparo Forest Reserve in Venezuela

1 Tree-ring analyses and dendrometer measurements were carried out on 37 tree species in a semi-deciduous forest of the Reserva Forestal de Caparo, Venezuela, where the mean annual rainfall is about 1700 mm and there is a dry season from December to March. The main purposes of the investigation were to show the seasonality of cambial growth, and the connection between precipitation patterns and tree-ring curves. Long-term rates of wood increment were also estimated.
2 Cambial markings in consecutive years showed that annual rings were formed by many species.
3 The distinctiveness of growth zones was usually greater in deciduous species than in evergreen species, although not all deciduous species had distinct rings.
4 Dendrometer measurements showed that the annual growth rhythm was related to precipitation patterns. Evergreen species tended to show only a short interruption of wood growth (during the later part of the dry season), whereas deciduous species stopped growth completely at the end of the rainy season.
5 For deciduous species, regression analyses showed close relations between tree-ring width and the sum of precipitation outside the rainy seasons (i.e. November to April). Evergreen species reacted to the total annual amount of precipitation.
6 Variation in longest available ring chronology (for Terminalia guianensis ) showed little correlation with the El Niño–Southern Oscillation effect.
7 On average trees from natural forests showed relatively constant growth over the entire life span. Plantation trees grew fast up to an age of 15–20 years, but annual increments then decreased to values seen in natural forest trees.     

Evaluating the annual nature of juvenile rings in Bolivian tropical rainforest trees

Knowledge on juvenile tree growth is crucial to understand how trees reach the canopy in tropical forests. However, long-term data on juvenile tree growth are usually unavailable. Annual tree rings provide growth information for the entire life of trees and their analysis has become more popular in tropical forest regions over the past decades. Nonetheless, tree ring studies mainly deal with adult rings as the annual character of juvenile rings has been questioned. We evaluated whether juvenile tree rings can be used for three Bolivian rainforest species. First, we characterized the rings of juvenile and adult trees anatomically. We then evaluated the annual nature of tree rings by a combination of three indirect methods: evaluation of synchronous growth patterns in the tree- ring series, ¹⁴C bomb peak dating and correlations with rainfall. Our results indicate that rings of juvenile and adult trees are defined by similar ring-boundary elements. We built juvenile tree-ring chronologies and verified the ring age of several samples using ¹⁴C bomb peak dating. We found that ring width was correlated with rainfall in all species, but in different ways. In all, the chronology, rainfall correlations and ¹⁴C dating suggest that rings in our study species are formed annually.     

Species-Specific Growth Responses to Climate Variations in Understory Trees of a Central African Rain Forest

Basic knowledge of the relationships between tree growth and environmental variables is crucial for understanding forest dynamics and predicting vegetation responses to climate variations. Trees growing in tropical areas with a clear seasonality in rainfall often form annual growth rings. In the understory, however, tree growth is supposed to be mainly affected by interference for access to light and other resources. In the semi-deciduous Mayombe forest of the Democratic Republic of Congo, the evergreen species Aidia ochroleuca, Corynanthe paniculata and Xylopia wilwerthii dominate the understory. We studied their wood to determine whether they form annual growth rings in response to changing climate conditions. Distinct growth rings were proved to be annual and triggered by a common external factor for the three species. Species-specific site chronologies were thus constructed from the cross-dated individual growth-ring series. Correlation analysis with climatic variables revealed that annual radial stem growth is positively related to precipitation during the rainy season but at different months. The growth was found to associate with precipitation during the early rainy season for Aidia but at the end of the rainy season for Corynanthe and Xylopia. Our results suggest that a dendrochronological approach allows the understanding of climate–growth relationships in tropical forests, not only for canopy trees but also for evergreen understory species and thus arguably for the whole tree community. Global climate change influences climatic seasonality in tropical forest areas, which is likely to result in differential responses across species with a possible effect on forest composition over time.     

The Potential of Tree Rings for the Study of Forest Succession in Southern Mexico

Studies of tropical secondary forest succession face strong limitations due to the slow pace of succession and the time-consuming task of monitoring processes. The occurrence of tree rings in secondary forest trees may help expand our knowledge on succession in these systems and may be useful for fallow dating in chronosequence studies. We examine here the potential of tree rings to study forest succession by sampling 70 species along chronosequences of dry and wet forests in southern Mexico. Based on wood anatomical features, we estimated that about 37 percent of the species presented distinct growth rings useful for ring studies. Overall, maximum number of rings matched well the interview-based fallow ages but, at some sites, trees had consistently higher numbers of rings, probably due to errors in fallow ages derived from interviews. Best fallow age estimations were obtained by examining rings in both pioneer and nonpioneer species. Reconstruction of species' establishment dates revealed that pioneer and nonpioneer species establish early during succession, and that species of both groups continue to recruit after many years. Our study clearly shows that tree ring analysis is a promising tool for studies on secondary forest succession in the tropics.     

Climatic signals in tree‐rings of Araucaria angustifolia in the southern Brazilian highlands

Araucaria angustifolia (Bertol.) O. Kuntze (Araucariaceae) is a Neotropical tree, widely distributed in subtropical mountain rain forests and nearby natural grasslands of Southern Brazil. This species produces annual growth rings, but its dendroclimatic potential is barely known. In the present paper, the long‐term growth patterns of A. angustifolia were investigated using annual growth ring time series and association to climate over the last century. Wood cores of A. angustifolia trees growing in forest and grassland habitats were obtained with an increment borer. The cores were surfaced, measured and cross‐dated. The dated ring‐width time series were standardized and submitted to correlation and principal component analysis to verify growth trends among sites and trees. Growth‐climate relationships were investigated using correlation and regression analyses, comparing the ordination axes scores to regional time series of precipitation and temperature. Due to anatomical irregularities, mainly partial rings, only 35 out of 60 trees were cross‐dated. The correlation and ordination analyses showed common tree‐growth trends within and between sites, indicative of a regional environmental force determining inter‐annual cambial activity variation. Despite growing in distinct habitats and disturbance regimes, A. angustifolia trees share a common long‐term growth pattern, which is significantly related to thermal conditions during the current and previous growing seasons. Moreover, site‐specific characteristics may have influenced opposite growth responses and association to climate conditions between forest and grassland trees.     

Age and radial growth pattern of four tree species in a subtropical forest of China

Subtropical forests are usually composed of many tree species. Knowledge of the age and radial growth variation of the dominant tree species is useful for understanding forest dynamics and community structure and function. The aims of this study are to explore whether there are identifiable annual growth rings in the main tree species and to examine the growth characteristics within and among the species in Mount Gutian subtropical forest of China. The results showed that four out of eight tree species from which samples were collected had visible and cross-datable rings. There were no stable relationships between the age and diameter for these subtropical trees. Significant differences existed in radial growth rate within and among the four species, suggesting a high spatial heterogeneity in the mixed-species subtropical forest. The common pattern in age distribution of multiple species suggests a stand-wide disturbance occurring around the 1960s. It is interesting to note that the growth rate at the same age intervals was different for trees younger than 40 years of age and older than 40 years of age, suggesting a change in climate or forest structure in the two time periods. The results obtained from this study help understand the growth dynamics in other subtropical forests having these tree species.     

Wood growth of Tabebuia barbata (E. Mey.) Sandwith (Bignoniaceae) and Vatairea guianensis Aubl. (Fabaceae) in Central Amazonian black-water (igapó) and white-water (várzea) floodplain forests

Tree growth is a fundamental indicator for conservation plans of Amazonian floodplain forests. In this study we use dendrochronology to analyze wood growth patterns of Tabebuia barbata and Vatairea guianensis, two tree species occurring in nutrient-rich white-water (várzea, Mamirauá Sustainable Development Reserve, MSDR) and nutrient-poor black-water (igapó, Amanã Sustainable Development Reserve, ASDR) floodplain forests of Central Amazonia. From 20 trees per species and floodplain system (total of 80 trees) growing under a similar flooding regime with a mean inundation height of about 4 m we measured diameter at breast height (dbh). We sampled two cores per tree with an increment corer at the height of dbh to determine wood density (WD), tree age and mean radial increment (MRI) rates. The wood samples were macroscopically analyzed. Both tree species show distinct annual tree rings characterized by marginal parenchyma tissues. MRI was measured by a digital measuring device and WD was determined by the ratio dry mass/fresh volume. MRI of both tree species was significantly higher in the várzea than in the igapó, which can be traced back to the contrasting nutrient status. WD showed no difference comparing both floodplain forest types. Tree ages of a species for the same diameter are more than twofold higher in the igapó than in the várzea. To insure a sustainable harvest, felling cycles in these forests should be adjusted according to rates of growth.     

Seeing the forest for its multiple ecosystem services: Indicators for cultural services in heterogeneous forests

The ecosystem service (ES) framework is gaining traction in ecosystem management as a means to recognize the multiple benefits that ecosystems provide. In forested ecosystems, many structural attributes (trees, understory plants and woody debris) create heterogeneous ecosystems that provide numerous ecosystem services, including many that are culturally important. However, application of the ES framework to forest management is challenged by difficulties measuring and comparing multiple ES across diverse and heterogeneous forest conditions. Indicators can help bring the ES approach to forest management by providing a means for accurate ES inventory and mapping. We measured 10 forest ES in contrasting forest types to investigate the effects of past forest harvesting in coastal temperate rainforest of Vancouver Island, BC, Canada. Our objectives were to build a systematic set of ES indicators for coastal temperate forests based on forest structural features, including trees, coarse woody debris, and understory plants. To achieve this, we 1) analyzed field data to compare the effects of forest age (old-growth vs. second-growth) and ecological site conditions (riparian vs. upland forest) on the bundle of ES provided by different forest types; and 2) worked with a local indigenous wood carver to identify attributes of cedar trees (Thuja plicata) essential for traditional uses, including canoe carving. Forest age and forest type had significant and major effects on bundles of ES. Old-growth forests provided three times higher carbon storage, nine times higher wood volume, and eighteen times higher canopy habitat services than recovering forests. Within old-growth forests, the proportion of trees suitable for traditional indigenous wood carving was significantly higher in riparian stands. Yet of 456 trees measured, only 17 were cedar with potential traditional uses. Of those, trees for canoe carving were the least frequent (n = 3), which we identified as large (>110 cm DBH) trees of exceptional quality. In general, old-growth riparian forests were a hotspot of ES, providing for example nearly three times as much carbon storage as old-growth forests on upland sites and 12 times the amount of carbon storage as found in second-growth forests on upland sites. These results indicate that typical inventories of forest ES, which usually generalize across heterogeneity, may oversimplify dramatic variations in ES bundles in forested landscapes. Our novel set of stand-level ES indicators can improve the accuracy of ES assessments, incorporate important cultural ES, and help address the role of landscape heterogeneity in influencing ES.     

An assessment of growth ring identification in subtropical forests from northwestern Argentina

Most subtropical forests in South America are located in regions with a marked seasonality in precipitation, which may induce the formation of annual bands in woody species. Due to the lack of precise information on tree-ring visibility, we evaluated the wood characteristics of 37 tree species in the subtropical Yungas and Chaco forests from northwestern (NW) Argentina. Anatomical features associated with the delimitation of growth bands were examined to establish the presence of tree rings. Different forest types reflect the precipitation gradients and wood anatomical features vary accordingly. Characteristics of wood structure are closely related to the dominant climatic patterns of each forest, revealing a common pattern of anatomical arrangements in terms of water transport and safety. In the Chaco and transitional forests, ring boundaries are related to marginal parenchyma whereas in montane forests growth ring boundary is mostly associated with the presence of thicker fibers at the end of the ring. The largest proportion of species with clearly marked growth rings occurs in the montane forest type of NW Argentina. Clear growth rings is a requisite for dendrochronological applications, hence the present work represents the first regional attempt to address the potential of subtropical species in South America to be used in dendrochronological studies.     

What tree rings can tell us about the competition between trees and lianas? A case study based on growth,anatomy, density,and carbon accumulation

In tropical forest, landscape fragmentation and the consequent degradation of disturbed forests increase the incidence of light and dry hot winds, causing a disturbance on natural regeneration. Under these conditions, lianas (woody vines) development is stimulated instead of other species, which are more suited to mature forest and under less influence of the edge effect. For this, lianas colonization is an important variable for assessing the disturbance level of a forest. In this context, it becomes important to understand the nature of the competitive relationships between hyper-abundant lianas and ring growth of the host trees. Here, we selected trees with occupation or absence of lianas from two tropical species – Pinus caribaea var. hondurensis (Caribbean pine) and Tectona grandis (teak) – localized in a semideciduous forest fragment in southeastern Brazil, aiming to compare growth, climatic response, anatomy (vessels and intra-annual density fluctuations), wood density and carbon, by tree-ring analysis. The results showed that the lianas caused a change in tree-ring anatomy of host trees in last 10 years, mainly. We observed that trees occupied by lianas had a decrease the radial growth and carbon in the two species, an increase of the vessels size in teak and a decrease of the IADF frequency in Caribbean pine. In teak, the climate-tree relationship indicated that trees with lianas had lower response to rainfall and higher response to temperature in the summer (rainy and hottest period); in Caribbean pine, we observed that trees with lianas had a 2-month delay in the radial growth response to rainfall in the dry season. In the teak group, we observed that host trees had higher wood density values than liana-free tree in the outer rings, and the opposite was showed for pine. These findings show that tree-ring growth of host trees are a strong bioindicator of forest disturbance caused by aggressive colonization of lianas. We believe that these methods are applicable to future studies relating to the effects of habitat fragmentation and forest degradation on biodiversity and ecosystem services, particularly in the context of global climate change.     

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.     

Millennium-Scale Crossdating and Inter-Annual Climate Sensitivities of Standing California Redwoods

Extremely decay-resistant wood and fire-resistant bark allow California’s redwoods to accumulate millennia of annual growth rings that can be useful in biological research. Whereas tree rings of Sequoiadendron giganteum (SEGI) helped formalize the study of dendrochronology and the principle of crossdating, those of Sequoia sempervirens (SESE) have proven much more difficult to decipher, greatly limiting dendroclimatic and other investigations of this species. We overcame these problems by climbing standing trees and coring trunks at multiple heights in 14 old-growth forest locations across California. Overall, we sampled 1,466 series with 483,712 annual rings from 120 trees and were able to crossdate 83% of SESE compared to 99% of SEGI rings. Standard and residual tree-ring chronologies spanning up to 1,685 years for SESE and 1,538 years for SEGI were created for each location to evaluate crossdating and to examine correlations between annual growth and climate. We used monthly values of temperature, precipitation, and drought severity as well as summer cloudiness to quantify potential drivers of inter-annual growth variation over century-long time series at each location. SESE chronologies exhibited a latitudinal gradient of climate sensitivities, contrasting cooler northern rainforests and warmer, drier southern forests. Radial growth increased with decreasing summer cloudiness in northern rainforests and a central SESE location. The strongest dendroclimatic relationship occurred in our southernmost SESE location, where radial growth correlated negatively with dry summer conditions and exhibited responses to historic fires. SEGI chronologies showed negative correlations with June temperature and positive correlations with previous October precipitation. More work is needed to understand quantitative relationships between SEGI radial growth and moisture availability, particularly snowmelt. Tree-ring chronologies developed here for both redwood species have numerous scientific applications, including determination of tree ages, accurate dating of fire-return intervals, archaeology, analyses of stable isotopes, long-term climate reconstructions, and quantifying rates of carbon sequestration.     

Climate response of cork growth in the Mediterranean oak (Quercus suber L.) woodlands of southwestern Portugal

In the Mediterranean climate regions, drought events are expected to affect the growth of forests ecosystems by changing trees growth rates and eventually inducing shifts in their growth patterns. Cork oak (Quercus suber L.) is a strictly western Mediterranean tree species periodically harvested for its bark, the cork. So far, cork oak has received limited attention for dendroclimatological studies due to its typical faint and erratic tree wood rings. Moreover, its distinct cork rings chronologies have been completely neglected. In this study we introduce an approach using cork ring chronologies dated back 9–10 years for climate response. Despite enhancing interannual variability and increasing statistical response to short-term climatic variability, still poorly understood, this study will possibly allow infer long-term climate response. We analyzed the cork ring chronologies of 55 cork samples collected in mature (under exploitation) trees in three distinct locations in southwestern Portugal. Cork growth recorded a high climate signal, with highly significant and coherent responses to the yearly climate-related sources of variation. We successfully assessed trends of cork growth via correlation analysis including selected climate variables among mean monthly temperature, monthly precipitation and, on an annual basis, eight precipitation indices. The high mean sensitivities and inter-series correlations found for cork ring chronologies combined with the significant variance explained by climate variables suggest that climate is likely one dominant signal that affects cork growth, but local environmental stresses can decisively affect this (climate) signal. Assuming cork growth as a proxy for cork oak growth, it seems conceivable that despite the trees being highly resistant to drought stress, cork oak woodlands in southwestern Portugal would have to face lesser growth in a global warming scenario.     

Stem growth rhythms in trees of a tropical rainforest in Southern Brazil

Key message

We demonstrate that tropical trees growing in wet climates can have a marked seasonality in cambium activity and stem growth associated with high temperature and day length of summer.

Abstract

Monitoring the rhythm of tree growth associated with the growth rings can contribute substantially to understanding forest dynamics and the management of tropical forests. In this study, we monitored the girth increment rhythm and described the wood characteristics (anatomy of growth rings, wood specific gravity) in 10 tropical tree species (103 individuals) naturally occurring in a wet and weakly seasonal region of Atlantic Forest in southern Brazil. We aimed to verify whether tree growth dynamics are associated with climate and woody anatomy in tropical trees with contrasting ecological characteristics. We installed permanent dendrometer bands and monthly assessed the girth increment for 22 months. We collected wood samples (non-destructive method), measured wood specific gravity and prepared permanent slides to characterize the growth ring markers. We found growth rings in all species (distinct in six species); deciduous species produced more distinguishable tree rings compared with semi-deciduous and evergreen tree species. Species varied in their accumulated girth growth (in average, from 1.83 to 62.64 mm), growth rates (1–15 %), and annual radial increment (0.16–5.44 mm). Girth increment was positively related to temperature and day length in five out of ten tree species, indicating the possible effects of these climatic variables in triggering cambial activity in these species. The growth pattern varied among species and was marginally associated to the tree deciduousness. We concluded that even in wet and less seasonal climates, there can be an association in the cambium activity and stem growth with the hotter and longer days of summer months.

     

Big eucalypts grow more slowly in a warm climate: evidence of an interaction between tree size and temperature

Large trees are critical components of forest ecosystems, but are declining in many forests worldwide. We predicted that growth of large trees is more vulnerable than that of small trees to high temperatures, because respiration and tissue maintenance costs increase with temperature more rapidly than does photosynthesis and these costs may be disproportionately greater in large trees. Using 5 00 000 measurements of eucalypt growth across temperate Australia, we found that high temperatures do appear to impose a larger growth penalty on large trees than on small ones. Average stem diameter growth rates at 21 °C compared with 11 °C mean annual temperature were 57% lower for large trees (58 cm stem diameter), but only 29% lower for small trees (18 cm diameter). While our results are consistent with an impaired carbon budget for large trees at warmer sites, we cannot discount causes such as hydraulic stress. We conclude that slower growth rates will impede recovery from extreme events, exacerbating the effects of higher temperatures, increased drought stress and more frequent fire on the tall eucalypt forests of southern Australia.     

The use of X-ray densitometry to evaluate the wood density profile of Tectona grandis trees growing in fast-growth plantations

Tectona grandis (teak) is an important commercial tree species that is widely used in tropical dendrochronology due to the formation of climate-sensitive annual growth rings. However, young trees growing in plantation conditions exhibit poor ring visibility during the first years of growth, limiting the dendrochronology application. In the present study, we use x-ray densitometry to determine the wood density profile between and within annual rings and at the sapwood-heartwood boundary in trees from fast-growth plantations. The resulting wood density profiles (WDP) can be categorized as uniform, stable growth, unstable growth, and false. The annual ring boundaries were indistinct in trees less than 8 years old. In mature trees, the annual ring boundaries are more defined. In relation to the sapwood-heartwood boundary, the WDP showed a decrease in the wood density; however, this decrease is influenced by the annual ring boundary when the two boundaries coincide. The identification of annual rings in trees growing in fast-growth plantations should be combined with X-ray densitometry and visual identification if wood density data are necessary for deriving other analysis, as climate change, from annual ring.     

The hydraulic limitation hypothesis revisited

We proposed the hydraulic limitation hypothesis (HLH) as a mechanism to explain universal patterns in tree height, and tree and stand biomass growth: height growth slows down as trees grow taller, maximum height is lower for trees of the same species on resource-poor sites and annual wood production declines after canopy closure for even-aged forests. Our review of 51 studies that measured one or more of the components necessary for testing the hypothesis showed that taller trees differ physiologically from shorter, younger trees. Stomatal conductance to water vapour (g(s)), photosynthesis (A) and leaf-specific hydraulic conductance (K L) are often, but not always, lower in taller trees. Additionally, leaf mass per area is often greater in taller trees, and leaf area:sapwood area ratio changes with tree height. We conclude that hydraulic limitation of gas exchange with increasing tree size is common, but not universal. Where hydraulic limitations to A do occur, no evidence supports the original expectation that hydraulic limitation of carbon assimilation is sufficient to explain observed declines in wood production. Any limit to height or height growth does not appear to be related to the so-called age-related decline in wood production of forests after canopy closure. Future work on this problem should explicitly link leaf or canopy gas exchange with tree and stand growth, and consider a more fundamental assumption: whether tree biomass growth is limited by carbon availability.